Song overproduction and selective attrition lead to song sharing in the field sparrow (Spizella pusilla)

Summary Male field sparrows settling on particular territories for the first time often sing two or more song types. Vocal experience with neighboring males then leads in most cases to the deletion from the male's repertoires of all but one song type. In 2 years, settling males retained the song type that most closely resembled that of a neighbor. Focal observations in 1 year revealed that the most actively singing neighbor was the one the new birds resembled. Songs were not selected on the basis of their absolute acoustic characteristics, but rather by their resemblance to neighbors' songs. After the attrition process is complete, males retain their song for the rest of their lives. Evidence supports the hypothesis that the actual acquisition of a song need not take place at the time and place as males occupy territories. Rather, males may acquire an abundance of song models earlier in life and then select songs from this repertoire that most closely resemble those of their neighbors. The development of mature repertoires and of local dialects is the outcome of one process in this species.     

Male's return rate, rather than territory fidelity and breeding dispersal, explains geographic variation in song sharing in two populations of an oscine passerine (Oreothlypis celata)

Males of some oscine passerines learn and share songs of neighboring males. This process can lead to the formation of song pattern neighborhoods or microhabitat song dialects. The degree to which song sharing occurs between populations and the spatial scale over which neighboring males share songs, however, can vary widely, and interpopulation comparisons have suggested that song sharing is more common in residents than in migrants. Here, we examine two populations of the orange-crowned warbler (Oreothlypis celata) to quantify patterns of song sharing at the northern (long-distance migrant) and southern (short-distance migrant) edges of the breeding distribution and to test if return rate, territory fidelity, and breeding dispersal explain the patterns found in the two populations. The southern population (O. celata sordida breeding on Santa Catalina Island, California; 33°N) had a higher annual return rate to their territories and exhibited higher song sharing in neighborhoods than their counterparts (O. celata celata breeding in Fairbanks, Alaska; 64°N). Year-to-year patterns of territory fidelity and breeding dispersal distances were similar between populations. Our results suggest that if migratory distance generally covaries with the proportion of returning males, this could explain different levels of song sharing between the short- and long-distance migrants.     

Vocal dialect and genetic subdivisions along a geographic gradient in the orange-tufted sunbird

At least four hypotheses have been suggested to explain the formation and maintenance of song dialects among birds: historic processes (epiphenomenon), genetic or local adaptation, acoustic adaptation, and social adaptation. We studied spatial and temporal distribution of dialect in the orange-tufted sunbird (Nectarinia osea), a small nectarivorous bird that expanded its breeding range in Israel during the past 100 years from the southern part of Rift Valley to the entire country. Sunbird range expansion was concurrent with the establishment of many small settlements with an ethos of gardening, which introduced many ornithophilous plants. We recorded songs and genetically screened individual sunbirds in 29 settlements distributed across a 380 km north–south gradient along the Rift Valley. We show that dialects cluster together into geographical regions in 70% of cases, a moderate concurrence to geography. Settlement establishment date, geographical position, and genetic distance between local populations (i.e., settlements) were all poor predictors for the variance among song dialects. The specific effect of habitat was not tested because all sampled localities were similar in their physical and acoustic properties. Using a network analysis, we show that dialects seem to aggregate into several network communities, which clustered settlement populations from several regions. Our results are best explained by either the epiphenomenon hypothesis or the social adaptation hypothesis, but at present our data cannot state unequivocally which of these hypotheses is better supported. Last, we discovered a negative association between network centrality and genetic diversity, a pattern that requires further examination in other systems.     

Song sharing correlates with lifetime social pairing success but not territory tenure in the Puget Sound white-crowned sparrow

Imitative song learning in birds often results in the formation of vocal dialects or local song neighborhoods in which males in close proximity share very similar songs. If song sharing confers a selective advantage due to intra-sexual competition or inter-sexual mate choice, song sharing should enhance the singer’s territory tenure or lifetime pairing success. We tested this in a migratory population of the Puget Sound white-crowned sparrow, Zonotrichia leucophrys pugetensis, in which some territory neighbors share whole songs. Song sharing with territory neighbors in the first year on territory did not correlate with territory tenure but correlated positively with social pairing success throughout a male’s lifetime when length of territory tenure was controlled. Thus, song sharing in this migratory population does not appear to confer a benefit in male–male interactions, at least as measured by the length of territory tenure. However, males that share songs do benefit from higher lifetime pairing success. Combined with the data available for other species, it appears that intra- and inter-sexual selection may have stronger effects on song sharing in sedentary and migratory populations, respectively.     

