SOUND PRODUCTION AND SOUND EMISSION IN SEVEN SPECIES OF EUROPEAN TETTIGONIIDS. PART II. WING MORPHOLOGY AND THE FREQUENCY CONTENT OF THE SONG

ABSTRACT

Sound production in seven species of bush crickets (Tettigonia cantans, T. virridissima, Decticus verrucivorus, D. albifrons, Psorodonotus illyricus, Ephippiger ephippiger, E. discoidalis) has been investigated. Aspects of wing morphology have been compared and show that areas of the dorsal fields and the mirror are correlated with the dominant frequencies of the songs. Tooth removal from the pars stridens produces gaps in the time structure of single syllables but no change in the song power spectra. The removal of the tegminal lateral field in long- and medium-sized wing species (T.c., T.v., D.a., D.v.) produces an increase in the ultrasonic components of caudally-emitted sound. This suggests an absorbing function for the lateral fields in intact animals. In all species removal of a small portion of the mirror frame or of the mirror membrane attenuates the whole stridulatory signal, but especially the ultrasonic components. The mirror therefore functions as an amplifier, especially for high frequencies. Manipulation of the dorsal fields of long- and medium-winged species, or the distal edges of tegmina of brachypterous species, deletes or shifts the songs' dominant frequency. Thus the different tegminal structures (and especially the dorsal fields) contribute to the time structures and power spectra of the stridulatory songs of these species.     

Behavioural adaptations of ground living bushcrickets to the properties of sound propagation in low grassland

Summary The acoustic behaviour of the closely related tetigoniid species Psorodonotus illyricus and Decticus verrucivorus have been invectigated by bioacoustical and behavioural methods. Both species show adaptations concerning the acoustic behaviour with respect to the biotope and the properties of sound propagation. These insects inhabit low grassland with an average vegetation height of about 20 cm which is also the general height for the song perches. Difficulties arise for efficient acoustic communication in such habitats. Sound propagation is influenced and limited by the strong ground attenuation and the excess damping by grass vegetation. Other limiting factors are the microclimatic conditions in the biotope. The two species counteract these difficulties by moving around in the biotope during stridulation. Both species mainly stridulate in the morning, avoiding problems of reduced sound transmission which often appear in the afternoon due to negative temperature gradients and resulting shadow zones. From the high mobility of these insects, it follows that individuals have no fixed territory and consequently no rivalry against conspecifics, which is very common among Orthopterans with a high degree of territoriality. It can be concluded that the preferred biotope influences and creates behavioural patterns in Orthopterans, especially here in the two investigated species of bushcrickets Psorodonotus illyricus and Decticus verrucivorus.     

SOUND PRODUCTION AND SOUND EMISSION IN SEVEN SPECIES OF EUROPEAN TETTIGONIIDS PART III. DETERMINATION OF THE MECHANISM OF SOUND PRODUCTION USING CEPSTRUM ANALYSIS

ABSTRACT

The mechanism of sound production in tettigoniids is examined by applying the method of ‘cepstrum’ analysis to insect calls. The power cepstrum is defined as the inverse Fourier transform of the logarithmic power spectrum. This analysis shows that the tettigoniid sound signal is a convolution in time of probably two components. The first is caused by the initial impact of teeth of the stridulatory file on the left wing against the plectrum on the right wing (termed the input pulse); the second is caused by the oscillating properties of the tegmina (these being a function of the intrinsic frequencies of dorsal fields and mirror and their damping properties). In the cepstrum each component appears as a varying number of peaks. The tooth impacts cause a very low quefrency peak probably representing the time in which the two tegmina are in contact during each impact and high quefrency peaks representing the impulse repetition rate. The oscillating properties of the tegmina cause two major quefrency peaks which can be clearly related to the size of the dorsal fields and of the mirror respectively, and therefore to their intrinsic frequencies. The high damping factor of the tegmina together with the transient shape of the tegminal input pulse causes a strong time limitation of the impulses and is therefore responsible for the broad frequency bands occurring in the power spectra of the tettigoniid songs. The impulse generation of a synthetic tettigoniid song is discussed.     

Acoustic Features Used in Song Discrimination by the Veery

To investigate which song features are used for discrimination by the veery (Catharus fuscescens), we synthesized songs by computer and manipulated or deleted various features. These songs were then played to territorial males in their natural habitat. Differences in response to these stimuli indicated the relative importance of the manipulated song components. We found that certain features must be maintained in order to elicit a territorial response similar to that elicited by the natural song. These features were the frequency changes between and within syllables, the dominant higher frequency band (or voice), the intra-syllable syntax, and the rapid repetitive amplitude and frequency modulations. Manipulations of inter-syllable syntax independent of frequency changes between syllables, broad changes in amplitude, and the lower frequency band, did not have a significant effect on response. We thus concluded that they are not essential as cues for discrimination or species identification. Re-recordings of songs broadcast along transects indicate that song components used in encoding species identity are those which transmit well across veery territories.     

