Song pattern recognition and an auditory time window in the female bushcricket Ancistrura nigrovittata (Orthoptera: Phaneropteridae)

Mate finding in the phaneropterid bushcricket Ancistrura nigrovittata is achieved by a duet, where the female replies with a short sound to the male song. In experiments with artificial song models we analysed the parameters necessary for eliciting a female response. A verse of the male song consists of a group of 5–9 syllabes which after an interval of about 400 ms is followed by a final syllable. The female response was shown to depend on two processes: (i) recognition of the syllable group as belonging to a conspecific male and (ii) perception of the final syllable as a trigger. Critical parameters for the recognition process are the duration of syllables and syllable pauses, as well as the number of syllables in a group. However, even with an optimal syllable group, the response probability still depends on the interval between the syllable group and the final syllable. The female only responds when the final syllable of the male song occurs within a 250 ms long time window begining approximately 250 ms after the end of the male's syllable group. Her reply consists of a single tick, which follows the male's final syllable with a latency of only 25 ms.     

Social status affects the degree of sex difference in the songbird brain

It is thought that neural sex differences are functionally related to sex differences in the behaviour of vertebrates. A prominent example is the song control system of songbirds. Inter-specific comparisons have led to the hypothesis that sex differences in song nuclei size correlate with sex differences in song behaviour. However, only few species with similar song behaviour in both sexes have been investigated and not all data fit the hypothesis. We investigated the proposed structure-function relationship in a cooperatively breeding and duetting songbird, the white-browed sparrow weaver (Plocepasser mahali). This species lives in groups of 2-10 individuals, with a dominant breeding pair and male and female subordinates. While all male and female group members sing duet and chorus song, a male, once it has reached the dominant position in the group, sings an additional type of song that comprises a distinct and large syllable repertoire. Here we show for both types of male-female comparisons a male-biased sex difference in neuroanatomy of areas of the song production pathway (HVC and RA) that does not correlate with the observed polymorphism in song behaviour. In contrast, in situ hybridisation of mRNA of selected genes expressed in the song nucleus HVC reveals a gene expression pattern that is either similar between sexes in female-subordinate male comparisons or female-biased in female-dominant male comparisons. Thus, the polymorphic gene expression pattern would fit the sex- and status-related song behaviour. However, this implies that once a male has become dominant it produces the duetting song with a different neural phenotype than subordinate males.     

Acoustic defence in an insect: characteristics of defensive stridulation and differences between the sexes in the tettigoniid Poecilimon ornatus (Schmidt 1850)

Many insects exhibit secondary defence mechanisms upon contact with a predator, such as defensive sound production or regurgitation of gut contents. In the tettigoniid Poecilimon ornatus, both males and females are capable of sound production and of regurgitation. However, wing stridulatory structures for intraspecific acoustic communication evolved independently in males and females, and may result in different defence sounds. Here we investigate in P. ornatus whether secondary defence behaviours, in particular defence sounds, show sex-specific differences. The male defence sound differs significantly from the male calling song in that it has a longer syllable duration and a higher number of impulses per syllable. In females, the defence sound syllables are also significantly longer than the syllables of their response song to the male calling song. In addition, the acoustic disturbance stridulation differs notably between females and males as both sexes exhibit different temporal patterns of the defence sound. Furthermore, males use defence sounds more often than females. The higher proportion of male disturbance stridulation is consistent with a male-biased predation risk during calling and phonotactic behaviour. The temporal structures of the female and male defence sounds support a deimatic function of the startling sound in both females and males, rather than an adaptation for a particular temporal pattern. Independently of the clear differences in sound defence, no difference in regurgitation of gut content occurs between the sexes.     

A Novel Context for Bird Song: Predator Calls Prompt Male Singing in the Kleptogamous Superb Fairy-wren,Malurus cyaneus

The cooperatively breeding superb fairy-wren, Malurus cyaneus, sings two types of song; a variable, complex song sung by both sexes, and a relatively invariant, stereotyped song sung only by males. Field observations and playback experiments revealed that the male song could be triggered by a variety of loud avian and non-avian vocalizations, but was triggered most frequently by the calls of predators or potential predators. The male song is sung both within the territory and in winter feeding flocks outside the territory. Acoustic properties of the song suggest that it is a long-range signal. A possible explanation for this unusual phenomenon is that the male song has evolved through sexual selection as an honest signal to females of male quality, for the purpose of obtaining extra-pair copulations.     

