Individually Specific Call Feature Is Not Used to Neighbour-Stranger Discrimination: The Corncrake Case

In various contexts, animals rely on acoustic signals to differentiate between conspecifics. Currently, studies examining vocal signatures use two main approaches. In the first approach, researchers search for acoustic characteristics that have the potential to be individual specific. This approach yields information on variation in signal parameters both within and between individuals and generates practical tools that can be used in population monitoring. In the second approach, playback experiments with natural calls are conducted to discern whether animals are capable of discriminating among the vocal signatures of different individuals. However, both approaches do not reveal the exact signal characteristics that are being used in the discrimination process. In this study, we tested whether an individual-specific call characteristic – namely the length of the intervals between successive maximal amplitude peaks within syllables (PPD) – is crucial in neighbour-stranger discrimination by males of the nocturnal and highly secretive bird species, the corncrake (Crex crex). We conducted paired playback experiments in which corncrakes (n = 47) were exposed to artificial calls with PPD characteristics of neighbour and stranger birds. These artificial calls differed only in PPD structure. The calls were broadcast from a speaker, and we recorded the birds'' behavioural responses. Although corncrakes have previously been experimentally shown to discriminate between neighbours and strangers, we found no difference in the responses to the artificial calls representing neighbours versus strangers. This finding demonstrates that even if vocal signatures are individual specific within a species, it does not automatically mean that said signatures are being crucial in discrimination among individuals. At the same time, the birds'' aggressive responses to the artificial calls indicated that the information transmitted by PPDs is important in species-specific call recognition and may be used by males and/or females to evaluate sender quality, similarly like sound frequency in some insect species.     

The Responses of Duetting Antbirds to Stereo Duet Playback Provide Support for the Joint Territory Defence Hypothesis

In multiple animal taxa, including many birds and primates, members of mated pairs produce coordinated acoustic displays known as duets. By observing the behaviour of territorial animals as they respond to playback‐simulated duets of rivals, we can gain insight into the behavioural significance of vocal duets. Playback experiments, however, have been conducted across a very narrow range of duetting animals. Furthermore, many studies have been conducted with single‐speaker playback, whereas stereo‐speaker playback offers more spatially realistic simulation of duets. Moreover, by evaluating the reactions of animals to separate loudspeakers broadcasting male and female duet contributions, we can study the interactions of both males and females with same‐sex vs. opposite‐sex rivals. We used a paired experimental design to broadcast duet stimuli through a single‐speaker and a stereo‐speaker apparatus to 30 pairs of duetting barred antshrikes Thamnophilus doliatus in Costa Rica. Our goals were (1) to evaluate whether territorial antbirds respond more aggressively to male vs. female duet components and (2) to assess aggressive responses of antbirds towards single‐speaker vs. stereo‐speaker playback. Neither males nor females differentiated between the loudspeaker simulating the male vs. female duet contribution during stereo‐speaker playback trials. Barred antshrikes displayed significantly stronger responses to stereo‐speaker playback compared with single‐speaker playback. Males displayed stronger playback responses than females with closer, quicker and more vocal responses. These results provide evidence for a joint resource defence function of antbird duets given that pairs responded together with equivalent intensity to male and female simulated intruders. This is the first study to show that although duetting is an aggressive territorial signal, birds do not necessarily respond to sex‐specific components of duets. Our results support the idea that spatially realistic stereo presentation of duet stimuli is critical for experimental duet research.     

