Collared Dove Responses to Playback: Slaves to the Rhythm

Temporal features are the most important parameters for differentiating among the perch coo vocalizations of doves in the genus Streptopelia . The role of temporal features in inducing territorial responses was studied for one of these species, the collared dove ( S. decaocto ), using playback experiments. The playback stimuli consisted of manipulated coos, which were tested against a preplayback period and against unmanipulated control coos. In a first set of three manipulations, the temporal structure was altered by removing an element from the species-typical three-element coos. A coo lacking the third element, which is a naturally occurring variant, was as equally effective as the control in eliciting a territorial response. The same applied to a coo lacking the second and longest element. In contrast, a coo lacking the first element did not elicit a significant increase in territorial activities compared with the preplayback period, and there was a significant reduction in response compared with the control coo. A second set of two manipulations concerned changes in the species-typical rhythm. The two variants both led to a significant reduction in response compared with the normal rhythm, although they were still effective in eliciting an increase in activities compared with the preplayback period. The results indicate that the rhythm of territorial cooing plays an important role in communicating to conspecifics. As different sympatrically living dove species differ in particular in temporal features, rhythm may be an important cue for species recognition and may contribute to reproductive isolation.     

Reaction to conspecific degraded song by the wren Troglodytes troglodytes: Territorial response and choice of song post

We investigated the response of the wren Troglodytes troglodytes to playback of a territorial song degraded by long-range propagation. It appears that the wren is sensitive to this degradation since the territorial reaction is less intense with the degraded song than with the undegraded one. However, the degraded song is still considered by the receiver as a specific territorial aggressive signal. This differential response suggests that the male wren can use the degradation characteristics of the signal to adapt its territorial reaction. Indeed, in response to this stimulus, the receiver wren chooses a higher song post. By so doing, the bird improves both the propagation distance of the emitted song and the receiver's ability to hear the opponent's song. This behavioural change may correspond to a communication strategy, counteracting the environmental constraints on sound propagation. Therefore, in response to sound degradation during long-range propagation, birds may have developed behavioural adaptations complementary to the various adaptations concerning song structure and coding-decoding processes.     

Effects of personality on territory defence in communication networks: a playback experiment with radio-tagged great tits

Individuals often differ consistently in behaviour across time and contexts, and such consistent behavioural differences are commonly described as personality. Personality can play a central role in social behaviour both in dyadic interactions and in social networks. We investigated whether explorative behaviour, as proxy of personality of territorial male great tits (Parus major), predicts their own and their neighbours'' territorial responses towards simulated intruders. Several weeks prior to playback, subjects were taken from the wild to test their exploratory behaviour in a standard context in the laboratory. Exploratory behaviour provides a proxy of personality along a slow–fast explorer continuum. Upon release, males were radio-tracked and subsequently exposed to interactive playback simulating a more or a less aggressive territorial intruder (by either overlapping or alternating broadcast songs with the subjects'' songs). At the same time, we radio-tracked a neighbour of the playback subject. Male vocal responses during playback and spatial movements after playback varied according to male explorative behaviour and playback treatment. Males with lower exploration scores approached the loudspeaker less, and sang more songs, shorter songs and songs with slower element rates than did males with higher exploration scores. Moreover, neighbour responses were related to the explorative behaviour of the subject receiving the playback but not to their own explorative behaviour. Our overall findings reveal for the first time how personality traits affect resource defence within a communication network providing new insights on the cause of variation in resource defence behaviour.     

Sensitivity of California Thrashers (Toxostoma redivivum) to song syntax

Many birds have songs with variable structure and sequences. In earlier studies, we characterized some features from the song structures of California Thrashers (Toxostoma redivivum). In the Thrashers, there were a large number of phrase types, dozens to hundreds and the songs that were sequences of these many phrases were not random, but show some structure referred to as syntax. For example, a syntactic rule might be that phrase type A can be followed by phrase type B, but not type C. We, along with others, have assumed that syntax is an important feature of songs. This paper describes an experimental attempt to determine that syntax is important to California Thrashers by recording the reaction of territorial thrashers to playbacks of other thrasher songs, some of which obeyed the syntax rules we had discovered while others violated those rules. We also obtained video recordings of their behaviour near the playback speakers. We observed differences in the reactions to the birds that heard these two types of playbacks. Resident males reacted to either playback type, but more strongly when the original order was preserved. We observed difference in their behavioural response to correct or altered syntax. This indicates that the syntax of their songs is perceived in territorial defence by the birds.     

