Effects of multiple-speaker playbacks on aggressive calling behavior in the treefrog <Emphasis Type="Italic">Dendropsophus ebraccatus</Emphasis>

In addition to producing signals, males of chorusing species also act as receivers by adjusting properties of their vocalizations in response to those of other nearby individuals. Although it is likely that males are responsive to more than one other individual, most playback studies investigating male response have involved dyads in which vocal responses are measured to stimuli presented from a single speaker. In this study, I explored changes in both the propensity to give aggressive calls and the temporal properties of those calls in response to the playback of multiple aggressive call stimuli in the treefrog Dendropsophus ebraccatus. I found that males were sensitive to both the number of simulated aggressive callers and their specific call characteristics. Males generally gave a highly aggressive response to the first stimulus presented, but their response to the modification of this stimulus by the addition or subtraction of a simulated competitor depended on the degree of aggressiveness of the stimuli. Males tended to decrease their aggressive responses when either a more aggressive call was silenced or a less aggressive call was added and to increase their aggressive responses in the opposite situation. Aggressive calling in this species is clearly affected by complex changes in the social environment and I suggest that future studies explore these issues in other species to improve the understanding of communication interactions.     

Size assessment in simulated territorial encounters between male green frogs (Rana clamitans)

We examined the ability of male green frogs to assess the size of an opponent based on the dominant frequency of their advertisement call, which is negatively correlated with size, using synthetic stimuli to simulate intruders of different sizes. In one field playback experiment, we broadcast a pair of stimuli representing a small and a large male; in a second experiment, we broadcast calls of a medium and a large male. In both experiments, males produced calls with significantly lower dominant frequencies in response to each stimulus. Contrast analyses revealed that males lowered the dominant frequency of their calls more in response to the large-male stimulus than in responses to the small- and medium-male stimuli. In the second experiment, males also responded to the large-male stimulus by calling at higher rates. There were no differences in mean note duration or the number of moves made toward or around the playback speaker in response to any stimulus. Thus, the frequency of an opponent's calls elicits a differential modification of calling behavior, primarily in the form of differential dominant frequency alteration, suggesting that males use dominant frequency to assess the size of opponents during aggressive encounters. Received: 17 April 1998 / Accepted after revision: 7 October 1998     

Aggressive thresholds in Dendropsophus ebraccatus: habituation and sensitization to different call types

Males in many chorusing anuran species use aggressive calls during defense of calling spaces from other males. The minimal intensity of another male’s vocalizations that elicits an aggressive call response has been termed the aggressive threshold. Previous studies of aggressive thresholds have shown that they are plastic: males habituated (increased their aggressive thresholds) in response to repeated presentation of stimuli above initial threshold levels. Habituation likely contributes to the stable chorus structure of these species, in which aggressive calling is rare compared to advertisement calls. I have observed high levels of aggressive calling in the treefrog Dendropsophus ebraccatus, suggesting that males of this species do not habituate. In this study, I investigated the plasticity of aggressive thresholds in D. ebraccatus. I measured the aggressive thresholds of males before and after suprathreshold stimulation by both advertisement and aggressive calls. I found that the different call types had different effects: males habituated to advertisement calls but lowered their aggressive thresholds in response to aggressive calls. I consider the latter response to be an example of sensitization, a behavior that has been documented infrequently in vocalizing anurans. Sensitization is a plausible mechanism responsible for the high levels of aggressive calling observed in this species. Given the high costs of aggressive calling, however, it is unclear why a mechanism that increases aggressive call output would be maintained.     

Diphasy in the advertisement calls of Geocrinia laevis (Anura: Leptodactylidae): vocal responses of males during field playback experiments

Summary The advertisement calls of Geocrinia laevis are diphasic, and consist of a single introductory note, followed by a series of repeated notes; diphasy is based on differentiation in the temporal structure of the two phases. In a series of field playback tests used to investigate the functional significance of diphasy, calling males were presented with one of three types of acoustic stimulus: complete calls (introductory and repeated notes), introductory notes only, or repeated notes only. There were no marked differences in the effects of these stimuli, and all resulted in a reduction in the rates of production of introductory notes and of repeated notes. Playback at levels of 110 dB PSPL and above caused virtual cessation of calling. The results suggest that one of the functions of the advertisement call, the communication of territoriality between males, does not reside in either the introductory note or the repeated notes, so that diphasy in the advertisement call has no apparent functional basis in this context. Whether the attraction of reproductively ripe females is a particular function of either phase remains untested. The advertisement calls of G. laevis are much less diphasic than those of a closely related species, G. victoriana, for which there is a clear partitioning of the functions of communication of territoriality and the attraction of conspecific mates between the introductory notes and the repeated notes, respectively. Furthermore, the acoustic repertoire of G. laevis is less complex than that of G. victoriana in lacking a vocalisation which functions specifically in close-range aggressive encounters between males.     