A review of acoustic playback techniques for studying avian vocal duets

ABSTRACT Playback experiments involve the broadcast of natural or synthetic sound stimuli and provide a powerful tool for studying acoustic communication in birds. Playback is a valuable technique for exploring vocal duetting behavior because it allows investigators to test predictions of the various hypotheses for duet function. Here, we adopt a methodological perspective by considering various challenges specific to studying duetting behavior, and highlighting the utility of different playback designs for testing duet function. Single‐speaker playback experiments allow investigators to determine how duetting birds react to different stimuli, but do not simulate duets in a spatially realistic manner. Multi‐speaker playback experiments are superior to single‐speaker designs because duet stimuli are broadcast with spatial realism and unique and additional predictions can be generated for testing duet function. In particular, multi‐speaker playback allows investigators to evaluate how birds respond to male versus female duet contributions separately, based on reactions to the different loudspeakers. Interactive playback allows investigators to ask questions about the time‐ and pattern‐specific singing behavior of birds, and to understand how singing strategies correspond to physical behavior during vocal interactions. Although logistically challenging, interactive playback provides a powerful tool for examining specific elements of duets (such as the degree of coordination) and may permit greater insight into their functions from an operational perspective. Interactive playback designs where the investigator simulates half of a duet may be used to describe and investigate the function of pair‐specific and population‐wide duet codes. Regardless of experimental design, all playback experiments should be based on a sound understanding of the natural duetting behavior of the species of interest, and should aim to produce realistic and carefully controlled duet simulations. Future studies that couple playback techniques with other experimental procedures, such as Acoustic Location System recordings for monitoring the position of birds in dense vegetation or multimodal techniques that combine acoustic with visual stimuli, are expected to provide an even better understanding of these highly complex vocal displays.     

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

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

Quiet violence: Savannah Sparrows respond to playback‐simulated rivals using low‐amplitude songs as aggressive signals

When animals compete over resources such as breeding territories, they often use signals to communicate their aggressive intentions. By studying which signals are associated with aggressive interactions, we gain a deeper appreciation of animal behaviour. We studied aggressive signalling in male Savannah Sparrows, Passerculus sandwichensis, focusing on signals that precede physical attack against territorial intruders. We simulated intruders using song playback and taxidermic models, and we determined which behaviours were associated with physical attack. Previous studies that have used this approach suggest that many species produce songs of dramatically lower amplitude, or “soft songs,” as signals of aggressive intention. Savannah Sparrows, however, are not known to produce soft songs, and therefore, they provided an interesting system for testing signals that predict attack. Of 93 playback subjects, 23 males attacked the simulated intruder and 70 did not. To our surprise, Savannah Sparrows produced soft songs, and the number of soft songs was a significant predictor of attack on the simulated intruder. Birds also showed a nonsignificant tendency to produce more “chip” calls prior to attack on the simulated intruder, whereas three other measured behaviours (aggressive calls, wing waving and passes over the model) did not predict attack. Our study contributes to the growing body of research on aggressive territorial signals and reveals that soft song is an even more widespread signal of aggression in songbirds than previously recognized.     

Soft songs in male ortolan buntings are used in an aggressive context but are not an aggressive signal

Many bird species produce low‐amplitude acoustic signals that have been poorly studied in comparison with loud, broadcast songs used for mate attraction and repelling rivals. In some birds, these soft signals were found to be emitted in an antagonistic context and were the most reliable predictor of a subsequent physical attack. The function of this signal is poorly understood, and several hypotheses have been proposed to explain the mechanisms and possible functions of such low‐amplitude signals. The subject of this study is the ortolan bunting, a small passerine species that produces soft songs during territorial defence. In this study, we aim to study whether the soft songs of the ortolan bunting are a signal of increased aggressiveness by testing if they meet the context, prediction and response criteria of aggressive signals. We simulated stranger male intrusion into a focal male territory with three different playback experiments. We found no significant differences in the male responses to the taxidermic model regardless of whether they were or were not producing soft songs in response. The males responded more strongly to loud songs than to soft songs during the simulated intrusions, and the males did not treat soft songs as a predictor of conflict escalation. Although soft songs clearly appeared during territorial encounters and were not present during spontaneous singing before the intrusions, our results did not support the hypothesis that soft songs indicate aggressive character. We suggest that soft songs in the ortolan bunting are intentionally used by birds to modify their intentions or target‐specific individuals within a close range.     