Female recognition of trills in the male calling song of the field cricket,<Emphasis Type="Italic"> Teleogryllus taiwanemma</Emphasis>

The calling song of the field cricket, Teleogryllus taiwanemma, is usually considered to consist of sequences of separate chirps. However, sometimes it comprises a phrase of several chirps in a row, with one long chirp (chirp) and a few short chirps (trills). In this study, I compared the phrase containing only chirps with that containing both chirps and trills by analyzing male songs and conducting playback experiments of male songs to females. The song analyses showed significant differences between chirps and trills for all song parameters except bandwidth. To test whether female preference differed with respect to the two phrases, I performed two-speaker playback experiments. When the same numbers of phrases were presented per unit time, females preferred the song with trills to that without trills. This result may reflect female preference for songs with greater sound density. In subsequent playback experiments, I equalized the total sound duration per unit time (duty cycle) in songs with and without trills. The numbers of females that preferred songs with and without trills did not differ significantly. This suggests that trills can attract females like chirps do, even though the two sounds have different components.     

Temporal parameters of male—female sound communication in Leptophyes punctatissima

ABSTRACT. Quantitative information is presented about acoustic communication between the sexes of the speckled bush cricket Leptophyes punctatissima Bosc (Orthoptera: Tettigoniidae). The male song elicits an acoustic response from a receptive female with a net delay (excluding the travelling time of the sound) of about 25 ms depending upon ambient temperature. Only when the female responses fall within the narrow temporal window between 20 and 50 ms after the onset of his song does the male perform phonotaxis. The precise timing of this duet is described, and its advantages and disadvantages are discussed.     

Sex role similarity and sexual selection predict male and female song elaboration and dimorphism in fairy‐wrens

Historically, bird song complexity was thought to evolve primarily through sexual selection on males; yet, in many species, both sexes sing and selection pressure on both sexes may be broader. Previous research suggests competition for mates and resources during short, synchronous breeding seasons leads to more elaborate male songs at high, temperate latitudes. Furthermore, we expect male–female song structure and elaboration to be more similar at lower, tropical latitudes, where longer breeding seasons and year‐round territoriality yield similar social selection pressures in both sexes. However, studies seldom take both types of selective pressures and sexes into account. We examined song in both sexes in 15 populations of nine‐fairy‐wren species (Maluridae), a Southern Hemisphere clade with female song. We compared song elaboration (in both sexes) and sexual song dimorphism to latitude and life‐history variables tied to sexual and social selection pressures and sex roles. Our results suggest that song elaboration evolved in part due to sexual competition in males: male songs were longer than female songs in populations with low male survival and less male provisioning. Also, female songs evolved independently of male songs: female songs were slower paced than male songs, although only in less synchronously breeding populations. We also found male and female songs were more similar when parental care was more equal and when male survival was high, which provides strong evidence that sex role similarity correlates with male–female song similarity. Contrary to Northern Hemisphere latitudinal patterns, male and female songs were more similar at higher, temperate latitudes. These results suggest that selection on song can be sex specific, with male song elaboration favored in contexts with stronger sexual selection. At the same time, selection pressures associated with sex role similarity appear to favor sex role similarity in song structure.     

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.     

The song structure and seasonal patterns of vocal behavior of male and female bellbirds (Anthornis melanura)

The bellbird (Anthornis melanura) is a honeyeater endemic to New Zealand, which uses song to defend breeding territories and/or food resources year round. Both sexes sing and the song structure and singing behavior have not yet been quantified. The number of song types, spectral structure, repertoire size, and singing behavior of male and female bellbirds was investigated for a large island population. Song types differed between the sexes with males singing a number of structurally distinct song types and females producing song types that overlapped in structure. Singing behavior also differed between the sexes; males often sung long series of songs while females sung each song at relatively long and variable intervals. Singing by both sexes occurred year round but the frequency of male and female singing bouts showed contrasting seasonal patterns. The frequency of female singing bouts increased as the breeding season progressed, whereas male singing bouts decreased. In contrast to almost all studied passerines, female bellbirds exhibited significant singing behavior and sung songs of complex structure and variety that parallel male song. These results provide a quantitative foundation for further research of song in bellbirds and in particular the function of female vocal behavior.     