Habituation and song repertoires in the great tit

Summary This paper is concerned with the idea that the song repertoires of passerine birds are an evolutionary adaptation to reduce habituation in listeners. In an experiment involving 16 territorial males I played either a single song or a repertoire of songs for 15 two-min trials through a single loudspeaker near the edge of the territory. In a second experiment with 10 birds I played the songs through one of two loudspeakers in different parts of the territory, alternating between loudspeakers on successive trials. The birds tended to habituate more rapidly to single song playback than to repertoires. In the second experiment the overall level of habituation was lower and the difference between the two treatments was less marked.Two features of song repertoire organisation are consistent with the habituation hypothesis (1) the avoidance of low recurrence intervals in switches between song types, and (2) the fact that within repertoire variability is as great or greater than between repertoire varability.The main problem with the habituation hypothesis is that habituation by listeners does not seem to be adaptive, so it is not clear why they should habituate. I suggest a hypothesis. Intruders may assess the density of birds in an area by listening to songs, so that habituation may be a mechanism by which this density assessment is achieved. Repertoires could be a mechanism by which resident birds cheat, through increasing the apparent density of singing birds.     

Song-type sharing in song sparrows: implications for repertoire function and song learning

One hypothesis for the function of song repertoires is that males learn multiple song types so that they may share songs with neighbors, allowing them to match during territorial interactions. In at least one song sparrow population, in Washington, territorial males share a high proportion of song types with their neighbors and use these shared songs in matching. We recorded song sparrows in Pennsylvania and quantified sharing of whole songs and song segments. We found that song sharing is an order of magnitude less common in the Pennsylvania population. We found sharing of song segments to be significantly more common than the sharing of whole songs in three of the five fields we examined, while we found no significant differences between whole and partial song sharing in the remaining two fields. Finally, we found no evidence that sharing is greater between birds in the same field compared to birds in different fields. Taken with the data from Washington song sparrows, these results provide evidence for intraspecific geographic variation in the organization of song repertoires, and suggest that song sharing has not been a strong selective force in the evolution of song repertoires in song sparrows as a species. Furthermore, Washington and Pennsylvania song sparrows differ in how they learn song, in that Washington birds copy whole songs, while Pennsylvania birds appear to copy and recombine song segments, as has been found in laboratory studies of song learning. Thus both song learning and the function of song repertoires differ between populations of song sparrows. Such intraspecific geographic variation offers a unique opportunity to explore the ecological and historical factors which have influenced the evolution of song. Received: 30 June 1997 / Accepted after revision: 8 March 1998     

Song of the humpback whale — Population comparisons

Summary Humpback whale songs recorded on tropical calving grounds exhibit different dialects depending on the oceanic basin. Songs sampled simultaneously from two populations in the North Pacific (Hawaii and Mexico) were essentially identical. These North Pacific songs were clearly different from the song type shared by two populations in the North Atlantic (Cape Verde Islands and West Indies). Songs from the Southern Hemisphere (Tonga) represent a third distinct dialect. Our evidence shows that, despite annual change in song organization, significant differences in humpback song occur between isolated ocean basins, while only subtle differences exist within an oceanic population (Hawaii and Pacific Mexico).     

Ranging of songs with the song type on use of different cues in Carolina wrens: effects of familiarity

Male territorial song birds are usually spaced far apart and most often hear conspecific song after it has been degraded by propagation through the environment. Their ability to use the degradation of songs to assess the distance of a singing rival without approaching (called ranging) presumably increases the efficiency of defending a territory. In order to assess degradation in a song the receiver needs to compare the characteristics of the received song to its characteristics at the source or at different distances. Earlier experiments on ranging in species with song repertoires have suggested that prior familiarity with the particular song type is necessary for ranging. Here I show that male Carolina wrens (Thryothorus ludovicianus) can use either temporal or spectral characteristics for ranging song types which they were unlikely to have heard previously. Playbacks consisting of only one song prevented subjects' close-range experience with the loudspeaker, and flights beyond the loudspeaker provided direct evidence for over-assessment of distance when songs were degraded. Because ranging of songs was not affected by the degree of familiarity with the song type, this experiment provides no evidence that song repertoires hinder ranging in Carolina wrens, as suggested by Morton's ranging hypothesis. Instead, at least approximate ranging of songs is evidently possible by assessment of degradation in general features of a species' songs. Received: 9 August 1996 / Accepted after revision: 15 February 1997     

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.     

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.     

