Phonology and Geographic Song Discrimination in Song Sparrows

We investigated whether song sparrows discriminate foreign from local songs using specific phonologic markers, a mechanism of geographic discrimination previously described for some other songbirds. Song sparrows from Linesville, Pennsylvania (PA) respond more strongly to local songs than to songs from Millbrook, New York (NY). We identify 61 note categories in PA and NY songs, of which 13 are unique to PA, 17 are unique to NY, and 31 are shared. The most common note category in PA song, ‘buzz BO5’, is present in 89% of PA songs and only in 8% of NY songs; this difference in percentage representation is the largest we found for any note category. Substituting this potential PA marker into NY songs, however, did not make those songs more salient to PA sparrows; instead, PA males tested with territorial playback responded significantly less aggressively to NY/PA hybrid songs than to NY songs. A series of control experiments showed that song sparrows do not detect substitution of PA notes into PA songs or of NY notes into NY songs. The results weigh against the hypothesis that geographic discrimination in song sparrows is accomplished simply by recognition of a small number of phonologic markers.     

Structure and Use of Song in the Yellow-headed Blackbird (Xanthocephalus xanthocephalus)

Analysis of yellow-headed blackbird songs indicated that accenting and buzzing songs differed in note structure, energy distribution, and loudness. Buzzing songs showed a shift in energy distribution as a result of propagation through marsh habitat whereas accenting songs did not. Results suggest that accenting songs should propagate further in marsh habitat than buzzing songs. Two experiments, done to test whether song types differed in functional distance, showed that the presence of a male song of either type, or a male or female model on their territories increased use of buzzing songs by males. In the absence of auditory or visual stimulation, males used mainly accenting songs. Results supported the hypothesis that buzzing songs are used primarily for short-range interactions whereas accenting songs are used primarily for long-range general advertising.     

Soft Song in Song Sparrows: Acoustic Structure and Implications for Signal Function

In many species of songbirds, males sometimes produce songs at distinctly lower amplitude than in normal singing. Depending on the species, these 'soft songs' may be sung in the context of female courtship, male–male aggression, or both. In song sparrows, males produce soft songs during aggressive interactions with other males, and the amount of soft song produced is the only singing behavior that can be used to reliably predict a subsequent attack by the singer. Although soft song is clearly an important signal in this species, little is known about the acoustic structure of soft song or about how that structure compares to the structure of normal 'broadcast song'. We recorded a large sample of soft songs and broadcast songs from 10 male song sparrows, and measured song amplitudes in the field while controlling the subject's distance to a calibrated microphone. We show that song sparrow males produce songs over a wide range of amplitudes, with soft songs in the range of 55–77 dB sound pressure level and broadcast songs in the range of 78–85 dB. We present evidence for two types of soft song: 'crystallized' soft songs that are broadcast repertoire song types sung at low amplitude, and 'warbled' soft songs that are not found in the broadcast repertoire. Although highly variable, warbled soft songs produced by individual birds could be grouped into song types based on spectrographic similarity. To our knowledge, a distinct repertoire of soft song types has not been previously reported for any songbird.     

Incidental effects and evolution of sound-producing organs in tree crickets (Orthoptera : Oecanthidae)

Male tree cricket (Orthoptera : Oecanthidae) calling songs are used by females to locate conspecific mates. Changes in these songs are crucial to speciation. Since certain features of cricket sounds are correlated with the structures of the stridulatory organs, changes in stridulatory organs must be correlated with changes in songs. Surprisingly, there is no established hypothesis to explain the mechanism of morphological change associated with changes in songs. Some alternative possibilities are explored, with reference to incidental effects. Available evidence suggests that stridulatory organs are genetically stable and that changes in these organs occur in steps rather than gradually.     

Is the Tonal Quality of Birdsong Learned? Evidence from Song Sparrows

Song sparrow (Melospiza melodia) songs are composed largely of pure-tonal sounds. This paper investigates the role that learning plays in the development of the tonal structure of song sparrow songs, as well as the role that tonal quality plays in determining the suitability of songs as models for learning. 20 birds were trained with both normal pure-tonal songs and modified songs that included harmonic overtones. The harmonic-modified songs were obtained from birds singing in a helium atmosphere, the result of which is to perturb vocal tract resonances and thus alter a song's tonal quality. Subjects learned equally well from normal and harmonic models. Birds that learned material from harmonic models reproduced some of this material with harmonic overtones, but the majority of notes learned from harmonic models were subsequently reproduced as pure-tonal copies. Thus, the tonal structure of songs does not influence young song sparrows in their selection of song models, but there is a strong tendency to reproduce songs in a pure-tonal fashion, even if learned from harmonic models.     