Variation in male wing song characters inDrosophila plantibia (Hawaiian picture-wingedDrosophila group)

The males of most species of the HawaiianDrosophila planitibia group produce songs when vibrating their wings during courtship. In four of the most recently evolved species,D. differens, D. planitibia, D. heteroneura, andD. silvestris, these songs are simple in structure and possess a higher and more variable carrier frequency than the songs of the more ancestral species of the group. In the present paper, we studied the variation in wing song production and song characters in aD. planitibia population. Some males vibrated their wings at a very low amplitude or slow speed, producing no detectable sound. Other males produced sound bursts varying in carrier frequency and burst length. The carrier frequency of the song depended mainly on the wing posture of the male during wing vibrations and was consistent for individual males. Variation between males of different isofemale progenies was not significant in any measured song trait. Songs of the males recorded in the present study differed, however, from songs of the males of a laboratory strain recorded earlier.     

蚱蝉(Cryptotympana atrata Fabricius)自鸣声音色的变化

蚱蝉自鸣声的音色分为单音色、双音色.及三音色等.本文进一步阐明每种音色的变化及高幅值脉冲对主音色能量的影响.蚱蝉自鸣声音色的变化主要是指频谱主音色频率(MTF)的显著改变、蚱蝉单色自鸣声的MTF主要在4.1—5.8kHz的频带内变化,双音色自鸣声的主次音色频率有相互颠倒现象,MTF主要在3.6—5.4kHz之间,三音色自鸣声的MTF虽然在3.5—4.5kHz比较窄的频带内,但三个音色峰的能量十分接近显示了三种音色成分.同只蚱蝉自鸣声,在不同的鸣声段具有近似相等的最大幅值,但高幅值脉冲个数的多少不同,相应主音色能量的大小与这些脉冲个数的多少对应.     

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.     

Selective phonotaxis in Tettigonia cantans and T. viridissima in song recognition and discrimination

The selectivity of female phonotaxis in Tettigonia cantans and T. viridissima was investigated on a Kramer treadmill, with respect to the specific differences in temporal pattern and spectrum of the songs of both species. In choice situations, both species preferred the conspecific song over the heterospecific one. The courses of both species were deflected by about 15–20° from the position of the conspecific song, that of T. viridissima being away from, that of T. cantans in the direction of the heterospecific song. In no-choice situations, song models with the temporal pattern of T. cantans did not attract T. viridissima. Models with the conspecific time pattern but heterospecific spectrum were as attractive as the conspecific model. In contrast, T. cantans was attracted by T. viridissima song presented alone. In choice situations, either spectral or temporal differences were sufficient for discrimination. The preference for the conspecific model gradually disappeared with stepwise reduction of its intensity and was reversed at −12 dB. Acoustic communication alone can serve species isolation in T. viridissima; however, premating isolation in T. cantans must involve other mechanisms. The orientation during the choice situations suggests a serial processing of song recognition and localization for the Tettigonia species. Accepted: 13 December 1997     

蝉的变音调复合声和发声机制的分析

蛙鸣蝉(Meimuna opalifera (Walk).ab.punctata Kato)的自然鸣声为“ji…guái”的重复单变调复合声.“ji”为主音频约4800Hz的准单音;“guái”的波形和主音频呈明显的演变,优势主频约2100Hz和2800Hz的变音调声.鸟鸣蝉(Meimuna opalifera (Walk)var.formosana Kato)的自然鸣声由重复的“jiū…ruǎ”和“jiū…gū…”合成的双变调复合声.“jiū”为基频和主频分别约625Hz和2100—2300Hz的准单音;“ruǎ”的波形和主音频呈明显的演变,基音和优势主频分别约575—625Hz和1550—1750Hz的变音调声.“gū”为优势主频约625Hz的准单音.变音调复合声不仅与腹部运动有关,而主要取决于发声肌的收缩特性和发声膜肋结构的振动特性.     

Calling song and selective phonotaxis in the field crickets,Gryllus firmus andG. pennsylvanicus (Orthoptera: Gryllidae)

The field cricket species, Gryllus firmusand G. pennsylvanicus,occur in a mosaic hybrid zone that roughly parallels the eastern slope of the Appalachian mountains in the northeastern United States. It is important to know what role, if any, the calling song plays in mate choice in sympatric and allopatric populations. In this report, we present results on the variability of calling song properties along transects across this hybrid zone. We also present the results of experiments on phonotactic selectivity of females from an allopatric population of G. firmus.The male calling song of allopatric G. firmuswas significantly slower in temporal rhythm (i. e., chirp and pulse repetition rates) and lower in pitch (i.e., dominant frequency) than that of allopatric G. pennsylvanicus.Calling song properties of males recorded in the hybrid zone varied considerably in temporal and spectral properties. In two-stimulus (choice) phonotaxis experiments, allopatric females of G. firmuspreferred synthetic calling songs with conspecific pulse repetition rates over songs that had lower and higher pulse rates. This preference persisted even when the sound pressure levels of alternative stimuli were unequal. Therefore, allopatric females of G. firmuscan discriminate between conspecific and heterospecific calling songs. Whether or not this same selectivity is present in sympatric populations remains unclear. Investigations of phonotactic selectivity in other allopatric and sympatric populations of both species are currently under way.     

