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.     

Song repertoire size varies with HVC volume and is indicative of male quality in song sparrows (Melospiza melodia)

Complex birdsong is a classic example of a sexually selected ornamental trait. In many species, females prefer males with large song repertoires, possibly because repertoire size is limited by the size of song control nuclei which reflect developmental success. We investigated whether song repertoire size was indicative of brain area and male quality in song sparrows (Melospiza melodia) by determining if repertoire size was related to the volume of song control nucleus HVC, as well as several morphological, immunological and genetic indices of quality. We found that males with large repertoires had larger HVCs and were in better body condition. They also had lower heterophil to lymphocyte ratios, indicating less physiological stress and a robust immune system as measured by the number of lymphocytes per red blood cell. Song repertoire size also tended to increase with neutral-locus genetic diversity, as assessed by mean d2, but was not related to internal relatedness. Our results suggest several mechanisms that might explain the finding of a recent study that song sparrows with large song repertoires have higher lifetime fitness.     

A test for repertoire matching in eastern song sparrows

Repertoire matching occurs when one songbird replies to another with a song type that the two birds share. Repertoire matching has previously been demonstrated to occur at well above chance levels in a western population of song sparrows, where it is hypothesized to serve as a low level threat in a hierarchy of aggressive signals. Here we test for repertoire matching in an eastern population of song sparrows. Previous work indicates that this eastern population differs from the western one in having lower levels of song sharing between neighboring males and in showing no association between song sharing and territory tenure. Here we confirm that males in this eastern population on average share few whole songs with their neighbors. The eastern males are familiar with their neighbors’ repertoires, as evidenced by a stronger singing response to stranger song than to neighbor song. Males in the eastern population did not repertoire match: when played an unshared song type from a specific neighbor, they did not reply with a song type shared with that neighbor more often than expected by chance or more often than in response to playback of a control song (an unshared stranger song). The results thus demonstrate a qualitative difference in vocal signaling strategies between two populations of the same species.     

Song Sharing in Neighboring and Non-Neighboring Thrush Nightingales (Luscinia luscinia) and its Implications for Communication

In many passerines, males have repertoires of different songs of which some songs are often shared with other males. Sharing of song repertoires among males can provide insights into the context in which songs were acquired and on the role of song repertoires in inter- and intrasexual communication. Here we studied repertoire sharing in male territorial thrush nightingales ( Luscinia luscinia ). We compared male vocal repertoires of the basic song components, full songs, and the sequencing of songs in a bout. The results show that males differed significantly in the size of their song repertoires but not in the size of the repertoire of basic song components. Moreover, they shared almost all (80%) the repertoire of song components but only 30% of their song types. Neighboring males shared significantly more song types than did non-neighboring males but interestingly they did not share more basic song components than non-neighboring males. These results show that the repertoire of basic song components is under much less sexual selection than the size of song repertoires. Sharing of song repertoires among neighbors presumably results from repertoire conversion over time and from males returning to their territories in the following season. Repertoire sharing then can be an indicator of territory tenure and thus it can be important in repelling rivals and in female choice.     

Element repertoire: change and development with age in Whitethroat Sylvia communis song

Song repertoires are often important determining factors in sexual selection. In several species, older males have larger repertoires than 1-year-old males. The development of large song repertoires by an individual is, however, poorly understood. We studied song element repertoire changes in five individual male Whitethroats Sylvia communis sampled as 1- and 2-year olds. These males increased the size of their element repertoire between their first and second year, but song length and number of different elements per song did not change. On average, 44.3% of the song elements in the first-year repertoire were also found in the second-year repertoire. Elements shared between years were found earlier in the songs and tended to occur in sequences. Sequences of shared elements also seemed to be conserved between years. The study suggests that the song element repertoire of the second year is partly based on the first-year repertoire, which may explain why large song repertoires are mainly expressed by males at least 2 years of age. It would appear, therefore, that song element repertoire size could be a reliable signal of male age.     

