Alternative forms of song matching in song sparrows

Song matching, replying to a song with a similar song, occurs in many songbird species. Almost all investigations of song matching have been of type matching, where one bird's reply is unambiguously similar to the other's song (i.e. the same song type). In many populations, however, neighbours do not share song types, and therefore cannot type-match. We hypothesized that a bird lacking a true type match could still song-match a stimulus song with a song from his repertoire that was similar in some way the birds recognized. We tested this hypothesis in song sparrows, Melospiza melodia, in two playback experiments. We played the subject a stranger song that was similar to one or more of his songs, but a type match to none of them. In the first experiment, we used playback songs that began with two buzzes (‘double-buzz’ songs). In the second experiment, we used songs that began with a slow trill that increased in tempo ('speed-up' songs). Birds replied at rates significantly above chance with their own double-buzz, or speed-up song match to the respective types of playback. The results suggest that birds who do not share true song types, can still song-match each other. This broad-sense form of song matching may also occur in populations with low song type sharing. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Species song discrimination in adult female song and swamp sparrows

Six female song sparrows (Melospiza melodia) and six female swamp sparrows (Melospiza georgiana) were treated with oestradiol, and their response to song measured by frequency and intensity of copulation-solicitation display. Test stimuli were synthetic songs composed of either song sparrow or swamp sparrow syllables assembled in either song-sparrow-like or swamp-sparrow-like temporal patterns. Female song sparrows responded preferentially to songs containing their own species' syllables and to songs containing their own species' temporal patterns. Swamp sparrows were also sensitive to both syllable type and temporal, pattern, in contrast to male swamp sparrows, which show no preference for swamp sparrow temporal patterns.     

Territory establishment,song learning strategies and survival in song sparrows

In most songbirds, the processes of song learning and territory establishment overlap in the early life and a young bird usually winds up with songs matching those of his territorial neighbors in his first breeding season. In the present study, we examined the relationships among the timing of territory establishment, the pattern of song learning and territorial success in a sedentary population of song sparrows (Melospiza melodia). Males in this population tend to learn their songs from their neighbors and consequently they show high song sharing with neighbors and use these shared songs preferentially in interactions with them. Males also show significant variation in the timing of territory establishment, ranging from their natal summer to the next spring. Using a three-year dataset, we found that the timing of territory establishment did not systematically affect the composition of the song repertoire of the tutee: early establishers and late establishers learned equally as much from their primary tutors and had a similar number of tutors and similar repertoire sizes, nor did timing of territory establishment affect subsequent survival on territory. Therefore, the song-learning program of song sparrows seems versatile enough to lead to high song sharing even for birds that establish territories relatively late.     

Role of gene regulation in song circuit development and song learning

The songbird has emerged as an important model for study of brain-behavior relationships by virtue of its rich natural advantages and from the pioneering efforts of explorers using anatomical and behavioral approaches. Now, molecular biology is providing a new and complementary paradigm for discerning songbird brain organization and function. Here, I review the work over the last 10 years that has laid the foundation for approaching songbird biology from the molecular perspective. As a result of this work, specific hypotheses can now be framed and tested regarding the mechanisms behind song circuit formation, behavioral plasticity, and the boundaries of adaptability. Age-related changes in more than 15 molecules have been observed in the song system of juvenile zebra finches, and these changes seem to define specific phases in circuit development. In adult songbirds, ordinary song-related activities such as singing and listening cause dramatic increases in gene expression in brain areas specific to each activity. The sensitivity of gene activation is modulated as a result of experience in adulthood and also changes during juvenile song learning. These studies have provided unexpected insights into the functional organization of the song circuit and the potential role of extrinsic modulatory systems in directing and limiting plastic change in the brain. With this rich base of knowledge, and techniques of gene manipulation on the horizon, answers to old questions seem within our reach: What sets the boundaries of neural plasticity? What limits learning? © 1997 John Wiley & Sons, Inc. J Neurobiol 33: 549–571, 1997     

Territorial song and song neighbourhoods in the Scarlet Rosefinch Carpodacus erythrinus