Male orientation on vocalization perches could optimize acoustic signal transmission in anurans

Acoustic signals are distorted by vegetation, wind currents, or other sounds when transmitted through the environment. Consequently, vocalizations with features that optimize sound transmission or behaviors that improve the efficacy of communication have evolved in many animal species. Among behavioral strategies, some species call from perches above the ground to increase the propagation distance of their acoustic signals. However, the orientation in the perch also influences the transmission of the vocalizations, so that frogs calling from different orientations (i.e., horizontal, upward, or downward) may affect differently the quality and efficacy of sound transmission. We implemented a sound transmission experiment to test for the effect of calling orientation (upward, downward, and horizontal) and distance on the attenuation and degradation of advertisement calls in the common dink frog Diasporus diastema. We broadcasted and re‐recorded advertisement calls at 2 m height, setting the speaker in three directions (upward, downward, and horizontal) to simulate different signaler orientations. We found that attenuation of the advertisement calls is significantly reduced when the speaker was directed either upward or downward, rather than horizontally. However, the degradation of call is lower when the speaker is direct horizontally. Since calls produced from either upward or downward orientations could travel farther, they could be used to signal male spatial location, while calls produced from a horizontal position could provide information on male quality at shorter distances at advanced phases of courtship.     

Reproductive isolation inProkelisia planthoppers (Homoptera: Delphacidae): Acoustic differentiation and hybridization failure

Males and females of Prokelisia marginata (Van Duzee) and Prokelisia dolusWilson communicate through substrate-transmitted vibrations. The acoustic signals (attraction and courtship calls) of these planthoppers are effective in mate location, attraction, and mate choice. Attraction calls are structurally distinct for both species and differ in pulse type, pulse repetition rate, and pulse duration. Using playback of prerecorded calls, individuals discriminated between conspecific and heterospecific signals. Depending on the sex and species, response calls were produced three to eight times more frequently to conspecifics than to heterospecifics. However, acoustic signals alone did not explain reproductive isolation and hybridization failure in these two congeners. Some heterospecific pairs called, courted, and attempted to join genitalia, but no connections were successful and no progeny were produced. Thus, acoustic behavior is not a guaranteed premating isolating mechanism in no-choice situations. Other courtship behaviors and possibly morphological differences in genitalia also contributed to their isolation. Females displayed a variety of rejection behaviors to conspecific and heterospecific males, suggesting that sexual selection (female choice), in addition to species recognition, may be an important force in the evolution of the acoustic signals of planthoppers. Although signal structure was not dependent on wing form (planthoppers exhibit wing dimorphism), the age when females first began to call was related to wing form. Brachypterous (flightless) females of both species began calling early in adult life (day 2), whereas macropterous (migratory) females began calling later in adult life (day 6). This pattern is consistent with the oogenesis-flight syndrome, in which reproductive maturity is delayed until after migration occurs.     

Effects of androgens and estrogens on crowings and distress callings in male Japanese quail, Coturnix japonica

In male Japanese quail, crowing behavior is considered to be strictly androgen-dependent. It was previously shown that in chicks, treatment with either testosterone or 5alpha-dihydrotestosterone (5alpha-DHT; a non-aromatizable androgen) induced crowing with motivation for distress calling in acutely isolated conditions. Many studies, however, have shown that the potencies of testosterone and 5alpha-DHT in activating crowing in castrated males are different. To clarify the effects of androgenic and estrogenic actions on the production of crows and distress calls, we injected quail daily from 11 to 42 days after hatching (Day 11 to 42) with testosterone propionate (TP), 5alpha-DHT, estradiol benzoate (EB) or vehicle and examined their calling behaviors both in a recording chamber (acutely isolated conditions) and in their home-cages (well-acclimated conditions). Both TP- and 5alpha-DHT-treated birds began to crow by Day 13 when isolated in the recording chamber. The TP-treated birds, however, crowed less frequently than 5alpha-DHT-treated ones. This, combined with the observations that distress calling was strongly inhibited in EB-treated birds, suggests that estrogen converted from testosterone may inhibit the motivation for distress calling. On the other hand, after chronic treatment of TP, but not of 5alpha-DHT, birds began to crow intensely in their home-cages earlier than vehicle treated controls, suggesting that estrogen is needed to initiate crowing behavior in sexually active males. Taken together, it is suggested that estrogenic actions affect the motivation underlying vocal behaviors, while the androgenic action is indispensable in generating crowing.     