Species‐level repertoire size predicts a correlation between individual song elaboration and reproductive success

Birdsong has long been considered a sexually selected trait that relays honest information about male quality, and laboratory studies generally suggest that female songbirds prefer larger repertoires. However, analysis of field studies across species surprisingly revealed a weak correlation between song elaboration and reproductive success, and it remains unknown why only certain species show this correlation in nature. Taken together, these studies suggest that females in numerous species can detect and prefer larger repertoires in a laboratory setting, but larger individual repertoires correlate with reproductive success only in a subset of these species. This prompts the question: Do the species that show a stronger correlation between reproductive success and larger individual repertoires in nature have anything in common? In this study, we test whether between‐species differences in two song‐related variables—species average syllable repertoire size and adult song stability over time—can be used to predict the importance of individual song elaboration in reproductive success within a species. Our cross‐species meta‐analysis of field studies revealed that species with larger average syllable repertoire sizes exhibited a stronger correlation between individual elaboration and reproductive success than species with smaller syllable repertoires. Song stability versus plasticity in adulthood provided little predictive power on its own, suggesting that the putative correlation between repertoire size and age in open‐ended learners does not explain the association between song elaboration and reproductive success.     

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.     

Immediate and long-term effects of testosterone on song plasticity and learning in juvenile song sparrows

Steroid sex hormones play critical roles in the development of brain regions used for vocal learning. It has been suggested that puberty-induced increases in circulating testosterone (T) levels crystallize a bird's repertoire and inhibit future song learning. Previous studies show that early administration of T crystallizes song repertoires but have not addressed whether new songs can be learned after this premature crystallization. We brought 8 juvenile song sparrows (Melospiza melodia) into the laboratory in the late summer and implanted half of them with subcutaneous T pellets for a two week period in October. Birds treated with T tripled their singing rates and crystallized normal songs in 2 weeks. After T removal, subjects were tutored by 4 new adults. Birds previously treated with T tended toward learning fewer new songs post T, consistent with the hypothesis that T helps to close the song learning phase. However, one T-treated bird proceeded to learn several new songs in the spring, despite singing perfectly crystallized songs in the fall. His small crystallized fall repertoire and initial lag behind other subjects in song development suggest that this individual may have had limited early song learning experience. We conclude that an exposure to testosterone sufficient for crystallization of a normal song repertoire does not necessarily prevent future song learning and suggest that early social experiences might override the effects of hormones in closing song learning.     

Interspecific Differences in the Response of Female Birds to Song Repertoires

The functional significance of the wide variation between bird species in the sizes of individual song repertoires is not understood. We have studied the effects of song repertoires on females. Song triggers copulation solicitation display in female sparrows treated with estradiol. Song sparrow males (Melospiza melodia) have repertoires of about 10 song types, and female song sparrows display significantly more to presentations of large repertoires than to a single song type. By contrast, male field sparrows (Spizella pusilla) and white-throated sparrows (Zonotrichia albicollis) each have only one song type, and their females show no significant increment in responsiveness to repertoires of several song types over a single song type. Swamp sparrows (Melospiza georgiana) fall in between, with male repertoires of three song types. Female swamp sparrows behave in intermediate fashion, responding more to several song types than to one, but the response increment is less than in song sparrows. Thus species differences that males exhibit in song repertoires are paralleled by differences in female responsiveness to multiple song types, implicating variations in female reactivity in the evolution of song repertoires. Female song sparrows respond preferentially to repertoires programmed in eventual variety rather than immediate variety, while field sparrows and white-throated sparrows show no discrimination.     