The functional significance of the diphasic advertisement call of Geocrinia victoriana (Anura: Leptodactylidae)

Summary The advertisement call of Geocrinia victoriana is markedly diphasic and consists of one introcuctory note (rarely two or three), followed by a series of repeated notes of similar carrier frequency, but much shorter duration and higher pulse rate. In two-choice field discrimination experiments with the two phases of the advertisement call presented as alternative stimuli, reproductively ripe females were attracted only to the repeated notes and contacted the source of these signals (a loudspeaker). Field playback experiments were carried out on residential males, with a complete call, an introductory note, or a series of repeated notes being used as the stimulus, and intensity varied upwards in 10 dB increments. With the complete call or the introductory note as the stimulus, calling behaviour of subjects altered significantly from the pre-stimulatory pattern, with a great reduction in output of repeated notes, and an increase in duration and decrease in pulse rate of introductory notes. The threshold intensity (lowest level for a significant change in calling behaviour) with the introductory note as the stimulus, 90–99.9 dB peak sound pressure level (PSPL), was at least 10 dB lower than that with the complete call as the stimulus (110–125 dB, PSPL). Playback of repeated notes had little effect on calling behaviour until the intensity exceeded 120 dB, PSPL. Playback of all three classes of stimulus at constant intensities below and above the threshold for the complete call (i.e. 99–99.9, and 110–125 dB, PSPL) indicated that changes in calling behaviour were due to the intensity of stimulus rather than to a cumulative effect based on the number of stimuli. Measurements of intensities of advertisement calls of nearest neighbours in natural breeding assemblages (modal class=104–106 dB, PSPL) were consistent with the experimental results. It was thus demonstrated that in G. victoriana the two functions of the advertisement call are partitioned: the repeated notes attract breeding females (hence are equivalent to the mating call), and the introductory notes are directed at other males in a territorial context. The increase in duration of the introductory note is graded rather than discrete, so that this phase of the advertisement call changes progressively from the equivalent of a long-range (first-order) to a short-range (second-order) territorial call, then to an encounter call.     

Territorial vocal behavior in hybrid smooth froglets, Geocrinia laevis complex (Anura: Myobatrachidae)

Males of the parapatically distributed myobatrachid frogs Geocrinia laevis and G. victoriana have highly divergent advertisement calls. Furthermore, the two species differ strongly in the complexity of their vocal repertoires, with males of G. victoriana possessing, and those of G. laevis lacking, a distinct territorial vocalization (encounter call). We investigated the territorial vocal behavior of males in a persistent natural hybrid population. Most hybrid males possessed encounter calls functionally equivalent to those of G. victoriana, that were produced following exposure to playback of recorded advertisement calls presented at >110 dB peak sound pressure level. The territorial acoustic responses were not associated with an index of hybridity derived from the structure of the advertisement call, suggesting genetic and functional decoupling of the two components of the vocal repertoire; i.e., advertisement calls and encounter calls. This decoupling may be the result of sexual selection favouring those hybrids with pronounced territorial behavior and the associated vocalization, regardless of the structure of their advertisement calls.Communicated by A. Mathis     

The function of call alternation in the African reed frog (Hyperolius marmoratus): precise call timing prevents auditory masking

The function of call alternation in the African painted reed frog, Hyperolius marmoratus was studied. Males actively avoided call overlap with neighboring males in their natural habitat. Advertisement calls produced by groups of two or three males showed less overlap than expected if they called at random. In addition, isolated males significantly reduced their calling rate to match the periodicity of a playback of periodic tone pulses, with vocalizations given during the silent intervals between tones. Gap-detection experiments showed that males suppressed vocalizations during playbacks of constant-frequency tone bursts and gave more calls during silent periods than expected by chance. Females discriminated against conspecific advertisement calls with many pulses in favor of calls with few or no pulses. This suggests that there would be little selection pressure on males to alternate calls so that pulses are not obscured. To test if call alternation functions to make calls more conspicuous, females were presented with identical conspecific advertisement calls from two speakers in four different temporal patterns: simultaneous, overlapping, abutting, and alternating. Females did not discriminate when calls were presented in a simultaneous, abutting, or alternating time pattern. When calls overlapped with the trailing call delayed from 0.5 to 70 ms females discriminated in favor of the leading call. When the intensity of the leading call was reduced by 6 dB the preference for the leading call was maintained when calls overlapped by 2 and 40 ms but was abolished or reversed at 0.5 and 70 ms, respectively. These results support the notion that the second of two partially overlapping calls was acoustically masked, rendering female painted reed frogs unwilling to approach or unable to locate such calls. It is suggested that either simultaneous masking or the precedence effect are responsible for the observed behavior. Acoustic masking of the second call by the first when calls overlapped was maintained even when the frequency of the first call was altered by 150 Hz above and below the more preferred frequency of the second call. Received: 13 April 1995/Accepted after revision: 5 November 1995     

