Changes in singing behavior of male black-capped chickadees (Parus atricapillus) following mate removal

We removed the mates of ten male black-capped chickadees (Pares atricapillus) during the nest-building period to determine the effect of female presence on dawn singing. During the first dawn chorus following mate removal, males sang significantly longer, increased movement within their territory, and increased the percentage of their territory covered while singing. After the female was returned, these parameters returned to the pre-removal values. Males did not alter the frequency range or modal frequency of their songs when the mate was removed, nor did they change the degree of frequency shifting in the fee-bee song. We conclude that dawn singing in the black-capped chickadee acts, in part, as an intersexual signal, and that the behavior of frequency shifting in the song may be directed more toward rival males than females. Correspondence to: K. Otter     

No evidence for acoustic mate-guarding in duetting buff-breasted wrens (Thryothorus leucotis)

There are few empirical tests of the acoustic mate-guarding hypothesis for the function of duetting in birds. This hypothesis states that when females are fertile, males initiate many songs or answer most of their mates solo songs to form duets and repel rival males seeking extra-pair copulations. We tested the hypothesis by comparing song initiation and answer rates of males and females in socially monogamous buff-breasted wrens (Throthorus leucotis) during pre-fertile and fertile periods. During pre-fertile periods, males often sang for short periods before being answered by their mates, yet first duets were formed earlier relative to dawn and more duets were given during the dawn chorus on pre-fertile than fertile mornings. Males initiated more songs during pre-fertile than fertile periods, whereas there was no difference between stages in female song initiation rates. The proportion of songs answered by individuals of both sexes did not differ between breeding stages. Other mate-guarding behaviours, such as frequent copulation and maintaining close proximity to mates when fertile, did not appear to be important in this species, as no copulations were observed and there was no difference in the time pairs spent in close proximity when females were fertile or not. Parentage analysis revealed that only 3% of 31 broods had young that were likely the result of extra-pair paternity. These findings do not support the acoustic mate-guarding hypothesis, and suggest that the low rate of extra-pair paternity in buff-breasted wrens was maintained without the use of acoustic or traditional paternity guards.Communicated by R. Gibson     

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.     

Song repertoires do not hinder neighbor-stranger discrimination

Summary Through playback experiments and a comparative study we tested the hypothesis that song repertoires hinder individual recognition by song in birds. Our playback tests take advantage of the peculiar singing behavior of two species of parulid warblers: the American redstart (Setophaga ruticilla) and the yellow warbler (Dendroica petechia). Males of both species sing either the same song repeatedly (repeat mode) or a repertoire of songs in series (serial mode). Subjects were played the repeat song and the serial songs from a territorial neighbor and from a stranger some territories distant. Counter to the prediction of the hypothesis, they discriminated equally well between neighbors and strangers when the several serial songs were used as when the single repeat song was played. Thus, neither the small repertoires used by redstarts in serial mode (2–8 songs) nor the larger ones used by yellows (an average of over 20 songs) appear to hinder neighbor-stranger discrimination. In our comparative test, we examined the relationship between neighbor-stranger discrimination and repertoire size among 20 species for which data were available. Using two comparative methods, we did not find the predicted significant negative relationship.     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

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.     

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.     

Energetic state and the performance of dawn chorus in silvereyes (Zosterops lateralis)

Stochastic dynamic programming (SDP) models predict that males singing to attract a mate should concentrate singing in what has been termed the dawn chorus. This is because male birds should have a variable surplus of fat in the morning that can be used to fuel singing, with the amount of fat available dependent upon such factors as his quality, foraging success and risk of predation. In this manner, the dawn chorus can act as an indicator of male quality in the context of female mate choice. We test a key prediction of SDP models of singing behaviour that males with greater fat levels should sing more. We conducted an experiment where we recorded the dawn chorus of male silvereyes (Zosterops lateralis) on three consecutive days. Each male received supplementary food on the second day, which enabled us to sample his dawn chorus before, during and after food supplementation. We also collected data on the effect of supplementary food on the body mass of silvereyes. As predicted by SDP models, we found that silvereyes sang for a greater proportion of the time after receiving supplementary food. Supplementary food also had a significant effect on the complexity of a male song, indicating that males not only increased the quantity of their song but also the quality of their song when they received extra food. As the provision of supplementary food significantly increased the mass of fed birds, our results support a causal link between male energy reserves and his ability to perform the dawn chorus.     

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.     