Age-related changes in song repertoire size and song type sharing in the Whinchat Saxicola rubetra

In many oscine passerines males’ songs, the repertoire size increases with age. At the same time it often remains unknown when and where males learn new songs. To infer the Whinchat Saxicola rubetra song learning strategy, we described and catalogued song-type repertoire, revealed age differences and examined song sharing strategies among neighbouring and distant males. We recorded song vocalizations of 40 males in a limited (104 ha) study plot during four years. Whinchats produce short and discrete songs with clear intersong pauses. In total 45 song types were allocated, the individual repertoire size averaged 23.5 ± 7.6 song types (range 9–34 song types). The males’ age significantly influenced the song-type repertoire size. The second calendar year (first breeding) males had a lower repertoire size than the older males. The majority of song types were shared by less than half of males in our sample. The Jaccard similarity indexes varied from 0.5 to 0.7. We could not find a relationship between males’ song sharing and geographic distances between their nests. We assume that Whinchat males learned new songs in the local population before territory establishment.     

Food-supplementing parents reduces their sons' song repertoire size

Food-supplemented parents typically produce more offspring, as numerous experiments on vertebrate populations have shown. ‘Propagule’ (egg or neonate) size and parental care may also be affected, with implications concerning the adult quality of offspring, although few experiments have addressed whether food-supplementing one generation affects adult quality in the next. We conducted a food supplementation experiment on song sparrows (Melospiza melodia) and tested whether song repertoire size, a demonstrated indicator of male quality, differed between the adult sons of fed (food-supplemented) and unfed (non-food-supplemented) parents. Counterintuitively, fed parents produced sons with smaller adult song repertoires, who may thus be expected to contribute fewer offspring, and fewer grand-offspring, to the population. Fed and unfed parents invested equally in the total biomass of their clutches and broods, and average nestling condition was comparable, but because fed parents produced more offspring, average egg and nestling sizes were reduced. Fed and unfed parents apportioned care differently within their broods, and we suggest compensatory growth of offspring emerging from light eggs, or egg size itself, may have affected adult repertoire size. Conceivably, the conservation benefits of food-supplementing populations could attenuate over time if fed parents produce offspring of poorer quality than themselves.     

Fitness correlates of song repertoire size in free-living song sparrows (Melospiza melodia)

Models of sexual selection propose that exaggerated secondary sexual ornaments indicate a male's own fitness and the fitness of his offspring. These hypotheses have rarely been thoroughly tested in free-living individuals because overall fitness, as opposed to fitness components, is difficult to measure. We used 20 years of data from song sparrows (Melospiza melodia) inhabiting Mandarte Island, British Columbia, Canada, to test whether a male's song repertoire size, a secondary sexual trait, predicted overall measures of male or offspring fitness. Males with larger song repertoires contributed more independent and recruited offspring, and independent and recruited grandoffspring, to Mandarte's population. This was because these males lived longer and reared a greater proportion of hatched chicks to independence from parental care, not because females mated to males with larger repertoires laid or hatched more eggs. Furthermore, independent offspring of males with larger repertoires were more likely to recruit and then to leave more grandoffspring than were offspring of males with small repertoires. Although we cannot distinguish whether observed fitness variation reflected genetic or environmental effects on males or their offspring, these data suggest that female song sparrows would gain immediate and intergenerational fitness benefits by pairing with males with large song repertoires.     

Innate differences in singing behaviour of sparrows reared in isolation from adult conspecific song

Innate differences in the singing behaviour of male swamp (Melospiza georgiana) and song (M. melodia) sparrows were identified by rearing males from the egg in the laboratory under identical conditions, in complete isolation from adult conspecific song. Isolation-reared males of both species displayed several abnormal song features, including reduced numbers of notes per song, longer durations of notes and inter-note intervals, and fewer notes per syllable. Despite these and other abnormalities, many species differences emerged that matched differences in the natural singing behaviour of the two species. These included differences in song repertoire size, song duration and degrees of segmentation, numbers of notes per song, durations of notes and inter-note intervals, and several measures reflecting the organization of songs into note complexes, syllables and trills. Although learning can influence all levels of organization of the motor patterns of song in swamp and song sparrows, its contribution to the achievement of normal song behaviour appears to be most crucial at the level of the fine structure of the notes and syllables from which the songs are constructed.     

