Female House Wrens may Use a Low‐Amplitude Call as an Aggressive Signal

The majority of work on aggressive signaling has focused on male–male contests. Although females in many species compete over important resources, female signals are understudied. In house wrens (Troglodytes aedon), females compete with other females to protect nesting cavities and eggs. We suggest that a high‐pitched, low‐amplitude call, hereafter abbreviated as ‘HI’, may be an aggressive signal used by female house wrens. Using playback of simulated female intruders, we found that females used HI calls significantly more often during playback of female house wrens compared to playback of heterospecifics. Additionally, HI calls were given significantly more often in the minute preceding physical attack. In comparison, song rates did not predict future attacks. Finally, we present pilot data that suggest listening females may respond more aggressively to female playback containing HI calls compared to playback containing only songs. This suggests the reliability of HI calls could be maintained by a receiver retaliation rule. HI calls bear a striking resemblance to the low‐amplitude songs of many male songbirds, in terms of both acoustic structure and social context. This is one of the few reports of a putative low‐amplitude aggressive signal in a female songbird. However, the nature of female competition may make low‐amplitude signaling an underappreciated signaling form in female animals.     

Male and female rufous‐and‐white wrens do not match song types with same‐sex rivals during simulated territorial intrusions

In birds with song repertoires, song‐type matching occurs when an individual responds to another individual's song by producing the same song type. Song‐type matching has been described in multiple bird species and a growing body of evidence suggests that song‐type matching may serve as a conventional signal of aggression, particularly in male birds in the temperate zone. Few studies have investigated song‐type matching in tropical birds or female birds, in spite of the fact that avian biodiversity is highest in the tropics, that female song is widespread in the tropics, and that female song is the ancestral state among songbirds. In this study of rufous‐and‐white wrens Thryophilus rufalbus, a resident neotropical songbird where both sexes sing, we presented territorial males and females with playback that simulated a territorial rival producing shared and unshared songs. In response, both males and females sang matched song types at levels statistically equal to levels expected by chance. Furthermore, males and females exhibited similar levels of aggression and similar vocal behaviours in response to playback of both shared and unshared songs. These results indicate that rufous‐and‐white wrens do not use song‐type matching in territorial conflicts as a conventional signal of aggression. We discuss alternative hypotheses for the function of song‐type sharing in tropical birds. In particular, we point out that shared songs may play an important role in intra‐pair communication, especially for birds where males and females combine their songs in vocal duets, and this may supersede the function of song‐type matching in some tropical birds.     

Do Blackbirds Signal Motivation to Fight with Their Song?

Conflicts about resources can be solved with physical fights, but animals will in general try to avoid them due to the costs and risks associated with fighting. Fight outcome depends on the fighting ability as well as the motivation to fight, which means that individuals should try to assess both during a pre‐escalation phase. In the present study, we investigated whether blackbirds (Turdus merula) acoustically signal their motivation to fight with their advertisement song, which consists of a motif part with pure‐toned low frequency elements followed by a twitter part with more complex and higher‐pitched elements. We tested (1) whether the temporal and spectral characteristics of spontaneous songs broadcasted by a territorial male could predict its subsequent aggressive behaviour to a playback and (2) if the song of territorial males changed in case they responded aggressively to the playback. We found no effects in the temporal song structure, which contradicts results from a previous study involving playback experiments with blackbirds. We did however find an interesting pattern in the spectral characteristics of the twitter, since aggressively responding males significantly increased the twitter frequency after the playback. The post‐playback songs of aggressive responders were nevertheless indistinguishable from spontaneous pre‐playback songs of non‐aggressive responders. Therefore, we conclude that changes in twitter frequency, rather than absolute twitter frequencies, are advertising information about motivation to fight.     