Individuality,kin similarity and experimental playback of contact calls in cooperatively breeding riflemen

Riflemen/tītipounamu (Acanthisitta chloris) are kin-based cooperatively breeding birds, which appear able to recognise their relatives. Here, we investigate the potential for vocalisations to act as recognition cues in riflemen. We identified an appropriate contact call and recorded it at the nest from 19 adult riflemen. Measurements of call characteristics were individually repeatable. In addition, call similarity was significantly correlated with relatedness among all birds and among males. Thus, in principle, these contact calls contain sufficient information for individual recognition of familiar kin, and some assessment of relatedness between unfamiliar birds. To test whether riflemen responded differently to calls of kin, we broadcast calls of relatives and non-relatives as separate treatments in a playback experiment. Focal birds rarely responded aggressively or affiliatively, and their tendency to do so was unrelated to treatment. We conclude that zip calls are suitable kin recognition cues, but whether they are used as such remains unknown.     

No need to shout: Effect of signal loudness on sibling communication in barn owls Tyto alba

In animal communication, signal loudness is often ignored and seldom measured. We used a playback experiment to examine the role of vocal loudness (i.e., sound pressure level) in sibling to sibling communication of nestling barn owls Tyto alba. In this species, siblings vocally negotiate among each other for priority access to parental food resources. Call rate and call duration play key roles in this vocal communication system, with the most vocal nestlings deterring their siblings from competing for access to the food item next delivered by parents. Here, we broadcast calls at different loudness levels and call rate to live nestlings. The loudness of playback calls did not affect owlets' investment in call rate, call duration or call loudness. The rate at which playback calls were broadcast affected owlets' call rate but did not influence their response in terms of loudness. This suggests that selection for producing loud signals may be weak in this species, as loud calls may attract predators. Moreover, given that owlets do not overlap their calls and that they communicate to nearby siblings in the silence of the night, loud signals may not be necessary to convey reliable information about food need.     

Not for Parents Only: Begging Calls Allow Nest‐Mate Discrimination in Juvenile Zebra Finches

The benefits of recognition of family members may range from inbreeding avoidance to cooperative and coordinated behaviors within the family group. In birds, recognition of family members has almost exclusively been studied between parents and offspring or within cooperatively breeding societies. Yet, recognition of nest‐mates could be of special importance in recently fledged birds of colonial species by helping in locating the nest, maintaining family group cohesion, or allowing detection of feeding opportunities by recognizing the begging calls nest‐mates produced on the return of a parent. Here we study nest‐mate discrimination based on begging calls in fledglings of domesticated zebra finches (Taeniopygia guttata), a gregarious songbird living in loose colonies in which juveniles may gather in crèches and are fed by parents up to 20 d after fledging. Using playback tests, we show that fledglings called more and spent more time near the loudspeaker in response to the begging calls of their nest‐mates than to the calls of other familiar individuals. Because each fledgling was exposed to the repeated association of the begging calls of its nest‐mates and the subsequent feeding of its parents, this preferential response to the nest‐mates' calls could be a conditioned response to the food reward. Whereas fledglings answered more to male fledgling calls than to female fledgling calls, response to playback was influenced neither by the sex of the subject nor by its brood size. Discriminant function analysis based on acoustic parameters showed that begging calls carried an individual signature as well as a brood signature which might account for such nest‐mate discrimination. Begging signals are major study systems of the evolution of communication in the face of conflicts of interest between signalers and receivers. Our results suggest that eavesdropping and communication networks may be other informative frameworks to understand the design of offspring solicitation signals.     

Gecko phonotaxis to cricket calling song: A case of satellite predation

Insectivorous Mediterranean house geckos, Hemidactylus tursicus, are found in the field close to the burrows of calling male decorated crickets, Gryllodes supplicans. In playback experiments where adult geckos were presented with calls of these crickets or calls of frogs as controls, geckos exhibited positive phonotaxis to broadcast cricket calling song. This indicates that geckos orient to and approach male cricket calls, even though the crickets call from burrows where they are protected from the geckos. However, this behaviour enables the geckos to intercept and consume female crickets that also respond phonotactically to the cricket calls. Thus geckos act as ‘satellite predators’, a situation which may impose sex-biased mortality on female crickets.     

