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

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

Graded Aggressive Calls in the African Painted Reed Frog Hyperolius marmoratus (Hyperoliidae)

The African painted reed frog, Hyperolius marmoratus, has a potentially complex communication system. Advertisement calls and aggressive calls, although distinct from each other, are in fact two ends of a continuum of graded calls. Playback experiments using standard advertisement calls showed that males increased the proportion of aggressive calls as the stimulus intensity was increased. In addition, three characteristics of the aggressive calls changed in response to higher playback levels. Males increased the number of pulses/call, increased call duration, and decreased dominant frequency. Aggressive calling occurred primarily during the early hours of the night, with considerable overlap with times when females were searching for mates in the chorus. Females tested in two-choice arena trials discriminated against aggressive calls in favor of advertisement calls. It is suggested that aggressive calls reduce a male's ability to attract a female and that a graded signalling system may enable males to escalate agonistic encounters with other males without rendering calls completely unattractive to females.     

Male Calling Behavior and Female Choice in the Neotropical Treefrog Hyla microcephala

The effect of stimulus call complexity and calling rate on the vocal responses of males and female mate choice was studied in Hyla microcephala in Panama. Males increased the number of notes in their calls in response to increases in stimulus call complexity during both playback of 1 to 8-note advertisement calls and during natural interactions. However, precise matching of the number of notes in stimuli and responses did not occur consistently. Males also increased calling rates if stimuli were presented above prestimulus rates. Two-stimulus choice experiments demonstrated that females prefer both higher calling rates and greater call complexity, indicating that the ways males change their vocal behavior during interactions increases their attractiveness to potential mates. Tests in which the relative intensity of a high and low rate stimulus was varied indicated that females prefer stimuli with higher total sound energy. In a natural chorus, it is likely that females simply approach males giving the most conspicuous calls.     

Males of <Emphasis Type="Italic">Hypsiboas goianus</Emphasis> (Anura; Hylidae) do not assess neighbor fighting ability through acoustic interactions

In frogs, acoustic signals are the most important communication mechanism, since they may be used in several social contexts. In many anurans, the dominant frequency of calls is negatively related to body size, and in such species, this spectral parameter may be considered a good predictor of fighting ability. We experimentally investigated the vocal behavior of 30 male Hypsiboas goianus in central Brazil to answer the following questions: (1) Do males change the acoustic parameters of their calls in response to conspecific intruders? and (2) Does the acoustic behavior of H. goianus depend on the simulated body size of their opponent? We used playback of synthesized calls with high (3573 Hz) and low (3123 Hz) dominant frequency to simulate small and large males, respectively. Males reduced the rate of advertisement calling in response to playback but did not change vocal behavior in response to low-frequency and high-frequency playback. So, while males adjust their calling activity in response to simulated conspecifics, there was no evidence that they assess the fighting ability of their opponents through acoustic interactions.     

Synchronized interdigitated calling in the Kuvangu running frog, Kassina kuvangensis

Male Kuvangu frogs show repetitive calling of pulsed advertisement calls in which up to seven calls are repeated in short succession. Recordings of pairwise interactions between males showed that calls were highly synchronized, with individual calls interdigitating with each other. Males frequently switched between the leader and follower role with neither male dominating the interaction. Interactive playback experiments using synthetic calls revealed that males slightly but significantly increased the number of calls per call group with increases in stimulus call number. Males also significantly increased call rate with the number of calls in the playback stimulus. Furthermore, when presented with shortened intercall intervals, males increased their own intercall intervals, thus ‘skipping’ a call and avoiding overlap with the playback. The low degree of call matching suggests that repetitive calling, apart from maintaining a male's attractiveness to females relative to rival males, may mediate male-male competition. In addition, synchronized interdigitated calling may serve to reduce predation, while maintaining species-specific temporal features of advertisement calls important to females. Kuvangu running frogs may have reduced the costs associated with synchrony and alternation by using a signal timing scheme that allows them to do both simultaneously.Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.       

Multisensory signals trigger approach behaviour in the fire-bellied toad Bombina orientalis: sex differences and call specificity

Animal communication often involves multimodal signals, and interactions between sensory modalities can trigger unique responses in receivers. Response to social signals was investigated in fire-bellied toads by exposing them to playback of male calls (advertisement and release calls) and a video clip of a male conspecific in the laboratory. The cues were presented in isolation and as a combined bimodal stimulus, and approach frequency, latency to approach and time spent around the stimulus source were measured. No positive phonotaxis was observed toward the advertisement call, both during the day and during a phonotaxis trial performed at night. However, females, but not males, approached with greater frequency, lower latency, and spent more time near the source of the bimodal stimulus in an experiment involving the advertisement call. Female response was specific to the advertisement call, as approach was not increased when the release call was used. Males, on the other hand, did not show increased approach in the advertisement call experiment, but approached with greater frequency the bimodal stimulus involving the release call within the first minute of stimulus presentation. The findings suggest that females orient toward calling males and that males eavesdrop on release calls, but in both cases a visual stimulus is also needed to trigger a response. Social approach in Bombina orientalis is thus dependent on multisensory cues, and the nature of the interaction between sensory modalities depends on receiver sex and call type.     