The role of distance,familiarity, and time of day in Carolina wrens responses to conspecific songs

Summary We tested two predictions of the ranging hypothesis (Morton 1982) which suggests that: (1) For distance estimation of singing intruders from them, birds compare degraded songs they hear to undegraded songs in their memory. (2) The strength of response of birds to songs depends on how well they can range the songs as coming from inside or outside the territory.In accordance with first prediction we found that, Carolina wrens discriminate significantly between songs played from outside to those played inside their territories only if they are familliar with the song type played (Fig. 1). In accordance with the second prediction, on the average, the birds in our study responded stronger to unfamiliar than to familiar songs when these were played from outside the territory, but stronger to familiar than to unfamiliar songs when played inside it (Fig. 2). We concluded that this was because they could range the familiar songs with greater certainty.The variance in the data was, in part, due to changes in the birds' responses during the day. We report for the first time, an increase in the strength of territorial defense of birds later in the morning, accompanied by a decrease in singing in response to song playback. We suggest that these changes reflect an increase in energy gain and worse acoustic conditions later in the day.     

Responses to playback of whistle songs and normal songs in male nightingales: effects of song category, whistle pitch, and distance

In most species of song birds, males develop song repertoires of several different songs. Among this variety, different songs may be used differently in communication and, thus, may have different functions. Here we studied vocal responses to playback of structurally different songs in male territorial nightingales (Luscinia megarhynchos). Nightingales have enormous vocal repertoires of about 200 song types. Among these songs, one category, whistle songs, sticks out syntactically and acoustically. Here we tested whether or not male nightingales match whistle songs with whistle songs and, if so, whether they also match the pitch of the broadcast whistles. Furthermore, we also tested if nightingales treat whistle songs as a separate category of songs. We conducted interactive playback experiments on nocturnal song in which each male received three playback treatments that differed in the number of whistle songs broadcast. Males responded differently during playbacks by singing significantly more whistle songs when the playback tapes contained many whistle songs than when they contained no whistle songs. Males also frequently matched the pitch of the broadcast whistle songs. In contrast to responses during playback, after the playback terminated males sang more whistle songs when no whistle songs were broadcast than when many whistle songs were broadcast. These findings suggest that whistle songs have a specific signal value and that nightingales treat them as a special song category. Electronic supplementary material to this paper can be obtained by using the Springer Link server located at http://dx.doi.org/10.1007/s00265-002-0511-1.     

Tutor choice and imitation accuracy during song learning in a wild population of the Puget Sound white-crowned sparrow

Songbirds learn to sing by modeling their songs on the songs of other males through a process of social learning. Models of social learning predict that animals should be selective in what and when they learn. In this study, we asked whether young males in a wild population of the Puget Sound white-crowned sparrow, Zonotrichia leucophrys pugetensis, were selective in their choice of tutor models and what factors influenced how accurately they imitated tutors’ songs. We first examined two strategies for tutor choice: whether pupils have a conformity bias and/or a preference for high-quality tutors. In keeping with a conformity bias, tutors that sang song types that were relatively common within a radius of about 500 m of their territory were more likely to be imitated than were tutors that sang rarer song types. Most potential tutors were not imitated by pupils. Aspects of tutor quality, such as age, pairing status, and survival to the next year had no effect on whether a tutor’s song was imitated. Secondly, we tested whether pupil repertoire size, pupil quality, and local abundance of tutor models affected the accuracy of song imitations. We found a trade-off between repertoire size and tutor imitation accuracy with males that sang two or more song types developing significantly poorer imitations than males that sang one type. We discuss possible functions of a conformity learning strategy and factors that could produce a trade-off between imitation accuracy and repertoire size.     

Soft song is a reliable signal of aggressive intent in song sparrows

Animals frequently use signals to modulate aggressive interactions. Establishing that a signal is aggressive or threatening requires demonstrating that it is more commonly used in agonistic contexts, that it predicts subsequent aggressive behaviors by the sender, and that receivers respond differently to this signal. Like many birds, song sparrows (Melospiza melodia) produce a low-amplitude “soft song” vocalization that has been hypothesized to be an aggressive signal. Soft song meets the first two criteria, but previous research has failed to demonstrate that soft song provokes aggression or that receivers even perceive soft song differently from normal loud song. We used a playback experiment with taxidermic mount presentation to test whether territorial male song sparrows respond differently to loud and soft song playbacks. Subjects reacted more strongly to the soft song playback by approaching the mount more closely, increasing wing wave displays, and increasing the proportion of their own songs that were soft songs, with further trends toward increasing the number of flights and attacks. These results confirm that soft song is a conventional signal of aggression in song sparrows and that increased receiver retaliation maintains its reliability.     

