Female response to song reflects male developmental history in swamp sparrows

One explanation for why female songbirds attend to male song is that the quality of a male's song is associated with the quality of his developmental history. We tested this hypothesis by playing back to female swamp sparrows (Melospiza georgiana) songs recorded from males of either inferior or superior developmental histories, as assessed by their rates of mass gain during the first 18 days post-hatching. Females showed significantly higher levels of courtship display in response to songs of males with superior growth than to songs of males with inferior growth. Out of nine song traits measured, only song duration correlated with variation in female response; duration was also the only trait that differed significantly in univariate comparisons between the superior growth songs and the inferior growth songs. In a multivariate analysis, however, inferior growth songs were best discriminated from superior growth songs by combining three song traits: trill rate, stereotypy, and the number of notes per syllable. We suggest that early developmental stress degrades song in many small ways, and that it is the cumulative effect of the resulting deficits that explains lower female response.     

Asymmetric mate choice,hybridization, and hybrid fitness in two sympatric grasshopper species

Hybridization between animal species has long been regarded as unusual, but is meanwhile accepted as a widespread phenomenon. Typically, sexual interactions among species are studied in secondary contact zones of closely related species (hybrid zones) or between invasive and native species, whereas hybridization between sympatric congeners has received little attention. Here, we present a study on the hybridization potential of two grasshopper species, Chorthippus parallelus and Chorthippus montanus, which occur sympatric in large parts of Eurasia. We performed a mate choice experiment with specimens from an area of local parapatry in northwestern Germany. Most copulations were conspecific, but males of both species and females of C. parallelus were indiscriminate in their mate choice, while females of C. montanus clearly preferred conspecific males. Although these results suggest asymmetric introgression, a no-choice hybridization experiment revealed that hybridization is possible in both directions. The hatching success of the hybrids was intermediate between the parental species. Female hybrids showed no clear mate preferences, indicating that back crossing in both directions is possible in principle, but the fertility of the hybrids and the fitness of the F2 generation remain unknown. Our study suggests that hybridization between sympatric species might occur more often than assumed. It might affect the ecology and local distribution of animals in a similar way as competition does.     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

Divergence in female calling song discrimination between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae)

Two species of closely related wood cricket, Gryllus fultoni (Orthoptera: Gryllidae) and Gryllus vernalis, occur together in some parts of the eastern United States and have a similar calling song structure, consisting of three-pulse chirps. A previous study revealed that chirp rate in G. fultoni was highest (greatest difference vis-à-vis chirp rate in G. vernalis) in sympatric populations, intermediate in near allopatric populations that were located close to the sympatric zone, and lowest in allopatric populations. A similar trend was observed in pulse rate, but the mean values of this trait showed much more convergence than chirp rate at the low end of the range of calling temperatures. In this study, we investigated the song discrimination of females from sympatric and allopatric populations of G. fultoni at about 23°C, which is near the middle of the normal range of calling temperatures. We used both single-stimulus and two-stimulus playback experiments to learn if geographical differences in song preferences paralleled those in calling songs. Stimuli presented were representative of calling songs in three classes of G. fultoni populations (sympatric, near allopatric, and far allopatric), a calling song of G. vernalis, and three calling songs with parameter values that were intermediate with respect to those of the songs of far allopatric G. fultoni and G. vernalis. In the single-stimulus playbacks, females of all G. fultoni populations responded poorly if at all to the heterospecific stimulus. Females of sympatric and near allopatric populations responded poorly to all intermediate stimuli, but females of far allopatric populations frequently responded to these sounds. In the two-stimulus playbacks, females of sympatric and near allopatric populations generally discriminated against intermediate and heterospecific stimuli. However, females of far allopatric populations often did not discriminate against intermediate stimuli, whose characteristics resembled the calling songs of G. vernalis. The divergent pattern of female phonotactic discrimination between sympatric and far allopatric populations was thus generally congruent with the pattern of divergence in chirp and pulse rates and would be expected to significantly reduce heterospecific mating in sympatry. These geographical patterns of female song discrimination and male calling songs conform to a commonly used definition of reproductive character displacement.     