Vibratory signals of the harlequin bug and their transmission through plants

Abstract. Males of the harlequin bug, Murgantia histrionica (Hahn), produce five different vibrational songs, whereas females produce one song. Songs differ from those of other stink bugs primarily in their species‐specific temporal characteristics. The broad band male courtship songs of M. histrionica are achieved by a combination of different frequency modulated and/or narrow band subunits, with several higher harmonic frequencies. Males rather than females initiate substrate‐borne vibrational communication, and the longer‐range calling songs found typically in other pentatomid species are lacking. Interindividual differences in song temporal and spectral characteristics are discussed. Transmission of vibrational songs through a cabbage head is more efficient along veins than along lamina. Attenuation of signals transmitted through veins is low and similar to that reported previously for plant stalks. On the leaf vein, distances between peak amplitude minima and maxima are different for the dominant and subdominant frequencies. At any distance from the vibration source, a different relationship between spectral peak amplitudes can be recorded. Resolution of these differences, together with velocity differences between signals recorded on the vein and lamina, may help small stink bugs to estimate distance and to locate each other on a plant.     

THE ROLE OF COURTSHIP SONGS IN REPRODUCTIVE ISOLATION AMONG POPULATIONS OF GREEN LACEWINGS OF THE GENUS CHRYSOPERLA (NEUROPTERA: CHRYSOPIDAE)

Male and female lacewings tremulate during courtship, establishing duets that always precede copulation. Three distinct courtship songs are found in populations of the green lacewing Chrysoperla plorabunda (P1, P2 and P3 song morphs). Analysis of five features of the songs for individuals collected from Connecticut, Idaho, Oregon and California showed few differences within song morphs, but sympatric song morphs differed significantly in temporal features of the songs and their mode of presentation. Playback experiments using recorded songs were performed on females with all possible sympatric and allopatric combinations of females and recorded songs. The results showed that females strongly prefer to duet with recordings of males of their own song type and usually showed no responses to songs of other types. Thus, song differences are effective barriers to reproduction between the sympatric morphs. Our results support the hypothesis that the three song morphs are true biological species.     

Extremely divergent song types in the genus Aerotegmina Hemp (Orthoptera: Tettigoniidae: Hexacentrinae) and the description of a new species from the Eastern Arc Mountains of Tanzania (East Africa)

The genus Aerotegmina comprises a group of African canopy-dwelling, predatory bushcrickets (katydids) that is known for its inflated tegmina and the unusually loud and low-frequency calling song of its type species. Here, we describe the songs of another two species of the genus which are much larger than the type species and have an even lower peak frequency. In addition, small and large species differ in song structure. While the type species and closely related forms have a broad banded, multi-peak song with strong components in the audio and ultrasonic range, the species treated here have an extremely narrow banded, resonant song with a carrier frequency of 2 kHz. With Q-values above 100 these sounds belong to the purest songs ever recorded in Orthoptera. Besides describing songs, stridulatory organs and a new species we discuss the possible functions of the song type. A key to the species of Aerotegmina is provided.     

Behavioral Sequence Leading to Sexual Isolation Between <Emphasis Type="Italic">Drosophila ananassae</Emphasis> and <Emphasis Type="Italic">D. pallidosa</Emphasis>

Drosophila ananassae and D. pallidosa are closely related, sympatric species that lack postmating isolation. Sexual isolation has been considered important in maintaining them as independent species. To clarify the behavioral processes leading to sexual isolation, we analyzed behavioral sequences and examined the effect of courtship song on mating success and on behaviors of both sexes by surgically removing male wings (song generators), female aristae (song receivers), or female wings (means of fluttering). We found that heterospecific courtship songs evoked female wing fluttering, whereas conspecific courtship song did not. Furthermore, female wing fluttering made courting males discontinue courtship. These findings suggest that strong sexual isolation is achieved through the following behavioral sequence: heterospecific song→female wing fluttering→courtship discontinuation.     

