Vocal interactions in common nightingales (<Emphasis Type="Italic">Luscinia megarhynchos</Emphasis>): males take it easy after pairing

Seasonal patterns of bird song have been studied intensively with a focus on individual males. However, little is known about seasonal patterns of singing during vocal interactions between males. Vocal interactions have been shown to be important in sexual selection as males may signal aspects of motivation or quality. Here, we investigated in nightingales (Luscinia megarhynchos) whether a male’s behaviour in vocal interactions at different stages of the breeding season is influenced by its mating status. We examined how males that differ in their subsequent mating success respond to a non-interactive, nocturnal playback presented during the period of mate attraction and subsequently during the egg-laying period. We found that mated males overlapped fewer songs and had a lower song rate during the egg-laying period compared to their responses during the mate-attraction period, whereas unpaired males did not vary in their responses between the two periods. Our results suggest that mating status is a key factor affecting singing behaviour in vocal interactions and that a time-specific singing pattern like song overlapping is used flexibly during vocal interactions. Because song overlapping is thought to be a signal of aggression in male–male vocal interactions, it seems that males vary the level of aggression in vocal interactions according to their mating status and to the stage in the breeding season.     

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.     

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.     

How a songbird with a continuous singing style modulates its song when territorially challenged

In songbirds of the temperate zone, often only males sing and their songs serve to attract females and to deter territorial rivals. In many species, males vary certain aspects of their singing behavior when engaged in territorial interactions. Such variation may be an honest signal of the traits of the signaler, such as fighting strength, condition, or aggressive motivation, and may be used by receivers in decisions on whether to retreat or to escalate a fight. This has been studied intensively in species that sing discontinuously, in which songs are alternating with silent pauses. We studied contextual variation in the song of skylarks (Alauda arvensis), a songbird with a large vocal repertoire and a continuous and versatile singing style. We exposed subjects to simulated territorial intrusions by broadcasting conspecific song and recorded their vocal responses. We found that males sing differently if they are singing spontaneously with no other conspecific around than if they are territorially challenged. In this last case, males produced lower-frequency syllables. Furthermore, they increased the sound density of their song: they increased the proportion of sound within song. They seem to do so by singing different elements of their repertoire when singing reactively. Furthermore, they increased the consistency of mean peak frequency: they repeated syllable types with less variability when singing reactively. Such contextual variation suggests that skylarks might use low frequencies, sound density, and song consistency to indicate their competitive potential, and thus, those song features might be important for mutual assessment of competitive abilities.     

Effects of different levels of song overlapping on singing behaviour in male territorial nightingales (Luscinia megarhynchos)

In signalling interactions, animals can directly address information to a specific individual. Vocal overlapping is such a signalling strategy used in songbirds, anurans, and insects. In songbirds, numerous studies using high rates of song overlap to simulate an escalating situation have shown that song overlapping is perceived as a threatening signal by interacting and by listening (eavesdropping) individuals, indicating a high social relevance of song overlapping. Here we present a playback experiment on nocturnally singing male territorial nightingales (Luscinia megarhynchos). Using three different rates of song overlap (1, 25, or 50%), we tested whether or not lower levels of song overlapping act as a signal of aggressive intent and if birds would increase the intensity of their response with increasing level of song overlapping. Subjects did not vary song duration in response to the different playback treatments but increasingly interrupted their singing with increasing overlap by the three playback treatments. The effects persisted even after the playback ceased to overlap and switched to an alternating singing mode. These results expand on previous studies by showing that song overlapping is interpreted as an aggressive signal even when it is used at low or moderate levels. They suggest that, within the range tested here, increasing levels of song overlapping are perceived to be increasingly aggressive.     

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

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

Repertoire sharing and song-post distance in nightingales (Luscinia megarhynchos B.)

Summary The relationship was studied between song-post distances and the extent of vocal repertoire sharing in 34 territorial nightingales settling in six homogeneously structured habitats. Repertoires were compared on the basis of shared song types and distances were measured between nocturnal song posts of first-order and higher-order neighbours. Our results showed that male nightingales shared fewer song types with very close and more distant neighbours than with neighbours at intermediate distances. This distribution is explained by the interaction betwee repelling and attracting components, which depends on the distance between song posts.     

