Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

An experimental study of song type function in the bobolink (Dolichonyx oryzivorus)

Summary We tested the hypothesis that the alpha and beta songs of male bobolinks (Dolichonyx oryzivorus) have separate intrasexual and intersexual functions by presenting caged males and caged females on the territories of male bobolinks. Although all males in our study population share both alpha and beta songs, we found no evidence for a specialized intersexual function of alpha song or a specialized intrasexual function of beta song. Territorial males increased their singing rate of both alpha and beta songs during both the male presentations and the female presentations. The proportions of the two song types did not change between control and experimental periods for either male or female presentations. We found no evidence in support of specialized intrasexual and intersexual functions of note types. Only 10% of the vocalizations initiated by territorial males were complete alpha or beta songs, while 87% were song fragments, and the remaining 3 % were compound songs. We did find that territorial males responded to simulated male intruders by increasing the proportion of fragmented songs and decreasing the proportion of compound songs. Moreover, males included fewer notes per song initiated and sang a higher proportion of songs that included only introductory notes during male presentations. We conclude that the vocal response of territorial male bobolinks to conspecific male intruders is to sing short, simple sequences of either song type. We found no consistent changes in vocal behavior of territorial males in response to female presentations other than increases in song rates.     

Effects of early auditory experience on the development of local song preference in female swamp sparrows

A growing number of studies show that learning about male mating signals can shape the way females discriminate among males and may influence the evolution of both female preferences and the male traits under selection. Female songbirds commonly prefer local songs over foreign songs from a different population. In some species, however, the extent of variation among songs sung by different males within a population is as great as the variation observed between populations, raising the question of how females are able to discriminate local from foreign songs. Here, we report that laboratory-reared female swamp sparrows (Melospiza georgiana) not only show a preference for the particular song types with which they were tutored as compared with both foreign songs and unfamiliar local songs but also show preference for unfamiliar local songs over unfamiliar foreign songs. An acoustic analysis comparing tutor songs and those presented as unfamiliar local and foreign songs suggests that female swamp sparrows might be attuned to the specifics of local note phonology when assessing the attractiveness of unfamiliar songs. Our results demonstrate that early auditory experience influences response to geographic song variation in female swamp sparrows, and suggest the possibility that female songbirds may generalize what they learn from songs early in life to novel songs heard in adulthood. Additional work is needed to evaluate the contribution of unlearned predispositions for local song.     

Females alter their song when challenged in a sex-role reversed bird species

Birdsong serves to attract mates and to deter territorial rivals. Even though song is not restricted to males, this dual function has almost exclusively been demonstrated for male song. To test the generality of hypotheses on birdsong, we investigated female song in the sex-role reversed, classically polyandrous African black coucal (Centropus grillii) in the context of female–female competition. We compared spontaneously vocalizing females with females vocally responding to a playback simulating a conspecific intruder. Females changed vocal parameters in response to playbacks: They lowered the pitch of their vocalizations and enhanced the duration of song elements when being challenged. Also, the composition of the vocalizations was altered. There was no significant correlation between pitch and body size parameters in spontaneous song, but there was for response songs, with larger females having a lower pitch. These changes in vocal properties suggest that the vocalizations are important for mutual assessment of competitive abilities in females. Our findings confirm the general role of intrasexual competition in vocal communication of birds.     

Influence of wing and tail morphology on the duration of song flight in skylarks

Summary Song flight, which is an aerial song display especially used by many open country bird species, is expected to be energetically very costly. Any morphological adaptation reducing the magnitude of this cost thus would be favored by selection. Male skylarks Alauda arvensis perform frequent song flights during a period of nearly half the year. Skylarks are sexually size dimorphic in most body traits, but particularly in wing area and wing span, which is absolutely and relatively larger in males than in females. Wing loadings, aspect ratios, and flight costs therefore are smaller in males than in females. I investigated the effect of wing area and aspect ratio on the duration of the song flights of individual birds by timing their duration before and after an experimental manipulation of wing area. Male skylarks were caught, ringed, and released (control I), had the tips of their wing feathers cut (control II), or had the tips of their wing feathers reduced by approximately 10 mm. There were no statistically significant differences in morphology or duration of songs between treatment groups before experimental treatments. However, males having the tips of their wing feathers reduced by ca. 10 mm performed only greatly abbreviated song flights. Original wing loading and aspect ratio also affected the duration of song flights, since male skylarks with low wing loadings and high aspect ratios performed longer song flights than did males with high wing loadings and low aspect ratios. This was the case both before and after experimental treatment. Wing area is suggested to reflect the ability of individual skylarks to invest in morphological structures allowing an increased song output.     