A field test of the audibility of urban versus rural songs in mountain chickadees

Anthropogenic noise can mask avian vocalizations, and several urban‐dwelling species adjust frequency or amplitude of vocalizations in ways that appear to compensate for increased noise levels. Playback studies have investigated whether receivers differentiate between signals produced by rural and urban males, but it is difficult to determine whether differential response to stimulus reflects differences in audibility versus perceived differences in male signals/condition that result from urban settlement. Here, we performed paired‐playback trials to determine whether mountain chickadees (Poecile gambeli) differentiate urban versus rural songs when both stimuli were broadcast within noise. For each playback, stimuli were played in short bouts starting at a low signal‐to‐noise ratio and increasing in relative amplitude with each successive bout. If the primary function of adjusted urban songs is propagation in noise, we hypothesized that focal males would respond sooner to urban versus rural playbacks (detect at lower signal‐to‐noise ratios). If urban songs solely encode information about the male's condition (such as increased aggression), then we predicted only a differential aggressive response to playbacks once detected. If urban songs both increase propagation and embed information on male condition, we predicted a combination of both response types. We found no difference in latency to first response in urban versus rural songs, but some evidence for differential aggression to playback dependent on both stimulus type (urban vs. rural) and local ambient noise levels; focal males in noisy (urban) sites responded aggressively to both stimulus types, whereas focal males in quiet (rural) sites responded more aggressively to urban than to rural stimuli. This context‐dependent discrimination may be the result of increased aggression in urban habitats, improved communication in noisy habitats by urban signallers and receivers, or some combination of the two.     

Sexual differentiation and seasonal variation in response to conspecific and heterospecific acoustic signals

Interspecific territoriality is frequently reported between closely related species; however, few studies have demonstrated interspecific territoriality between distantly related species living in sympatry. We conducted playback experiments to investigate territorial behaviour in male and female White‐bellied Wrens (Uropsila leucogastra) in response to simulated conspecific and heterospecific intruders during the breeding and non‐breeding seasons. We explored whether heterospecific songs of the Happy Wren (Pheugopedius felix), a distantly related species and ecological competitor, elicited antagonistic responses from focal White‐bellied Wrens, and whether such responses differed between the sexes. We also examined whether male and female responses to conspecific and heterospecific rivals varied with season. We found that male White‐bellied Wrens always responded to conspecific song, and responded significantly more to heterospecific song compared to a control stimulus (Tropical Parula, Setophaga pitiayumi). In contrast, although female White‐bellied Wrens responded strongly to conspecific song, their response to heterospecific song did not differ significantly from the control stimulus. The proportion of males that responded to heterospecific songs and the proportion of females that responded to conspecific songs varied seasonally, showing significantly lower responses during the breeding season. The intense responses of male White‐bellied Wrens to playback of heterospecific songs suggest that they recognise ecological competitors based on their vocal signals. Furthermore, the decrease in agonistic interactions during the breeding season is in line with the hypothesis that aggressive behaviour may be detrimental to reproductive and parental activity, and the hypothesis that heterospecific animals pose less of a threat during the breeding season.     

A Test of the Effect of Male Song on Female Nesting Behaviour in the Serin (Serinus serinus): a Field Playback Experiment

It is well established, through laboratory experiments, that male song in birds can stimulate female reproductive activity, affecting their behaviour and physiology, such as follicular growth, nest building and egg‐laying. However no clear demonstration has yet been provided that this effect works under natural conditions. Previous work in natural populations of serins showed that female nest‐building behaviour correlated with male singing time. Furthermore male serin song peaked exactly in the day that rapid follicular growth was estimated to start in females, suggesting that in this species song may also serve to stimulate the female's reproductive development. Direct causal evidence, however, was lacking. We conducted field playback experiments to investigate how song can influence female nesting activity during nest building. Our results show that females who listened daily to playbacks of serin songs, during the nest‐building stage, spent more time nest building than females that were not exposed to additional songs. Moreover, the singing behaviour of the mated males was not affected by the playbacks, suggesting that the song playback treatment had a direct positive effect on female nesting behaviour.     

