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.     

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.     

Mating preference functions of individual female barking treefrogs, Hyla gratiosa, for two properties of male advertisement calls

Abstract.— A mating preference function describes the relationship between variation in a trait in potential mates and the strength of the preference for that trait. Few studies have measured mating preference functions either at a population level or for individuals. We used two-choice playback experiments to determine the mating preference functions of individual female barking treefrogs for two call characteristics: call-repetition rate and fundamental frequency. We tested each female four times with each pair of stimuli and with three to six pairs of stimuli. Individual females exhibited directional preferences for higher call rates and stabilizing preferences for intermediate fundamental frequency. These individual preference functions were similar to population-level preferences documented in previous studies. Within a stimulus type (call rate or fundamental frequency), nearly all females exhibited the same general preference function. However, females varied in the minimum difference between stimuli necessary to elicit a unanimous choice for the higher call rate, and they differed in both the intermediate fundamental frequency they preferred most and the minimum difference in fundamental frequency that elicited a unanimous choice for one of the two alternatives. The variation we observed among females was not repeatable; repeatabilities were in general low and statistically nonsignificant. The observed variation in the preferences of females, even if unrepeatable, should weaken selection on male traits relative to selection in the absence of such variation.     

Female Preference Intensities on Different Call Characteristics and Symmetry of Preference Above and Below the Mean in the Iberian Midwife Toad Alytes cisternasii

The intensity of female preference for call characteristics is measured in two‐speaker playback tests with female Iberian midwife toads ‘Alytes cisternasii’ using an experimental design that allows quantitative comparisons across characters. Female preferences do not appear to be symmetrical above and below the population mean, female preference being more efficient at discriminating against low quality competing males that at selecting optimum males among good ones. Female preference is significantly more effective selecting males calling at higher call rates (a highly variable call characteristic probably related to the physical state of the male) rather than selecting calls of low dominant frequency (a characteristic of low intra‐individual variation, related to male size and age).     

The Advertisement Call of the European Treefrogs (Hyla arborea): A Multilevel Study of Variation

In this study, we analysed the patterns of variation of the European treefrog's advertisement call at four levels of organization: within individuals, within populations, among populations of the same species, and among different species of the same clade. At the within-individual level, call acoustic properties are distinguished into static and dynamic properties. At the within-population level, two sources of call variation were analysed: temperature and body size. Temperature affects both temporal and spectral properties of the call. Body size mostly affects spectral properties. At the between-population level, calls do not show significant differences with respect to temporal properties, but they do differ in two spectral (and stereotyped) properties: the fundamental frequency and the difference in amplitude between dominant and fundamental frequencies. Finally, at the between-species level, call differences are much more conspicuous: they involve both spectral and temporal and both static and dynamic properties. At all four levels, body size is associated with call variation, explaining 11% of the differences among populations of the same species and 73% of the differences among species of the same clade. On the basis of these results, we hypothesize that patterns of variation of call acoustic properties, their constraints, and their biological functions are intrinsically associated. We discuss the role that such an association might play in the evolution of acoustic signals.     

The spectral structure of vocalizations match hearing sensitivity but imprecisely in Philautus odontotarsus

It is generally thought that for species using vocal communication the spectral properties of the sender’s calls should match the frequency sensitivity of the receiver’s auditory system. Nevertheless, few studies have investigated both sender and receiver characteristics in anuran species. In the present study, auditory brainstem responses (ABRs) were recorded in the serrate legged treefrog, Philautus odontotarsus, in order to determine if male call spectral structure and hearing sensitivity in males and females have co-evolved in this species. The results showed that the spectral structures of male vocalization match both male and female hearing sensitivity, even though the dominant frequencies of male calls (2.5 kHz) are mismatched with the regions of best frequency sensitivity (1.4 and 2.8 kHz). In addition, the results show that, in contrast with most previous ABR studies in non-human animals, but consistent with human studies, there are noticeable sex differences in peripheral auditory sensitivity in Philautus insofar as females exhibit lower auditory thresholds than males across the entire 1.8–18 kHz frequency range. The results also show that the dominant frequency of male calls is negatively correlated with body size, indicating that call characteristics reflect body size in this species which may be used by females during mate choice.     

Female call preferences in tree-hole frogs: why are there so many unattractive males?

In tree-hole frogs, Metaphrynella sundana, the fundamental call frequency varies widely between males. In field playback experiments, females strongly preferred calls from the lower range of frequencies found in the population. There was no correlation, however, between male size and call frequency, as is normally the case for anurans, so large males were not necessarily more attractive to females. Presence or absence of upper harmonics in the call had no effect on female choice. Tree holes with shallow air columns were more often used by calling frogs, and were presumably more common, than deep holes. Since male M. sundana actively exploit the resonant properties of tree holes for mate attraction, and high frequencies match comparatively shallow holes, the benefits of attaining acoustic matching probably select for high-frequency calls. In addition, males with high-frequency calls may be heard from a greater distance in the vicinity of torrent streams. Since the level of such noise in the forest varies in time and space, different frequencies may prove optimal in different contexts, thereby preserving the observed variation within the population. Having an ‘unattractive’ high-frequency call should be potentially beneficial only when calling males do not congregate, a condition that our data suggest is fulfilled in this system.     

