Song characterization in the spectacled warbler (Sylvia conspicillata): a circum-Mediterranean species with a complex song structure

The spectacled warbler (Sylvia conspicillata) is a small passerine with a patchy distribution throughout the circum-Mediterranean region, including the North Atlantic archipelagos of Madeira, Canary Islands and Cape Verde. Here we characterize the species song structure on the island of Fuerteventura, quantifying repertoire size, inter- and intra-individual spectrographic variation, to determine whether acoustic variation occurred within an island population. Male song display was organized in song bouts of a variable number of song phrases, which in turn were made up of 4–69 syllables. We classified syllable types to derive a measure of repertoire size (number of different syllables) per song bout, and then used rarefaction methods to calculate the estimated repertoire size for our population of males. Three categories of song bout length were considered in analyses: short song bouts of 10 phrases, average bouts of 19 phrases and long bouts of ≥ 29 phrases. The observed and estimated repertoire size per male (between 43 and 126 syllables per male) increased with song bout duration, although the relationship was not significant for the estimated values. To test whether songs could be individually specific, we measured 11 spectrotemporal parameters of the song. A discriminant analysis using these variables performed poorly in classifying songs to the individuals that uttered them, but we found less variation in the individual than in the population for three out of the 11 variables. These individually specific variables, involving the first or the most common syllable of the song, the trill, were the duration of the first syllable of the phrase, the duration and the dominant frequency of the trill syllable. Our study emphasizes the complexity of spectacled warbler songs, in which males continuously add novel syllables over the entire song bout. This complexity appears to be determined by individual innovation capabilities rather than by the behaviour of copying neighbour repertoires, since songs of close birds were not more similar than songs from far-away territories.     

Evidence for High Variability in Temporal Features of the Male Coda in Müller’s Bornean Gibbons (<Emphasis Type="Italic">Hylobates muelleri</Emphasis>)

Most pair-living primate species engage in duets, wherein males and females produce coordinated vocalizations. Previous analyses of male gibbon contributions to the duet have shown that calls are individually distinct. Here we investigate variation in the temporal and spectral parameters in the male contribution to the duet, also known as the coda, of wild, nonhabituated male Müller’s Bornean gibbons (Hylobates muelleri), recorded both opportunistically and as a response to playbacks at the Stability of Altered Forest Ecosystems site in Sabah, Malaysia. We used linear discriminant function analysis to estimate the intra- and interindividual variation in 13 spectral and temporal parameters of the vocalizations (N?=?337) of 31 male gibbons. To further understand how call features vary within and between individuals we used a multivariate, variance components model to investigate how variance in features was partitioned at these two levels. We could identify males with a 66% accuracy using leave-one-out cross-validation, a relatively low score compared to female Müller’s Bornean gibbons and males of other species. We found that for some features (such as maximum frequency of the notes) most of the variance occurred between males, but for others (specifically total duration of the call and duration of rest in between notes) most of the variance occurred within a single male. Overall, male Müller’s Bornean gibbon coda vocalizations showed greater variability relative to their female counterparts, raising questions about potential differences in the function of the male and female contributions to the duet in Müller’s Bornean gibbons and the gibbon taxon as a whole.     

Acoustic features contributing to the individuality of wild agile gibbon (Hylobates agilis agilis) songs

We examined acoustic individuality in wild agile gibbon Hylobates agilis agilis and determined the acoustic variables that contribute to individual discrimination using multivariate analyses. We recorded 125 female-specific songs (great calls) from six groups in west Sumatra and measured 58 acoustic variables for each great call. We performed principal component analysis to summarize the 58 variables into six acoustic principal components (PCs). Generally, each PC corresponded to a part of the great call. Significant individual differences were found across six individual gibbons in each of the six PCs. Moreover, strong acoustic individuality was found in the introductory and climax parts of the great call. In contrast, the terminal part contributed little to individual identification. Discriminant analysis showed that these PCs contributed to individual discrimination with high repeatability. Although we cannot conclude that agile gibbon use these acoustic components for individual discrimination, they are potential candidates for individual recognition.     