Temporal call changes and prior experience affect graded signalling in the cricket frog

We investigated how male cricket frogs Acris crepitans, alter their advertisement calls in response to broadcasts of synthetic calls that were either 'attractive' or 'aggressive'. The stimulus calls differed in temporal but not spectral characteristics. Male cricket frogs produced a more aggressive call when presented with the aggressive stimulus, indicating that they perceived the temporal differences between the two call categories. The direction and degree of temporal and spectral changes depended on the relative dominant frequency of the resident and opponent. If the resident's dominant frequency was initially higher than the stimulus frequency, the pattern of change in dominant frequency mirrored that seen for the temporal call characters. In contrast, if the resident's initial dominant frequency was below that of the stimulus, then the temporal and spectral changes were in opposite directions. Furthermore, stimulus order influenced whether males responded differently to playbacks of aggressive and attractive calls; males that received the aggressive call first produced more aggressive calls during the aggressive stimulus, while males that received the attractive call first produced similar calls in response to the two stimuli. This suggests that experience with different types of signals influences the subsequent calling behaviour of male cricket frogs. Copyright 1999 The Association for the Study of Animal Behaviour.     

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.     

Sound production in the whitemouth croaker and relationship between fish size and disturbance call characteristics

The whitemouth croaker produces two different sounds using extrinsic sonic muscles: (1) male advertisement calls during the spawning season and (2) disturbance calls, produced by both sexes. The advertisement call, related to courtship, was recorded in the field and from two marked spawning males of 28 and 30.5 cm L_T in the laboratory. It consists of individual pulses with average durations of 19.7 ms and 17.8 ms for the two males respectively, interpulse intervals of 496 ms and 718 ms, and dominant frequencies of 280 Hz and 316 Hz. Pulses are emitted in bouts of one to three min duration. Disturbance calls consist of a burst of pulses produced at short intervals, and the pulse duration, interpulse interval and dominant frequency of the pulses average 19.8 ms, 17.1 ms, and 363 Hz, respectively. Dominant frequency and interpulse interval decrease and pulse duration increases with fish size. Sound characteristics change markedly in young of the year individuals (lower than 25 cm L_T) after which they appear to stabilize. Higher dominant frequency in the advertisement than in the disturbance call and the relationship of dominant frequency to pulse duration suggest that dominant frequency is determined as a forced response to muscle contraction parameters rather than by the natural frequency of the swimbladder.     

Male green frogs lower the pitch of acoustic signals in defense of territories: a possible dishonest signal of size?

In responses to broadcasts of conspecific advertisement calls,male green frogs (Rana clamitans) lower the dominant frequencyof their calls. Because dominant frequency is negatively correlatedwith male body size in green frogs, frequency alteration providesa means of potentially exaggerating size during territorialcontests. In field playback experiments, we broadcast syntheticstimuli representing small, medium, or large intruders to territorialresidents. We tested the hypotheses that males use frequency alteration to provide honest signals of their size or theirsize-independent fighting ability, or to dishonestly signalsize. Dominant frequency did not better predict male size inresponse calls than in unsolicited calls. The magnitude offrequency alteration was not related to body size, general condition, or an indirect measure of fighting ability. Thus,males did not use frequency alteration to provide honest informationabout body size or size-independent fighting ability. However,males significantly increased their apparent size by producinglower frequency calls. Small males produced relatively lowerfrequency calls in response to the large-male stimulus (comparedto the small-male and medium-male stimuli), but large malesdid not. Further, the magnitudes of frequency alteration weresignificantly greater in responses to the large-male stimulus,primarily because small males responded with a greater decreasein frequency to the large-male stimulus than to the small-maleand medium-male stimuli. These results support several predictions of the dishonest signal hypothesis and suggest that dishonestymay be a conditional strategy used by small males.     