Individual recognition of neighbors by song in a suboscine bird, the alder flycatcher Empidonax alnorum

The ability of territorial males to discriminate between songs of their neighbors and songs of strangers has been demonstrated in 27 species of songbirds. Such experiments test only the ability of a subject to discriminate between two classes of stimuli, familiar (neighbors) and unfamiliar (strangers) songs. Individual recognition of neighbors is a finer, more complex type of discrimination. The ability of territorial males to recognize individual neighbors by song has been documented in 12 species of oscine passerines (Passeriformes, Passeri), but has never been demonstrated in suboscine passerines (Tyranni). We investigated recognition of songs of individual neighbors in a suboscine, the alder flycatcher (Empidonax alnorum). We performed a series of song playback experiments and recorded responses of territorial males to songs of neighbors and songs of strangers broadcast from two locations, the neighbor boundary and an opposite boundary. Subjects responded more aggressively to songs of a neighbor when played from the opposite boundary than when played from the neighbor boundary. They responded with equal aggression to songs of strangers regardless of location of playback. The difference in response to neighbor songs between speaker locations and the lack of a difference in response to stranger songs indicate that territorial males associate a particular song with a particular location (territory), and thus recognize individual neighbors.Communicated by I. Hartley     

Female choice for complex song in the European starling: a field experiment

Male European starlings Sturnus vulgaris sing long complex songs that appear to be important in the courtship of females but which also influence competitive interactions between males. We tested the hypothesis that females choose mates on the basis of the complexity of their songs, rather than on the quality of the territories the males defended. In order to determine whether certain territories were preferred over others, the first set of birds to settle in the experimental nest-boxes was removed and a second set allowed to settle. Consistent preferences for certain nest-boxes were indicated by correlations between the settlement patterns of the first and second sets of birds. However, males with the most complex song did not necessarily occupy the most preferred nest sites. Males with more complex song acquired mates faster. This relationship remained significant when nest-site preference was statistically controlled, indicating that female starlings chose males with complex song rather than those that defended preferred nest sites. A number of morphological variables were also found to be uncorrelated with female choice. Song complexity in European starlings increases with age, and the evolution of song complexity in this species is consistent with an age-indicator model of sexual selection. Males with larger repertoires were also in better condition, indicating that females obtain high-quality mates by choosing on the basis of male song. Received: 29 April 1995/Accepted after revision: 9 September 1995     

The territorial flight of male houseflies (Fannia canicularis L.)

Summary The heart rate (HR) of three male and five female European blackbirds (Turdus merula) was monitored by radiotelemetry under three conditions: in a dark cage, in a lighted cage, and in an outdoor aviary. In all three, the response to recorded bird song was tested.The resting HR in the cage ranged from 4.8 to 6.3 beats per second. The HR of all the birds changed in response to playback of conspecific song as well as of the songs of other species. In 68% of trials a typical biphasic HR curve was obtained, with acceleration followed by deceleration (Fig. 2). The time from stimulus onset to the second curve inflection (t _iII) averaged 15–18 s; this was significantly longer for conspecific than for heterospecific song. The parameter t _iII was used as a basic measure of the response.The reaction of the males was longer-lasting than that of the females. The possibility that this reflects the territorial role of the songs is discussed. Response durations were longest in the dark cage, and shortest in the aviary. The reason is thought to be the gradation in total stimulus input available to the bird.The results suggest that the HR response consists of an unspecific component (acceleration) plus a component specific to bird sounds (deceleration). At this fundamental level of stimulus processing it is possible to study reactions to single strophes not only in males, as with more traditional methods, but also in females and young birds.     

Do male plumage and song characteristics influence female off-territory forays and paternity in the hooded warbler?