Calling in wild silvery gibbons (Hylobates moloch) in Java (Indonesia): behavior, phylogeny, and conservation

Hardly any behavioral data are available for the silvery gibbon (Hylobates moloch), an endangered primate that is endemic to the island of Java, Indonesia. We studied the singing behavior of the easternmost population of this species in the Dieng mountains, central Java, in 1998-1999. We aimed to document the timing of singing, quantify the amount of singing by the respective sexes, and explore the role of bioacoustics in density estimation. A total of 122 song bouts in at least 12 groups were monitored. No duet songs were heard. Most of the song bouts (91.5%) were female solo song bouts or female scream bouts. In contrast to an earlier study on the westernmost population of silvery gibbons, during which few if any male songs were heard, at least 8.5% of the song bouts in our study were male solo song bouts. They were significantly longer in duration than the female songs. All male song bouts uttered before dawn (0520 hr) were produced in a chorus fashion, with at least three individuals participating. Choruses occurred about once every 8.5 days, and lasted longer and occurred earlier than female solo song bouts. Most male songs (60%) started between 0355-0440 hr, when it was still dark. All female songs, in contrast, started after 0500 hr, and female singing activity peaked around 0600. Regular male singing, male chorusing, and regular predawn singing have not previously been reported for silvery gibbons. Similarly separated periods of male and female solo songs and the absence of duetting have been observed in Kloss's gibbons (H. klossii) on the Mentawai Islands, and may represent synapomorphies shared by both species. The pronounced individual-specific song characteristics of silvery gibbons allow accurate mapping of groups. The density of gibbons at our study site was established to be 1.9-3.7 groups/km2, corresponding to 6.7-13.1 individuals/km2. We reassess the suitability of gibbon songs as a means of estimating the density and size of gibbon populations, and discuss the proximate causes for the absence of duetting in silvery gibbons.     

Species Recognition by Song in the Veery (Catharus fuscescens: Aves)

Territorial male veeries (Catharus fuscescens) were presented with three series of reorganised songs to determine the functional properties of song structure. Songs consist of three parts: a low frequency initial phrase, a higher frequency middle phrase, and again a lower frequency end phrase. The latter two consist of vibrato syllables. Three series of experiments were performed, using various combinations of song syllables. In Series I, songs consisted of repetitions of a single syllable, one from each part of the song. Veeries responded to syllables from the middle and end phrases as to normal songs, both in localisation and in vocal responses. However, they showed little localisation and vocalisation responses to the introductory syllables. In Series II and III, veeries responded significantly more to songs in which syllables from the middle and end phrases were presented in proper sequence, rather than not. The minimum requirements for intraspecific recognition were determined in relation to the number and order of syllables in the song.     

Advertising vocalization of the clamorous reed warbler (<Emphasis Type="Italic">Acrocephalus stentoreus</Emphasis>, Sylviidae)

The advertising vocalization of the clamorous reed warbler, one of the least studied representatives of the genus in the Palearctic, has been analyzed. In this species, advertising vocalization consists of compact acoustic constructs (songs) separated by pauses. The duration of songs (median 3.6 s) and pauses (median 2.9 s) varies only slightly. Each song consists of three to four different or identical syllables. Syllables (stereotyped acoustic constructs) consist of two to four notes. Each male has from 5 to 25 syllables in his repertoire. A sharp contrast between low-frequency and high-frequency notes and strict sequence of these notes in syllables (low-frequency notes come first, then follow high-frequency notes) are characteristic of this song. All types of syllables in the song of a given male can be divided into two groups: initiator syllables and other syllables. Initiator syllables most often occur at the beginning, while others occur in the middle or at the end of the song. Stable links between syllables have been revealed: the majority of them occur in the song together with certain syllables (one or two) of other types. The whole range of songs of each individual can be divided into several classes. A certain type of initiator syllable and a certain set of subsequent syllables are characteristic of each class. However, the variability of songs within each class is quite high. Modes of immediate and eventual variety alternate in the sequence of songs from different classes. Thus, 47.4% of songs are followed by a song from the same class (eventual variety), while 52.6% of songs are followed by a song from a different class (immediate variety). The advertising vocalization of the clamorous reed warbler is compared with that of other representatives of the genus Acrocephalus.     