Signal Interception and the Use of Soft Song in Aggressive Interactions

Song sparrows, like many species of songbirds, produce songs of especially low amplitude during aggressive contests. Such ‘soft songs’ have been shown to be reliable signals of intention to attack in song sparrows. Low intensity is a paradoxical feature in a highly aggressive signal, in that it seems likely to make the signal less intimidating to an opponent. A hypothesis that resolves this paradox is that use of soft songs benefits a signaler by limiting the interception of his signals by third party receivers. Here, we test this hypothesis with respect to song sparrows and two classes of third party receivers, predators and conspecific males. We tested whether selection to avoid predation is responsible for use of soft song by examining whether male song sparrows increase production of soft song in an aggressive context when they also receive signals (alarm calls) that indicate a predator is present. We found that the proportion of soft song produced by male song sparrows was actually significantly lower in the predator context than in a control context, directly contradicting the prediction. We tested whether avoiding eavesdropping by conspecific males is a benefit of soft song by removing territory owners and simulating interactions on their territories using playback from two loudspeakers. We found that intrusions by third party males were more likely in trials in which the simulated owner countered an intruder's songs using soft songs than in trials in which the simulated owner countered with loud song, again directly contradicting the hypothesis. We conclude that limiting interception by predators and conspecific males does not provide an explanation for use of soft song by song sparrows.     

Multiple signaling functions of song in a polymorphic species with alternative reproductive strategies

Vocal traits can be sexually selected to reflect male quality, but may also evolve to serve additional signaling functions. We used a long‐term dataset to examine the signaling potential of song in dimorphic white‐throated sparrows (Zonotrichia albicollis). We investigated whether song conveys multifaceted information about the vocalizing individual, including fitness, species identity, individual identity, and morph. We also evaluated whether song traits correlate differently with fitness in the two morphs, as the more promiscuous strategy of white, relative to tan, morph males might impose stronger sexual selection. Males with high song rates achieved higher lifetime reproductive success, and this pattern was driven by white morph males. In addition, males that sang songs with many notes survived longer, but this pattern was less robust. Thus, song traits reflect differences in fitness and may more strongly affect fitness in the white morph. Song frequency was unrelated to fitness, body size, or morph, but was individual specific and could signal individual identity. Songs of the two morphs displayed similar frequency ratios and bandwidths. However, tan morph males sang songs with longer first notes, fewer notes, and higher variability. Thus, song could be used in morph discrimination. Variation in frequency ratios between notes was low and could function in conspecific recognition, but pitch change dynamics did differ between four different song types observed. Our results support a multiple messages model for white‐throated sparrow song, in which different song traits communicate discrete information about the vocalizing individual.     

Song matching in a long-lived,sedentary bird with a low song rate: The importance of song type,song duration and intrusion

Territorial songbirds often match the song features or singing patterns of rivals, commonly as an aggressive signal. Most studies of song matching have been on Northern Hemisphere species with short lifespans and high song rates, but vocal matching is predicted to be affected both by longevity and territorial stability. We studied song matching in males of the white-browed scrubwren, Sericornis frontalis, a long-lived, sedentary, territorial Australian songbird. We quantified natural song rate and diversity, and then conducted three playback experiments to test: (a) whether males match by song type; (b) how they respond physically and vocally to territorial intrusion; and (c) whether they match by song length, and use it as an agonistic signal. Males naturally had very low song rates, singing on average less than three times per hour, and moderate repertoires, with an estimated mean of 17.5 song types for individual males. Males did not engage in extended counter-singing bouts. The first experiment showed that males matched the song type of immediate neighbours almost 90% of the time, if that type was in their repertoire. The remaining experiments revealed that song-type matching was an aggressive signal; males responded more aggressively to, and were more likely to match, playback simulating a neighbour's territorial intrusion than song from their shared boundary. Males did not match songs by length, but they produced longer songs after simulated intrusion. Males also responded more aggressively to playback of longer songs that simulated intrusion, but less aggressively to longer songs from the territory boundary. Overall, we show that sedentary, long-lived songbirds with low song rates, can use song-type matching as an aggressive signal to communicate with neighbours and intruders. Song length had a different role in communication, possibly related to individual quality or territory ownership.     