Responses of golden-backed uakaris, <Emphasis Type="Italic">Cacajao melanocephalus</Emphasis>, to call playback: implications for surveys in the flooded Igapó forest

We investigated the potential use of call playback for surveying the elusive golden-backed uakari (Cacajao melanocephalus), a diurnal social primate that typically inhabits the relatively inaccessible black-water swamp forests of the upper Amazon (Igapó forest) during the wet season. Furthermore, our objective was to understand better the functions of vocalisations of these rare and threatened monkeys. We focussed on the “tchó” call, a loud vocalisation that varies in temporal and frequency characteristics according to behavioural context and individual signaller (Bezerra et al. in Int J Primatol, 2010b). We investigated the vocal and behavioural responses of golden-backed uakaris from three Igapó areas to three different playback stimuli: own group calls (OGC); neighbouring group calls (NGC); and control (background noise). The call stimuli were used to simulate the unexpected presence of monkeys from the monkeys’ own and neighbouring Igapó areas. Playback of OGC and NGC increased calling and altered behaviour in golden-backed uakaris, whereas no noticeable response occurred to the control stimuli. Furthermore, there was no evidence of habituation to the call stimuli and the natural behavioural pattern of the groups was not obviously affected. Hence, call playback seems to be a valuable tool for locating these elusive monkeys in river surveys. Most of the agonistic-related behaviour observed after NGC playback was also observed after OGC playback. As golden-backed uakaris are not territorial, the presence of strange conspecifics to the area may not necessarily represent a more threatening situation. Our methods and results may also be useful for surveys of other primate species living in the Igapó forest and in other habitat types.     

Fighting talk: complex song elicits more aggressive responses in a vocally complex songbird

Song complexity in many songbirds is a trait subject to sexual selection. It is often associated with male territorial defence. Empirical studies testing differential male responses to rival song in vocally complex songbirds have, however, been scarce. We conducted playback experiments of the endemic New Zealand Tui Prosthemadera novaeseelandiae to test the aggressive response of territorial male Tui to rival songs with differing complexity levels. Overall, complex songs evoked significantly stronger responses from territorial males than did simple songs. Following playback of complex songs, focal males approached the playback more closely and rapidly, and responded with songs of higher complexity than they did to playback of simple songs. This suggests males could both distinguish between different levels of complexity within the Tui repertoire, and perceive a more complex song as a greater territorial threat. Our study is one of the first to demonstrate strong aggressive responses to increased levels of song complexity in a songbird species with highly complex vocalizations.     

Sex-specific territorial responses in Tawny Owls Strix aluco

A territorial intruder often poses more of a threat to the territory holder of the same sex as itself. As territorial aggression is associated with costs, aggression shown by birds towards territorial intruders of the opposite sex deserves investigation. This behaviour could arise due to the reproductive value of a mate or through mutualism between members of a pair. We investigated these hypotheses by presenting mated pairs of Tawny Owls Strix aluco with playback of male calls, female calls and a male and female duetting, and recording the number and intensity of responses by the male and female territory holders. Females responded significantly more often to female than to male calls. Males responded equally often to male and female playback. Males which had previously bred successfully with their mate were significantly more likely to respond to female playback in the spring, which suggested males were responding to female playback due to the reproductive value of their mate. There was no evidence of mutualism between members of a pair.     

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.     

Mutual Wattle Ornaments in the South Island Saddleback (Philesturnus carunculatus) Function as Armaments

Many birds have ornamental traits that are expressed in both sexes. Wattles—colorful fleshy structures that hang from the lower bill—are frequent among birds, but remain poorly understood and are generally presumed to be under sexual selection. The South Island saddleback (Philesturnus carunculatus) is an endangered bird endemic to New Zealand in which both males and females possess wattles. We used behavioral observations, morphological measures, and a playback experiment to investigate the role of wattles in saddlebacks during territorial encounters. Wattles were monomorphic when controlled for body mass, and became engorged with blood in both sexes during visual displays. In a playback experiment using male song, wattle engorgement was significantly associated with territorial intrusions in males but not in females. However, female wattle engorgement was significantly more likely during playback experiments in the absence of their mate, suggesting wattles are used in territorial defense by both sexes. The markedly similar use of wattles by both males and females in visual displays as a response to territorial intrusions supports the hypothesis that these elaborate structures function in part as armaments.     