Throughout the range of the Scarlet Rosefinch, its territorial song consists of 3–9 (usually 4–5) elements, of which there are 5 different types. The differences lie in the way the pitch of the element changes in time (frequency "slope") and the width of the frequency band. Within a given type of song, the various elements can be present in almost any combination. Therefore, so many song types can be formed that the songs in even small parts of the species' area are clearly distinct from one another. Despite this capacity for variation, however, by chance identical songs may be sung in widely separated parts of the area, in some cases by different subspecies.
The species has not developed large-scale dialects or regiolects based on a song tradition acquired during an early imprinting phase. Scarlet Rosefinches tend to breed in small colonies, groups of up to about 15 pairs characterized by the same type of song (song neighbourhoods, formed by the development of a microlect).
Microlects develop by a founder effect. When males, near one-year old or older, join one another to form isolated colonies after arrival in the breeding region, they adopt ("learn") the song type that will eventually characterize the colony from the first male to arrive at the site. After the colony has been founded, in most cases each male uses only one type of song during a breeding season, with practically no variation of the temporal and frequency parameters.
Singing the same type of song, the members of a colony accept one another sufficiently to allow the breeding territories to be closely packed. It appears that a long-lasting capacity for acoustic learning, in combination with colony-like breeding and great ecological flexibility, has allowed the Scarlet Rosefinch to become the most successful species of the genus Carpodacus .     

Seasonal changes in avian song control circuits do not cause seasonal changes in song discrimination in song sparrows

In seasonally breeding songbirds, brain nuclei of the song control system that act in song perception change in size between seasons. It has been hypothesized that seasonal regression of song nuclei may impair song discrimination. We tested this hypothesis in song sparrows (Melospiza melodia), a species in which males share song types with neighbors and must discriminate between similar songs in territorial interactions. We predicted that song sparrows with regressed song systems would have greater difficulty in discriminating between similar songs. Sparrows were housed either on short days (SD) and had regressed song circuits, or were exposed to long days and implanted with testosterone (LD+T) to induce full growth of the song circuits. We conducted two experiments using a GO/NO-GO operant conditioning paradigm to measure song discrimination ability of each group. Birds learned four (experiment 1) or three (experiment 2) pairs of song types sequentially, with each pair more similar in the number of shared song elements and thus more difficult to discriminate. Circulating T levels differed between the SD and LD+T groups. The telencephalic song nuclei HVc, RA, and area X were larger in the LD+T birds. The two groups of sparrows did not differ, however, in their ability to learn to discriminate between shared song types, regardless of the songs' similarity. These results suggest that seasonal changes in the song control system do not affect birds' ability to make difficult song discriminations.     

Quality of song learning affects female response to male bird song

Bird song is unusual as a sexually selected trait because its expression depends on learning as well as genetic and other environmental factors. Prior work has demonstrated that males who are deprived of the opportunity to learn produce songs that function little if at all in male-female interactions. We asked whether more subtle variation in male song-learning abilities influences female response to song. Using a copulation solicitation assay, we measured the response of female song sparrows (Melospiza melodia) to songs of laboratory-reared males that differed in the amount of learned versus invented material that they included and in the degree to which learned material accurately matched the model from which it was copied. Females responded significantly more to songs that had been learned better, by either measure. Females did not discriminate between the best-learned songs of laboratory-reared males and songs of wild males used as models during learning. These results provide, to our knowledge, a first experimental demonstration that variation in learning abilities among males plays a functionally important part in the expression of a sexually selected trait, and further provide support for the hypothesis that song functions as an indicator of male quality because it reflects variation in response to early developmental stress.     

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.     

Is song special?

Akutagawa and Konishi (2001)([this issue of Neuron]) describe the spatial and temporal pattern of SNAg (song system nuclear antigen) expression within a subset of song-associated forebrain nuclei of grass finches. The timing and estrogen inducibility of SNAg expression suggest that it may function in establishing neural connections key to vocal learning.     

Seasonal changes in song nuclei and song behavior in Gambel's white-crowned sparrows

In males of several songbird species, the morphology of forebrain nuclei that control song changes seasonally. The only seasonally breeding songbird in which seasonal changes in the structure of song control nuclei have been reported not to occur is the nonmigratory Nuttall's subspecies of white-crowned sparrow. In the present study, we manipulated photoperiod and plasma testosterone concentrations in captive male white-crowned sparrows of the migratory Gambel's subspeices. Males exposed to photoperiods and plasma testosterone concentrations typical of those experienced by wild breeding males had larger song control nuclei than males held on a winter photoperiod. We also found seasonal change in stereotypy of spectral and temporal parameters of song in wild Gambel's white-crowned sparrows. We hypothesize that seasonal changes in song control nuclei may correlate with seasonal changes in song stereotypy. © 1995 John Wiley & Sons, Inc.     