Heterospecific song matching in two closely related songbirds (Parus major and P. caeruleus): great tits match blue tits but not vice versa

To date, song research has focused primarily on the interactions of conspecifics. However, frequent interactions of songbirds with heterospecifics may necessitate adequate communication outside the species boundary. In this study, we focus on heterospecific communication behaviour of two small sympatric congeneric passerines, great and blue tits (Parus major and Parus caeruleus), which breed in overlapping territories and compete for food and nesting cavities. By means of a first playback experiment, we show that (1) heterospecific matching (imitating songs of the other species) is a strategy frequently used by great tits but not by blue tits, (2) both blue tit trilled and untrilled song can be accurately matched by great tits and that (3) almost half of the great tits in our study population match at least one blue tit song across all studied breeding stages, indicating that this heterospecific matching behaviour is a common feature in this population. A second playback experiment showed that these great tit imitations of blue tit songs do not function in intraspecific communication between male great tits. Hence, these heterospecific imitations appear to be designed for interspecific communication with blue tits. These findings suggest a strong heterospecific influence on the vocal learning process, repertoire composition and repertoire use in great tits and provide a possible mechanism that can drive song convergence in songbirds.     

Song rates of dark-eyed juncos do not increase when females are fertile

Frequency of singing by birds may vary with reproductive stage in ways that reflect variation in the functions of song in intersexual and intrasexual communication. In dark-eyed juncos (Junco hyemalis) high-amplitude song is produced only by males. To investigate the function of this song, we tested whether fertility of females affected singing by their mates or by neighboring males. Using focal observations, song censuses, and radiotracking data, we determined whether song production varied between and among periods when females were fertile and non-fertile. Our findings show that males do not increase song production when their mates are fertile, nor do they increase song production when neighboring females are fertile. These results suggest that male juncos do not signal their intent to defend territories (or mates) more when females are fertile and that they do not use song to advertise to specific potential participants in extra-pair fertilizations. Received: 13 February 1997 / Accepted after revison: 2 May 1997     

Song amplitude and body size in birds

Bird song is a sexually selected multidimensional signal. A fundamental question regarding the evolution of sexually selected signals is what information they convey and how their honesty is maintained. Song amplitude is a performance-related signal trait that varies considerably between individuals, but this signal dimension has been neglected in past studies. I found that median song amplitude in male nightingales (Luscinia megarhynchos) and zebra finches (Taeniopygia guttata) did not vary significantly with body size or residual body mass. In contrast, I found a significant negative correlation between body size (and also residual mass) and the maximum song amplitude during interactive singing in nightingales. However, the function of these more subtle differences in song amplitude remains to be investigated. By and large, the results of this study suggest that mean song amplitude is unlikely to indicate a bird’s body size or current condition (measured as residual mass).     

Reproductive strategy and singing activity: blue tit and great tit compared

The costs and benefits of bird song are likely to vary among species, and different singing patterns may reflect differences in reproductive strategies. We compared temporal patterns of singing activity in two songbird species, the blue tit (Cyanistes caeruleus) and the great tit (Parus major). The two species live side by side year round, and they have similar breeding ecology and similar rates of extra-pair paternity. However, they differ in two aspects of reproductive strategy that may have an influence on song output: blue tits are facultatively polygynous and have a fairly short breeding season with almost no second broods, whereas great tits are socially monogamous but more commonly raise second broods. We found that great tit males continued singing at high levels during the egg-laying and incubation periods, while monogamously paired blue tit males strongly reduced singing activity after the first days of egg-laying by their female. Since males of both species sang much more intensely shortly before sunrise than after sunrise, at midday or in the evening, this difference was most conspicuous at dawn. No differences in singing activity were found within species when testing for male age. We suggest that in contrast to blue tits, great tit males continued singing after egg-laying to defend the territory and to encourage the female for a possible second brood.     

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.     

Do male barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) experience a trade-off between the expression of multiple sexual signals?