Species-specificity and individual variation in the song of male Nathusius’ pipistrelles (<Emphasis Type="Italic">Pipistrellus nathusii</Emphasis>)

Sequences of the advertisement calls produced by male Nathusius’ pipistrelles (Pipistrellus nathusii) during the autumn mating period were recorded from individuals at two separate sites in Antrim, Northern Ireland, in August 2004. Several male roosts were found at these sites in close proximity to a single maternity roost, each containing approximately 200 adult females and their young. Analysis of measured parameters of four identified call types revealed that there were significant differences in call structure between sites and between individuals. Playback experiments, performed outside the adult female and juvenile roost sites, comprised of experimental advertisement call sequences of P. nathusii, Pipistrellus pygmaeus and Pipistrellus pipistrellus and control sound recorded without bats present (silence). Response was measured by simultaneously recording ultrasound during playbacks and counting the number of echolocation pulses identified as those of P. nathusii above a predetermined amplitude threshold. Significantly greater numbers of P. nathusii echolocation pulses were recorded during playback of male P. nathusii advertisement calls than during playback of congeners’ advertisement calls and control sound. The number of echolocation pulses recorded was similar during playback of P. pipistrellus and P. pygmaeus advertisement calls and silence. We suggest that, due to call complexity, male P. nathusii advertisement calls should be classified as ‘song’. Species-specificity and individual variation suggests that the songs of male P. nathusii have the potential to play a role in mate attraction and mate assessment.     

Interspecific acoustic interactions of the neotropical treefrog Hyla ebraccata

Summary Hyla ebraccata, H. microcephala and H. phlebodes commonly occur together in Panama. The three species have calls exhibiting broad frequency overlap and call during the same season and time of day from similar microhabitats, frequently in close proximity. The vocal repertoires of the three species are structurally and functionally similar. All employ multi-part advertisement and aggressive calls which consist of a primary note followed by a variable number of clicks. H. ebraccata males often responded to heterospecific calls with multi-note synchronized responses, and calls with primary notes greater than 150–200 ms were most effective in eliciting synchrony. Playback experiments with synthetic 1-note advertisement calls of different durations and both synthetic 1-note advertisement calls and 200 ms tones of different frequencies demonstrated that H. ebraccata males will synchronize with stimuli which are similar in frequency and duration to conspecific calls. Data from a two-choice experiment with female H. ebraccata demonstrate that calls of individual H. microcephala can reduce the attractiveness of a H. ebraccata male's calls if primary notes overlap. By synchronizing response calls to those of H. microcephala, a H. ebraccata may reduce the chances that his calls are rendered less attractive to potential mates.Aggressive calls of these species are graded and are characterized by higher pulse repetition rates and often longer durations than advertisement calls. H. ebraccata males respond to aggressive calls of H. microcephala and H. phlebodes as they do to their own calls. Heterospecific aggressive interactions probably occur because the species interfere acoustically. Our results demonstrate that H. ebraccata males behave in ways which enhance their ability to communicate in a noisy assemblage of conspecific and heterospecific males.     