Nightingales in space: song and extra-territorial forays of radio tagged song birds

Summary Most temperate zone passerines defend territories during the breeding season. Commonly the size of these territories is estimated by plotting the singing locations of the males. However, an individual's activities need not be restricted to the area used for singing. So far, only little quantitative information has been available to determine how the singing territory relates to the activity range of male songbirds. Here, we present a study in which we used radio-tracking techniques to collect quantitative data on the spatial behaviour of 11 male territorial Nightingales (Luscinia megarhynchos). The results show that the singing territories made up only about 50% of the activity ranges. Nevertheless, males spent over 90% of the time within the singing territory. Singing territories almost never overlapped but activity ranges overlapped in all cases with the activity range of at least one neighbour. Males intruded into neighbouring territories throughout the season but the longest excursions were made before and during the female fertile period. The time spent for forays correlated significantly with song rate and territories of males with higher song rates in turn were less frequently the object of forays by other males. Song rate can be indicative of male quality, so that our findings strongly suggest that foray behaviour is related to male quality in nightingales.

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Das Raum- und Gesangsverhalten von männlichen Nachtigallen in der Brutsaison: radiotelemetrische Untersuchungen

Zusammenfassung Männliche Singvögel markieren mit ihrem Gesang ein Revier. Empirische Studien zeigen allerdings, dass Männchen und Weibchen das Revier verlassen und in Reviere von Nachbarn eindringen. Solche Ausflüge sind vor allem in dichteren Habitaten schwierig zu beobachten, so dass für die meisten Vogelarten kaum Datenmaterial für die Raumnutzung außerhalb der Gesangsreviere vorliegt. In dieser Studie haben wir über den Verlauf der Brutsaison 11 männliche territoriale Nachtigallen telemetriert. Die Ergebnisse zeigen, dass die Aktionstradien im Mittel doppelt so groß waren, wie die Gesangsreviere und dass die Männchen ca. 8% der Zeit außerhalb der Gesangsreviere verbracht haben. Dabei überlappten die Aktionsräume benachbarter Männchen deutlich. Die Männchen drangen vor allem vor und während der fertilen Phasen der Weibchen in Nachbarreviere vor. Darüber hinaus drangen Männchen mit höherer Gesangsaktivität signifikant länger in Nachbarreviere ein, als Männchen mit geringerer Gesangsaktivität. Reviere von Männchen mit höherer Gesangsaktivität wurden dabei auch signifikant seltener von anderen Männchen besucht. Da die Gesangsaktivität mit der Qualität eines Männchens zusammenhängen kann, weisen die Ergebnisse darauf hin, dass qualitativ bessere Männchen mehr in Nachbarreviere eindringen als qualitativ schlechtere Männchen und die Reviere der qualitativ besseren Männchen auch weniger häufig von Rivalen besucht werden. Dass solche Ausflüge von Männchen für Versuche zu außerpaarlichen Kopulationen genutzt werden, ist anzunehmen, muss aber noch geklärt werden. Die räumlichen Beziehungen territorialer Männchen reflektieren darüber hinaus vermutlich Aspekte der sozialen Beziehungen.

     

Species song discrimination in choice experiments with territorial male swamp and song sparrows

Playbacks of synthetic and normal songs were used to determine what song parameters are important to species song recognition by territorial male swamp sparrows (Melospiza georgiana) and song sparrows (Melospiza melodia). Syllable morphology influences the discrimination of conspecific song from song sparrow song in adult swamp sparrows, but temporal pattern does not; this result duplicates that found in studies of song learning of the young in this species. Species song recognition is influenced by both syllable morphology and temporal pattern in adult song sparrows, in contrast to the situation in young sparrows, which seem not to distinguish their own species' song from that of swamp sparrows on either cue.     

Assessing natural dispersal of New Zealand bellbirds using song type and song playbacks

New Zealand?s managed offshore islands provide sanctuary to endangered and rare fauna but also benefit common native species. These productive islands may facilitate the expansion of mobile species back to the mainland. In northern New Zealand, many mainland protected sites are located on coastal headlands within short distances of these offshore islands. Bellbirds (Anthornis melanura), locally extinct on the mainland of this region for >100 years, are capable of dispersing these distances and are occasionally sighted along the coast. Nonetheless, it was unknown whether they had established breeding populations. Natural dispersal events are difficult to assess in terms of their source, structure and likelihood of succeeding. Females are generally more difficult to detect but when present provide conservation practitioners with confidence that a population may establish. Here we test a non-invasive monitoring method for a self-reintroduced population of bellbirds at Tawharanui, a managed coastal headland situated equal distances (20 km) from two potential source populations, Little Barrier and Tiritiri M?tangi islands (LBI and Tiri, respectively). Bellbird song playbacks effectively confirmed the presence of both male and female bellbirds. The male and female song types recorded at Tawharanui were not found on Tiri but matched those of LBI and we propose this as the source population. We tested our playback protocol at other coastal parks and advocate annual playback surveys for detecting new populations at potential mainland sites.