Sex-specific spectral tuning for the partner's song in the duetting bushcricket Ancistrura nigrovittata (Orthoptera: Phaneropteridae)

The song of the male bushcricket Ancistrura nigrovittata consists of a sequence of verses. Each verse comprises a syllable group, plus, after about 400 ms a single syllable serving as a trigger for the female response song. The carrier frequency of the male song spectrum peaks at around 15 kHz, while the female song peaks at around 27 kHz. The thresholds of female responses to models of male songs are lowest for song frequencies between 12 and 16 kHz and therefore correspond to the male song spectrum. The threshold curve of the female response to the trigger syllable at different frequencies has the same shape as the tuning for the syllable group. Phonotactic thresholds of male Ancistrura nigrovittata to synthetic female responses at different frequencies are lowest between 24 and 28 kHz and thereby correspond to the female song spectrum and clearly differ from female response thresholds. Activity of the tympanic fibre bundle of both sexes is most sensitive between 15 and 35 kHz and therefore not specifically tuned to the partner's song. Individual behavioural thresholds have their minimum within 10 dB of the values of tympanic thresholds.     

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。     

The structure and function of songs emitted by southern green stink bugs from Brazil, Florida, Italy and Slovenia

Nezara viridula (L.) (Pentatomidae: Heteroptera) from Brazil, Florida, Italy and Slovenia, communicate by vibratory songs associated with long‐range calling and close‐range courting, rivalry and repelling. Each song is composed of spectrally and temporally different units. Spectrally different pulses of duration less than 300 ms are present in the male calling song. The female calling song is characterized by pulse trains composed of pulses shorter than 150 ms and pulse trains composed of a longer (> 700 ms) and shorter (< 250 ms) pulse. Shorter and longer pulses have different spectral characteristics. The male and female courtship songs are characterized by fusion of shorter (< 150 ms) pulses into a pulse train usually followed by a shorter (< 200 ms) postpulse in the case of the male courtship song. The female repelling song is a several seconds long vibration of irregular temporal structure. The short (< 400 ms) male rival song pulses are frequency modulated. The dominant frequency peaks of the songs investigated lie between 70 and 130 Hz. The dominant frequency and the microstructure of song spectra show no population specificity. The average duration varies more in calling than in courtship songs. The repetition time varies extensively in songs of different populations. Normal communication followed by copulation was observed between mates from Slovenia and Brazil and between mates from Florida and Italy. The potential role of different temporal and spectral parameters for species recognition and mate location is discussed in view of the expected distortion of the characteristic signal structure during transmission through plants.     

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.     

Paradoxical hypermasculinization of the zebra finch song system by an antiestrogen

Exogenous estrogens, when administered to hatchling female zebra finches, masculinize the morphology and function of their neural vocal control system. The first of two experiments evaluated whether tamoxifen citrate is an antiestrogen in zebra finches, and the second determined whether it would block the masculinization hypothesized to be caused in hatchling males by the males' endogenous estradiol. In the first experiment adult female zebra finches were ovariectomized and injected for 10 days with estradiol benzoate (EB), tamoxifen, EB and tamoxifen combined, or vehicle (control). The dependent variable was oviduct weight. The EB-stimulated growth of the oviduct was blocked by tamoxifen, which had no effects when administered alone. Thus, tamoxifen acts as an antiestrogen in the zebra finch oviduct. In Experiment 2, male and female zebra finches were treated with tamoxifen or vehicle for the first 20 days after hatching. The males were castrated at 20 days. At 60 days we compared the song control regions of experimental and control males and females. In both sexes tamoxifen increased the somatic areas of neurons in RA (robust nucleus of the archistriatum), HVc (caudal nucleus of the ventral hyperstriatum), and MAN (magnocellular nucleus of the anterior neostriatum). Tamoxifen also increased the volumes of HVc, RA, MAN, and Area X in males. Thus, tamoxifen failed to block masculinization of males, but masculinized females and hypermasculinized males. Tamoxifen's hypermasculinization of the male and masculinization of the female song system is paradoxical given that (1) estradiol does not have similar effects on the male song system, and (2) tamoxifen antagonizes the effects of EB in the oviduct.     

Einfluß der Konzentration der Zuckerlösung auf den Gesang und das Balzverhalten des Gelbbauchnektarvogels (Nectarinia venusta)*

Different concentrations of a sucrose solution vary the courtship song and behaviour of the male yellow-bellied sunbird Nectarinia venusta- the duration of subsong, total singing duration, and the absolute number of full song phrases. With high concentrations the sunbird sings more full song phrases but less subsong during the courtship season than otherwise. The various effects are described.     

