Vocal Responsiveness to Interfering Sounds By a Frog From The Southern Temperate Forest,Batrachyla leptopus

Interfering sounds from biotic and abiotic origins are likely to shape the responsiveness of sound communicating animals. Among these sources of interference, interactions among acoustically active species have been studied to quite a limited extent. The vocal responses of 20 male frogs Batrachyla leptopus from the temperate austral forest in Chile were tested with conspecific calls and with the calls of two sympatric species: Batrachyla taeniata and Batrachyla antartandica, broadcast at amplitudes of 73, 79, 85, 91 and 97 dB SPL peak. Also, the vocal activity of the subjects during exposure to a 3‐min continuous broadband noise presented at 67 dB SPL RMS was monitored. The subjects gave higher responses on average to the conspecific relative to the heterospecific calls, but in most comparisons, these differences did not reach levels of significance. In addition, the vocal activity of males of B. leptopus did not increase in the presence of the continuous broadband noise. The lack of clear preferential responses for conspecific signals contrasts with the sharp selectivities that B. taeniata and B. antartandica have shown for their own calls in previous studies. Such different vocal behavior could be related to the extensive geographic overlap of B. leptopus with the two other species in the temperate austral forest, where mixed choruses of this species with each of the two congeneric taxa have been reported to occur occasionally. The lack of vocal activation in the presence of continuous noise also contrasts with the increased vocal output with which the other two taxa respond to this intrusion and is likely to result from a relatively high spontaneous vocal activity in B. leptopus.     

Vocal Strategies in Confronting Interfering Sounds by a Frog from the Southern Temperate Forest,Batrachyla antartandica

Animals are communicating by sound face interference from biotic and abiotic sources. Contrasting strategies have been reported in different taxa in the presence of prolonged noises, but in particular, interactions among acoustically active species have been studied to a very limited extent. In addition, reactions of a single species to interferences having contrasting structural patterns have not been explored systematically. The vocal responses of 16 male frogs Batrachyla antartandica from the temperate austral forest in Chile were tested with conspecific calls and with the calls of two sympatric species: B. taeniata and B. leptopus, broadcast at amplitudes of 73, 79, 85, 91, and 97 dB peak sound pressure level (SPL). Also, the vocal activity of the subjects during exposure to a 3‐min continuous broadband noise presented at 67 dB root mean square (RMS) SPL was monitored. The subjects gave significantly higher responses to the conspecific relative to the heterospecific calls but increased their vocal output in the presence of continuous noise. The preference of vocal responses for the conspecific relative to heterospecific signals bears a general resemblance to those exhibited by B. taeniata in a previous study and potentially contributes to the typical segregation in conspecific choruses observed in areas where these frogs breed in sympatry. Such lack of vocal responses to heterospecific stimuli contrasts with the increase in vocal activity during exposure to continuous noise lacking a temporal fine structure and points to identify different strategies in confronting interferences of diverse origin.     

Restricted responsiveness to noise interference in two anurans from the southern temperate forest

Animals adopt different strategies to communicate by means of sound in noisy environments such that some species increase, while others decrease their vocal activity in the presence of interference. Anuran amphibians from diverse latitudes exhibit both kinds of responses. Recent studies have shown that males of Batrachyla taeniata and Batrachyla antartandica from the temperate austral forest do not call in response to the presentation of advertisement calls of sympatric congeneric species. In contrast, Batrachyla leptopus responds to these signals in a similar way as to conspecific calls. The responsiveness of B. taeniata to natural abiotic interference has also been tested and found that noises of such sources produce strong increases in vocal activity. To assess the diversity in responsiveness to acoustic intrusion in this group, we exposed males of B. leptopus and B. antartandica to prolonged pre‐recorded natural abiotic noises of wind, creek, rain, and to a band‐pass noise centered at 2,000 Hz, at 67 dB Sound Pressure Level (SPL). The subjects did not increase their vocal activity significantly when exposed to these sounds and to band‐pass noise at increasing intensities (55–79 dB SPL). These results contrast with the increase in vocal activity observed previously in B. taeniata to continuous abiotic noise and point to the existence of diverse strategies to confront acoustic intrusion among related species. The lack of vocal activation observed also contrasts with the responsiveness of B. leptopus to heterospecific signals, but parallels the lack of response to such sounds in B. antartandica. Furthermore, the results obtained contrast with the responsiveness of these species to synthetic prolonged sounds observed in previous studies, suggesting that the modes of responses to acoustic intrusion may depend on previous experience, rather than having a species‐specific nature.     