Synchronous waving in an ocypodid crab, Ilyoplax pusilla : analyses of response patterns to video and real crabs

In several species of ocypodid crabs, including Ilyoplax pusilla (De Haan), waving displays tend to be synchronized among neighboring individuals. However, no quantitative study has yet been conducted. In this study, responses of I. pusilla males to waving of video and real crabs were examined. Spontaneous, periodic waving was observed even when no stimulus waves were present. When single isolated waves of the video crab were randomly delivered (single pulse perturbation), crabs waved either immediately after the stimuli or after some time had elapsed. Assuming that waves immediately after the stimuli were triggered prior to those stimuli, crabs typically responded after a roughly constant delay from the onset of each stimulus. The delay (response time) was close to, but slightly shorter than, the spontaneous wave period. Dependency of response time upon timing of the stimulus (stimulus time) was at most weak. Responses of crabs to periodic waves of the video crabs and waves of␣the real crabs were basically consistent with the results of single pulse perturbation, although there was a consistent tendency for stimulus time and response time to be negatively correlated near stimulus time = 0 s. Overall, these results demonstrate that males of I. pusilla adjust signal timing relative to a neighbor in a manner of phase delay synchrony. The similar response pattern has also been noted in synchronous flashing of fireflies and the calling of katydids. Moreover, these experiments with I.␣pusilla demonstrate the value of video playback as a research tool in this species. Received: 4 April 1997 / Accepted: 17 December 1997     

The effect of social interactions on calling energetics in the gray treefrog (Hyla versicolor)

Summary The vocal behavior of Hyla versicolor was studied in the field by means of behavioral observations and playback experiments, and these data were coupled with measurements of oxygen consumption in calling frogs to estimate the effect of social interactions on calling energetics. Male gray treefrogs have intense calls (median peak SPL=109 dB, fast RMS SPL=100 dB at 50 cm). At an air temperature of 23° C, males produced an average of 1,200–1,300 calls/h for 2–4 h per night. Calling rates and call durations differed among individuals, but were relatively constant for each male during periods of sustained calling. Males in dense choruses gave calls about twice as long as isolated males, but produced calls at about half the rate. Consequently, total calling effort and estimated aerobic costs were largely independent of chorus density. Playbacks of recorded calls to males in the field elicited increases in call duration and decreases in calling rate, regardless of the rate or duration of the stimulus. Males gave longer calls in response to long calls or to stimuli presented at high rates, but they did not precisely match either stimulus rate or duration. Calling effort and estimated oxygen consumption changed only slightly during stimulus playbacks. These results indicate that male-male competition elicits pro-found changes in the vocal behavior of calling males, but these changes have little effect on energy expenditure. We estimated that most calling males had metabolic rates of about 1.7–1.8 ml O₂/(g\h), or about 280 J/h for an average size (8.6 g) male at 20° C. Although changes in call duration and calling rate did not affect aerobic costs of calling, males producing long calls at slow rates called for fewer hours per night than males producing shorter calls at higher rates. This suggests that calling time may be limited by the rate at which muscle glycogen reserves are depleted.     

Female mate selection in the natterjack toad: active choice or passive atraction?

Summary Variation in seasonal mating success among male natterjack toads (Bufo calamita) was influenced by the number of nights that males spent at the pond and by male body size. Large males produced louder and lower frequency calls than small males, and maintained larger acoustic territories. After arriving at the pond, one half of all observed females mated with the first male they encountered. The remainder visited several males before initiating amplexus, but no criteria could be identified that females might have used when deciding whether to accept or reject a male. Movements between several males seem to be best explained by low female responsiveness to male advertisement calls on cold nights which were nonoptimal for oviposition. Females attempted to reject non-calling males both before and after amplexus, but this may be a mechanism to avoid mismating with males of the common toad (Bufo bufo), an explosive breeder that utilised the same pond. In two-choice playback experiments using synthetic advertisement calls, females showed no preference for calls based on their frequency. Females preferred calls of intermediate pulse repetition rate equivalent to those produced by a male at the same body temperature. Pulse rate is thus potentially subject to stabilishing selection and may be an important character for species-recognition. Females preferred fast rather than slow call rates, but only when the alternative rates were extreme. They also preferred calls which they perceived at the highest sound pressure level, but did not discriminate between absolute sound pressure levels of alternative stimuli at different distances. Since females that delay mating and oviposition may suffer predation, it is suggested that female preference for loud, rapidly repeated calls may be adaptive in the sense of minimizing the costs of locating conspecific males, rather than maximizing the probability of obtaining a high quality mate. Competition between males to maintain large acoustic territories and produce calls that can be easily detected by females would seem to be a sufficient mechanism to explain the evolution of the striking calls produced by male natterjacks.     