Soft songs in male ortolan buntings are used in an aggressive context but are not an aggressive signal

Many bird species produce low‐amplitude acoustic signals that have been poorly studied in comparison with loud, broadcast songs used for mate attraction and repelling rivals. In some birds, these soft signals were found to be emitted in an antagonistic context and were the most reliable predictor of a subsequent physical attack. The function of this signal is poorly understood, and several hypotheses have been proposed to explain the mechanisms and possible functions of such low‐amplitude signals. The subject of this study is the ortolan bunting, a small passerine species that produces soft songs during territorial defence. In this study, we aim to study whether the soft songs of the ortolan bunting are a signal of increased aggressiveness by testing if they meet the context, prediction and response criteria of aggressive signals. We simulated stranger male intrusion into a focal male territory with three different playback experiments. We found no significant differences in the male responses to the taxidermic model regardless of whether they were or were not producing soft songs in response. The males responded more strongly to loud songs than to soft songs during the simulated intrusions, and the males did not treat soft songs as a predictor of conflict escalation. Although soft songs clearly appeared during territorial encounters and were not present during spontaneous singing before the intrusions, our results did not support the hypothesis that soft songs indicate aggressive character. We suggest that soft songs in the ortolan bunting are intentionally used by birds to modify their intentions or target‐specific individuals within a close range.     

The effects of calling song spacing and intensity on the attraction of flying crickets (orthoptera: Goryllidae: Nemobiinae)

Previous studies have shown that sexually signaling males across different taxa show stereotyped spacing behavior that may be related to aspects of their signals, such as intensity. However, few studies have shown that the separation between signaling males affects their relative attractiveness. Using two sound traps broadcasting the calling song of the cricketEunemobius carolinus, we show that the separation, relative intensity, and absolute intensity of the calling songs influence calling song attractiveness. For calling songs separated by 5 m, the proportion of individuals attracted to the higher intensity song increased as the relative intensity difference of the two songs increased from 3 to 6 dB. For calling songs that differed by 6 dB, relative attraction to the less intense song decreased with decreasing song separation. These two results are consistent with the predictions of a model (Forrest and Raspet, 1994) that suggests that dense spacing is more costly for less powerful singers and that this cost increases with increasing differences in relative intensity. When the relative intensity of the songs was held constant (6 dB), we found that discrimination between songs decreases as the song absolute intensity increases. In particular, a greater proportion of individuals was attracted to the high-intensity song when the songs were broadcast at 103 and 97 dB than when the songs were broadcast at 109 and 103 dB. Unlike mammals and birds, the ability ofE. carolinus to discriminate between songs that differ in intensity may decrease as the absolute intensity increases. This may mean that females are less discriminating when they are closer to singing males.     

Recurring patterns in the songs of humpback whales (Megaptera novaeangliae)

Humpback whales, unlike most mammalian species, learn new songs as adults. Populations of singers progressively and collectively change the sounds and patterns within their songs throughout their lives and across generations. In this study, humpback whale songs recorded in Hawaii from 1985 to 1995 were analyzed using self-organizing maps (SOMs) to classify the sounds within songs, and to identify sound patterns that were present across multiple years. These analyses supported the hypothesis that recurring, persistent patterns exist within whale songs, and that these patterns are defined at least in part by acoustic relationships between adjacent sounds within songs. Sound classification based on acoustic differences between adjacent sounds yielded patterns within songs that were more consistent from year to year than classifications based on the properties of single sounds. Maintenance of fixed ratios of acoustic modulation across sounds, despite large variations in individual sounds, suggests intrinsic constraints on how sounds change within songs. Such acoustically invariant cues may enable whales to recognize and assess variations in songs despite propagation-related distortion of individual sounds and yearly changes in songs.     

黑蚱蝉(Cryptotympana atrata Fabricius)鸣声的波谱特征

本文研究了双气囊黑蚱蝉的Click声、自鸣声和群鸣声.其鸣声波形兼有调幅与鸣声主频变化特征.1.Click声主峰频率分布在2.64—5.73KHz之间.2.自鸣声主峰频率在3.6—6KHz之间,比Click声的分布宽度略窄.3.群鸣声的主峰频率分布在4—7KHz范围内.     