Behavioural response evoked by conspecific distress calls in two neotropical treefrogs

Anurans emit distress calls when attacked by predators as a defensive mechanism. As distress calls may trigger antipredator behaviour even in individuals that are not under attack, we tested whether this defensive behaviour induced behavioural changes in neighbouring conspecifics. We compared the behavioural responses of two species of Neotropical hylid frogs (genus Boana) to conspecific distress calls and white noise. Individuals of both species interrupted their vocal activity and decreased call rate after hearing the distress call. Natural variation on signal intensity calibrated among the nearest neighbours did not influence the response and we did not observe negative phonotaxis after any acoustic stimulus. Despite the fact that many predators are acoustically oriented, we could not determine if such response (reduced call rate) was induced by risk assessment or by the masking effect on advertisement calls. Boana faber responded similarly to white noise and distress calls, while B. bischoffi responded more intensely to distress calls. Duration of silence after playbacks in B. faber was longer than B. bischoffi. We suggest that, if the signals are interpreted as a risk cue by neighbouring conspecifics, each species may be preyed upon by different predators, as they may have led to distinct defensive strategies and different responses to distress calls. If risk assessment information is included in distress calls, it triggers behavioural responses only in the nearest neighbours, as we did not observe responses on the vocal activity of the interspecific chorus. Our results add relevant data about acoustic communication and interpretations by anurans, highlighting the importance of considering cues within common and widespread signals.     

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.     

Species differences of male loud calls and their perception in sulawesi macaques

Playback experiments were conducted to investigate interspecific discrimination of male loud calls in Sulawesi macaques. Loud calls of four macaque species living in Sulawesi (Macaca tonkeana, M. maurus, M. hecki, andM. nigrescens) and a control stimulus (an 8-sec frequency modulated sound) were played back to semi-free-ranging Tonkean macaques (M. tonkeana). A preliminary acoustic analysis indicated that the calls of these four species differ in some spectral and temporal features. In the playback experiments, Tonkean macaques responded in a similar manner to conspecific calls and calls of two other species,M. maurus andM. hecki. In contrast, animals responded more weakly to the call ofM. nigrescens and the control stimulus. Males responded more strongly than females to all stimuli, while females appeared to be more discriminating for species differences than males. Analyses on the acoustic features of loud calls suggested that high frequency, wide frequency range, and repetition of sound units at a high rate elicit quick responses from animals.     

Phonotaxis to male’s calls embedded within a chorus by female gray treefrogs, Hyla versicolor

During the reproductive season, male Hyla versicolor produce advertisement calls to attract females. Females exhibit phonotaxis and approach the individual callers, resulting in amplexus. For frogs that call from dense choruses, the extent to which and the range from which a male’s advertisement call within a chorus can be heard by a receptive female leading to phonotaxis is unclear. We investigated females’ responses to natural choruses in the field and found that they were attracted and showed directed orientation to breeding choruses at distances up to 100 m. To assess the role of acoustic cues in the directed orientation, we conducted acoustic playback experiments in the laboratory using conspecific call and noise as stimuli, as well as chorus sounds (that contained calls from a focal male) recorded at various distances, all played at naturalistic intensities. Using two response metrics (females’ normalized response times and their phonotaxis trajectories) we found that, unlike the field experiments, females oriented and were attracted to chorus sounds from 1 to 32 m only, but not from >32 m, or to band-limited noise. Possible reasons for the observed difference in phonotaxis behavior in the two experimental conditions were discussed.     