The Development of Local Song Preferences in Female Cowbirds (Molothrus ater): Flock Living Stimulates Learning

We carried out two experiments across 2 yr on song perception in female cowbirds (Molothrus ater). In the first experiment, juvenile and adult female brown‐headed cowbirds, living in same‐sex flocks in outdoor aviaries, were periodically tutored with recordings of local male cowbirds’ songs. In the spring, four adult male cowbirds were placed with half of the females for a 12‐d period. We then tested song preferences of all females by measuring copulation solicitation displays during the breeding season. We found that the females exposed only to tape‐tutor songs preferred those songs to those of the unfamiliar males used as companions and that the females allowed to interact with males preferred their songs over the familiar tape‐tutor songs. These data establish the modifiability of female cowbirds’ song preferences at the level of local song. In a second experiment, we studied the playback responses of juvenile females, hand‐reared from the egg, who were tape‐tutored only in the spring in the presence or absence of adult females. There were no differences between the responses of juveniles housed with or without adult females and the hand‐reared juveniles were significantly less responsive to song than adult females. Adult females responded more to familiar songs than to the unfamiliar songs: juvenile females made no such distinction. Taken as a whole, these data are the first to document that female cowbirds’ song preferences for local song can be reshaped by post‐natal experience. These data complement recent study in cowbirds and other species showing that socially more complex contexts reveal plasticity in female song preferences that are not apparent when learning opportunities are constrained by impoverished laboratory settings.     

Song Amplitude of Rival Males Modulates the Territorial Behaviour of Great Tits During the Fertile Period of Their Mates

Bird song is a widely used model in the study of sexual selection. Variation in the expression of sexually selected traits is thought to reflect variation in male genetic and/or phenotypic quality. Vocal amplitude is a song parameter that has received little attention in the context of sexual selection, but there is some evidence that the intensity of bird song affects female preferences. Here, we tested whether the amplitude of broadcast song plays a role in male–male competition. We used song playback with varying song amplitude (within the natural amplitude range of the species) and a dummy bird taxidermy to simulate territorial intrusions in the great tit, Parus major, during the fertile period of the female and measured the response of the local male. The results show that playback amplitude significantly affected the subjects’ behaviour: after approaching to within 25 m around the loudspeaker, territorial males stayed longer within that perimeter after the playback of high‐amplitude songs compared with low‐amplitude songs. Our findings add to the small but growing body of evidence suggesting that vocal amplitude may be a sexually selected song trait.     

Function of pair duets in the eastern whipbird: cooperative defense or sexual conflict?

Paired male and female eastern whipbirds, Psophodes olivaceus,sing precisely coordinated, male-led duets. Four broad explanationshave been proposed for the function of duets: 1) cooperativeresource defense, 2) prevention of partner usurpation, 3) defenseof an individual's own position within the partnership, or 4)mate identification and localization. These 4 hypotheses makedifferent predictions about how male and female residents shouldrespond to simulated intrusion by other pairs or individuals.We compared the behavioral and vocal responses of 20 pairs ofeastern whipbirds to simulated territorial intrusions by: 1)a solitary singing male, 2) a solitary singing female, and 3)a duetting pair. Males and females did not coordinate theirapproach to the playback speaker and showed sex-specific responsesto playback. Males did not respond differently to duetting versussolo singing intruders. By contrast, females approached moreclosely during solo female song than during solo male song orduet playback. Females also produced specific vocalizationsonly in response to duet and solo female playback. Both sexesapproached the speaker more closely and quickly during playbackof same-sex solo songs than opposite-sex solo songs. Finally,females answered more of their mate's songs during simulatedintrusion by a lone female than during simulated intrusion bya lone male. Our results suggest that duets in this speciesprimarily function to allow females to defend their exclusiveposition in a partnership. Mate defense by females is unusualin birds but may be promoted in eastern whipbirds by a female-biasedsex ratio and the need for exclusive access to male care. Thus,duets result from independent and conflicting strategies ofmate and territory defense in males and females.     

