ABSOLUTE AND RELATIVE PITCH PRODUCTION IN THE SONG OF THE WHITE-THROATED SPARROW (ZONO TRICHIA ALBICOLLIS)

ABSTRACT

The songs of white-throated sparrows consist of 5 notes of very pure tonal quality. In ascending song, the pitch increases substantially from Note 1 to Note 2; whereas, in descending song, pitch decreases substantially from Note 2 to Note 3. Variability in the absolute pitch of each note is considerable among birds, but much less within individuals. Analysis of the major pitch changes in these songs shows that the pitch interval (ratio of the higher to the lower frequency), a measure of relative pitch constancy, predicts the frequency of the higher note in the pitch change more precisely than does the difference between the frequencies of the two notes, a measure of absolute pitch constancy. We conclude that white-throated sparrows produce relative rather than absolute pitch constancy during the major frequency changes in their songs and suggest that pitch interval may be an important cue in species recognition.     

MICRODIALECT AND GROUP SIGNATURE IN THE SONG OF THE SKYLARK ALAUDA ARVENSIS

ABSTRACT

The Skylark Alauda arvensis is a territorial species of open landscape in which pairs settle in stable and adjacent territories during the breeding season. Due to the heterogeneity of the habitat, territories are gathered in patches spaced by a few kilometres, in which each male produces very long and complex flight songs as a part of the territorial behaviour. We showed that, in a given patch, all the males (neighbours) share some particular sequences of syllables in their songs, whereas males settled in different patches (strangers) have almost no sequences in common. Such a phenomenon is known as microdialect. To test the hypothesis that these shared sequences support a group signature, we made playback experiments with “chimeric” signals: songs of strangers where the sequences shared by neighbours were artificially inserted. Behavioural responses to playbacks indicated a neighbour-stranger discrimination consistent with the dear enemy phenomenon, i.e. a reduced aggression toward neighbours compared to strangers. Furthermore, the same level of responses, observed when a “chimeric” song and a neighbour song were broadcast, indicated that shared sequences are recognised and identified as markers of the neighbourhood identity.     

SONG POPULATIONS AND DISPERSAL IN STEELBLUE AND PURPLE WIDOWFINCHES

Payne, R. B. 1985. Song populations and dispersal in Steelblue and Purple Widowfinches. Ostrich 56:135-146.

Songs of Steelblue Widowfinches Vidua chalybeata were recorded over eight years at Lochinvar National Park, Zambia. Neighbouring males share their song-type repertoires. Individual birds changed the structural details of all song types in their repertoire from year to year, and over 4–5 years the songs accumulated changes so that a song type could scarcely be recognized as the same. A few males in addition switched their song repertoires when they dispersed from one song population to another, or when their old neighbours disappeared and new neighbouring males sang a different song repertoire. Individual marked adult birds moved from one song neighbourhood to another. Purple Widowfinches had a similar song behaviour but only one song dialect was found in the study area.

An estimate of the proportion of immigrants was made from the proportion of birds with songs unlike their neighbours, and from birds that had a mixed song repertoire or that switched their song repertoires. Song populations were sampled at several localities in the Transvaal, Botswana, Zambia, Kenya, Tanzania, Cameroon, and Nigeria. An overall minimal estimate of dispersal between song neighbourhoods was 18 %. The incidence of observed dispersal and the movements indicated by the song differences among local widowfinches were considerable and more than sufficient to prevent genetic isoloation and differentiation of local song populations.     