Differential Response to Interspecific and Intraspecific Signals Amongst Chickadees

Black‐capped chickadees (Poecile atricapillus) and mountain chickadees (P. gambeli) have a similar vocal repertoire and share many other life history traits; yet, black‐capped chickadees are socially dominant to mountain chickadees where populations overlap. Previous research suggested that in contact zones, both species respond weakly to heterospecific songs during the breeding season, and have suggested minimal interspecific competition. However, both black‐capped and mountain chickadees discriminate between conspecific and heterospecific chick‐a‐dee calls, suggesting attention is paid to interspecific signals. We compared the responses of both black‐capped and mountain chickadees to conspecific and heterospecific chick‐a‐dee calls during the winter, when both species compete for the same food resources. We conducted an aviary playback experiment exposing both species to playback composed of heterospecific and conspecific chick‐a‐dee calls, which had been recorded in the context of finding food sources. Responses from the tested birds were measured by recording vocalizations and behaviour. Black‐capped chickadees responded significantly more to conspecific than to heterospecific stimuli, whereas the subordinate mountain chickadees responded to both mountain and black‐capped chickadee calls. Based upon the reactions to playbacks, our results suggest these two closely related species may differ in their perception of the relative threat associated with intra‐ versus interspecific competitors.     

Robust calling performance in frogs infected by a deadly fungal pathogen

Reproduction is an energetically costly behavior for many organisms, including species with mating systems in which males call to attract females. In these species, calling males can often attract more females by displaying more often, with higher intensity, or at certain frequencies. Male frogs attract females almost exclusively by calling, and we know little about how pathogens, including the globally devastating fungus, Batrachochytrium dendrobatidis, influence calling effort and call traits. A previous study demonstrated that the nightly probability of calling by male treefrogs, Litoria rheocola, is elevated when they are in good body condition and are infected by B. dendrobatidis. This suggests that infections may cause males to increase their present investment in mate attraction to compensate for potential decreases in future reproduction. However, if infection by B. dendrobatidis decreases the attractiveness of their calls, infected males might experience decreased reproductive success despite increases in calling effort. We examined whether calls emitted by L. rheocola infected by B. dendrobatidis differed from those of uninfected individuals in duration, pulse rate, dominant frequency, call rate, or intercall interval, the attributes commonly linked to mate choice. We found no effects of fungal infection status or infection intensity on any call attribute. Our results indicate that infected males produce calls similar in all the qualities we measured to those of uninfected males. It is therefore likely that the calls of infected and uninfected males should be equally attractive to females. The increased nightly probability of calling previously demonstrated for infected males in good condition may therefore lead to greater reproductive success than that of uninfected males. This could reduce the effectiveness of natural selection for resistance to infection, but could increase the effectiveness of selection for infection tolerance, the ability to limit the harm caused by infection, such as reductions in body condition.     

Generalization in response to mate recognition signals

Females usually exhibit strong and unequivocal recognition of conspecific mating signals and reject those of other sympatric heterospecifics. However, most species are allopatric with one another, and the degree to which females recognize mating signals of allopatric species is more varied. Such mating signals are often rejected but are sometimes falsely recognized as conspecific. We studied the dynamics of mate recognition in female túngara frogs (Physalaemus pustulosus) in response to a series of calls that were intermediate between the conspecific and each of five allopatric-heterospecific calls: two that elicited recognition from females in previous studies and three that did not. This study shows that females perceive variation in allopatric mating signals in acontinuous manner with no evidence of perceptual category formation. The strength of recognition is predicted by how different the target stimulus is from the conspecific call within a series of calls. But the differences in recognition responses among call series are not predicted by the similarity of the call series to the conspecific call. The latter result suggests that the strength of recognition of allopatric signals might be influenced by processes of stimulus generalization and past evolutionary history.     