Broad Selectivity for Courtship Song in the Cricket Gryllus bimaculatus

Various characteristics of a long‐distance acoustic signal have been shown to vary to different degrees. It has been suggested that female preferences based on stable song parameters are stabilising or weakly directional, and preferences based on variable parameters are strongly directional. We tested this hypothesis based on a short‐distance signal (courtship song) produced by the field cricket, Gryllus bimaculatus. We studied the degree of variability of different courtship song parameters and the behavioural importance of several parameters using synthesised song models in playback experiments. We found that most of the courtship song elements of G. bimaculatus were quite variable (coefficient of variation, CV, in the range of 20–53%). The most variable parameter of the courtship song was the relative amplitude of two elements: high‐amplitude ticks and low‐amplitude pulses. Because songs containing only ticks (of rare occurrence) appeared to be more effective than songs with both ticks and pulses (of frequent occurrence), we consider female preferences to be directional. Alteration of less variable traits, such as the carrier frequency and duration of ticks (CV = 20–25%), had a different effect on female responsiveness. The synthesised songs with different carrier frequencies of ticks were as attractive to females as the positive control (courtship of muted males accompanied by playback of the recorded song). Altering the duration of ticks had a crucial effect on the female response rate, decreasing female responsiveness to the level observed in the negative control (courtship of muted males). Thus, we did not find a strong relationship between the variability of individual song parameters and their potential importance in song recognition and the evaluation of male quality. The partial inconsistency of our results with the data of other authors may be due to different patterns of past and current selection on long‐distance and short‐distance acoustic signals.     

ASYMMETRIC RESPONSE PATTERNS TO SUBSPECIES‐SPECIFIC SONG DIFFERENCES IN ALLOPATRY AND PARAPATRY IN THE GRAY‐BREASTED WOOD‐WREN

Song divergence between closely related taxa may play a critical role in the evolutionary processes of speciation and hybridization. We explored song variation between two Ecuadorian subspecies of the gray‐breasted wood‐wren (Henicorhina leucophrys) and tested the impact of song divergence on response behaviors. Songs were significantly different between the two subspecies, even between two parapatric populations 10 km apart. Playback experiments revealed an asymmetric response pattern to these divergent subspecies specific songs; one subspecies responded more to songs of its own subspecies than to the other subspecies’ songs, whereas the second responded equally strongly to songs of both subspecies. While song parameters revealed a mixed pattern of divergence between allopatric and parapatric populations, the majority of spectral characteristics showed increased divergence in parapatry, suggestive of character displacement. This increased song divergence in parapatry appeared to affect behavioral responses to playback as discriminating responses were most prominent in parapatry and against parapatric songs. The clear behavioral impact of subspecies‐specific song differences supports a potential role for song as an acoustic barrier to gene flow. The asymmetric nature of the responses suggests that song divergence could affect the direction of gene flow and the position of the subspecies‐specific transition.     

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.     

Female Songs of the Nonduetting Javan Gibbons (<Emphasis Type="Italic">Hylobates moloch</Emphasis>) Function for Territorial Defense

Duets in territorial, pair-living primates may function to maintain intragroup cohesion, promote intergroup avoidance, and assist in territorial and resource defense, as well as advertising and reinforcing pair bonds. Despite the absence of duetting in Javan gibbons (Hylobates moloch), recent playback experiments suggested that Javan gibbon songs also play a role in pair-bond advertisement as well as territorial and mate defense. However, playback experiments only assess motivations of the listener, which may not reflect the motivations of the caller. We conducted an observational study of naturally occurring female songs in two groups of Javan gibbons from July 2009 to March 2010 and from March to November 2011 in Gunung Halimun–Salak National Park, Indonesia. We investigated female singing rates in relation to singing location, daily path length, occurrence of intergroup encounters, feeding rate, allogrooming rate, and distance between pair mates. The two females produced 47 songs during 164 observation days. Females in the area of their home range that overlapped with neighboring groups sang more frequently than expected based on time spent in the area of overlap vs. the home range interior. Groups also had longer daily path lengths on days when females sang than on nonsinging days, and on days when they visited the area of overlap than on nonvisiting days. Our findings indicate that, like the duets of other pair-living territorial primates, female Javan gibbon songs function for territorial defense, but we found no support for other functions such as intergroup avoidance, resource defense, and pair-bond reinforcement.     