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.     

Population differences and acoustic stability in male songs of wild western black crested gibbons (Nomascus concolor) in Mt. Wuliang, Yunnan

Population differences and acoustic stability in male songs of the western black crested gibbon (Nomascus concolor) at Mt. Wuliang, Yunnan, were investigated using data collected over 4 years. Detectable vocal differences were found between the western and eastern populations within Mt. Wuliang. Discriminant functions calculated using the phrases recorded in 2007-2008 correctly classified phrases recorded in 2009, and discriminant functions produced from phrases recorded in 2007-2009 could correctly classify phrases recorded in 2010 for 4 of 5 males. Results showed that the male songs of western black crested gibbons remain relatively stable over years. In light of the acoustic individuality in male songs of this gibbon species, which has recently been proved, and the acoustic stability shown in this research, we suggest that the song of male individuals may be useful in monitoring individuals or family groups of this species.     

Individuality in male songs of wild black crested gibbons (Nomascus concolor)

This is the first study of vocal individuality in male songs of black crested gibbons. The sound recordings were carried out at two field sites, Pinghe, Ailao Mountains, and Dazhaizi, Wuliang Mountains, both located in Yunnan province, China. A total of 127 coda phrases of 38 male songs bouts of eight individual male gibbons were analyzed. Stepwise discriminant function analysis was used to examine the acoustic individuality of the males. We found that individuality among neighbors was very pronounced. Moreover, individuality within a site (i.e. among neighbors) is higher than among individuals between sites. Our finding suggests that black crested gibbons may actively increase their degree of vocal individuality against that of their immediate neighbors by vocal adjustment.     

Individuality in the songs of wild agile gibbons (Hylobates agilis) of Peninsular Malaysia

Recordings were made and analyzed of the female dominated duet songs by eight adult pairs of wild agile gibbons from two sites in Peninsular Malaysia. A statistical analysis of their songs revealed that individuality occurred throughout the females' songs from both sites. Individuality in gibbon songs may allow singing individuals to more efficiently and effectively locate and identify neighboring nonmate conspecifics.     

丹顶鹤性活动的声行为研究

丹顶鹤繁殖期的性活动可分为雄鹤求偶、雌鹤对雄性求偶的应答、两性交配和交配完结4个阶段,其相应的鸣声模式分别为雄性的求偶鸣声、雌性对雄性求偶的应答声和两性的对鸣声、两性对唱的交配声和两性的高声合唱。4个阶段鸣声都是以基本音的主频率(PF)为主音的单音调声,前3个阶段都带数个近似fn=nf0(f0=FP)关系的低幅值谐频成分。第4个阶段带数个近似fn=nf0(f0=FP)关系的高幅值谐频成分;品质因数(Q3dB)多半为4～6,声脉冲重复频率(RFP)一般为150～180Hz,而第2阶段声的RFP一般为180～260Hz。雄性鸣声的每个单次叫声中含有的音节数较少,一般不超过4个;而雌性鸣声比较复杂。每个单次叫声中含有的音节数较多,一般都在7～8个以上;但雌雄鸣声的每个音节都是由3个声脉冲组成。雄鹤鸣唱声频率变化范围较小,而雌鹤鸣唱声频率变化形式是由低到高达到高峰后又开始下降。4个阶段的鸣声都具有较好共鸣。只有第2阶段发声运动较快。而且发现雄鹤鸣唱单次鸣叫声的音节数“增多”。各阶段鸣声特性均存在差异,不同配偶间均存在显著差异,研究结果表明丹顶鹤雌雄都具有不同的鸣声,且其性活动过程中不同的鸣声行为具有较高的个体识别信号潜能。另外,求偶鸣叫声和求偶应答与对鸣声在性活动鸣声中起着决定性的作用。     