Avian vocalisations: the female perspective

Research on avian vocalisations has traditionally focused on male song produced by oscine passerines. However, accumulating evidence indicates that complex vocalisations can readily evolve outside the traditional contexts of mate attraction and territory defence by male birds, and yet the previous bias towards male song has shaped – and continues to shape – our understanding of avian communication as a whole. Accordingly, in this review we seek to address this imbalance by synthesising studies on female vocalisations from across signalling contexts throughout the Aves, and discuss the implications of recent empirical advances for our understanding of vocalisations in both sexes. This review reveals great structural and functional diversity among female vocalisations and highlights the important roles that vocalisations can play in mediating female-specific behaviours. However, fundamental gaps remain. While there are now several case studies that identify the function of female vocalisations, few quantify the associated fitness benefits. Additionally, very little is known about the role of vocal learning in the development of female vocalisations. Thus, there remains a pressing need to examine the function and development of all forms of vocalisations in female birds. In the light of what we now know about the functions and mechanisms of female vocalisations, we suggest that conventional male-biased definitions of songs and calls are inadequate for furthering our understanding of avian vocal communication more generally. Therefore, we propose two simple alternatives, both emancipated from the sex of the singer. The first distinguishes song from calls functionally as a sexually selected vocal signal, whilst the second distinguishes them mechanistically in terms of their underlying neurological processes. It is clear that more investigations are needed into the ultimate and proximate causes of female vocalisations; however, these are essential if we are to develop a holistic epistemology of avian vocal communication in both sexes, across ecological contexts and taxonomic divides.     

Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)

Many gomphocerine grasshoppers communicate acoustically: a male's calling song is answered by a female which is approached phonotactically by the male. Signals and recognition mechanisms were investigated in Chorthippus biguttulus with regard to the cues which allow sex discrimination. (1) The stridulatory files on the hindfemur of both sexes are homologous in that they are derived from the same row of bristles, but convergent with respect to the “pegs”. In males the pegs are derived from the bristles, and in females from the wall of the bristle's cup. (2) Male and female songs are generated by similar, probably homologous motor programs, but differ in the duration, intensity, “gappyness” of syllables, risetime of pulses, and the frequency spectra. The hindleg co-ordination during stridulation and the resulting temporal song patterns are less variable in males than in females. (3) For both sexes, recognition of a mate's signal depends on species-specific syllable structure. For males it is essential that the female syllables consist of distinct short pulses, whereas females reject “gappy” syllables. Males strongly prefer “ramped” pulses, females respond to syllables irrespective of steeply or slowly rising ramps. Males react only to the low-frequency component, whereas females prefer spectra containing both, low and high frequency components. Accepted: 20 November 1996     

Acoustic behaviour of both sexes of the speckled bush cricket Leptophyes punctatissima

As in other phaneropterine bush-crickets, both male and female Leptophyes punctatissima can stridulate. Stridulation in the female, which had previously been overlooked, occurs in response to the male call. The main frequency of the very brief chirps is around 40 kHz. The male is stimulated to chirp more frequently by receiving response chirps from the female. Unlike most Tettigoniidae the male moves toward the female and is thus dependant on her response signal. Communication can occur over distances of at least 5 m. Some structures involved in stridulation in the female differ from that described in other phaneropterines. The insects used in these experiments were all reared in the laboratory from wild-caught parents. Between forty and fifty different insects were used at various times for laboratory studies on the song, and four different males and three females were used on the field communication experiments. The sexes were kept separate from the moult to adult or for at least 10 days before experiments.     

Male courtship song frequency as an indicator of male genetic quality in an insect species,Drosophila montana.

Most theoretical models on evolution of male secondary sexual characters and female preferences for these characters suggest that the male characters evolve in response to female preferences that may themselves evolve in response to direct or indirect benefits of choice. In Drosophila montana (a species of the D. virilis group), females use male song in their mate choice, preferring males that produce songs with short sound pulses and a high carrier frequency. We demonstrate here that the females get indirect benefits from their choice: in our data the frequency of the male song correlated with the survival rate of the male''s progeny from egg to adulthood (indirect benefit for the female), but not with the fecundity of his mating partner (no direct benefit for the female). Male wing centroid asymmetry did not correlate with male wing song characters, nor with female egg production nor the fitness of her progeny, suggesting that fluctuating asymmetry in male wings does not play a major role in sexual signalling. The fact that the male song gives the female information on the male''s condition/genetic quality in D. montana suggests that in this species the evolution of female preferences for male song characters could have evolved through condition-dependent viability selection presented in some ''good genes'' models.