Independent evolution of song structure and note structure in American wood warblers

This study addresses the issue of how evolutionary convergence within shared environments shapes some features of bird song while leaving others unaffected, using as an example the songs of 51 North American wood warblers (Parulinae). I combined published information on breeding habitats and evolutionary relationships to show that the structure of warbler songs is correlated with habitat, whereas the structure of the notes that comprise the songs is relatively unaffected by habitat and more closely related to phylogenetic history. The results confirm known relationships between bird song and habitat, including correlations between song frequency and the type and density of canopy foliage, and between the number and arrangement of notes in the song and foliage density and moisture. More importantly, the results suggest that individual notes and whole songs are to some extent functionally independent, because the configuration of notes shows more evidence of evolutionary constraint than does the way notes are assembled into songs.     

Behavioral responses of hybrid lacewings (Neuroptera: Chrysopidae) to courtship songs

The study of hybrid courtship songs and the behavioral responses of hybrids and parental individuals to hybrid songs can be useful in understanding the origin of reproductive isolation among species that differ mainly in their courtship songs. Here we test the hypotheses (a) that hybrid lacewings prefer hybrid songs to either of the parental songs from a cross betweenChrysoperla plorabunda andC. johnsoni, and (b) that parental individuals prefer their own song over those of hybrids. Analysis of songs showed that most features of hybrid songs are intermediate between the two parents. Hybrids organize their songs with a series of simple volleys like those ofC. plorabunda. Female hybrids from two reciprocal crosses and females and males of the parental species were presented with choices of hybrid and parental songs. Hybrids responded more to recordings of hybrid songs than to recordings ofC. plorabunda but did not differ in the responses given toC. johnsoni and hybrid songs. In contrast, males and females of both parental lines preferred to duet with recordings of their own song types and did not respond to hybrid songs. Our results demonstrate that hybrids would be at a disadvantage in nature, because neitherC. plorabunda norC. johnsoni will respond to their songs.     

NORTH PACIFIC HUMPBACK WHALE SONGS: A COMPARISON OF SOUTHEAST ALASKAN FEEDING GROUND SONGS WITH HAWAIIAN WINTERING GROUND SONGS

Humpback whales sing long, complex songs on their wintering grounds. On 25 August 1979 and 3 September 1981, we made recordings of humpback whale songs in southeastern Alaska, showing that humpback whales also sing on the summer feeding grounds. Both these Alaskan samples are songs in that they are repeating cyclical sound patterns and follow the known structure for humpback whale song. The Alaskan songs contain all the same material sung in the same order as that heard off Mexico and Hawaii in the surrounding wintering seasons. However, song, theme and some phrase durations are abbreviated in the Alaskan songs. The recording of these two songs represents the full sample of song recorded from 155 days over five years of attempting to record humpback whale song in Alaskan waters.     

Sexual Differences in the Responses of Captive Great Reed Warblers (Acrocephalus arundinaceus) to Variation in Song Structure and Repertoire Size

The function and evolution of repertoires in songbirds is still not clearly understood. This study involves the great reed warbler Acrocephalus arundinaceus and playback of recorded song to both captive males and females under controlled laboratory conditions. Females were implanted with oestradiol and the copulation solicitation display used as an index of sexual response. Males were not implanted and crest-raising used as an index of aggressive response. Two types of songs were used (short and long) and two sizes of repertoire (high and low). Females only responded to long songs, and responded significantly more to high than low repertoires. It was concluded that long songs have a sexual function and that high repertoires are the result of intersexual selection through female choice. Males responded to short and long songs, but did not discriminate between high and low repertoires. Adding a live male to the experiment made no difference to female responses but males responded significantly more. When field studies are also taken into consideration the overall conclusion is that song quality and repertoire size is particularly important in female choice, and that short songs and a male presence are more important in aggressive behaviour. Integration of both field and laboratory studies on males and females should lead to more rigorous testing of hypotheses concerning the function and evolution of bird songs.     