Some sexual selection models envisage exaggerated male secondary sexual characters to be costly and therefore reliable indicators of the quality of potential mates to choosy females. If male secondary sexual characters have a natural selection cost, they may be linked to each other by reciprocally constraining relationships that would prevent individual males from increasing their level of multiple signaling. Barn swallow (Hirundo rustica) males have at least two costly signals relevant to socio-sexual interactions: tail length and song. Under the hypothesis that a trade-off exists between male signals, we manipulated the maintenance cost of tail ornaments to test whether this reduced the quantity and quality of song, a condition-dependent, phenotypically plastic signal. Contrary to our expectation, tail elongation had no effect on singing activity and song complexity. However, tail-elongated males produced songs with longer terminal parts ('rattles'). Long rattles are associated with highly competitive social contexts and high circulating levels of testosterone, suggesting that tail-elongated males were more frequently involved in either male-male aggressive or inter-sexual interactions. Therefore, this study shows that males are not displaying multiple signals at the maximum possible level, implying that this system is open to unreliable communication. However, long-term trade-offs between signal expression and viability may prevent males from displaying both signals at higher rates.Communicated by: M. Webster     

Discrimination of vocal performance by male swamp sparrows

In aggressive communication, the interests of signalers and receivers are directly opposed, presenting a challenge to the maintenance of reliable signaling. Index signals, whose production is constrained by physical ability, offer one solution to the reliable signaling problem. Vocal performance, the ability to produce physically challenging songs, is likely such a signal in swamp sparrows. Maximum vocal performance varies between males and is correlated with aspects of quality. However, vocal performance can be modulated in aggressive contexts by increasing the frequency bandwidth and trill rate of songs. This study examines receiver response to (1) differences in performance of the same song types by different signalers and (2) individual modulation of performance between contexts. Results demonstrate that male receivers show differential response to between-male differences in song type performance, but do not show differential response to the smaller scale modulations of performance produced by individuals singing the same song type at different times. This pattern suggests that vocal performance cannot be effectively cheated and may therefore serve as a good example of an index signal.     

An attractive male trait and aggressiveness are negatively correlated in wild field crickets, but uncorrelated in lab-reared crickets

It is generally thought that attractive male traits are positively correlated with dominance (aggressiveness). However, growing evidence suggests that this is not necessarily the case. We investigated whether calling song, a male mating display used by females to evaluate potential mates in a field cricket (Gryllus integer), is correlated with aggressiveness. In this species, females prefer males with longer durations of singing time. We measured singing time by measuring song over three continuous days using a custom-designed audio-monitor and assessed aggressiveness by pairing males in agonistic interactions. Our results showed that for males caught in the field, the percentage of time spent singing was negatively correlated with aggressiveness. However, in males that were raised in the laboratory, the percentage of time spent singing was not correlated with aggressiveness. Since calling duration is an attractive male trait in these crickets, our data show that the attractiveness of males can be either negatively related to aggressiveness (field) or unrelated to aggressiveness (lab). Neither of these results fit the traditional view that preferred male mating cues should be positively correlated with dominance (aggressiveness). These results also suggest that measurements of aggressiveness made in the lab do not necessarily correspond with aggressiveness in the field.     

Song length variation serves multiple functions in the white-crowned sparrow

A wide variety of hypotheses has been proposed to explain the structural diversity in bird song repertoires. Song diversity is frequently described in terms of song “types” or within-type “variants.” Male Puget Sound white-crowned sparrows, Zonotrichia leucophrys pugetensis, produce variants of their single adult song type by altering the number of repetitions of syllables in the terminal trill. We tested whether variation in trill length correlated with distance to the receiver and with signaling context as predicted by the eavesdropping avoidance hypothesis and the strategic signaling hypothesis. In accordance with the eavesdropping avoidance hypothesis, males sang variants with shorter trills, and sang quieter and less frequently when near their mate during the incubation phase than during spontaneous singing while unpaired. Males also sang variants with short trills, but at a high rate and variable amplitude when within 10 m of an opponent during close male–male territorial interactions. In agreement with the strategic signaling hypothesis, males decreased trill length immediately before chasing an opponent, but did not change length consistently prior to flight. We conclude that the occurrence of short quiet songs sung near the mate agrees with predictions of the eavesdropping avoidance hypothesis, while short songs sung near other males are best explained by the strategic signaling hypothesis. Trill length variation may be a conventional signal of aggressive intentions in male–male contests stabilized by receiver-imposed retaliation costs.     