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

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

Evolutionary and neurobiological implications of selective phonotaxis in the spring peeper (Hyla crucifer)

Males of the spring peeper (Hyla crucifer) in central Missouri produce frequency-modulated, sinusoidal advertisement calls with a duration of 90–250 ms, and a mid-point frequency of 2800–3360 Hz. The frequency of the call is inversely correlated with body size. In playback experiments with synthetic stimuli, females did not prefer a frequency-modulated call to a call of constant frequency. Females preferred a call with a duration of 150 ms to sounds with durations of 40, 75 and 400 ms; a call of 300 ms was just as attractive as the 150-ms call. Females preferred a call of 2875 Hz to alternatives of 4000 Hz and 2600 Hz. The auditory system of H. crucifer is thus only roughly tuned to the temporal and spectral properties of the advertisement call. The female's specificity with respect to duration alone is adequate for species recognition, but intraspecific mate choice based on call frequency is extremely unlikely.     

Pacific treefrogs use temporal integration to differentiate advertisement from encounter calls

Male Pacific treefrogs, Hyla regilla, use advertisement and encounter calls in their reproductive communication. Encounter calls are produced when a male hears either advertisement or encounter calls at amplitudes that exceed his aggressive threshold for that call type, and they are believed to play a role in male spacing in a chorus. Aggressive thresholds of males for advertisement and encounter calls are plastic; males resume advertisement calling (i.e. accommodate) after repeated exposure to either advertisement or encounter calls that exceed their aggressive thresholds for that call type. Presentation of one call type does not elevate a male's aggressive threshold for the other call type, indicating that discrete neural ‘channels’ exist for processing advertisement versus encounter calls. Because these calls are spectrally highly similar, it is likely that they are differentiated in the central auditory system on the basis of their different temporal structure; pulses are repeated in advertisement and encounter calls at rates of approximately 100 pulses/s and 30 pulses/s, respectively. The present study tested whether consecutive advertisement-type interpulse intervals (IPIs) are required to activate the filter for advertisement calls. Males were accommodated to synthetic stimuli that differed in the proportion of advertisement versus encounter IPIs. Aggressive thresholds of males for the encounter call and advertisement call (monophasic type) were elevated 450% and 58%, respectively, following accommodation to a stimulus that had nine of each IPI type, arranged in an alternating sequence. Aggressive thresholds for the monophasic advertisement call were elevated 151%, however, when males were accommodated with stimuli that had two sets of eight to nine consecutive advertisement IPIs; consistent with previous work, aggressive thresholds for the encounter call were not significantly elevated. Aggressive thresholds for the advertisement call are typically elevated approximately four-fold following accommodation with that call type. These findings are consistent, therefore, with the hypothesis that several consecutive advertisement IPIs are required to activate the ‘advertisement channel’; if consecutive advertisement IPIs were not required, the stimulus that consisted of the same number of advertisement and encounter IPIs should have induced comparable elevations of the aggressive thresholds of males for advertisement and encounter calls. Selective filters for the advertisement call, therefore, appear to integrate information across many consecutive ‘correct’ IPIs. This integration process may contribute to a female's call preference.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Arginine Vasotocin Injection Increases Probability of Calling in Cricket Frogs, but Causes Call Changes Characteristic of Less Aggressive Males

Male cricket frogs,Acris crepitanscommunicate to males and females using advertisement calls, which are arranged into call groups. Calls at the middle and end, but not beginning of the call group, are modified in response to male–male aggressive interactions. We found in this field study of male cricket frogs in natural breeding choruses that the peptide hormone arginine vasotocin (AVT) not only increased the probability that males called after injections, but also caused modifications in middle and end calls to produce calls characteristic of less aggressive males. Moreover, AVT-injected males showed significantly greater increases in call dominant frequency than saline-injected males, again, a characteristic of less aggressive males. Cricket frog calls are used to both repel males and attract females, thus call changes may relate to male–male and/or male–female interactions. Saline-injected males also demonstrated significant changes in several call traits, including changes that occurred in the beginning and middle calls of the call groups, but not the end calls. AVT appeared to block some call changes produced through handling. These data suggest that AVT can influence acoustic communication in frogs in several ways, including effects on call characteristics and dominant frequency, as well as potentially blocking some handling effects.     