Extra-pair fertilizations are common in many socially monogamous species, and paternity studies have indicated that females may use male vocal performance and plumage ornaments as cues to assess male quality. Female off-territory forays may represent a key component of female choice and male extra-pair mating success, and female foray behaviour is expected to be strongly influenced by indictors of male quality. In this study, we examined how male song and ornamentation affect how often females left their territories, which males they visited and extra-pair paternity in a socially monogamous passerine, the hooded warbler (Wilsonia citrina). We radiotracked 17 females during the fertile period and quantified male vocal performance (song output and rate) and plumage characteristics (size of the black melanin hood and colour of the black hood, yellow cheeks and breast areas). We obtained blood samples and determined paternity at 35 nests including those of 14 females that we radiotracked. Eleven (65%) of the 17 females forayed off-territory, whilst fertile and female foray rate was positively correlated with the number of extra-pair young in the nest. Females that left their territories more frequently were paired with males that sang at a low rate. In addition, extra-pair mates had higher song rates than the social mates they cuckolded (5.3 songs/min vs. 4.4 songs/min). Female off-territory forays or extra-pair paternity were not significantly related to male plumage characteristics. Our results indicate that a high song rate influences both the foray behaviour of a male’s social mate and the likelihood that he will sire extra-pair offspring with neighbouring females.     

Mate choice by female white-crowned sparrows in a mixed-dialect population

Summary It has been argued that the song dialects of white-crowned sparrows, Zonotrichia leucophrys, may function to restrict gene flow between dialect populations, thus promoting adaptation to local environmental conditions. One proposed mechanism whereby this may occur is if a female mates preferentially with males singing her natal dialect, typically the song type of her father. Previous field and laboratory studies of the role played by song differences in mate choice by female white-crowned sparrows have produced equivocal results. These investigations were conducted in populations in which a single song dialect was predominant. We examined mate choice in a population in which two song dialects are equally common. We found that songs of mates of individual females in successive years were no more likely to be of the same dialect than expected by chance. In addition, individual females did not mate preferentially with males whose songs matched those of their fathers. We conclude that, in this population, females choose mates on the basis of characteristics other than song type.     

Territoriality and monogamy among agile gibbons (Hylobates agilis)

Summary I simulated territorial intrusions by mated pairs, solitary males and solitary females through song playback to investigate the behaviors that agile gibbons (Hylobates agilis) use to maintain territoriality and monogamy. Mated male and female gibbons approached and duetted in response to a simulated territorial intrusion by a neighboring group. Strange solitary females elicited similar approach and singing responses from mated pairs. In contrast, the simulated presence of a solitary male consistently evoked approaches by mated males only. Females and males did not contribute equally to pair movements or to answering songs; males led the majority of approaches, while females initiated all duets. I hypothesize that carrying small, vulnerable infants may inhibit females from leading approaches. These results are consistent with previous studies which suggest that female territoriality prevents male gibbons from becoming polygynous.     

Mate choice plasticity in the field cricket <Emphasis Type="Italic">Teleogryllus oceanicus</Emphasis>: effects of social experience in multiple modalities

Social experience can elicit phenotypically plastic changes in mate choice, but little is known about the degree to which social information from one modality can influence mating decisions based on information from a different modality. I used the field cricket Teleogryllus oceanicus to test whether experience of chemical cues mimicking a high density of sexually mature males causes changes in mate choice based on acoustic signals. T. oceanicus males produce long-range calling songs to attract females for mating, but they also produce waxy, non-volatile hydrocarbons on their cuticle (CHCs) which, when deposited on a substrate, can be detected by females and may provide demographic information. I manipulated female experience of substrate-bound male CHCs and then performed acoustic mate choice trials. When CHCs were present on the substrate during trials, females showed greater motivation to respond to male calling song. This effect diminished with repeated exposure to male songs, demonstrating that the importance of olfactory cues in altering acoustic mate choice decreased with increasing exposure to acoustic signals. However, the temporal nature of CHC experience mattered: previous experience of CHCs did not alter subsequent female choice for male calling song traits. Exposure to male song increased the threshold of mate acceptance over time, and individuals varied considerably in overall levels of responsiveness. Taken together, the results demonstrate that mate choice is dependent on social context mediated by multiple modalities in T. oceanicus, but they do not support the idea that prior experience of social cues in one modality necessarily influences later mating decisions based on other signalling modalities.     