Dealing with urban noise: vermilion flycatchers sing longer songs in noisier territories

In noisy conditions, several avian species modulate their songs in amplitude and in the temporal or frequency domains, presumably to improve communication. Most studies on how passerine birds perform such adjustments have been carried out in oscines, a group well known for the importance of learning in the development of their songs. On the other hand, suboscines, in which learning appears to have little influence on the development of their songs, have been largely neglected. We evaluated song adjustment to noise in the vermilion flycatcher (Pyrocephalus rubinus), a suboscine bird. We conducted song recordings and noise measurements at several territories within Mexico City during the length of the dawn chorus. Males living in noisier places sang long songs, while those males inhabiting quieter places sang both short and long songs. We also found evidence of individual song plasticity, as males sang less versatile songs (i.e., songs with more introductory elements) later in the morning when noise levels were higher. This individual shift in song seems to be more associated to time of the day rather than to the observed rise in noise. However, we cannot discard an effect of noise, which should be evaluated with an experiment. We discuss our results in the context of other studies with oscine passerines and other taxa and consider implications for signaling in intra- and intersexual contexts.     

Experimental manipulation of the dawn and dusk chorus in the blackbird Turdus merula

Summary We present experimental and correlational data that demonstrate that both the nutritional state of the male and his mate's fertility affect the timing and intensity of dawn and dusk song in the blackbird (Turdus merula). Food supplementation leads to paired males singing earlier, for longer, and at higher peak rate at both dusk and dawn succeeding the treatment. We argue that the timing, not just the intensity, of song conveys information about the male's phenotypic condition. Both dawn and dusk song start earlier and have longer duration as a male's mate reaches peak fertility indicating that, in agreement with literature on the great tit (Parus major), song at this time has particular significance in mate guarding for paired males of monogamous species. We discuss the relative importance of these factors as functional explanations for the dawn chorus. Offprint requests to: I.C. Cuthill     

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.     

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.     

Geographic variation in songs of the Bewick's wren: a search for correlations with avifaunal complexity

Summary Song characteristics of the Bewick's wren (Thryomanes bewickii) are compared from nine localities in the western United States. Character shifts, i.e., a difference in means, are evident for all song characters: Arizona and Colorado songs are especially short and long, respectively (Figs. 1, 2); songs of insular (Santa Cruz Is.) and nearby mainland populations in California are very dissimilar (Table 1); excluding the insular population, the frequency range of song phrases is positively correlated with latitude (Fig. 3). Variance shifts, i.e., a difference in repertoire size or Coefficients of Variation (CV's) of measured song characters, are also present; most notably, Arizona males have exceptionally stereotyped songs, with small song phrase repertoires (Table 2) and low CV's. Population densities and/or habitat structure undoubtedly influence signal design, but correlations reported here suggest that the avifaunal complexity and the corresponding vocal milieu should also be examined rigorously as possibly important influences.     

Temporal performance roles during vocal interactions in nightingales (Luscinia megarhynchos B.)

Summary The temporal patterning of vocal interactions between territorial nightingales (Luscinia megarhynchos B). was investigated during nocturnal dyadic singing. We distinguished three forms of temporal performance roles (Table 1; Fig. 3): inserter (preferred song start 0.5–1.0 s after offset of a neighbour's song), overlapper (preferred song start 0.5–1.0 s after onset of a neighbour's song), and Autonomous bird (no evidence for temporal responses adjusted to the timing of a neighbour's singing; Fig. 4). With the beginning of the daily dawn chorus, mutual temporal adjustment of song performances could no longer be ascertained (Fig. 2). To test the flexibility of song onset timing, we presented subjects with three playback programs (I, II, III) simulating a non-flexible conspecific bird. Sequences of songs recorded from a stranger that shared no song types with the subjects were used; intersong pauses were unaltered in I, lengtened in II, and shortened in III. Results confirmed that the timing of song onset can be influenced by the temporal patterning of auditory stimuli (Table 1; Fig. 5): Inserters and overlappers sang at a slower rate during program II than during program I. During program III, which by accelerated stimulus succession caused frequent overlap of songs, inserters and overlappers interrupted their singing behaviour (Fig. 6). This suggests that overlapping, which results in signal interference, is of some functional significance in vocal interactions.     

Sexual selection and the evolution of bird song: A test of the Hamilton-Zuk hypothesis

Summary Hamilton and Zuk (1982) suggested that secondary sexual characters evolve because they allow females to assess a potential mate's ability to resist parasites. A prediction of this theory is that the degree of elaboration of secondary sexual characters should be positively correlated with parasite load across species. In support of their hypothesis, Hamilton and Zuk reported a correlation across North American passerine species between haematozoa prevalence and both brightness and song complexity and variety, scored on a subjective six point scale. Here we show that this relationship is confounded by phylogenetic associations. We use quantitative data on song duration, inter-song interval, song continuity, song rate, song versatility, and song and syllable repertoire size for 131 species of European and North American passerines to test the Hamilton-Zuk hypothesis. Across species, there are significant negative relationships between haematozoa prevalence and song continuity, contrary to the direction predicted by Hamilton and Zuk. In accordance with their prediction, there is a positive correlation with song versatility. However, these relationships come about through taxonomic associations: within taxa there are no consistent relationships between any of the song variables and haematozoa prevalence. None of the other song variables correlate with haematozoa prevalence. We conclude that there is no evidence of an association between song elaboration and parasites. Offprint requests to: A.F. Read     