Sound-pressure Levels and Song Preferences in Female Red-winged Blackbirds (Agelaius phoeniceus) (Aves,Emberizidae)

Female red-winged blackbirds (Agelaius phoeniceus) were tested for response to songs of male red-winged blackbirds differing in sound-pressure level (SPL) of playback. Subjects gave significantly more copulation-solicitation displays in response to playback of songs at 85 dB SPL than to playback of the same songs at 72 dB SPL. The strength of the preference, as judged by the ratio of response to high and low SPL playback (1.15:1), was lower than that of preferences for high SPL shown by insects and anurans. Female red-winged blackbirds responded preferentially to a conspecific song relative to a heterospecific imitation, even when the imitation was presented at an SPL 13 dB higher than that of the conspecific song. By contrast, female redwings did not maintain a preference for multiple conspecific song types over single types when the single song types were presented at the higher SPL. These results are compatible with Klump & Gerhardt's (1987) suggestion that the intensity independence of female preferences varies with the relative benefit females obtain from each preference.     

Immediate early gene (ZENK, Arc) expression in the auditory forebrain of female canaries varies in response to male song quality

In male songbirds, the song control pathway in the forebrain is responsible for song production and learning, and in females it is associated with the perception and discrimination of male song. However, experiments using the expression of immediate early genes (IEGs) reveal the activation of brain regions outside the song control system, in particular the caudomedial nidopallium (NCM) and the caudomedial mesopallium (CMM). In this study on female canaries, we investigate the role of these two regions in relation to playback of male songs of different quality. Male canaries produce elaborate songs and some contain syllables with a more complex structure (sexy syllables) that induce females to perform copulation solicitation displays (CSD) as an invitation to mate. Females were first exposed to playback of a range of songs of different quality, before they were finally tested with playback of songs containing either sexy or nonsexy syllables. We then sectioned the brains and used in situ hybridization to reveal brain regions that express the IEGs ZENK or Arc. In CMM, expression of ZENK mRNA was significantly higher in females that last heard sexy syllables compared to those that last heard nonsexy syllables, but this was not the case for NCM. Expression of Arc mRNA revealed no differences in either CMM or NCM in both experimental groups. These results provide evidence that in female canaries CMM is involved in female perception and discrimination of male song quality through a mechanism of memory reconsolidation. The results also have further implications for the evolution of complex songs by sexual selection and female choice.     

Riverside wren pairs jointly defend their territories against simulated intruders

Duets are a jointly produced signal where two or more individuals coordinate their vocalizations by overlapping or alternating their songs. Duets are used in a wide array of contexts within partnerships, ranging from territory defence to pair bond maintenance. It has been proposed that pairs that coordinate their songs might also better coordinate other activities, including nest building, parental care and defending shared resources. Here, we tested in the riverside wren (Cantorchilus semibadius), a neotropical duetting species that produces highly coordinated duet songs, whether males and females show similar responses to playback. During territorial disputes in songbird species, individuals tend to direct their attention towards same‐sex territorial intruders, but this bias might be less pronounced in duetting species. We performed a dual‐speaker playback experiment to examine how mated individuals respond to speakers broadcasting female‐versus‐male duet contributions. We found that riverside wrens have high levels of converging behaviour by duetting and remaining in close proximity to one another when responding to simulated paired intruders. Males and females spent more than 80% of their time less than 1 m apart while defending their territory. Both individuals in a pair aggressively engaged with both male and female simulated trespassers by approaching equally close and spending equal time near the two speakers. These results suggest that both sexes perceive a paired territorial intrusion as a similar threat and that both partners are highly invested in defending the shared resources. This study is one of the few to demonstrate equal attention and aggression from mated pairs towards simulated same‐sex and opposite‐sex intruders. We suggest that pairs responding together, in close proximity to one another, might be favourable in duetting species when defending the territory because maintaining a close distance between partners facilitates the extreme coordination of their joint territorial signals.     