Predator-specific alarm calls in Campbell's monkeys, Cercopithecus campbelli

One of the most prominent behavioural features of many forest primates are the loud calls given by the adult males. Early observational studies repeatedly postulated that these calls function in intragroup spacing or intergroup avoidance. More recent field experiments with Diana monkeys (Cercopithecus diana) of Taï Forest, Ivory Coast, have clearly shown that loud male calls function as predator alarm calls because calls reliably (1) label different predator classes and (2) convey semantic information about the predator type present. Here, I test the alarm call hypothesis another primate, the Campbell's monkey (C. campbelli). Like Diana monkeys, male Campbell's monkeys produce conspicuous loud calls to crowned hawk eagles (Stephanoaetus coronatus) and leopards (Panthera pardus), two of their main predators. Playback experiments showed that monkeys responded to the predator category represented by the different playback stimuli, regardless of whether they consisted of (1) vocalisations of the actual predators (crowned hawk eagle shrieks or leopard growls), (2) alarm calls to crowned hawk eagles or leopards given by other male Campbell's monkeys or (3) alarm calls to crowned hawk eagles or leopards given by sympatric male Diana monkeys. These experiments provide further evidence that non-human primates have evolved the cognitive capacity to produce and respond to referential labels for external events.     

Communication by changing signals: call switching in red-winged blackbirds

Summary Territorial male red-winged blackbirds use many different alert calls which overlap broadly in the context of their use and which are often given in continuous, repetitive fashion. Males give one call type many times before switching to another type, and call during all daily activities. In this field study we demonstrate that males tend to give the same call types as their neighbors and change types to match the changes of their neighbors. Individual males change call types in response to the appearance of a mounted hawk and also match call type changes broadcast from a loudspeaker. The various call types are not associated with particular behavioral contexts. We suggest that red-winged blackbirds operate a general acoustic alert system by calling repetitiously and changing call types when detecting environmental changes such as appearances of predators. Evidence is presented that communication is acheived primarily during switching among different call signals, and not by specific calls that refer to particular stimuli or states of alertness.     

The effect of perceived intruder proximity and resident body size on the aggressive responses of male green frogs, Rana clamitans (Anura: Ranidae)

We examined variation in aggressive responses within and among individuals in the green frog, Rana clamitans. We tested the hypothesis that resident males respond in a graded fashion to changes in perceived intruder proximity (stimulus intensity). We also investigated how response level varied with responder body size. We found that green frogs differentially alter aggressive responses (increasing movement and calling rates while decreasing the duration and dominant frequency of their calls) with an increase in stimulus intensity. Body size did not appear to be a significant influence on most response levels. We suggest that aggressive responses are graded, and advertisement and aggressive calls represent opposite ends of a continuum. Green frogs do not exhibit clearly defined thresholds for responses like those reported for some other frogs. By using graded responses rather than discrete all-or-nothing responses, males may reduce the potential costs of aggressive encounters.     

Acoustically mediated neighbor recognition in the North American bullfrog,Rana catesbeiana

Summary Playback experiments were conducted to determine whether territorial male bullfrogs (Rana catesbeiana) were capable of discriminating between advertisement calls of neighbors and strangers. Territorial males adjacent to subject males were removed and replaced with a speaker which broadcast calls of a stranger or the removed male. Ten of 11 males responded more strongly to calls of strangers than to calls of neighbors. When advertisement calls of neighbors were broadcast to subject males from positions within the removed neighbor's territory and from positions opposite the subject male, all 9 males tested responded more strongly to calls of a neighbor broadcast from a new position than to playbacks of the same call broadcast within the removed neighbor's territory. Because male bullfrogs recognize familiar calls and associate these calls with a particular direction or location, these data provide the first evidence for acoustically mediated neighbor recognition in frogs.     

Advertisement-call modification,male competition,and female preference in the bird-voiced treefrog <Emphasis Type="Italic">Hyla avivoca</Emphasis>

Senders and receivers influence dynamic characteristics of the signals used for mate attraction over different time scales. On a moment-to-moment basis, interactions among senders competing for a mate influence dynamic characteristics, whereas the preferences of receivers of the opposite gender exert an influence over evolutionary time. We observed and recorded the calling patterns of the bird-voiced treefrog Hyla avivoca to assess how the dynamic characters of calls vary during interactions among groups of males in a chorus. This question was also addressed using playback experiments with males. Playback experiments with females showed how changes in dynamic call properties are likely to affect male mating success. Frogs calling in pairs, groups, or in response to playbacks produced longer calls than did isolated males. During call overlap, males often increased the duration of the silent interval (gaps) between the pulses of their calls so that the pulses of the calls of two neighbors interdigitated. This change resulted in increased variability of pulse rate, a traditionally static acoustic property; however, males also produced high proportions of non-overlapped calls in which variability in pulse rate was low and had species-typical values. Females preferred long calls to short- and average-duration calls, and non-overlapped calls to overlapped calls. Given a choice between pairs of overlapped calls, females preferred pairs in which the proportion of overlap was low and pairs in which the pulses of such calls interdigitated completely. The observed patterns of vocal competition thus reflect the preferences of conspecific females, which have influenced the evolution of the calling behavior of H. avivoca. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Territorial defense in the bannertail kangaroo rat (Dipodomys spectabilis): footdrumming and visual threats