Sound Spectrum Characteristics of Songs of Hainan Gibbon (Nomascus hainanus)

Gibbons are characterized by their species-specific calls, or songs. There are few studies of songs of Hainan gibbon (Nomascus hainanus). To study the sound spectrum characteristics and test for intergroup differences in Hainan gibbon song, we studied the singing behavior of Hainan gibbons in Bawangling National Nature Reserve, Hainan Province, China, intermittently from August 2002 to February 2013, collecting 184 recordings. Our results show that: 1) Hainan gibbon song bouts occur mainly 0–4 h after dawn. 2) The songs of adult males living in groups are composed mainly of one to three short notes and one to five long notes, while solitary adult male songs consist only of long frequency modulated notes and no short or single notes. 3) The song chorus is dominated by adult males, while females add a great call. Males do not have a great call, unlike those in other gibbon species. There are no female solos. 4) The sound spectrum frequency is similar in adult males living in two different groups, but the duration of the first long note differed significantly between the groups. The sonic frequencies of male and female songs are lower than those of other gibbons: no more than 2 kHz. Hainan gibbon sound structure is simple, although females participate in the chorus, reflecting their primitive status among gibbon species.     

Species song discrimination in adult female song and swamp sparrows

Six female song sparrows (Melospiza melodia) and six female swamp sparrows (Melospiza georgiana) were treated with oestradiol, and their response to song measured by frequency and intensity of copulation-solicitation display. Test stimuli were synthetic songs composed of either song sparrow or swamp sparrow syllables assembled in either song-sparrow-like or swamp-sparrow-like temporal patterns. Female song sparrows responded preferentially to songs containing their own species' syllables and to songs containing their own species' temporal patterns. Swamp sparrows were also sensitive to both syllable type and temporal, pattern, in contrast to male swamp sparrows, which show no preference for swamp sparrow temporal patterns.     

Acoustic preference functions and sexual selection on the male calling song in the grasshopper Chorthippus biguttulus

Male courtship songs have two functions in species recognition and intraspecific mate choice. Female preference might thus exert different types of selection pressure on male song traits. We used a combination of acoustic mate choice experiments and statistical analyses to examine how traits of the calling songs of male nightingale grasshoppers,Chorthippus biguttulus , are influenced by different sexual selection pressures. We recorded calling songs of males and tested their attractiveness to females in acoustic mate choice experiments. The attractiveness values were a good estimate of the potential male mating success. In experiments with a pair of males, females copulated significantly more often with the male that had the higher attractiveness value. To detect directional, stabilizing, disruptive or correlative selection acting on male song properties we used linear and nonlinear regressions between male song traits and female response behaviour. Three signal traits were revealed to be under directional selection: song loudness, pause to syllable ratio and the mean duration of gaps within syllables. A nonlinear regression testing for correlative selection showed that a fourth song trait, rhythm, in combination with mean gap duration was also important for female mate choice. With these traits and trait combinations we were able to explain 42% of the variance in attractiveness between males. Since we found no evidence for stabilizing selection, but ample evidence for directional selection, we conclude that selection on the traits examined is related to mate choice mainly in the context of intraspecific sexual selection and probably less so in species recognition. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Innate differences in singing behaviour of sparrows reared in isolation from adult conspecific song

Innate differences in the singing behaviour of male swamp (Melospiza georgiana) and song (M. melodia) sparrows were identified by rearing males from the egg in the laboratory under identical conditions, in complete isolation from adult conspecific song. Isolation-reared males of both species displayed several abnormal song features, including reduced numbers of notes per song, longer durations of notes and inter-note intervals, and fewer notes per syllable. Despite these and other abnormalities, many species differences emerged that matched differences in the natural singing behaviour of the two species. These included differences in song repertoire size, song duration and degrees of segmentation, numbers of notes per song, durations of notes and inter-note intervals, and several measures reflecting the organization of songs into note complexes, syllables and trills. Although learning can influence all levels of organization of the motor patterns of song in swamp and song sparrows, its contribution to the achievement of normal song behaviour appears to be most crucial at the level of the fine structure of the notes and syllables from which the songs are constructed.     