Two Novel Vocalizations Are Used by Veeries (Catharus fuscescens) during Agonistic Interactions

Avian vocalizations are common examples of the complex signals used by animals to negotiate during agonistic interactions. In this study, we used two playback experiments to identify agonistic signals in a songbird species with several acoustically complex songs and calls, the veery. In the first experiment, we compared veery singing behavior in response to simulated territorial intrusions including playback of three variations of veery song: 1) song alone as a control, 2) songs with added whisper calls, and 3) songs with introductory notes removed. In the second experiment, we used multimodal stimuli including songs, whisper calls and songs with introductory notes removed, along with a robotic veery mount. Focal males readily responded to all of the playback stimuli, approached the speaker and/or robotic mount, and vocalized. Male veeries gave more whisper calls, and sang more songs without the introductory note in response to all types of playback. However, veeries responded similarly to all types of stimuli presented, and they failed to physically attack the robotic mount. These results indicate that rival veeries use two different types of novel vocalizations: whisper calls and songs lacking the introductory note as agonistic signals, but do not allow us to discern the specific functions of these two vocalizations.     

Female recognition of trills in the male calling song of the field cricket,<Emphasis Type="Italic"> Teleogryllus taiwanemma</Emphasis>

The calling song of the field cricket, Teleogryllus taiwanemma, is usually considered to consist of sequences of separate chirps. However, sometimes it comprises a phrase of several chirps in a row, with one long chirp (chirp) and a few short chirps (trills). In this study, I compared the phrase containing only chirps with that containing both chirps and trills by analyzing male songs and conducting playback experiments of male songs to females. The song analyses showed significant differences between chirps and trills for all song parameters except bandwidth. To test whether female preference differed with respect to the two phrases, I performed two-speaker playback experiments. When the same numbers of phrases were presented per unit time, females preferred the song with trills to that without trills. This result may reflect female preference for songs with greater sound density. In subsequent playback experiments, I equalized the total sound duration per unit time (duty cycle) in songs with and without trills. The numbers of females that preferred songs with and without trills did not differ significantly. This suggests that trills can attract females like chirps do, even though the two sounds have different components.     

Geographic song discrimination in relation to dispersal distances in song sparrows

Whether geographic variation in signals actually affects communication between individuals depends on whether discriminable differences in signals occur over distances that individuals move in their lifetimes. We measure the ability of song sparrows (Melospiza melodia) to discriminate foreign from local songs using foreign songs recorded at a series of increasing distances and compare the results with previous measurements of dispersal distances. We test discrimination in males using playback of songs on territories and measuring approach and in females using playback to estradiol-treated captives and measuring courtship display. Females fail to discriminate against foreign songs recorded at 18 km but do discriminate against foreign songs recorded at 34, 68, 135, and 540 km. Males fail to discriminate against foreign songs recorded at 18, 34, 68, 135, and 270 km but do discriminate against foreign songs from 540 km. Females are more discriminating, but even they do not discriminate at a distance three times the root-mean-square dispersal distance, as estimated from mtDNA variation. We suggest that female preference for local songs benefits females not because it allows them to reject foreign males but because accurate production of local song serves as a test of song-learning ability.     

Social Guidance of Vocal Learning by Female Cowbirds: Validating its Functional Significance

Juvenile male M. a. ater cowbirds, who have never heard other male cowbirds sing, develop distinctively different repertoires when housed with M. a. ater females from their own area versus M. a. obscurus females from a distant population. Because female cowbirds do not sing, the differences in the males' songs do not arise through vocal imitation. Here we provide data demonstrating that the songs of female-housed males are functionally, as well as acoustically, distinctive. The songs of 8 groups of males were tested where the groups differed by age of singer, acoustic experience, and identity of social companion. The playback results demonstrate that non-singing female cowbirds not only stimulate the male to modify song content, but song potency. As such, they demonstrate the critical role female cowbirds may assume in the proximate and ultimate regulation of vocal development.