Male quality,reproduction, and survival in the great tit (Parus major)

Summary Three possible measures of male quality (social dominance, song, and size), reproduction, and survival were studied in a single population of great tits. Winter dominance position on a feeder was related to strophe length (number of phrases per strophe), inversely related to positive drift (decrease of the singing rate of the phrases in a strophe), but not related to song repertoire size. Neither winter dominance position nor song were related to size (wing length, tarso-metatarsus length, weight).Singing capacity was not correlated with individual reproductive success in a single breeding season, using a rather limited data set. However, better singers (males which sing longer strophes, show less positive drift, and have larger song repertoires) survived better and had a higher individual lifetime reproductive success (on the basis of a male's recruited offspring of all breeding seasons). Our results show that there exist measurable differences whereby birds that are dominant in winter sing better, survive longer, and produce more surviving offspring during their life time. We suggest that differences in male quality are the common cause (direct and indirect) of all these effects.     

Acoustic communication in phaneropterid bushcrickets: species-specific delay of female stridulatory response and matching male sensory time window

Summary The stridulatory sounds and movements produced by the females of various bushcricket species (Tettigonoidea: Phaneropteridae: Barbitistini) are compared with those of the males. Behavioral experiments are carried out to determine the significance of the female sounds in acoustic communication. Selection factors involved in the evolution of female stridulation are discussed. The morphological apparatus for sound production has evolved independently in males and females. Whereas males rub a toothed file on the underside of the left wing over the inner edge of the right wing, the plectrum, the females stridulate by rubbing a thickened vein on the underside of the left wing over modified spines on the upper surface of the right wing (Fig. 2). Similarly, the movements responsible for sound production are not homologous in males and females. In the male the audible closing movement is always preceded by wing opening, whereas the female in general initiates the closing movement when the wings are in the resting slightly opened position, and abruptly produces complete closure (Fig. 3, 4 and 5). The female responds to male singing by emitting one to several highly damped sound pulses each lasting less than one ms. The interval between the song of a conspecific male and this response is a very precise species-specific characteristic (Fig. 7). In species with male songs that are complicated in structure or continuous, the females respond only at specific time-after patticular markers in the song of the male. The time interval between male song and female response is an important criterion by which the male identifies conspecific female song (Fig. 8). Because the response delay of the female and corresponding neuronal time window in the male are distinct they may be important in species discrimination.     

Role-reversal in katydids: Habitat influences reproductive behaviour (Orthoptera: Tettigoniidae,Metaballus sp.)

Summary Mating behaviour of the katydid Metaballus sp. varies. At sites QS and DB (in 1982) females competed for access to calling males and males chose mates by mating with heavier, more fecund females. At another site (BR) there was no evidence of role-reversal in reproductive behaviour, and males were observed to compete for mates. This species has a large spermatophore, a product of male reproductive glands, that is eaten by the female after mating. Males at the DB site had small reproductive glands. This suggests that some aspect of the QS and DB environments decreases spermatophore production; spermatophores become a limiting resource for females resulting in the reversal in reproductive roles observed at these sites. A field experiment that involved moving individuals from site BR to QS in 1983 determined that mating system was influerced by site (Table 1). At BR, males produced a continuous calling song, a third of the males observed attracted mates, and called for about 30 min before the female arrived; courtship duration was short. Males that were moved from BR to QS encountered a higher density of receptive females as all males attracted females after an average of just 3 min of calling. They changed their behaviour by producing short periodic bursts of song (zipping), and by courting females for long periods of time. The long courtship period may function as as a mate-assessment period for males. The reproductive behaviour of BR males moved to QS differed from that of native QS males only in the length of time spent in copula.     

Behavioral correlations with aberrant patterns in humpback whale songs

Summary Humpback whales emit long, patterned vocalizations referred to as songs. We analyzed songs from 22 recording sessions whick took place from January to April, 1980 on Silver Bank, Dominican Republic. The sequence of patterns (which we call themes) in these songs is remarkably regular. In this sample, there were 427 transitions from one theme to another. Only 20 of these did not follow the expected sequence. We present evidence that some themes are sung out of their usual order far more often than others. We also show that when themes are sung out of sequence, the whale is at the surface, blowing. This finding provides a link between the song structure and humpback behavior. Because aspects of the song structure can be linked to breathing, we hypothesize that a function of the songs is to show how long a whale can hold its breath, and that the duration of a song may be an indication of stamina and physical condition.