Spatial and age-related variation in use of locally common song elements in dawn singing of song sparrows <Emphasis Type="Italic">Melospiza melodia</Emphasis>: old males sing the hits

In many songbirds, individuals have repertoires of multiple song types, some of which may be shared with others in the local area. Hypotheses about the evolution of song repertoires differ as to whether selection acts primarily on repertoire size itself or the ability to match songs of neighbours. We used a 16-channel acoustic location system to record neighbourhoods of song sparrows (Melospiza melodia melodia) during the dawn chorus. We asked whether males sing all songs with similar frequency as predicted by the Repertoire Size Hypothesis, whether males preferentially sing highly shared songs as predicted by the General Sharing Hypothesis, or whether use of highly shared songs is associated with phenotype as predicted by the Conditional Sharing Hypothesis. Contrary to the Repertoire Size Hypothesis, most males did not sing all songs equally often. Contrary to the General Sharing Hypothesis, we found no general tendency to overproduce highly shared songs. The degree to which males overproduced highly shared songs was repeatable across days, indicating consistent individual differences, and varied across neighbourhoods. Moreover, and consistent with the Conditional Sharing Hypothesis, older males were more likely to overproduce highly shared songs. If highly shared song is a conventional signal of aggression, with the threat of receiver retaliation maintaining honesty, older males may be more willing or able to risk conflict. Alternatively, males may learn which songs are effective signals for an area. Finally, age-related variation in vocal performance may shape the adaptive value of highly shared song.     

Songs of isolation-reared sparrows function in communication,but are significantly less effective than learned songs

Summary Songs of male song sparrows (Melospiza melodia) and swamp sparrows (M. georgiana) isolated before hatching from conspecific song were played to males and females of both species. Isolate songs of these sparrows resemble natural conspecific song in several aspects of gross structure, but differ from natural song in note structure. Male territory owners of both species responded more to conspecific isolate song than to heterospecific normal song. Captive females of both species, previously treated with estradiol, courted in response to isolate song but not to heterospecific song. We conclude that there is sufficient speciesspecific information in isolate song of both species to allow a degree of normal function. To assess the importance of the structural refinements added through learning, we compared response to isolate and natural conspecific songs. Male territory owners and captive females responded more to natural than to isolate songs in both species. Learning thus significantly increases the potency of song in both intersexual and intrasexual communication.     

Bachelor and paired male rufous-and-white wrens use different singing strategies

To attract a breeding partner, males may behave differently when they are bachelors compared to when they are paired. Comparisons between groups of paired males versus groups of unpaired males in temperate-breeding animals have revealed such differences in signalling behaviour. Few studies, however, have explored how individual males alter their signalling behaviour with changes in pairing status, and very few investigations have explored paired versus unpaired male behaviour in tropical animals. During a 5-year study in Costa Rica, we analysed changes in the singing behaviour of male rufous-and-white wrens (Thryothorus rufalbus) when they were paired and when they were bachelors. We compared three aspects of male vocal behaviour: gross differences in song output, variation in repertoire use and differences in song structure. Males as bachelors had significantly higher song output and switched song types less frequently. Contrary to our expectation, bachelors sang significantly fewer song types from their repertoire compared to when those same males had a breeding partner. Songs sung by bachelor males were higher in syllabic diversity and had broader-bandwidth terminal syllables than the songs those males sang only when paired. Within song types, the fine structure of songs remained consistent across pairing status. Our results demonstrate that males change their singing behaviour with pairing status, delivering songs at a higher rate but with less variety when they are bachelors. Rufous-and-white wrens are renowned for their vocal duets, and we discuss the pattern of repertoire use in light of their duetting behaviour. These results enhance our understanding of how male behaviour varies with pairing status and the importance of vocal signalling behaviour in socially monogamous tropical animals.     

Song output and stimulation of the female in white-throated sparrows

Summary We investigated the importance of song length and singing rate in stimulating female white-throated sparrows, Zonotrichia albicollis. In November 1988 ten female and one male white-throated sparrow were captured during the fall migration. To enhance the expression of sexual receptivity, the females were implanted with 17-beta-estradiol silastic pellets. Using the visual stimulus of a non-singing male to further enhance the expression of sexual receptivity in the females, we found that females gave a greater response, in the form of copulation displays, to a five-note versus a two-note version of a typical male white-throated sparrow song. Each version was played back at four songs per minute. In May 1990 birds were captured and treated as before, except that the two-note song was played at ten songs per minute and the five-note song at four songs per minute. Therefore, each female heard twenty notes per minute. In the second experiment, the females did not show a statistical difference in response to the two- and five-note song playbacks. We interpret the results of the second experiment as indicating the song rate as well as song length is important in stimulating female white-throated sparrows because increasing the rate of the two-note song brought the response up to the level of response we obtained to the five-note song played at the slower rate. We conclude that in white-throated sparrows, song output is important for female stimulation.     