The function of duetting in magpie-larks: conflict, cooperation, or commitment?

Avian duetting is a poorly understood phenomenon despite many hypotheses as to its function. Contrary to the recent view that duetting functions for mate guarding and is a result of conflict between the sexes, Australian magpie-larks, Grallina cyanoleuca, do not use duetting as a paternity guard. I used a playback experiment to investigate the role of antiphonal duetting in territorial defence and pair bond maintenance, two traditional hypotheses about the function of duetting. The experiment showed that, like many nonduetting species, magpie-larks recognize neighbours on the basis of song. It also provided evidence of functional differences between duetting and solo singing which indicate that temporal coordination of song between partners is used to maintain the territory and pair bond. Duets were more threatening territorial signals than solo songs: males initiated more vocalizations in response to playback of duets than playback of solos. Simulated intrusion also caused males and females to approach the speaker together and coordinate more of their vocalizations to form duets. Females did not engage in sex-specific territorial defence, responding equally strongly to playback of male and female song, and maintaining both territory and pair bond by attempting to exclude intruders of either sex. Males initiated more vocalizations in response to playback of male than female song, and their likelihood of duetting appeared to be related more to threats to the pair bond, in particular desertion by their partner. Copyright 2000 The Association for the Study of Animal Behaviour.     

Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales

Animal communication often occurs in communication networksin which multiple signalers and receivers are within signalingrange of each other. In such networks, individuals can obtaininformation on the quality and motivation of territorial neighborsby eavesdropping on their signaling interactions. In songbirds,extracting information from interactions involving neighborsis thought to be an important factor in the evolution of strategiesof territory defense. In a playback experiment with radio-taggednightingales Luscinia megarhynchos we here demonstrate thatterritorial males use their familiar neighbors' performancein a vocal interaction with an unfamiliar intruder as a standardfor their own response. Males were attracted by a vocal interactionbetween their neighbor and a simulated stranger and intrudedinto the neighbor's territory. The more intensely the neighborhad interacted with playback, the earlier the intrusions weremade, indicating that males eavesdropped on the vocal contestinvolving a neighbor. However, males never intruded when wehad simulated by a second playback that the intruder had retreatedand sang outside the neighbor's territory. These results suggestthat territorial males use their neighbors' singing behavioras an early warning system when territorial integrity is threatened.Simultaneous responses by neighboring males towards unfamiliarrivals are likely to be beneficial to the individuals in maintainingterritorial integrity.     

Alternative antipredator responses of two coexisting Daphnia species to negative size selection by YOY perch

Our study showed for the first time in nature that two coexistingDaphnia adopted alternative life history and behavioural strategiesto cope with negative size-selection predation by gape-limitedyoung-of-the-year (YOY) perch. We evaluated the phenotypic plasticityin life history and behavioural traits of two coexisting Daphniaspecies, D. pulicaria (2 mm) and D. galeata mendotae (1.4 mm),in response to seasonal changes in predation by YOY yellow perch(Perca flavescens) in a mesotrophic lake. We expected that thelarge-sized D. pulicaria, the most likely subjected to size-selectivepredation by YOY perch, will show stronger antipredator responsesthan the small-sized D. galeata mendotae. To test this hypothesis,we examined changes in life history and behavioural traits injuveniles and adults of both species during four YOY fish predationperiods that were selected based on the presence of YOY perchin the pelagic zone and the relative abundance of Daphnia preyin their gut contents. Our study supports the scenario of negativesize-selective predation by gape-limited YOY perch on both Daphniaspecies. The electivity index indicated that no daphnids witha body length > 1.75 mm were predated by YOY yellow perch.Coexisting Daphnia exhibited phenotypic plasticity in theirantipredator defenses based on their vulnerability to seasonalchanges in size-selective predation of YOY perch. Juvenile Daphniawere the targeted prey and they responded by a decreased bodylength. Behavioural defenses were the dominant strategy usedby both adult Daphnia populations to withstand high predation.A decreased size at maturity was not employed by Daphnia, exceptat the very end of the predation period. Behavioural defensesare short-term strategy adopted to avoid predation. Both antipredatordefenses became unnecessary expenses and were no longer sustainedafter the predation period.     