Testing the ability of males and females to respond to altered songs in the dueting green lacewing, Chrysoperla plorabunda (Neuroptera: Chrysopidae)

Green lacewings in the carnea group of Chrysoperla engage in species-specific heterosexual duets using low-frequency substrate-borne signals. Within each species, both sexes sing nearly identical songs. Songs are the principal barriers to hybridization between sympatric species in the complex. Here, we investigated the responsiveness of males and females of Chrysoperla plorabunda to synthesized, prerecorded songs that differed from the species mean in the period between repeated volleys of abdominal vibration. We tested 15–16 males and 15–16 females using playbacks of two signals that gradually increased or decreased in volley period, starting at the species mean. We found that (1) duets during courtship are accurate, interactive, and adjustable by each participant; (2) in staged duets, both sexes respond best to song tempos near the mean volley period of their population, but can nonetheless maintain duets with signals of nearly twice, or half, the normal volley period; (3) individuals fine-tune their adjustments to signals of different volley periods by changing their own volley duration and latent period, or less often by inserting extra volleys or skipping every other volley; (4) males are significantly better at matching signals of changing tempo than females; and (5) the range of song responsiveness of C. plorabunda does not overlap the natural range of volley periods found in Chrysoperla adamsi, an acoustically similar sibling species, thus reaffirming strong behavioral isolation. In sum, the precise, almost unbreakable heterosexual duets characteristic of song species of the carnea group result from tight mutual feedback between partners. Effective reproductive isolation between species can be based on song differences alone.     

Soft song in song sparrows: response of males and females to an enigmatic signal

Low-amplitude “soft song” is used by a variety of songbirds; in some species during aggressive encounters, in others during courtship, and yet others in both these contexts. In song sparrows (Melospiza melodia), soft song has thus far been observed only in aggressive encounters, where its production is a more reliable predictor of attack than any other signaling behavior. We used song playback to test the response of both male and female song sparrows to soft song. The design of the playback experiments took into account the existence of two classes of soft song: crystallized soft song, which consists of song types also found in the broadcast repertoire, and warbled soft song, which consists of less-structured song types not found in the broadcast repertoire. Female song sparrows responded with significantly less courtship display to the playback of crystallized soft song than to that of normal broadcast song, and response to warbled soft song was if anything lower than to that of crystallized soft song. Male song sparrows responded equally aggressively to normal broadcast song as to crystallized soft song, and equally aggressively to warbled soft song as to crystallized soft song. The female results support the conclusion that neither form of soft song functions in courtship. The male results suggest that the reliability of soft song as a signal of aggressive intent is not maintained by a receiver retaliation rule.     

Female choice for complex song in the European starling: a field experiment

Male European starlings Sturnus vulgaris sing long complex songs that appear to be important in the courtship of females but which also influence competitive interactions between males. We tested the hypothesis that females choose mates on the basis of the complexity of their songs, rather than on the quality of the territories the males defended. In order to determine whether certain territories were preferred over others, the first set of birds to settle in the experimental nest-boxes was removed and a second set allowed to settle. Consistent preferences for certain nest-boxes were indicated by correlations between the settlement patterns of the first and second sets of birds. However, males with the most complex song did not necessarily occupy the most preferred nest sites. Males with more complex song acquired mates faster. This relationship remained significant when nest-site preference was statistically controlled, indicating that female starlings chose males with complex song rather than those that defended preferred nest sites. A number of morphological variables were also found to be uncorrelated with female choice. Song complexity in European starlings increases with age, and the evolution of song complexity in this species is consistent with an age-indicator model of sexual selection. Males with larger repertoires were also in better condition, indicating that females obtain high-quality mates by choosing on the basis of male song. Received: 29 April 1995/Accepted after revision: 9 September 1995     

Local adaptation and sexual selection: a reciprocal transfer experiment with the grasshopper Chorthippus biguttulus

Immigration into locally adapted populations has been suggested, among other potential causes, to maintain genetic variance in fitness necessary for good-genes models. Using a reciprocal transplant experiment we examined whether females prefer native to transferred males in the grasshopper Chorthippus biguttulus. On average, native and transferred males did not differ in their attractiveness, measured as female response rate to playbacks of male acoustic courtship signals. In line with this result, we found no significant effect of transfer on body size, condition, fluctuating asymmetry or song traits. However, the reciprocal transplant experiment showed that environmental conditions did influence body condition and maximum loudness of the calling song, but that the genetic origin of male grasshoppers had no significant effect on any of the analysed traits.Communicated by L. Simmons     

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.     