An experimental investigation of the bioacoustics of cowbird song

Summary Female cowbirds (Molothrus ater ater), maintained in isolation from males during the breeding season, respond to the playback of male song with copulatory postures. They respond to some songs more than to others. Cowbird song potency can thus be operationally defined by the proportion of copulatory postures a song elicits across multiple playbacks. The purpose of the present study was to explore whether song potency changes with distance in the field. No field recordings elicited high levels of responding by the females. When songs of known high potency are systematically degraded, the results indicate that female cowbirds are sensitive to small changes in signal to noise ratio and to atmospheric attenuation. The data suggest that cowbird song potency degrades very rapidly with transmission distance in the field.     

Testing the developmental stress hypothesis in canaries: consequences of nutritional stress on adult song phenotype and mate attractiveness

The complex songs of songbirds are thought to have evolved through sexual selection. Sexually selected signals must be associated with costs in order to ensure their honesty as indicator of male quality. Costs may relate to the development of the neural substrate underlying song learning, which develops already very early in life. Song may, therefore, serve as an indicator of the early developmental history. This nutritional stress hypothesis has initially been confirmed for a variety of species, but recent studies using zebra finches as a model species reported somewhat inconsistent effects, and the functional consequences of changes in adult song phenotype remain unclear. We tested the nutritional stress hypothesis in canaries by manipulating either the brood size or the food quality postfledging. The brood size manipulation had a significant effect on early development, and low food quality postfledging led to a transient reduction in body mass. However, we did not find evidence that any of the song traits measured reflected the early developmental conditions, which is in conflict with the nutritional stress hypothesis. Canaries may be less vulnerable to nutritional stress or are able to compensate stressful conditions during early development. However, if males compensated, this compensation may have come at a survival cost. Female mate choice decisions were independent of the developmental history of a male. Instead, females preferred males singing longer song bouts, a trait that may contain a heritable component.     

Habitat-induced changes in song consistency affect perception of social status in male chickadees

The ability of male black-capped chickadees to maintain consistent internal structure between successive iterations of their songs is affected by both their social rank and the quality of their habitat. Lab studies reveal that female chickadees discriminate between songs of dominant and subordinate males, which vary in acoustic structure. We investigate whether males also rely on acoustic structure to assess rival quality during agonistic interactions, and whether habitat-induced differences in song consistency affect the perception of male rank. We conducted a playback experiment to simulate territorial intrusions by dominant, stimuli males into the territories of dominant, subject males; stimuli males were recorded either in low-quality (young forest) or high-quality (mature forest) habitats. Stimuli from low-quality habitat had lower song consistency than those from high-quality habitats, despite being recorded from males of equivalent social rank. Subject males for playbacks (also socially dominant males) were chosen from either habitat type. Subject males in mature forest responded less to young-forest stimuli compared to mature-forest stimuli, despite the stimuli in both cases being recorded from dominant males. Conversely, male subjects in young forest did not differentiate between stimuli, but their response to both stimuli was lower than that of mature-forest subject males to mature-forest stimuli. We demonstrate that the ability to maintain internal song structure in the black-capped chickadee constitutes a signal that appears to be used by males to assess the level of threat of intruders, and that this perception is affected by habitat from which the stimulus males were recorded.     

The behavioral regulation of monogamy in gibbons (Hylobates muelleri)

Summary Responses of mated individuals to playbacks of the songs of solitary females and males permit evaluating the behavioral mechanisms maintaining monogamy in gibbons. When female songs are played back from the centers of their ranges, mated female gibbons typically initiate duets and group approaches toward playback sites. Female songs played back from range boundary locations elicit duetting responses. Responses to solitary female songs do not differ from responses to song duets used by established mated pairs to mediate patterns of intergroup spacing. Mated males lead silent group approaches toward the sites of male song playbacks. These results suggest that range defense by female gibbons forces males into accepting monogamous mating relationships and that monogamy in gibbons is regulated by intersexually-supported, intrasexual aggression.