CORRELATED EVOLUTION OF BEAK MORPHOLOGY AND SONG IN THE NEOTROPICAL WOODCREEPER RADIATION

Mating signals may diversify as a byproduct of morphological adaptation to different foraging niches, potentially driving speciation. Although many studies have focused on the direct influence of ecological and sexual selection on signal divergence, the role of indirect mechanisms remains poorly understood. Using phenotypic and molecular datasets, we explored the interplay between morphological and vocal evolution in an avian radiation characterized by dramatic beak variation, the Neotropical woodcreepers (Dendrocolaptinae). We found evidence of a trade-off between the rate of repetition of song syllables and frequency bandwidth: slow paced songs had either narrow or wide frequency bandwidths, and bandwidth decreased as song pace increased. This bounded phenotypic space for song structure supports the hypothesis that passerine birds face a motor constraint during song production. Diversification of acoustic characters within this bounded space was correlated with diversification of beak morphology. In particular, species with larger beaks produced slower songs with narrower frequency bandwidths, suggesting that ecological selection on beak morphology influences the diversification of woodcreeper songs. Because songs in turn mediate mate choice and species recognition in birds, these results indicate a broader role for ecology in avian diversification.     

BEHAVIOURAL RESPONSE TO SONG PLAYBACK BY MALE AND FEMALE WHITE-CROWNED SPARROWS OF TWO SUBSPECIES

ABSTRACT

Nuttall's white-crowned sparrow Zonotrichia leucophrys nuttalli and the Puget Sound white-crowned sparrow Z. i. pugetensis intergrade through a zone of hybridization in northern California. The songs of the two differ in the sequence of syllable types and in the phonology of the complex syllables. We did a playback experiment to males of both subspecies in the field and to females of both subspecies in the laboratory to determine their response to the two subspecies songs. By making alterations of features of the songs, we also attempted to identify characteristics that account for subspecific recognition. Males of both subspecies were more responsive to their own subspecies song than to that of the other subspecies. Substitution of Puget Sound complex syllables into the Nuttall's song caused a reduction in response by male Nuttall's, but alteration of a Nuttall's song to a Puget Sound sequence of syllables had no effect. Females of both subspecies gave more sexual displays when played a song of their own subspecies than when played that of the other subspecies. Female Nuttall's gave an intermediate response when played Nuttall's songs with Puget Sound complex syllables or when played Nuttall's songs altered to a Puget Sound sequence. Puget Sound females did not discriminate between their own subspecies song and either of the altered stimulus songs separately. They did respond less to a song that combined both alterations than to their own subspecies song.     

IMPROVISATION OF NEW NOTES DURING SINGING BY MALE SARDINIAN WARBLERS SYLVIA MELANOCEPHALA

ABSTRACT

The songs of Sardinian warblers Sylvia melanocephala are extremely complex and variable, and even consecutive songs uttered by the same male usually differ. To provide a better understanding of the nature of the variation recorded, an attempt has been made to determine size and composition of the note repertoire of three selected individuals. The analysis was carried out on samples each consisting of several dozen songs, and the various notes were sorted on the basis of the visual similarity of their spectrographs patterns. Each male turned out to have a very large note repertoire which does not show any overlap with those of other males, with the only exception a particular note found in the songs of all the males. In two birds out of three, however, the graphs of the occurrence of new notes in consecutive songs show no tendency to become asymptotic. This suggests that the assessment of note repertoire size in these birds is not accurate, as they continue inserting new notes in their songs even after they ha ve uttered several dozen songs. A table recording the presence or absence of the various notes in consecutive songs revealed that at least two of the three birds used different sets of notes in different bouts of songs, switching from one to another after short pauses made between two consecutive bouts. All this makes the attempt to assess the actual size of the note repertoire in these birds very difficult, and makes it probable that Sardinian warblers do not possess a finite repertoire of notes. The pattern of note selection observed for the three males in this study is best explained by assuming that these birds improvise new notes as they sing.     

A SYSTEM FOR DESCRIBING BIRD SONG UNITS

ABSTRACT

Bird song researchers have not agreed on a common set of units of analysis by which birds' songs of various different species might be described. Analysis of 50 papers reveals 28 unit designations and considerable variation in their application despite only three different methods for identifying units. The lack of consensus on units arises from the fact that units generated by the same methods at different levels of organization are given different names. A method for designating bird song units is offered for discussion which uses the concept of level of organization to stress the fundamental unanimity of method. It is hoped that consideration of this method will lead ultimately to greater standardization in the protocols by which researchers generate and name bird song units.     