Acoustical communication and the mating system of the Australian whistling moth Hecatesia exultans (Noctuidae: Agaristinae)

Males of the Australian whistling moth Hecatesia exultans produce ultrasonic acoustical signals while perched on low vegetation. Some males call more or less continuously for several hours during midday with individuals occupying the same general calling area for up to several weeks. The nearest neighbour of calling males is typically 15 to 25m distant, at the outer edge of the estimated range at which neighbours can detect each other's ultrasonic signals. Calling male intruders occasionally enter an occupied territory, resulting in aerial clashes with nearly continuous signalling by both combatants. Males respond to playback of taped signals by flying toward the speaker and sometimes by calling while perched on or near the speaker. Females sometimes visit calling males, with copulation following very soon after the female alights on vegetation near the male's perch. Males increase the rate of sound production by about 11% when presented with moving pinned specimens or paper models of conspecifics. These observations and experiments indicate that males use ultrasound as long-distance communication signals designed to attract sexually receptive females and to establish territorial residency in competition with other males.     

Elevated stress hormone diminishes the strength of female preferences for acoustic signals in the green treefrog

Mate selection can be stressful; time spent searching for mates can increase predation risk and/or decrease food consumption, resulting in elevated stress hormone levels. Both high predation risk and low food availability are often associated with increased variation in mate choice by females, but it is not clear whether stress hormone levels contribute to such variation in female behavior. We examined how the stress hormone corticosterone (CORT) affects female preferences for acoustic signals in the green treefrog, Hyla cinerea. Specifically, we assessed whether CORT administration affects female preferences for call rate — an acoustic feature that is typically under directional selection via mate choice by females in most anurans and other species that communicate using acoustic signals. Using a dual speaker playback paradigm, we show that females that were administered higher doses of CORT were less likely to choose male advertisement calls broadcast at high rates. Neither CORT dose nor level was related to the latency of female phonotactic responses, suggesting that elevated CORT does not influence the motivation to mate. Results were also not related to circulating sex steroids (i.e., progesterone, androgens or estradiol) that have traditionally been the focus of studies examining the hormonal basis for variation in female mate choice. Our results thus indicate that elevated CORT levels decrease the strength of female preferences for acoustic signals.     

Variable effect of playback of chickadee mobbing calls on detection probability of boreal forest birds

Modification of the point count survey method to include playback of songbird mobbing calls in an attempt to increase detection probabilities has met with mixed success. We compared detection probabilities for boreal forest songbirds using traditional point count methods and counts using broadcasts of the mobbing calls of Black‐capped Chickadees (Poecile atricapillus) in an attempt to increase detection probability. We conducted 594 point counts during the 2010 breeding season in Newfoundland, Canada. Each point count consisted of an 8‐min silent observation period followed by an 8‐min broadcast of Black‐capped Chickadee mobbing calls. Occupancy model results showed that response to playback broadcast varied across species, with detection probabilities higher for seven of 17 species during the silent portions of point counts and three species more likely to be detected during playback intervals. For all species, the number of visual detections increased during periods of playback and, averaged across species, individuals were >6 times more likely to be seen during the playback period than during the silent period. Differences in detection probability among observers were apparent during both silent and playback periods. We suggest that using playback of chickadee mobbing calls during point count surveys of common boreal forest songbird species may be most beneficial when visual detection is important. However, playback may also be useful for species‐specific surveys during periods when birds are less likely to be vocal or for studies of less common species with chronically low detection probabilities. A combined silent and playback approach could also be useful, although observer and species differences should be accounted for if comparing data across species or studies.     

Sexual Differences in Parental Investment in Response to Parent‐Absent Calls

Begging in birds is a complex behaviour used by nestlings to solicit feeds from caregivers. Besides calling when parents are present, nestlings of some species also perform less conspicuous repeat calls when parents are absent. The fact that these calls are produced when parents are not at the nest does not mean that parents cannot hear them when they approach the nest or forage in its vicinity. In this study, we experimentally investigated the relationship between parent‐absent repeat calls (ARC) and frequency of parental visits, considering parent/offspring communication as a possible implication of these acoustic signals. A playback experiment was conducted to detect changes in parental investment in response to increases in parent‐ARC, expecting a differential sexual response. Results showed that females clearly responded to repeat calls, increasing their visit rate significantly with respect to females that received the control treatment. Males, on the contrary, did not change their visit rate in response to the treatment. This result provides evidence for a role of parent/offspring communication in parent‐absent repeat calling, an additional function to sibling negotiation processes. The sex‐specific response that we found is in agreement with previous studies that have found that females are more responsive than males to variation in solicitation and hunger signals performed by nestlings.     