Anti‐predator response of naïve and experienced common bully to chemical alarm cues

Histological analysis of the skin of common bully Gobiomorphus cotidianus , a New Zealand native eleotrid fish, revealed the presence of club cells in the epidermis. Epidermal club cells are frequently associated with the production of alarm substance (Schreckstoff). The behavioural responses of perch‐naïve and perch‐experienced common bullies to either conspecific skin extract or chemical cues from an introduced predator, perch Perca fluviatilis , were then examined. Both perch‐naïve and perch‐experienced common bullies exhibited a behavioural response when exposed to conspecific skin extract, indicating the probable presence of an alarm substance. In contrast, only perch‐experienced common bullies recognized and exhibited a subsequent behavioural response to the odour of perch. This study is the first to document the presence of epidermal club cells and a behavioural response to a conspecific chemical alarm signal for fishes in the Eleotridae. The results indicate that common bully can learn to recognize perch odour as a threat, and that this ability may be a result of previous predator labelling involving a conspecific alarm substance.     

Parasites reduce territorial behaviour in red grouse (Lagopus lagopus scoticus)

Hypotheses for the cause of population cycles have focused on the role of intrinsic mechanisms such as spacing behaviour and extrinsic mechanisms such as parasitism and predation. This paper examines the interaction between the two dominant hypotheses in the cycles of red grouse, parasitism and spacing behaviour. The influence of the caecal nematode, Trichostrongylus tenuis, on male aggressiveness was investigated using two approaches. First, the territorial behaviour of a group of male grouse experimentally treated with an anthelmintic to reduce parasite intensities was compared with a control group with natural intensities of infection. Second, the response of treated and control males to a novel conspecific territorial intruder were recorded using playback tests. Treated males, with reduced levels of parasitism, won significantly more territorial contests than control males and produced more aggressive behaviour in response to the playback recordings. These results show that parasite removal enhanced aggressive behaviour. Implications of these results on red grouse population dynamics are discussed.     

The Response of Corn Buntings to Playback of Dialects

The response to playback of three different dialects by 10 territorial male corn buntings was investigated; the dialects played were own dialect, the dialect of a neighbouring population (NEAR) and a dialect from 40 km away (FAR). A significant effect of dialects was found for two measures of response, latency and singing rate. Response strength varies in the pattern OWN > NEAR > FAR for these measures. This pattern of response would not maintain the observed homogeneity of dialect populations. The results are similar to those obtained by experiments on other dialect species.     

Cautious response of inexperienced birds to conventional signal of stronger threat

Several studies demonstrated that bird song functions as a first line of territorial defence. The efficiency of deterring rivals depends strongly on the strategy of singing used (e.g. alternating/overlapping singing, singing with low/high rate, matching song type of a rival or singing different type). Causes of between males variation during countersinging are still not fully understood, especially when different signals have similar production costs and their meaning is assigned by arbitrary convention (conventional signalling). We tested whether an oscine bird with small repertoire size, the ortolan bunting Emberiza hortulana , differentiate strategy of responding to song of an intruder in relation to its age and threat value of signals. We performed playback experiments to measure response of second year (SY) and after second year (ASY) males to a song of low (eventual variety singing) and high (immediate variety singing) threat value. We found substantial differences in response to playback, which were related both to the type of stimuli used and age of responding males. Both SY and ASY males gave more calls than songs in response to immediate variety playback, which suggest stronger vocal response to the signal of higher threat value. Approaching loudspeaker was similar for both age classes when lower threat value signal was played back, while simultaneously SY males clearly avoided approaching loudspeaker when stronger threat values signal was played back. We conclude that ortolan bunting differentiate response to signal of different threat value and that the strength of response depends on the age of a male. This study provides experimental evidence that age of receiver affects its response to a territorial intruder. It also demonstrates that observed in many studies variation in response to playback may be an effect of age differences between males, which rarely is controlled.     