Comparison of the physiology of the auditory receptor organs in Gryllus bimaculatus and Ephippiger ephippiger: CSD Recordings within the auditory neuropiles

The syllables of the song of the tettigoniid Ephippiger ephippiger consist of a series of short sound impulses with a broad-banded frequency spectrum. Syllables of the song of the gryllid species Gryllus bimaculatus are nearly pure tones with sharply tuned frequency maxima. A comparison of the physiology of the audiotory receptor organs of both species was carried out by using acoustical stimuli with different carrier frequencies and time-amplitude patterns. The neuronal ensemble activity of receptor cell groups of the tympanal organ was measured within the prothoracic ganglion using the CSD technique. In E. ephippiger, response maxima were found at carrier frequencies mirroring the broad frequency content of the conspecific song. The receptor cells of E. ephippiger are highly sensitive to transient sound impulses. In G. bimaculatus, the receptor cell population is more sharply tuned to the basic frequencies of the natural songs; pure tones represent more effective stimuli than transient sound signals. The causes for these speciesspecific differences are discussed with regard to probable adaptations of the receptor organs to the parameters of the conspecific songs. © 1993 John Wiley & Sons, Inc.     

Local habitat complexity correlates with song complexity in a vocally elaborate honeyeater

Song complexity is an important behavioural trait in songbirds, subject to sexual selection. Elucidation of intraspecific variation in song complexity can provide insights into its evolution. In this study, we investigated song complexity variation in tūī (Prosthemadera novaeseelandiae), a vocally complex songbird endemic to New Zealand. At two separate nature reserves, we recorded male songs in two habitat types: forest remnants with high habitat complexity, and open habitats with lower habitat complexity. Analyses indicated strong evidence that song complexity was higher in forest habitats. Possible explanations for this divergence include: (i) competition between individuals results in higher quality, dominant males with more complex songs occupying forest habitats, and less competitive males occupying open habitat zones; (ii) forest habitats provide more abundant resources therefore higher tūī density, resulting in more complex songs; and (iii) a higher abundance of food in dense forest habitats may reduce nutritional stress during development resulting in full development of song nuclei. However, these hypotheses on the drivers of habitat effects on tūī song complexity remain to be tested.     

Causes of the latitudinal gradient in birdsong complexity assessed from geographical variation within two Himalayan warbler species

Aspects of birdsong complexity, such as the number of distinct notes in a song, commonly increase along latitudinal gradients, a pattern for which at least 10 explanations have been suggested. In two Himalayan warblers, songs are more complex in the northwest than in the southeast. In Grey‐hooded Warbler Phylloscopus xanthoschistos, high complexity results from increased note diversity within song types, sung across a higher bandwidth. In Blyth's Leaf Warbler Phylloscopus reguloides, high complexity is a consequence of increased variation between song types. The hypothesis with strongest support is that songs evolved to be more complex in species‐poor, demonstrably less noisy environments. We consider geographical variation to be an outcome of sexual selection favouring complexity across environments, where detection of the signal varies. Sexual selection favouring complexity may be resolved in different ways, because complexity has multiple features (repertoire size, song switching, etc.). We argue this has led to the great diversity in song that we have documented among five Phylloscopus species.     

CALLING SONG DISPLACEMENT IN A ZONE OF OVERLAP AND HYBRIDIZATION

The ground crickets Allonemobius fasciatus and A. socius meet in a mosaic zone of overlap and hybridization stretching from the East Coast to at least Illinois. To test whether male calling song differences were enhanced in sympatry, we analyzed the songs of crickets from inside and outside the zone of overlap along two transects. No evidence of calling song displacement was found in A. socius males from populations within the zone of overlap. On the other hand, A. fasciatus displayed calling song displacement in three populations. Our results are consistent with the hypothesis that the selective pressure exerted by the challenge from a related species is frequency dependent. While not a conclusive demonstration, the observed shifts in calling song are strongly suggestive of reproductive character displacement.     

Temporal patterns of intra- and interspecific acoustic signals differ in two closely related species of Acanthoplus (Orthoptera: Tettigoniidae: Hetrodinae)

Males of the closely related African tettigoniids Acanthoplus discoidales and Acanthoplus longipes produce a long-lasting calling song and a short disturbance sound. The temporal patterns of the sounds were analysed in respect to species differences and song type differences. The calling songs of both species consist of impulses which are separated into verses of two syllables, with fewer impulses in the first syllable. A. longipes produces more impulses in each syllable than A. discoidales and has longer verse durations, verse intervals and syllable intervals. Also, the disturbance sounds, produced after mechanical stimulation, contain distinct verses of impulses. The disturbance sound of A. longipes has a higher number of impulses per verse than that of A. discoidales. The frequency spectra of the songs in both species have similar peak frequencies (around 12.5 kHz) and both species have their greatest hearing sensitivity in the range between 5 and 10 kHz. Females of A. longipes perform phonotaxis only to songs with a species-specific temporal pattern. By contrast, females of A. discoidales react positively to calling songs of both species.