Choice of singing sites by male bushcrickets (Tettigonia viridissima) in relation to signal propagation

Summary The spatial dispersion of singing male bushcrickets (Tettigonia viridissima) in a marshland habitat was found to be significantly clumped. Males clustered in patches of taller vegetation, buth within such clusters, males were regularly spaced with a mean distance of 6 m between nearest-neighbours. Males occupied perches on plants that were, on average, 0.3 m above the top of surrounding vegetation. Physical and acoustical interactions between males were observed more frequently when males were singing from higher sites. Excess attenuation of the male song was found to increase with frequency but decreased markedly with increasing elevation of the singing male above the ground. The maximum detection range of the song, realized when the insect was singing > 1 m above the surrounding vegetation, was estimated as 60 m for the fundamental frequency (10 kHz), 38 m for the 1st harmonic (20 kHz) and 14 m for the second harmonic (30 kHz). By contrast, when males sang from the middle of dense reed beds, the estimated detection distance was only 8 m, 6 m and 4 m for each frequency band, respectively. Males could have increased the detection range of their songs almost three fold by singing from higher positions than those usually observed in the field. This suggests that there may be a cost of singing at higher elevations such as an increased risk of predation and/or increased aggression from neighbours. We suggest the spacing strategy adopted by males reflects a compromise between maximizing the range over which their songs can be detected and accurately localized by females and minimizing interference from competing males. Offprint requests to: T. Eiriksson     

Singing performance,frequency matching and courtship success of long-tailed manakins (Chiroxiphia linearis)

We examined the relationship between song performance and courtship success in the lek-mating long-tailed manakin (Chiroxiphia linearis) of Costa Rica. Teams of male manakins form cooperative partnerships consisting of a dominant alpha male and a beta male partner with a variable number of subordinate affiliates. Each alpha and beta partnership performs a joint song virtually in unison, which attracts females to their lek arena. We quantified four aspects of song performance, including synchrony, degree of consistency in the alpha male's singing, degree of consistency in the beta male's singing, and degree of sound frequency matching between partners. When all four variables were included in a discriminant function analysis, average frequency matching alone was useful in discriminating between teams with high rates of visitation by females and those with low visitation rates. Only frequency matching was correlated with the rate at which females visited display lek arenas. We propose that the degree of frequency matching provides an unambiguous ideal against which dual-male courtship performance can be assessed. This aspect of song performance may provide information useful to females in assesing a potential mate's ability to form a cooperative long-term partnership.     

Seasonal variation in the duetting behaviour of rufous-and-white wrens (<Emphasis Type="Italic">Thryothorus rufalbus</Emphasis>)

Seasonal variation in animal signalling behaviour has been well documented and has contributed much to our understanding of male signals. In contrast, we know little about seasonal variation in female signals or signals produced jointly by males and females, such as the vocal duets of birds. Here, we examine how singing behaviour changes in relation to time of year and breeding stage in rufous-and-white wrens (Thryothorus rufalbus), neotropical songbirds where both males and females sing and where breeding partners coordinate songs to produce vocal duets. We recorded a colour-marked population of birds over an extended time period encompassing multiple breeding stages. Across all time frames and breeding stages, males sang at higher rates than females and male solos were more common than duets or female solos. Males and females showed divergent seasonal patterns of singing. Females sang more often early in the year, during the pre-breeding season, and female song tapered off as the breeding season progressed. Duetting followed a parallel pattern, which resulted from females showing less duet responsiveness to their partner’s songs later in the year. Male independent song rate peaked at the onset of the rainy season – a time when females become fertile – and males showed the highest level of duet responsiveness during this period. Our results suggest that early in the year, duets appear to be cooperative displays, functioning in joint territory defence and/or the coordination of breeding activities. When females are fertile, however, increased duet responsiveness by males is consistent with mate or paternity guarding.     

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.