Typical versions of learned swamp sparrow song types are more effective signals than are less typical versions

The learned songs of songbirds often cluster into population-wide types. Here, we test the hypothesis that male and female receivers respond differently to songs depending on how typical of those types they are. We used computational methods to cluster a large sample of swamp sparrow (Melospiza georgiana) songs into types and to estimate the degree to which individual song exemplars are typical of these types. We then played exemplars to male and female receivers. Territorial males responded more aggressively and captive females performed more sexual displays in response to songs that are highly typical than to songs that are less typical. Previous studies have demonstrated that songbirds distinguish song types that are typical for their species, or for their population, from those that are not. Our results show that swamp sparrows also discriminate typical from less typical exemplars within learned song-type categories. In addition, our results suggest that more typical versions of song types function better, at least in male–female communication. This finding is consistent with the hypothesis that syllable type typicality serves as a proxy for the assessment of song learning accuracy.     

Whip it good! Geographic consistency in male songs and variability in female songs of the duetting eastern whipbird Psophodes olivaceus

Geographic variation in male bird songs has been studied extensively, but there have been few investigations of geographic variation in female songs or sex differences in patterns of geographic variation. We compared patterns of variation in male and female songs of eastern whipbirds Psophodes olivaceus by analyzing recordings from 16 populations across the species’ geographic range in eastern Australia. We found remarkably different patterns of geographic variation between the sexes. Female eastern whipbird songs are easily categorized into discrete song types. Song types are shared between nearby females, but female songs show pronounced differences at a continental scale. In contrast, male eastern whipbird songs show high consistency throughout the species’ geographic range. All recorded males share the ability to transpose the frequency of the introductory whistle and most recorded males share the ability to vary the direction of the slope of the terminal whip crack. For eight of nine measured variables, female songs show significantly higher levels of variation than male songs. We discuss whether sex differences in dispersal, song learning strategies, and song function may explain these sex differences in patterns of song variation. We suggest that eastern whipbirds have experienced a decoupling of male and female song learning strategies and that the songs of each sex have responded to different selective pressures in the context of countersinging interactions. We speculate that consistency in male songs throughout the geographic range of eastern whipbirds may arise through female preference for males that perform large bandwidth whip cracks.     

Imitation and Invention in Song Learning in Nightingales (Luscinia megarhynchos B., Turdidae)

Song learning in oscine birds is often defined solely as a process of song imitation; nonetheless, not all songs produced by laboratory‐tutored birds are imitations of the model songs. If song learning were strictly a process of imitation, these non‐imitated songs (inventions) would be expected to contain no learned attributes. To determine whether species‐typical song attributes can be learned in the absence of imitation, we compared the imitations and inventions of laboratory‐tutored nightingales (Luscinia megarhynchos B.) with the songs of wild nightingales and the songs of laboratory‐reared, untutored nightingales. The species‐typical song attribute measured was stereotypy. We quantified stereotypy by four variables: (1) percentage of notes shared between two renditions of the same song type (2) difference in repetition rates of the same trill in two renditions of the same song type (3) acoustic similarity of the same note in two renditions of the same song type, and (4) acoustic similarity of the same note repeated within a trill. Wild songs and imitated songs were significantly more stereotypical than the songs of untutored birds for all measures. For the percentage of notes shared (1), and the acoustic similarity of notes in two renditions of the same song type (3), invented songs did not differ from the songs of untutored birds, suggesting that imitation is necessary for the acquisition of these song characteristics. However, invented songs were significantly more stereotypical than the songs of untutored birds for measures of stereotypy related to trills (2 and 4), and neither imitated nor invented songs differed significantly from the songs of wild birds in terms of trill rate stereotypy (2). Thus, it appears that the process of learning to produce trills may differ from the process of learning non‐repetitive song components: increased stereotypy in trills occurs even when the trills themselves are not copied from song models. Strict imitation does not fully account for the acquisition of some learned song attributes.     

Complementary neural systems for the experience-dependent integration of mate-choice cues in European starlings

Choice of a particular mate phenotype may arise out of experience with the very phenotypes under consideration. Female European starlings (Sturnus vulgaris) prefer males that sing predominantly long-bout songs over males that sing predominantly short-bout songs, and thus, song-bout length is a phenotypic parameter instrumental in releasing the female's mate choice. The preferred long-bout songs induce higher expression of the immediate early gene (IEG) ZENK in the female auditory telencephalon than short-bout songs do, but this sensitivity to song length depends on the female's recent song experience. Here, we compared the experience-dependent modulation of ZENK with that of another IEG, FOS, and report that ZENK and FOS expression in the caudomedial mesopallium and caudomedial nidopallium show different modulation properties that complement natural variation in song-bout length. As reported previously, ZENK expression was greater in response to novel long-bout than to novel short-bout songs following a 1-week experience with long-bout but not short-bout songs. In contrast, FOS expression was greater in response to novel long-bout than to novel short-bout songs following a 1-week experience with short-bout but not long-bout songs. Thus, the ZENK and FOS signaling pathways are made sensitive to variation in song length by experiences with songs at opposite ends of the starling song-variation continuum, suggesting the presence of complementary neural systems made sensitive in register with the natural axis of phenotypic variation fundamental to the female's mate choice.     