白头鹎方言的初步研究

对浙江省乘泗岛，普陀岛，温岭，长兴，杭州和龙游等６个地区白头鹎的鸣声进行了研究，发现它们鸣声主句的语调，音节数，持续时间，频谱特征和频率范围等均有差异，说明白头鹎鸣声中普遍存在“方言”，但它们作为同一个种在鸣声主句上有其共同特征：主要是鸣声多在１．５－３ｋＨｚ的低中频段内变化，单音节的鸣叫声等都极为相似。     

A GENETIC AND BEHAVIORAL ANALYSIS OF MATE CHOICE AND SONG NEIGHBORHOODS IN INDIGO BUNTINGS

Neighboring males of indigo buntings (Passerina cyanea) share songs in southern Michigan. We sampled polymorphic enzymes to compare the genetic variation between mates and the variation among contiguous song neighborhoods. Mate choice was independent of the genetic and morphometric similarity of female and male, and these measures were independent of each other. The incidence of extrapair copulations and fertilizations was independent of the song of cuckolding males. Breeding success of the mated pairs was independent of their genetic or morphological similarity. Males characterized by different song dialects did not differ in mean lifetime reproductive success. We found no significant genetic differences among the neighborhoods. Most birds that bred in one song neighborhood were born in another, and neighborhoods were not isolated demes. Bunting songs may provide no information to a female about genetic quality of males. The results are consistent with a neutral model of no mate choice for genes.     

Acoustic communication in the mandarin vole (<Emphasis Type="Italic">Lasiopodomys mandarinus</Emphasis>, Rodentia)

Acoustic communication in the mandarin vole is described, including sharp and quiet squeals (in males and females) and male songs. All vocal signals show individual variation, which can be used for recognizing individual animals. Female quiet squeals and male songs are emitted not only during sexual behavior but also during friendly interactions, forming a dialogue between the animals. The role of social organization of the species in imparting communicative significance to these vocal signals is discussed.     

SONG STRUCTURE AND SINGING BEHAVIOUR OF WILLOW WARBLER PHYLLOSCOPUS TROCHILUS ACREDULA IN POPULATIONS OF LOW AND HIGH DENSITY

ABSTRACT

Song structure and singing behaviour in two nesting populations of Willow Warbler Phylloscopus trochilus acredula of different density were studied in Moscow and Kostroma Regions, Russia, from 1994–1997. In total 520 songs of 9 males from the low-density population and 566 songs of 12 males from the high density population were analysed. Singing periods in the high density Willow Warbler population (11 males, 758 min.) were longer (58% of the time) than in the low density population (13 males, 866 min.) (58%). In the high density population, 51% of singing time consisted of “singing duels” (singing in gaps between the songs of a neighbouring male, and singing during the singing of a neighbouring male), while in the low density population only 7% did. The initial phrases of Willow Warbler songs were significantly longer in the high density population, with up to 9–13 elements, while there were no occurrences of this length in the low density population. Songs as a whole were longer and more variable in the high density population. Thus changes in structure, singing activity and behaviour are linked to male-male interaction and density of the nesting community.     

Ecomorphological patterns in the Blackcap Sylvia atricapilla: insular versus mainland populations

Capsule Blackcap Sylvia atricapilla populations from the Azores archipelago show morphological differences to continental birds which are consistent with the ‘Island Rule’.

Aims The morphology of insular vertebrates is usually the result of the evolution in their particular environment and leads to predictable morphological patterns, according to the Island Rule. We test the predictions of the Island Rule, using the Blackcap of the Azores archipelago as our model.

Methods We compared morphological variation (body size and wing shape) of populations from the nine islands of the Azores to continental birds, using multivariate indexes. Also, we looked at the relationship between these patterns and possible insular ecological drivers of morphological divergence.