Antiphonal duetting in Steere’s liocichla (Liocichla steerii): male song individuality and correlation between habitat and duetting behavior

We studied duet song and vocal duetting behavior in an endemic Taiwanese passerine, Steeres liocichla (Liocichla steerii). We found that the leading male song in duets was highly individualistic. Also, we found duetting behavior varied significantly across different habitat types. Females were more likely to answer male songs in densely vegetated, steep forest habitat compared to open agricultural habitat. These findings provide quantitative evidence for vocal individuality for a duet song and provide tentative support for the idea that females are duetting to reveal their location to their mates.     

Socially induced brain differentiation in a cooperatively breeding songbird

Birds living in social groups establish dominance hierarchies, and taking up the dominant position influences behaviour and physiological parameters. In cooperatively breeding white-browed sparrow weavers (Plocepasser mahali), the transition from subordinate helper to dominant breeder male induces the production of a new type of song. This song contains a large number of new syllables and differs in temporal pattern from duet songs produced by all other group members. Here we show that this change in social status of adult males affects the morphology of a behavioural control circuit, the song control system of songbirds that is composed of large neuron populations. The volume of the song control areas HVC and RA and their gene-expression levels depend on males' social status. Dominant males have several times larger testes than subordinates, which is not reflected in circulating androgen and oestrogen levels. Our findings suggest a remarkable differentiation of adult vertebrate brains in relation to changing social cues.     

杭州市区白头鹎鸣声的微地理差异

2002年5—8月,对中国东部浙江省杭州市区的白头鹎鸣声的微地理差异进行研究。在约60 km2的 研究区内,选择8个调查点(4个城区,4个丘陵山地),录制了80个雄性白头鹎的511个鸣声样本,并随机选 取每一调查点的20个鸣声样本进行分析。其结果显示,白头鹎在杭州市区至少有8种微地理鸣声方言,每个微 地理鸣声方言都有一典型鸣句;它们在听觉上、波形结构、音节组成、音节频谱特征等方面均不相同。有的一 路之隔的相邻微地理鸣声方言之间存在明显差异;部分区域出现鸣声混合现象;有的个体还具有“多语”功 能。白头鹎鸣声产生微地理差异的原因可能与鸟类个体的扩散和城市中鸟类栖息地的人为改变有关。     

Singing by male and female Kloss gibbons (Hylobates klossii) in the Peleonan Forest, Siberut Island, Indonesia

Kloss gibbons (Hylobates klossii) are endemic to the Mentawai Islands in Indonesia and are one of only two gibbon species in which mated pairs do not sing duets. This is the first long-term study of the factors influencing the singing activity of Kloss gibbons within a northern Siberut Island population and follows two previous studies in central Siberut nearly 30 years ago. We collected data on the presence/absence of male and female singing within the study area on 198 days and within a focal group on 47 days. Rainfall during the time period in which they normally sing inhibits singing in both males and females. Our study supports the hypothesis that male and female songs function in intrasexual resource defence, as singing is associated with singing by same-sex neighbours, and same-sex choruses are more likely to occur after one or more days of silence (from that sex), suggesting there is pressure for individuals to communicate with same-sex neighbours regularly. Singing was not coordinated within a mated pair, suggesting that vocal coordination of the pair has been lost with the loss of the duet and that Kloss gibbon songs do not convey information to neighbours about the strength of the pair bond. On days when males sang predawn, females were more likely to sing after dawn and earlier in the morning. Additionally, the number of groups singing in female choruses was positively associated with the number of males that had sung in the predawn male chorus. We suggest that female songs have an intersexual territory defence as well as an intrasexual function.