Song Functions in Nonduetting Gibbons: Evidence from Playback Experiments on Javan Gibbons (<Emphasis Type="Italic">Hylobates moloch</Emphasis>)

Territorial, pair-living primates usually perform long-distance calls as duets in which adult males and females coordinate their calls. Previous studies using playback experiments have shown that gibbon duets convey information about the status of the caller (location, familiarity, sex of the caller, and paired status) and gibbons use this information to respond to achieve several nonmutually exclusive functions, including intragroup contact, territorial defense, and pair-bond advertisement and strengthening. However, not all pair-living gibbons duet, and it is unclear whether the same results should be expected in nonduetting species. We conducted song playback experiments (N = 47 trials) to test hypotheses about song functions in nonduetting gibbons on two groups of wild Javan gibbons (Hylobates moloch) in the Gunung Halimun-Salak National Park, Indonesia. Javan gibbons initiated movement toward the speaker more quickly in response to songs broadcast in the center of the territory, stranger songs, and songs of unpaired individuals than to songs at the border, neighbor songs, and songs from paired individuals. These results suggest that Javan gibbons can localize songs, and that Javan gibbon songs transmit information about the identity and paired status of the caller. Our results imply that Javan gibbon solo songs are likely to function for territorial defense and pair-bond advertisement like duets in other primates.     

Acoustic characteristics, early experience, and endocrine status interact to modulate female zebra finches' behavioral responses to songs

Female songbirds use male songs as an important criterion for mate selection. Properties of male songs are thought to indicate the male's quality as a potential mate. Song preferences in female zebra finches are known to be influenced by two factors--early auditory experience and the acoustic characteristics of males' songs. Studies often investigate song preferences by priming females with estrogen. However, estrogenic influences on song preferences have not been studied. We investigated the relative influence of early auditory experience, acoustic features of songs, and estrogen availability on song responsiveness in female zebra finches. Juvenile female zebra finches were tutored for 10 days with 40 songs per day with one of three acoustically different song types--simple songs, long-bout songs or complex songs. A fourth group of females was untutored. Aside from this brief song exposure, females were raised and maintained without exposure to male songs. During adulthood, females' behavioral responses to the three song types were tested under three hormone conditions--untreated, estradiol-treated and 1,4,6-androstatriene-3,17-dione (ATD)-treated (to lower endogenous estrogen). Based on the results of our study, four conclusions can be drawn. First, song responsiveness in female zebra finches is strongly affected by minimal early acoustic experience. Second, inexperienced female zebra finches are inherently biased to respond more to complex songs over other song types Third, although female zebra finches are inherently biased to respond more to complex songs, early acoustic experience may either reinforce or weaken this inherent responsiveness to complex songs. Fourth, estrogen selectively accentuates song responsiveness in acoustically-experienced female zebra finches.     

No Effect of Body Size on the Frequency of Calling and Courtship Song in the Two-Spotted Cricket,Gryllus bimaculatus

The relationship between body size and vocalization parameters has been studied in many animal species. In insect species, however, the effect of body size on song frequency has remained unclear. Here we analyzed the effect of body size on the frequency spectra of mating songs produced by the two-spotted cricket, Gryllus bimaculatus. We recorded the calling songs and courtship songs of male crickets of different body sizes. The calling songs contained a frequency component that peaked at 5.7 kHz. On the other hand, courtship songs contained two frequency components that peaked at 5.8 and 14.7 kHz. The dominant frequency of each component in both the calling and courtship songs was constant regardless of body size. The size of the harp and mirror regions in the cricket forewings, which are the acoustic sources of the songs, correlated positively with body size. These findings suggest that the frequency contents of both the calling and courtship songs of the cricket are unaffected by whole body, harp, or mirror size.