Results Our findings are concordant with Island Rule predictions, as in general birds from the Azores are larger than continental populations, especially birds from the most distant islands. Wing shape also differs significantly, as Azorean Blackcaps tend to have rounder wings than continental birds with a migratory-like phenotype.

Conclusion Overall, we conclude that the observed morphological patterns in Blackcap in the Azores conform in general to the Island Rule predictions.     

SONGS OF BLUE TITS PARUS CAERULEUS PALMENSIS FROM LA PALMA (CANARY ISLANDS)—A TEST OF HYPOTHESES

ABSTRACT

Parus major and Parus caeruleus palmensis have some song types in common. They consist of whistled element-types in different frequency ranges. Both species show territorial reactions to playback of the song-types belonging to the other species. P.c. palmensis does not share any songtypes with Central European Parus caeruleus. This is confirmed by playback experiments with the different song types, which do not elicit any territorial behaviour. The songs of P.c. palmensis also differ from those of blue tits from any other Canary Island. They may be derived from contact calls of the Lanzarote population. They have been developed by young birds that had not yet learnt their songs when they colonized La Palma from Lanzarote without reaching any of the other Canary Islands. It is suggested that the uniform song repertoire was developed by adapting the already learned contact calls to each other. The similarities to song types of P. major are due to convergence rather than character displacement. All other Canary Island tit populations share the main features of their songs, which are characterized by quick frequency changes. It seems most likely that these islands have been colonized by birds which had already learnt their songs. Differences in song repertoires are mainly caused by drift events which happen to occur because of population bottlenecks during the colonization process.     

Divergence in mating signals correlates with genetic distance and behavioural responses to playback

Animals use acoustic signals to defend resources against rivals and attract breeding partners. As with many biological traits, acoustic signals may reflect ancestry; closely related species often produce more similar signals than do distantly related species. Whether this similarity in acoustic signals is biologically relevant to animals is poorly understood. We conducted a playback experiment to measure the physical and vocal responses of male songbirds to the songs of both conspecific and allopatric‐congeneric animals that varied in their acoustic and genetic similarity. Our subjects were territorial males of four species of neotropical Troglodytes wrens: Brown‐throated Wrens (Troglodytes brunneicollis), Cozumel Wrens (T. beani), Clarion Wrens (T. tanneri) and Socorro Wrens (T. sissonii). Our results indicate that birds respond to playback of both conspecific and allopatric‐congeneric animals; that acoustic differences increase with genetic distance; and that genetic divergence predicts the strength of behavioural responses to playback, after removing the effects of acoustic similarity between subjects’ songs and playback stimuli. Collectively, these results demonstrate that the most distantly related species have the most divergent songs; that male wrens perceive divergence in fine structural characteristics of songs; and that perceptual differences between species reflect evolutionary history. This study offers novel insight into the importance of acoustic divergence of learned signals and receiver responses in species recognition.     

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.     

ACOUSTIC COMMUNICATION SIGNALS OF MYSTICETE WHALES

ABSTRACT

Mysticete (baleen) whales produce a variety of vocalizations and sounds, but relatively few of these have been well described with accompanying behavior. This review concentrates on the vocalizations consistently associated with behavioral interactions or acoustic exchanges between or among conspecifics. These communication “signals” have been categorized for this review as contact calls of single animals outside of the breeding season (including cow-calf pairs), vocalizations reported during the breeding season (often designated as “songs”), and calls produced by active groups of whales that may or may not have a reproductive function. While much remains unknown, the data obtained thus far indicate that the social vocalizations of baleen whales have structural/functional similarities with those of other mammals and birds.     