Song‐type sharing in a population of Song Sparrows in the eastern United States

Male Song Sparrows (Melospiza melodia) sometimes interact with neighboring territory owners by song‐type matching or repertoire matching. In some Song Sparrow populations, levels of song sharing are high and most neighbors can interact by matching, but levels of song sharing are much lower in other populations, limiting the degree to which males can match their neighbors. One explanation for variation in sharing levels is that the importance of song‐type and repertoire matching, and therefore the extent of song sharing, varies geographically in North America, being greater in western populations than eastern populations. However, to date, two studies of eastern populations have provided conflicting evidence concerning levels of song sharing by Song Sparrows. Thus, we measured sharing of whole songs and introductory phrases of songs between males with adjacent territories in another population of Song Sparrows in the eastern United States (North Carolina), near the eastern and southern limits of the species’ breeding range. Males (N = 17) in our study shared an average of only 8.7% of their song types with neighbors, and more than half of neighbor pairs shared no whole songs. Sharing of introductory phrases was more common (mean = 22.8%). The level of whole song sharing in our study is the second lowest yet reported for any Song Sparrow population, supporting the hypothesis that sharing is generally lower in eastern than in western populations.     

Rufous‐capped Warblers Basileuterus rufifrons show seasonal,temporal and annual variation in song use

In the majority of songbird species, males have repertoires of multiple song types used for mate attraction and territory defence. The wood‐warblers (family Parulidae) are a diverse family of songbirds in which males of many migratory species use different song types or patterns of song delivery (known as ‘singing modes’) depending on context. The vocal behaviour of most tropical resident warblers remains undescribed, although these species differ ecologically and behaviourally from migratory species, and may therefore differ in their vocal behaviour. We test whether male Rufous‐capped Warblers Basileuterus rufifrons use distinct singing modes by examining song structure and context‐dependent variation in their songs. We recorded multiple song bouts from 50 male Warblers in a Costa Rican population over 3 years to describe seasonal, diel and annual variation in song structure and vocal behaviour. We found that Rufous‐capped Warbler songs are complex, with many syllable types shared both within and between males’ repertoires. Males varied their song output depending on context: they sang long songs at a high rate at dawn and during the breeding season, and shortened songs in the presence of a vocalizing female mate. Unlike many migratory species, Rufous‐capped Warblers do not appear to have different singing modes; they did not change the song variants used or the pattern of song delivery according to time of day, season or female vocal activity. Our research provides the first detailed vocal analysis of any Basileuterus warbler species, and enhances our understanding of the evolution of repertoire specialization in tropical resident songbirds.     

Female choice for song characteristics in the house finch

Song has been proposed to function in mate choice, and such a role has been demonstrated experimentally in a number of species. We presented captive female house finches, Carpodacus mexicanus, with a choice of two songs, each of which was identical except for a single parameter that had been either accentuated or minimized. To ensure that our playback tapes offered choices within the normal range of variation in house finch song, we recorded wild male finches, then created playback tapes using values for the song parameters under consideration that were above or below the mean for the local population, but within the natural range of variation. We presented females that had not previously mated with choices between songs that (1) were long or short, (2) were given at fast or slow rates, and (3) included a large or small repertoire of unique song elements. Females showed significant preferences for long songs and for songs presented at a faster rate, but there was no significant preference for large or small repertoires of unique song elements. Song length and rate each seem likely to indicate a male's energy reserves, and thus could be important sources of information for females choosing mates.