Summary Footdrumming and visual displays associated with territorial defense in the bannertail kangaroo rat (Dipodomys spectabilis) were studied in the field in Arizona. We predicted that foot-drumming functioned like bird song in territorial advertisement. Rats were tested in their territories for their responses to the presentation of four different stimuli: a playback of airborne footdrumming patterns (audio); a stuffed and mounted kangaroo rat in an aggressive posture (visual); a combination of both (audio-visual); and a playback of cricket stridulation as a control. Kangaroo rats investigated all experimental stimuli equally by becoming alert, approaching, and retreating (Fig. 2). Rats, therefore, seemed to assess the amount of threat and footdrummed at higher rates and sooner in response to the more threatening visual stumulus of a stuffed rat than to the apparently less threatening ones of a airborne footdrumming playback or cricket stridulation (Fig. 1). We conclude that rats footdrum at long ranges to repel a potential intruder and also at close ranges when territories are threatened by a persistent intruder. Chase and attack, as in territorial birds, are the final lines of defense.     

A test of the "dear enemy effect" in the strawberry dart-poison frog (<Emphasis Type="Italic">Dendrobates pumilio</Emphasis>)

Some territorial animals exhibit a form of social recognition, commonly termed the "dear enemy effect", in which territory residents display lower levels of aggression toward familiar neighbors compared to unfamiliar individuals who are non-territorial "floaters". Despite the widespread occurrence of territorial social systems and use of acoustic signals for communication in anuran amphibians, only two previous studies have demonstrated vocally mediated dear enemy behavior in a territorial frog. In this study, I conducted neighbor-stranger discrimination playback experiments in a third species of territorial frog, the strawberry dart-poison frog, Dendrobates pumilio (Anura, Dendrobatidae). In the first experiment (n=24), I broadcast the calls of a subject's nearest neighbor and the calls of an unfamiliar individual from the approximate midpoint between the subject's and the neighbor's territories. Although males responded to the stimuli, they did not exhibit differential responses to the calls of neighbors and strangers. In a second experiment (n=22), I broadcast the calls of a neighbor and a stranger to subjects through a speaker located in the approximate center of the neighbor's territory. Males also responded to the playback, although less intensely than in the first experiment, but no discrimination between the calls of neighbors and strangers was found. Thus, territorial males of the strawberry dart-poison frog appear not to discriminate behaviorally between the advertisement calls of neighbors and strangers. Several proximate and ultimate-level hypotheses for this lack of vocally mediated neighbor-stranger discrimination are discussed.Communicated by T. Czeschlik     

Site specificity in advertisement calls and responses to playbacks of local and foreign call variants in satin bowerbirds, <Emphasis Type="Italic">Ptilonorhynchus violaceus</Emphasis>

Avian vocalisations often show patterns of geographic variation. Previous work on the satin bowerbird has shown that although spatial variation in this species’ advertisement calls is strongly associated with habitat structure, some variation is apparent within habitat types. Seventeen populations located throughout the species’ distribution were used to examine whether spatial call variation could be influenced by other processes such as random drift or the presence of fine-scale vocal traditions; if this were the case, differing call variants would be expected at geographically discrete sampling sites both within and among habitat types. There were population-specific call variants at each of the sites sampled, with different variants apparent even within habitat types. At most sites, individuals gave only a single variant of advertisement call, and the call variant at one site, sampled after a 5-year interval, appears to have been relatively stable. Playback experiments were conducted at three populations to examine whether local call variants invoked a greater response than several non-local variants differing in their degree of similarity to the local variant. Birds responded strongly to local call variants but not to either of two foreign variants, one of which was similar to their local variant and one of which was very different. A pattern of geographic variation across populations, the fact that local and non-local variants evoke different responses and circumstantial evidence indicating that individuals can learn new calls all suggest that factors affecting song learning and the ability of males to establish and defend a bower site may have contributed to the establishment of geographically variable vocal cultures in this species.