Song discrimination learning in zebra finches induces highly divergent responses to novel songs

Perceptual biases can shape the evolution of signal form. Understanding the origin and direction of such biases is therefore crucial for understanding signal evolution. Many animals learn about species-specific signals. Discrimination learning using simple stimuli varying in one dimension (e.g. amplitude, wavelength) can result in perceptual biases with preferences for specific novel stimuli, depending on the stimulus dimensions. We examine how this translates to discrimination learning involving complex communication signals; birdsongs. Zebra finches (Taeniopygia guttata) were trained to discriminate between two artificial songs, using a Go/No-Go procedure. The training songs in experiment 1 differed in the number of repeats of a particular element. The songs in experiment 2 differed in the position of an odd element in a series of repeated elements. We examined generalization patterns by presenting novel songs with more or fewer repeated elements (experiment 1), or with the odd element earlier or later in the repeated element sequence (experiment 2). Control birds were trained with only one song. The generalization curves obtained from (i) control birds, (ii) experimental birds in experiment 1, and (iii) experimental birds in experiment 2 showed large and systematic differences from each other. Birds in experiment 1, but not 2, responded more strongly to specific novel songs than to training songs, showing 'peak shift'. The outcome indicates that learning about communication signals may give rise to perceptual biases that may drive signal evolution.     

SONG INHERITANCE AND MATING IN A SONGBIRD WITH LOCAL DIALECTS

ABSTRACT

In this paper we document the pattern of geographic variation in song of the Corn Bunting in a marked population in Sussex. Song variation is best described as a system of local dialects with three song types in each dialect. We examine the inheritance of dialects from father to son; sons sing the same dialect as their nearest neighbour, rather than inheriting the dialect of the father. Therefore songs seem to be learned after dispersal. We also compare the dialects of mates and fathers of females: our results suggest that females do not rely on dialects when pairing. These results are discussed in the context of the current controversy surrounding other species with dialects and hypotheses relating dialects to the genetic structure of populations.     

Rock wrens preferentially use song types that improve long distance signal transmission during natural singing bouts

When animals are capable of producing variable signals they may preferentially use some signal types over others. Among songbirds, individuals are known to alter song type form and usage patterns in contest and mating situations, but studies have not examined how song choice improves signal efficacy during broadcast song. For this study we investigated rock wren Salpinctes obsoletus song type use rates during natural singing bouts. We tested three hypotheses for adaptive song use during broadcast song: 1) birds improve signal content by increasing the use of high quality songs, 2) birds optimize for signal propagation by preferentially using songs that transmit well, and 3) birds maintain energy by reducing the use of costly songs. The study included 19 058 songs sung by 12 individuals, each of which had a measured song repertoire of between 52 and 117 song types which were produced at highly variable rates. Results indicated that rock wrens did not preferentially sing song types with shorter durations or fewer frequency switches, as would be expected if they selected song types to minimize delivery costs. They also did not favor songs with more rapid trills or more frequency switches, as would be expected if they adjusted song use primarily to indicate quality. Focal birds did preferentially sing significantly longer songs with lower bandwidths, lower frequencies, and slower trill rates. Results suggest that natural broadcast singing patterns are shaped more by the benefits of long distance transmission than by the benefits of advertising performance ability or the costs of song production.     

THE COMMUNICATIVE SIGNIFICANCE OF TONALITY IN BIRDSONG: RESPONSES TO SONGS PRODUCED IN HELIUM

ABSTRACT

“Pure tones” are a distinctive acoustic feature of many birdsongs. Recent research on songbird vocal physiology suggests that such tonal sounds result from a coordinated interaction between the syrinx and a vocal filter, as demonstrated by the emergence of harmonic overtones when a bird sings in helium. To investigate the communicative significance of vocal tract filtration in the production of birdsong, we used field playback experiments to compare the responses of male swamp sparrows Melospiza georgiana to normal songs and those same songs recorded in helium. We also measured responses to pure tone songs that had been shifted upward in frequency to match the average spectra of those songs with added harmonics. Male sparrows were significantly more responsive to the playback of normal songs than to either helium songs with added harmonics or frequency- shifted pure tone songs. Songs with harmonics retained a high degree of salience, however. We conclude that explanations for the occurrence of tonal sounds in birdsongs must consider perceptual attributes of songs as communicative signals, as well as problems of song production and transmission.