Individual recognition of song repertoires in two wood warblers

We examined recognition of individual neighbors in hooded warblers (Wilsonia citrina) and Kentucky warblers (Oporornis formosus) by comparing responses to playbacks of neighbors' songs from appropriate and inappropriate boundaries of the subjects' territories. Kentucky warblers, which sing one song type each, responded more strongly to the songs of neighbors broadcast from incorrect than from correct boundaries. Hooded warblers use their repertoires of three to nine song types in two distinct modes of singing: repeat mode involves repeated presentation of one song type; mixed mode involves presentation of two or more song types in irregular sequence. Playbacks of neighbors' mixed-mode songs from appropriate and inappropriate boundaries indicated capabilities of individual recognition similar to those reported previously for repeat-mode songs (Godard 1991) and to those of Kentucky warblers. Repertoires of moderate size, therefore, have no pronounced influence on eventual recognition of individual neighbors in hooded warblers.     

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.     

Song sharing in two populations of song sparrows (Melospiza melodia)

Sharing song types with immediate neighbors is widespread in birds with song repertoires, and sharing songs may confer a selective advantage in some cases. Levels of song sharing vary between different geographical populations of several bird species, and ecological differences often correlate with differences in singing behavior; in particular, males in migratory subspecies often share fewer songs than males in resident subspecies. The song sparrow (Melospiza melodia) appears to fit this pattern: resident song sparrows in western North America generally share 20–40% of their repertoire (of about eight songs) with each neighbor, while migratory subspecies from eastern North America often share 10% or less. We compared song sharing in two populations within a single subspecies of song sparrow (M. m. morphna) in Washington State. These populations, separated by only 120 km, nonetheless differ in migratory tendencies and several other ecological and life history variables. We recorded complete song repertoires from 11 male song sparrows in a high-elevation, migrating population at Gold Creek in west-central Washington, and compared them to two samples (n = 15 and n = 36) from a coastal, resident population at Discovery Park, Seattle, Washington. Despite major differences in habitat, population density, and migratory tendencies, song sharing among Gold Creek males was as high as that among Discovery Park males. In both populations, sharing was highest between immediate neighbors, and declined with distance. We conclude that at the within-subspecies level, neither migration nor population density affect song sharing in song sparrows, a song repertoire species. Received: 26 November 1998 / Received in revised form: 1 May 1999 / Accepted: 29 May 1999     

Temporal patterning of within-song type and between-song type variation in song repertoires

In many songbird species, males sing a repertoire of distinct song types. Song sparrows typically are described as having repertoires of about a dozen song types, but these song types are themselves quite variable and some songs are produced that appear intermediate between two types. In this study we quantify the similarity between successive songs produced by song sparrows in order to determine if differences between song types are emphasized or deemphasized in bouts of continuous singing. In spite of the high degree of variation within song types and similarity between song types observed in this species, we show that transitions from one song type to the next are distinct as compared to transitions within sequences of the same type (Figs. 4 and 5). Variation does not accumulate across sequences of the same song type, and the average variation observed within a continuous sequence of the same song type is significantly less than is predicted from the total variation recorded for that type across many different bouts (Fig. 6). These results support the view that song sparrows include two levels of variation in their singing: differences between song types as is commonly observed in species with song repertoires, and an independent level of variation observed for songs of the same type.     

The song of the humpback whale Megaptera novaeangliae in the West Indies

Songs of the humpback whale Megaptera novaeangliae were recorded and analyzed from Grand Turks in the Bahamas to Venezuela. The design features of the song are as follows. The basic song evolves through a series of different sounds in a fixed order. The song is produced only in the winter tropical calving grounds, just before the whales arrive on the banks. Redundancy is high in that syllables, motifs, phrases and the entire song are repeated. Low, intermediate, and high-frequency sounds are scattered throughout the song. One sound is associated with blowing. The song appears to be partially different each year and there are some differences within a year between banks which may indicate that dialects are present. It is suggested that songs from other populations are quite different. The apparent yearly changes do not occur at one point in time. Only single individuals produce the song and they are hypothesized to be young, sexually mature males. The implications of these various design features are discussed.     

Sound degradation as a distance cue in great tit (Parus major) song

Summary We report an experiment designed to test the ideas that: 1. male songbirds can use cues from the distortion of song by environmental factors (degradation) to estimate the distance of another singing male; 2. song degradation is assessed by reference to an internal standard. Great tits respond more strongly to undegraded than to degraded songs when both are played at the same amplitude and from the same position in the territory. This difference in response is shown only if the playback song is familiar to the test bird; familiar songs being those sung either by the test bird or neighbours of the test bird. We interpret these results as evidence that cues from song degradation can be used to estimate the distance of a singing conspecific and that degradation assessment is only possible if the bird has an internal representation of the song (because either it and/or a neighbour sings the song). We discuss the implications of these results for Morton's (1982) ranging hypothesis, and for the distinction between learning and performance in bird song. Our results partially support the ranging hypothesis, but question the nature of unrangeable songs sensu Morton. The finding that birds can assess the degree of degradation of songs that they do not sing, supports the idea that birds learn more songs than they sing.