Responses of male cricket frogs (Acris crepitans) to attenuated and degraded advertisement calls

We examined the vocal and non‐vocal responses of male cricket frogs (Acris crepitans) to conspecific advertisement calls that had been attenuated or degraded by reducing the depth of amplitude modulation (AM). Both are characteristic of changes to the call as it is transmitted through natural habitats. As stimulus calls became more intense or less degraded, male cricket frogs gradually decreased their call rate and increased the number of call groups and pulse groups in their calls, changes indicative of increased aggressive interactions. At the higher intensities and lower degradation levels, the probability that males would shift to one of two non‐vocal behavioral responses, attacking the perceived intruder or ceasing calling and abandoning the call site, gradually increased. The results show that differences in signal attenuation and AM degradation levels are perceived by males and trigger both vocal and non‐vocal behavioral responses consistent with their use in evaluating the distance to a challenging male. Furthermore, the results indicate that the male responses are graded, increasing as intensity rises and degradation falls, and hierarchical, with vocal responses preceding behavioral responses over the range of intensities and degradation levels presented.     

Neural Activity Patterns in Response to Interspecific and Intraspecific Variation in Mating Calls in the Túngara Frog

Background

During mate choice, individuals must classify potential mates according to species identity and relative attractiveness. In many species, females do so by evaluating variation in the signals produced by males. Male túngara frogs (Physalaemus pustulosus) can produce single note calls (whines) and multi-note calls (whine-chucks). While the whine alone is sufficient for species recognition, females greatly prefer the whine-chuck when given a choice.

Methodology/Principal Findings

To better understand how the brain responds to variation in male mating signals, we mapped neural activity patterns evoked by interspecific and intraspecific variation in mating calls in túngara frogs by measuring expression of egr-1. We predicted that egr-1 responses to conspecific calls would identify brain regions that are potentially important for species recognition and that at least some of those brain regions would vary in their egr-1 responses to mating calls that vary in attractiveness. We measured egr-1 in the auditory brainstem and its forebrain targets and found that conspecific whine-chucks elicited greater egr-1 expression than heterospecific whines in all but three regions. We found no evidence that preferred whine-chuck calls elicited greater egr-1 expression than conspecific whines in any of eleven brain regions examined, in contrast to predictions that mating preferences in túngara frogs emerge from greater responses in the auditory system.

Conclusions

Although selectivity for species-specific signals is apparent throughout the túngara frog brain, further studies are necessary to elucidate how neural activity patterns vary with the attractiveness of conspecific mating calls.     

Vocal response of male European water frogs (Rana Esculenta complex) to mating and territorial calls

The responses of male European water frogs (the two species Rana lessonae and Rana ridibunda and their hybrid, Rana esculenta) to playback of their mating and territorial calls were studied during the mating season.In order to select biologically relevant intensities for the presentation of the recorded calls, the sound pressure of the calls produced by the frogs themselves was established prior to the experiment. At a distance of 1 m the most intense calls were those of R. ridibunda, with a sound pressure of 110 dB (peak SPL). The smaller males of R. esculenta gave calls about 5 dB lower in intensity. The calls of R. lessonae, the smallest phenotype, were still less intense, 10 dB lower than those of R. ridibunda.The territorial calls of all three phenotypes elicited territorial calls in all of the males tested, as a rule accompanied by approach to the sound source. The sound pressure required to elicit a vocal response was nearly the same for each of the three different territorial calls. Sometimes encounter calls and warning calls were given in addition to territorial calls.When the mating calls were presented at low intensity, in some cases the males responded with their own mating calls. Mating calls at higher intensity elicited the same behavior that appeared following presentation of territorial calls, but significantly higher sound pressures were required to elicit such a response to mating calls than to territorial calls. The males of R. lessonae and R. esculenta did not respond to the mating calls of R. ridibunda, and each of them had significantly lower thresholds to the mating call of its own phenotype than to that of the other. The males of R. ridibunda responded only to conspecific mating calls.The vocal-response thresholds are compared with those of the electrodermal response reacting to the same stimuli. The significance of the different calls of the European water frogs is discussed.     