Modality interactions alter the shape of acoustic mate preference functions in gray treefrogs

Sexual selection takes place in complex environments where females evaluating male mating signals are confronted with stimuli from multiple sources and modalities. The pattern of expression of female preferences may be influenced by interactions between modalities, changing the shape of female preference functions, and thus ultimately altering the selective landscape acting on male signal evolution. We tested the hypothesis that the responses of female gray treefrogs, Hyla versicolor, to acoustic male advertisement calls are affected by interactions with visual stimuli. We measured preference functions for several call traits under two experimental conditions: unimodal (only acoustic signals presented), and multimodal (acoustic signals presented along with a video‐animated calling male). We found that females were more responsive to multimodal stimulus presentations and, compared to unimodal playbacks, had weaker preferences for temporal call characteristics. We compared the preference functions obtained in these two treatments to the distribution of male call characteristics to make inferences on the strength and direction of selection expected to act on male calls. Modality interactions have the potential to influence the course of signal evolution and thus are an important consideration in sexual selection studies.     

Acoustic Competition in Male Midwife Toads Alytes obstetricans and Alytes cisternasii: Response to Neighbor Size and Calling Rate. Implications for Female Choice

Effects of neighbor on male calling behavior was studied through playback experiments of synthetic calls to males of two species of midwife toads. The responses of resident males were scored considering two temporal parameters (call duration and calling rate) and one spectral parameter (dominant frequency). The sounds used for the playback tests included two levels of fundamental frequency (correlated with male size) and two levels of call repetition rate. In both species, resident males only changed their calling rate in the presence of an intruder, and the response was different for synthetic calls with two levels of dominant frequencies and with two calling rates. Resident size was not significantly correlated with the magnitude of the change in the calling rate. On the other hand, resident calling rate was significantly and positively correlated with the magnitude of the increase in calling rate of the stimulus. The maximum relative increase in calling rate was observed in A. cisternasii. In phonotaxis tests, females are preferentially attracted to calls emitted at a higher rate confirming the importance of changes in calling rate for female attraction.     

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.     

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.     

Melodic males and flashy females: Geographic variation in male and female reproductive behavior in red‐eyed treefrogs (Agalychnis callidryas)

Geographic variation in courtship behavior can affect reproductive success of divergent phenotypes via mate choice. Over time, this can lead to reproductive isolation and ultimately to speciation. The Neotropical red‐eyed treefrog (Agalychnis callidryas) exhibits high levels of phenotypic variation among populations in Costa Rica and Panama, including differences in color pattern, body size, and skin peptides. To test the extent of behavioral premating isolation among differentiated populations, we quantified male advertisement calls from six sites and female responses to male stimuli (acoustic and visual signals) from four sites. Our results show that both male advertisement calls and female behavior vary among populations: Discriminant function analyses can predict the population of origin for 99.3% ± 0.7 of males based on male call (dominant frequency and bandwidth) and 76.1% ± 6.6 of females based on female response behavior (frequency and duration of visual displays). Further, female mate choice trials (n = 69) showed that population divergence in male signals is coupled with female preference for local male stimuli. Combined, these results suggest that evolved differences among populations in male call properties and female response signals could have consequences for reproductive isolation. Finally, population variation in male and female behavior was not well explained by geographic or genetic distance, indicating a role for localized selection and/or drift. The interplay between male courtship and female responses may facilitate the evolution of local variants in courtship style, thus accelerating premating isolation via assortative mating.     

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.     

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.     

Developmental plasticity of mating calls enables acoustic communication in diverse environments

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

Independent Mating Preferences for Male Body Size and Coloration in Female Trinidadian Guppies

Although females in numerous species generally prefer males with larger, brighter and more elaborate sexual traits, there is nonetheless considerable intra‐ and interpopulation variation in mating preferences amongst females that requires explanation. Such variation exists in the Trinidadian guppy, Poecilia reticulata, an important model organism for the study of sexual selection and mate choice. While female guppies tend to prefer more ornamented males as mates, particularly those with greater amounts of orange coloration, there remains variation both in male traits and female mating preferences within and between populations. Male body size is another trait that is sexually selected through female mate choice in some species, but has not been examined as extensively as body coloration in the guppy despite known intra‐ and interpopulation variation in this trait among adult males and its importance for survivorship in this species. In this study, we used a dichotomous‐choice test to quantify the mating preferences of female guppies, originating from a low‐predation population in Trinidad, for two male traits, body length and area of the body covered with orange and black pigmentation, independently of each other. We expected strong female mating preferences for both male body length and coloration in this population, given relaxation from predation and presumably relatively low cost of choice. Females indeed exhibited a strong preference for larger males as expected, but surprisingly a weaker (but nonetheless significant) preference for orange and black coloration. Interestingly, larger females demonstrated stronger preferences for larger males than did smaller females, which could potentially lead to size‐assortative mating in nature.     