Bushcricket song structure and predation by the acoustically orienting parasitoid fly Therobia leonidei (Diptera: Tachinidae: Ormiini)

Males of most bushcricket species produce acoustic advertisement signals to attract females for mating. These signals can also increase conspicuousness to predators. In the genus Poecilimon (Orthoptera: Phaneropterinae) males are attacked by the parasitoid fly Therobia leonidei (Diptera: Tachinidae: Ormiini) which locates males by their calls. In Greece T.leonidei parasitizes several Poecilimon species with different song structures: we examined whether host choice is related to song structure by comparing parasitism rates in two closely related Poecilimon species. One of these species produces monosyllabic songs, the songs of the other species being polysyllabic. The tachinid fly parasitized the polysyllabic species to a greater extent. We demonstrate in a field-experiment that this preference for the polysyllabic species does not depend on local adaptations of the fly. The most probable explanation for the preference of the fly for the polysyllabic singing species seems to be better detection of longer songs. This result is discussed in the context of male song evolution. Received: 4 November 1997 / Accepted after revision: 22 March 1998     

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

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

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.     

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.     

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.     

Individual recognition of neighbors by song in a suboscine bird, the alder flycatcher Empidonax alnorum

The ability of territorial males to discriminate between songs of their neighbors and songs of strangers has been demonstrated in 27 species of songbirds. Such experiments test only the ability of a subject to discriminate between two classes of stimuli, familiar (neighbors) and unfamiliar (strangers) songs. Individual recognition of neighbors is a finer, more complex type of discrimination. The ability of territorial males to recognize individual neighbors by song has been documented in 12 species of oscine passerines (Passeriformes, Passeri), but has never been demonstrated in suboscine passerines (Tyranni). We investigated recognition of songs of individual neighbors in a suboscine, the alder flycatcher (Empidonax alnorum). We performed a series of song playback experiments and recorded responses of territorial males to songs of neighbors and songs of strangers broadcast from two locations, the neighbor boundary and an opposite boundary. Subjects responded more aggressively to songs of a neighbor when played from the opposite boundary than when played from the neighbor boundary. They responded with equal aggression to songs of strangers regardless of location of playback. The difference in response to neighbor songs between speaker locations and the lack of a difference in response to stranger songs indicate that territorial males associate a particular song with a particular location (territory), and thus recognize individual neighbors.Communicated by I. Hartley     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

Function of male courtship vocalizations in red-winged blackbirds

Summary During courtship, male red-winged blackbirds (Agelaius phoeniceus) produce long sequences of simple, single-note precopulatory calls, punctuated with occasional songs. Male song has previously been shown to stimulate copulation solicitation in captive, estradiol-implanted females of this species (Searcy and Brenowitz 1988; Searcy, in press). Here I show that sequences of precopulatory calls also stimulate copulation solicitation in female redwings, but that they are not as stimulatory as song. In tests contrasting mixed bouts of song and precopulatory calls with bouts of song alone, the mixed bouts elicited significantly more female solicitation in one instance, more but not significantly more in a second, and an exactly equal amount as song alone in a third. Neither song nor precopulatory calls could be shown to affect proximity of females to the speaker.     

Duetting and mate-guarding in Australian magpie-larks (Grallina cyanoleuca)

A recently favored hypothesis is that duetting in birds has a mate-guarding function: a male responds vocally to his partner’s song, thereby forming a duet that repels males who are attracted to her song. Previous studies have not provided unambiguous tests of the mate-guarding hypothesis because: (1) the probability of a male answering his partner’s song has not been shown to increase specifically when the female is fertile, and (2) the probability of a male answering his partner’s song has not been assessed separately from simply a higher song initiation rate. We investigated extra-pair paternity, mate-guarding, and duetting in the socially monogamous Australian magpie-lark (Grallina cyanoleuca). DNA fingerprinting revealed that 3% of young were the result of extra-pair paternity, and we found that males guarded fertile females by staying close to them. However, males did not initiate songs at a higher rate when females were fertile and actually reduced their probability of replying to female song during this period. We conclude that although male magpie-larks did guard fertile females in an attempt to prevent extra-pair copulations, they did not use duetting for this purpose. Received: 10 May 1999 / Received in revised form: 27 September 1999 / Accepted: 2 October 1999     

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.