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

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

The mixed-species chorus as public information: tungara frogs eavesdrop on a heterospecific

Multispecies choruses represent a promising but uninvestigatedforum for public information. Although frogs exposed to a potentialpredator call more readily in the presence of conspecific callsthan in their absence, none are known to make comparable useof heterospecific calls. To test for heterospecific eavesdropping,we isolated calling male túngara frogs (Physalaemus pustulosus),presented them with a potential predator, and recorded theirresponses to playbacks of 1 of 4 stimuli: calls of a conspecific,a sympatric heterospecific (Leptodactylus labialis), an allopatriccongener (Physalaemus enesefae), or silence. We found that malescalled more in response to the L. labialis call than to eitherthe silent stimulus or the P. enesefae call. In contrast, theP. enesefae call did not result in significantly more callingthan the silent stimulus. The conspecific call was the mosteffective at promoting calling. The data indicate that túngarafrogs selectively attend to the call of a heterospecific. Wehypothesize that such heterospecific eavesdropping contributesto the emergent behavior of mixed-species choruses.     

Complex Vocal Responses to Conspecific Calls in Whistling Frogs Eleutherodactylus johnstonei: Playback Experiments in the Field

We simulated the presence of an acoustic competitor by broadcasting conspecific playbacks to males of Johnstone's whistling frog, Eleutherodactylus johnstonei, in the field. We broadcast calls that differed in duration (short, typical, and long), dominant frequency (high, typical, and low), and period (short, typical, and long), and analyzed male vocal responses. We tested the hypothesis that males respond by escalating vocally when they are exposed to female‐attractive calls and by ignoring unattractive ones. At the population level, males responded to playbacks in ways that would potentially increase their attractiveness with regard to solo calling: males increased the duration, reduced the dominant frequency, and increased their calling effort (duty cycle), despite an increase in call period. The modification of call duration occurred only in response to playbacks of low‐frequency calls, long calls, and short‐period calls (selective response), while the modification of the dominant frequency was independent of the characteristic of the playback (fixed response). Contrary to the expected, males did not reduce the call period when they were exposed to attractive playbacks. At the ultimate level, the results suggest energy‐saving strategies. In addition, males seem to trade off call period for the avoidance of acoustic interference with attractive calls as calling effort was typically increased by increasing call duration but only rarely by reducing the call period. Interactive playbacks are necessary to better understand the calling strategies of males of E. johnstonei.     

Ravens adjust their antipredatory responses to con‐ and hetero‐specific alarms to the perceived threat

Heterospecific alarm calls are typically found in situations where multiple species have a common predator. In birds, they are particularly common in mixed mixed‐species flocks. In species with highly developed social and cognitive abilities like corvids, there is the potential for differential responses to heterospecific vs. conspecific calls according to the riskiness of the habitat. We tested the responses of free‐ranging ravens (Corvus corax) to conspecific alarm calls and compared them to heterospecific alarm calls of jackdaws (Corvus monedula). We observed the proportion of ravens leaving the feeding site after the con‐ or hetero‐specific playback was presented in a situation of low threat (wild boar—Sus scrofa enclosure) and high threat of predation (wolf—Canis lupus enclosure). We show that ravens responded to conspecific calls more intensively at the wolves than at the wild boar, but the response to conspecific calls was in both enclosures stronger than to the control (great tit—Parus major song). The response to the heterospecific alarm was also stronger in the wolves’ enclosure, but it did not differ from control in the wild boar enclosure. These findings suggest that ravens are aware of the meaning of the jackdaw alarm calls, but they respond to it only in a situation of high predatory threat (wolves are present). In the wild boar enclosure, the ravens probably consider jackdaws warning against some other predator, very probably harmless to ravens. This interpretation requires further testing, as both enclosures differ also in respect to other parameters like food quality and shelter availability.     