Variability of Female Responses to Conspecific vs. Heterospecific Male Mating Calls in Polygynous Deer: An Open Door to Hybridization?

Males of all polygynous deer species (Cervinae) give conspicuous calls during the reproductive season. The extreme interspecific diversity that characterizes these vocalizations suggests that they play a strong role in species discrimination. However, interbreeding between several species of Cervinae indicates permeable interspecific reproductive barriers. This study examines the contribution of vocal behavior to female species discrimination and mating preferences in two closely related polygynous deer species known to hybridize in the wild after introductions. Specifically, we investigate the reaction of estrous female red deer (Cervus elaphus) to playbacks of red deer vs. sika deer (Cervus nippon) male mating calls, with the prediction that females will prefer conspecific calls. While on average female red deer preferred male red deer roars, two out of twenty females spent more time in close proximity to the speaker broadcasting male sika deer moans. We suggest that this absence of strict vocal preference for species-specific mating calls may contribute to the permeability of pre-zygotic reproductive barriers observed between these species. Our results also highlight the importance of examining inter-individual variation when studying the role of female preferences in species discrimination and intraspecific mate selection.     

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.     

MATE CHOICE IN THE SAILFIN MOLLY,POECILIA LATIPINNA

We examined both female and male mate choice in the sailfín molly, Poecilia latipinna. Female mollies preferred larger males over smaller ones when comparing males from their own populations. Although the expression of this preference depends on a female's receptive state, the level of female preference does not appear to be associated with any other attribute of the female or of the males. When presented with males of the same size from different populations, females preferred native over foreign males in some but not all population combinations. These results cannot be explained by a bias for higher size-specific rates of courtship displays that is shared by all females. Males preferred larger over smaller females; larger males exhibited stronger preferences, and preference for the larger female also increased as the disparity in size between the two object females increased. We found no evidence that males preferred native over foreign females when encountered singly or in size-matched combinations. These results indicate that discrimination among populations arises because females exercise divergent directional preferences for size-specific trait values that are associated with differences among males in these values. This result implies an active role for sexual selection in contributing to the maintenance of the behavioral or morphological distinctions among males observed within and among populations.     

Evidence of an emerging female preference for an artificial male trait and the potential for spread via mate choice copying in Poecilia latipinna

The initial purpose of the project described herein was to assess the preference of female Poecilia latipinna for an artificial novel male trait—an orange‐colored distal fringe added to the caudal fin of an otherwise wild‐type conspecific. Analysis of the preliminary data revealed consistent individual differences in the strength of female preference for either orange‐tailed or wild‐type males. This finding inspired the study's second aim—to evaluate whether the preference for orange‐tailed males observed among a subset of females could spread via mate choice copying to others in the population that initially preferred wild‐type males. Two experiments and a control were conducted wherein females were simultaneously presented with an orange‐tailed and a wild‐type dummy male using a standard dichotomous choice design. In the first experiment, female preference was assessed on two separate occasions in order to characterize the variability and consistency in preference for orange‐tailed versus wild‐type males. The second experiment addressed mate choice copying: Female preference was again assessed on two separate occasions, but involved pairing a model female with the non‐preferred male for a period of time between preference tests. A third set of control tests were conducted using the same protocol as the copying experiment except that subject females were unable to see the model paired with the non‐preferred male. Results showed that, although females collectively preferred neither the orange‐tailed nor the wild‐type dummy male in the first round of preference tests, the majority showed relatively strong individual preferences. The subset of females that preferred the orange‐tailed over the wild‐type male in the first round of testing all maintained their preferences in the second round whether or not they had observed a model in association with the non‐preferred wild‐type male between tests. However, females that preferred the wild type over the orange male in the first round of testing copied the model's choice of the non‐preferred orange‐tailed male in their second round of preference testing. These results highlight the importance of recognizing the likelihood that only a subset of females will express a preference as it first emerges within a population. In such instances, the preference may not be detected at the population level—a point frequently overlooked in studies of mate choice. Additionally, these data highlight the importance of assessing the preferences of individual females and their capacity to drive evolutionary change within populations. Lastly, this study offers evidence of a possible mechanism by which a novel male trait might spread via mate choice copying by exploiting an emerging sensory bias within a subset of females in the population.