Male territoriality,fighting and assessment of fighting ability in the Australian frog Uperoleia rugosa

Males of Uperoleia rugosa form aggregations around the breeding pond. Within the aggregations, each calling male maintains an exclusive zone in which it does not tolerate another calling male. However, attacks on silent intruders, satellite males or females were never observed. Territorial males warned calling intruders by changing from advertisement calls to encounter calls (a display); if the intruder persisted in calling the resident would either attack or retreat and call elsewhere. Fights were usually won by the heavier males, and males of similar weight were more likely to fight, whereas males retreated from opponents that were substantially heavier. Calling was energetically costly and territorial males lost weight and thus their fighting abilities declined, whereas silent satellite males gained weight and were able to oust lighter territorial males. There was thus a dynamic interchange between territorial and satellite males depending on their relative weights. The playback of recorded advertisement calls showed that males can assess each others' fighting abilities on the basis of their vocalizations. Males retreated from a loudspeaker broadcasting the calls of a larger male but they attacked the loudspeaker if the calls of smaller or similar sized males were played (N=39, P<0·001, one-tailed binomial test). Dominant frequency was probably the call parameter used in assessment as it was correlated with the weight (r=−0·71, P<0·001, N=100) and hence the fighting ability of the caller, but this hypothesis needs further testing.     

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.     

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

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

Multimodal Signaling in Male and Female Foot-Flagging Frogs Staurois guttatus (Ranidae): An Alerting Function of Calling

In multimodal communication, individuals use several sensory modalities for information transfer. We report on novel observations of foot‐flagging in the Bornean ranid frog Staurois guttatus that is temporally linked to advertisement calling. In addition, we document the first case of foot‐flagging in a female anuran as well as additional visual displays in both males and females including arm‐waving, vocal‐sac pumping and open‐mouth display. In males, advertisement calls and foot‐flags were given throughout most of the day, suggesting that acoustic and visual signals form a multicomponent and multimodal display. We tested the efficacy‐based alerting signal hypothesis of multimodal communication using acoustic playback experiments with males. This hypothesis predicts that an initial signal draws the receiver's attention to the location of a subsequent more informative signal. Several lines of evidence supported the alerting hypothesis. First, the latency between foot‐flags and advertisement calls was significantly higher than that between advertisement calls and foot‐flags, suggesting a functional linkage with calls drawing attention to foot‐flags. Secondly, advertisement calling had a signaling function with males responding significantly more often with both calls and foot‐flags compared with pre‐ and post‐playback control periods. Finally, and most notably, all males tested turned towards the playback stimulus, suggesting that the advertisement call serves to focus their attention on subsequent signals. We discuss the potential of multimodal signaling for conspecific and heterospecific communication and the circumstances under which such a multimodal communication system could evolve.     

Advertisement Call Variation in the Golden Rocket Frog (Anomaloglossus beebei): Evidence for Individual Distinctiveness

Individual distinctiveness in acoustic signals can allow receivers to respond appropriately to different individuals. The aim of this research was to examine signal variation and to investigate the relative importance of different acoustic properties for coding individual distinctiveness in the advertisement calls of male golden rocket frogs (Anomaloglossus beebei). We examined patterns of within‐individual and among‐individual variability in 760 advertisement calls of 40 males as well as repeatability in 16 additional males that were recorded on two different occasions. We examined eight call properties, and all properties exhibited significant among‐individual variation and moderate to high repeatability across days. We employed discriminant function analysis to examine individual distinctiveness statistically. These analyses assigned 79% of calls to the correct individual. Fine temporal properties, including pulse duration, pulse rate, and pulse interval, as well as the spectral property of dominant frequency, contributed most toward discrimination among individuals. These results indicate that individual male golden rocket frogs can be distinguished statistically by their advertisement calls. We use these findings to evaluate results of previous playback studies in A. beebei and to generate testable predictions regarding the potential for specific call properties to function in mate selection and social recognition.     

Do Male White-Lipped Frogs Use Seismic Signals for Intraspecific Communication?

Modern frogs and toads possess a structurally unique saccule,endowing them with seismic sensitivity greater than that observedso far in any other group of terrestrial vertebrates. In synchronywith their advertisement calls, approximately half of the callingmales of one frog species, the Puerto-Rican white-lipped frog(Leptodactylus albilabris), produce impulsive seismic signals(thumps). The spectral distribution of power in these seismicsignals matches precisely the spectral sensitivity of the frog'ssaccule. The signals have sufficient amplitude to be sensedeasily by the frog's saccule up to several meters from the source—wellbeyond the typical spacing when these frogs are calling in agroup. This circumstantial evidence suggests that white-lippedfrogs may use the seismic channel in intraspecific communication,possibly as an alternative to the airborne channel, which oftenis cluttered with noise and interference. Using the frog's vocalizationsas our assay, we set out to test that proposition. In responseto playback calls, the male white-lipped frog adjusts severalof its own calling parameters. The most conspicuous of theseinvolves call timing—specifically the tendency for a gapin the distribution of call onsets, precisely timed with respectto the onsets of the playback calls. When the airborne componentis unavailable (e.g., masked by noise), approximately one infive animals produces the calling gap in response to the seismicsignals alone.