Choosing a Mate in a Cocktail Party‐Like Situation: The Effect of Call Complexity and Call Timing between Two Rival Males on Female Mating Preferences in the Túngara Frog Physalaemus pustulosus

Signal detection, recognition, and localization are hampered when multiple signalers coincide in time and space, a problem known as ‘cocktail party effect’. In many taxa, senders utter complex calls consisting of two or more elements which often vary in the ease with which they can be assessed in different signaling environments. Receivers’ selective attention to different cues may increase the probability of correctly assigning a signal to its source (localization) in face of conspecific interference. Túngara frogs, Physalaemus pustulosus, produce complex calls consisting of an initial whine, followed by zero up to seven broad‐banded, amplitude‐modulated chucks. Under ideal conditions (without interference or noise), females prefer whines followed by chucks over whines alone, but the preference is not linear; females do not discriminate between whines with one or two chucks. When whines lack chucks, call overlap elicits random responses in females, with no preference for leading calls. In this study, I explored the combined effect of call timing and call complexity on female preferences in a two‐choice paradigm—a simplification of the cocktail party scenario. I tested the hypothesis that the effect of call overlap can be reduced when the calls of one of the two rivals have chucks, specifically more chucks than those of the rival. I gave females a choice between whines alone and with chucks (one or two) presented at three time relations (alternated, abutted, and partially overlapped) and two emission orders (whine with less chucks leading and whine with more chucks leading). I found that the preference for one chuck over no chuck was preserved in all the experimental treatments, but when a w + 2chk preceded a w + chk, either overlapped or abutted, a preference existed for the whine with more chucks. Therefore, an interaction between call order and the number of chucks was obtained. The results only partially supported the hypothesis, and call order emerges as an opportunistic component of signaling in P. pustulosus.     

Variation in chick‐a‐dee calls of bridled titmice (Baeolophus wollweberi): Frequent use of non‐combinatorial calls in a combinatorial calling system

Chick‐a‐dee calls of Poecile (chickadee) and Baeolophus (titmouse) species are complex in terms of the structural composition of note types and the diversity of messages. Studies so far have mainly focused on the calls of various chickadee and just one titmouse species—the tufted titmouse (B. bicolor). To begin to address this lack of titmouse data, our study investigated variation in note composition of calls of bridled titmice (B. wollweberi). We obtained calls from 26 flocks in the Chiricahua Mountains of Arizona in the overwintering flocking period. Bridled titmice produce proportionally more non‐combinatorial call variants than combinatorial call variants. The number of the single noted calls furthermore exceeded the number of multinote calls. In general, structural variation in the combinatorial calls appears to be comparable to calls of better‐studied chickadees and of tufted titmice, although bridled titmice appear to have a unique call length distribution. We also analyzed some behavioral associations with call variation and found that flight behavior and close interactions between individuals were associated with use of specific note types. Finally, we found microgeographic variation in note type use in these calls. We discuss some possible explanations for call complexity in this species.     

Dual Function of Chip Calls Depending on Changing Call Rate Related to Risk Level in Territorial Pairs of White‐Eared Ground‐Sparrows

Different mechanisms have been proposed for encoding information into vocalizations: variation of frequency or temporal characteristics, variation in the rate of vocalization production, and use of different vocalization types. We analyze the effect of rate variation on the dual function of chip calls (contact and alarm) produced by White‐eared Ground‐sparrows (Melozone leucotis). We conducted an acoustic playback experiment where we played back 1 min of four chip call rates (12, 36, 60, 84 calls/min). We measured the response of territorial pairs using behavioral responses, and fine structural features of calls produced in response to those playbacks. White‐eared Ground‐sparrows showed more intense behavioral responses to higher than lower call rate playbacks. Both individuals of the pair approached the source of the playback stimulus faster, produced the first vocalization faster, produced more vocalizations, and spent more time close to the stimulus in higher call rate than in lower call rate playbacks. Frequency and duration characteristics of calls (chip and tseet) were similar in response to all call rate playbacks. Our playback experiment elicited different intensity of behavioral responses, suggesting that risk‐based information is encoded in call rate. Our results suggest that variation in the rate of chip call production serves a dual function in this species; calls are used at lower rates for pair contact and at higher rates for alarm/mobbing signals.