Overlapping and matching in the song contests of black-capped chickadees

During vocal contests, animals alter both the timing and the patterning of responses to their opponents. Time-specific responses (such as overlapping an opponent's song) and pattern-specific responses (such as matching the type of an opponent's song) may reveal information about interacting animals. Here we explore the consequences of overlapping and frequency matching during song contests of male black-capped chickadees, Poecile atricapillus. Using interactive playback, we engaged birds of high and low dominance status in vocal interactions with a simulated territorial intruder. The playback intruder either overlapped or avoided overlapping the subjects' songs and either matched or avoided matching the frequency of the subjects' songs. Individuals who were overlapped by the playback intruder showed higher variability in their song length and song timing than individuals who were not overlapped. Individuals who were frequency-matched by the playback intruder responded with more agitated responses (more flights and passes over the speaker and closer distances of closest approach) and spent more time farther away from the loudspeaker. We argue that the timing of song delivery and the choice of song type are distinct functional components of vocal interactions, where overlapping and matching songs are threatening signals that have separate consequences for opponent behaviour and song performance. High-ranking males responded at greater distances from the loudspeaker in all treatments and responded with lower agitation levels than low-ranking males. We demonstrate that males of different quality show different behavioural responses to territorial intruders, where males of high status appear reluctant to engage an intruder as intensely as males of low status.     

Song duration mediates responses of territory owner in a songbird species with a small song repertoire

Song is a sexually selected trait that is involved in mate attraction and territory defence in birds. Songs may convey information about different male quality components. They are flexible in terms of frequency, amplitude, and duration. Although changes in song duration are common, the function of this behaviour has been studied less strongly. It is known that song duration changes within a singing bout and may provide information about aggressive motivation. We tested whether the elongation and shortening of songs affects the responses of territorial ortolan bunting males to neighbour song playback. If changing song duration signals level of aggressiveness, then songs differing solely in duration may elicit behavioural responses of different strength. We performed two tests with different males assigned to two experimental groups and measured approaching and vocal response. In Experiment 1, we tested 18 males, which responded to the playbacks of elongated and normal (control) neighbour songs. In Experiment 2, we tested 17 males, which responded to the playbacks of shortened and normal (control) neighbour songs. Males responded significantly stronger to longer songs in both experiments as measured by the approach variable. Vocal response was not affected by treatment, but it was affected by the order of playback presentation. Our results indicate that song duration might be used for signalling current aggressive motivation during close interactions with rivals.     

Memory-dependent adjustment of vocal response latencies in a territorial songbird

Vocal interactions in songbirds can be used as a model system to investigate the interplay of intrinsic singing programmes (e.g. influences from vocal memories) and external variables (e.g. social factors). When characterizing vocal interactions between territorial rivals two aspects are important: (1) the timing of songs in relation to the conspecific’s singing and (2) the use of a song pattern that matches the rival’s song. Responses in both domains can be used to address a territorial rival. This study is the first to investigate the relation of the timing of vocal responses to (1) the vocal memory of a responding subject and (2) the selection of the song pattern that the subject uses as a response. To this end, we conducted interactive playback experiments with adult nightingales (Luscinia megarhynchos) that had been hand-reared and tutored in the laboratory. We analysed the subjects’ vocal response latencies towards broadcast playback stimuli that they either had in their own vocal repertoire (songs shared with playback) or that they had not heard before (unknown songs). Likewise, we compared vocal response latencies between responses that matched the stimulus song and those that did not. Our findings showed that the latency of singing in response to the playback was shorter for shared versus unknown song stimuli when subjects overlapped the playback stimuli with their own song. Moreover birds tended to overlap faster when vocally matching the stimulus song rather than when replying with a non-matching song type. We conclude that memory of song patterns influenced response latencies and discuss possible mechanisms.     

Acoustic discrimination of sympatric morphs in Darwin's finches: a behavioural mechanism for assortative mating?

Populations with multiple morphological or behavioural types provide unique opportunities for studying the causes and consequences of evolutionary diversification. A population of the medium ground finch (Geospiza fortis) at El Garrapatero on Santa Cruz Island, Galápagos, features two beak size morphs. These morphs produce acoustically distinctive songs, are subject to disruptive selection and mate assortatively by morph. The main goal of the present study was to assess whether finches from this population are able to use song as a cue for morph discrimination. A secondary goal of this study was to evaluate whether birds from this population discriminate songs of their own locality versus another St Cruz locality, Borrero Bay, approximately 24 km to the NW. I presented territorial males with playback of songs of their own morph, of the other morph, and of males from Borrero Bay. Males responded more strongly to same-morph than to other-morph playbacks, showing significantly shorter latencies to flight, higher flight rates and closer approaches to the playback speaker. By contrast, I found only minor effects of locality on responsiveness. Evidence for morph discrimination via acoustic cues supports the hypothesis that song can serve as a behavioural mechanism for assortative mating and sympatric evolutionary divergence.