Experience-based plasticity of acoustically evoked aggression in a territorial frog

Territorial male bullfrogs ( Rana catesbeiana) display lower levels of aggression toward familiar territorial neighbors compared to unfamiliar individuals based on the perception of individual differences in vocalizations. Stimulus-specific habituation elicited by repeated exposures to a neighbor's vocalizations is believed to play a role in mediating the low levels of aggression between neighbors. The present study describes habituation in multiple components of the bullfrog territorial aggressive response and examines whether these separate response components habituate at similar or different rates. In response to repeated broadcasts of synthetic bullfrog advertisement calls in a field playback experiment, the numbers of aggressive calls and movements, the latency to the first aggressive call, and the distance approached toward the playback speaker exhibited significant response decrements that developed at nearly identical rates. All four responses exhibited recovery upon subsequent broadcasts of a novel stimulus simulating a different individual. In contrast, the number of advertisement calls did not exhibit significant decrements with repeated playbacks. The patterns of stimulus-specific response decrement were inconsistent with explanations such as sensory adaptation, effector fatigue, non-specific changes in motivation, and a simple form of auditory habituation. Possible mechanisms for the observed plasticity in aggression are discussed in the context of anuran communication.     

Escalation of aggressive vocal signals: a sequential playback study

Rival conspecifics often produce stereotyped sequences of signals as agonistic interactions escalate. Successive signals in sequence are thought to convey increasingly pronounced levels of aggressive motivation. Here, we propose and test a model of aggressive escalation in black-throated blue warblers, presenting subjects with two sequential and increasingly elevated levels of threat. From a speaker outside the territorial boundary, we initiated an interaction (low-threat level), and from a second speaker inside the territory, accompanied by a taxidermic mount, we subsequently simulated a territorial intrusion (escalated threat level). Our two main predictions were that signalling behaviours in response to low-threat boundary playback would predict signalling responses to the escalated within-territory threat, and that these latter signalling behaviours would in turn reliably predict attack. We find clear support for both predictions: (i) specific song types (type II songs) produced early in the simulated interaction, in response to boundary playback, predicted later use of low-amplitude ‘soft’ song, in response to within-territory playback; and (ii) soft song, in turn, predicted attack of the mount. Unexpectedly, use of the early-stage signal (type II song) itself did not predict attack, despite its apparent role in aggressive escalation. This raises the intriguing question of whether type II song can actually be considered a reliable aggressive signal. Overall, our results provide new empirical insights into how songbirds may use progressive vocal signalling to convey increasing levels of threat.     

Song matching, overlapping, and switching in the banded wren: the sender's perspective

Interpreting receiver responses to on-territory playback ofaggressive signals is problematic. One solution is to combinesuch receiver-perspective experiments with a sender-perspectiveexperiment that allows subjects to demonstrate how their choiceof singing strategies is associated with their approach behavior.Here we report the results of a sender-perspective study onthe banded wren (Thryothorus pleurostictus) and combine informationon context and results of previous receiver-perspective experimentsto clarify function. Territorial males were presented with a5-min playback consisting of song types present in their repertoire.We assessed the degree to which the subjects' song-matchingrate, overlapping rate, and song-type versatility were correlatedwith their approach latency, closeness of approach, latencyto first retreat, and time spent close to the speaker. Maleage, breeding stage, and features of the playback stimuli werealso considered. Song matching was associated with rapid andclose approach, consistent with the receiver-perspective interpretationof type matching as a conventional signal of aggressive motivation.Overlapping was associated with earlier retreat and togetherwith the aversive receiver response to our previous overlappingplayback experiment suggests that overlapping is a defensivewithdrawal signal. High versatility was associated with slowerfirst retreat from the speaker and high levels of reciprocalmatching between subject and playback. Males with fledglingssang with particularly low versatility and approached the speakeraggressively, whereas males with nestlings overlapped more andretreated quickly. Finally, older males matched more but overlappedless.