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.     

SOUND PRODUCTION AND SOUND EMISSION IN SEVEN SPECIES OF EUROPEAN TETTIGONIIDS. PART I. THE DIFFERENT PARAMETERS OF THE SONG; THEIR RELATION TO THE MORPHOLOGY OF THE BUSHCRICKET

Abstract

Comparative studies of sound production and sound emission in seven species of European tettigoniids have been carried out. The species chosen were two Tettigoniines (Tettigonia cantans, Tettigonia viridissima), two Ephippigerines (Ephippiger discoidalis, Ephippiger ephippiger), and three Decticines (Decticus albifrons, Decticus verrucivorus, Psorodonotus illyricus). The factors which determined the choice of species were the different morphology (for example body shape and weight, and wing size) of the three subfamilies. The parameters of the different songs (e.g. dominant frequency, intensity) are normally not correlated to any of the investigated morphological characteristics of the animals. In the brachypterous species intraspecific correlations exist between wing size and the dominant low frequency band of the call. This frequency band is also observable at related higher frequencies in the ultrasonic range (20–60 kHz), the observed band width increasing with frequency. Sound emission in all species is to some extent directional. This directionality is related to body size and wing structure. The song structure of the different species does not appear to be related to any observable characteristic of the habitat of die animals. A possible exception may be the song of Psorodonotus illyricus with a particularly low dominant frequency band. The phylogenetic development of the songs seems to be determined by relationships between the different species rather than to any factors contributed by the habitat.     

Characterization and Generation of Male Courtship Song in Cotesia congregata (Hymenoptera: Braconidae)

Background

Male parasitic wasps attract females with a courtship song produced by rapid wing fanning. Songs have been described for several parasitic wasp species; however, beyond association with wing fanning, the mechanism of sound generation has not been examined. We characterized the male courtship song of Cotesia congregata (Hymenoptera: Braconidae) and investigated the biomechanics of sound production.

Methods and Principal Findings

Courtship songs were recorded using high-speed videography (2,000 fps) and audio recordings. The song consists of a long duration amplitude-modulated “buzz” followed by a series of pulsatile higher amplitude “boings,” each decaying into a terminal buzz followed by a short inter-boing pause while wings are stationary. Boings have higher amplitude and lower frequency than buzz components. The lower frequency of the boing sound is due to greater wing displacement. The power spectrum is a harmonic series dominated by wing repetition rate ∼220 Hz, but the sound waveform indicates a higher frequency resonance ∼5 kHz. Sound is not generated by the wings contacting each other, the substrate, or the abdomen. The abdomen is elevated during the first several wing cycles of the boing, but its position is unrelated to sound amplitude. Unlike most sounds generated by volume velocity, the boing is generated at the termination of the wing down stroke when displacement is maximal and wing velocity is zero. Calculation indicates a low Reynolds number of ∼1000.

Conclusions and Significance

Acoustic pressure is proportional to velocity for typical sound sources. Our finding that the boing sound was generated at maximal wing displacement coincident with cessation of wing motion indicates that it is caused by acceleration of the wing tips, consistent with a dipole source. The low Reynolds number requires a high wing flap rate for flight and predisposes wings of small insects for sound production.     

BUZZING BEES: COMMUNICATION BETWEEN BUMBLE BEE SOCIAL PARASITES (HYMENOPTERA: APIDAE) AND THEIR HOSTS

ABSTRACT

Although sounds are produced by highly eusocial bees in a variety of contexts, their meaning and evolution are poorly understood. In this study we examined the communicative function of sound during dominance disputes in primitively eusocial bumble bees (Hymenoptera: Apidae). The interaction between sound production and dominance behaviour was studied in the context of parasitism by obligate bumble bee social parasites (Psythyrus spp.). Females of Psithyrus bohemicus and P. vestalis produced sound during dominance interactions in bumble bee host colonies. Parasites mauled and pushed host bees, after which they buzzed with their wings folded. The sounds were broad band, with a mean frequency of 820 Hz for the loudest harmonic produced by a P. bohemicus female. The frequency range of recorded sounds matched that of the vibratory stimuli to which isolated bumble bee workers responded. Buzzing by some bumble bee social parasites may assist them in advertising a position of dominance in egg-laying hierarchies established in the absence of pheromonal inhibition of ovarian development.     

A NEW ACOUSTIC SIGNAL OF THE FRUIT-FLIES DROSOPHILA SIMULANS AND D. MELANOGASTER

ABSTRACT

We found a new acoustic signal in Drosophila simulans (si) and D. melanogaster (me). It is a ‘rejection signal’ (RS) produced by adult males and young males and females in response to the courting behaviour of mature males who emit ‘pulse songs’ (i.e. love song: LS). It occurs most frequently in si, less in adults me except if the interacting males belong to different chemical morphs (i.e. temperate or equatorial population). There are no differences in the LS characteristics directed to various sexes and ages. The RSs produced by adult males or by young animals do not differ significantly either. They are emitted by neither virgin nor fecundated adult me females but a few times by virgin adult si females. The RS (like the LS) is a multipulse signal but intervals between pulses are about twice those of LS, around 90 ms for si and 80 ms for me. They are very irregular, as is the distribution of energy along the bandwidth mainly between 300 and 800 Hz for si and 200 and 600 Hz for me. The sound level of the RS is from 10 to 20 dB less than the LS. The RS seems to be linked to the ‘flicking’ behaviour produced by both wings, while the LS always corresponds to the so-called ‘wing vibration’.     

Revised concept of Chrysoperla mediterranea (Hölzel), a green lacewing associated with conifers: courtship songs across 2800 kilometres of Europe (Neuroptera: Chrysopidae)

The Holarctic carnea-group of Chrysoperla green lacewings consists of numerous cryptic, sibling species, distinguished principally by vibrational courtship songs. A European member of this species-group, C. mediterranea (Hölzel), was originally diagnosed by its dark green coloration, red band across the front of the head, narrow wings, minute basal dilation of the pretarsus and ecological association with pines. Here we describe its courtship and mating song, which uniquely distinguishes the species from all other members of the carnea-group in Europe. Males and females both produce a single type of song, characterized by a long train of short, closely spaced volleys and nearly constant carrier frequency. Measured song features are nearly invariant across a large geographical range that extends well outside that previously known for C. mediterranea. Principal components analysis confirms the uniqueness of the mediterranea song in comparison with five other sympatric song species of the carnea-group. Populations which show the mediterranea song phenotype do not necessarily retain all of the traditional morphological features of the species, particularly in eastern parts of the range. Specifically, the green coloration may be less intense, the wings broader and the pretarsal basal dilation larger, resulting in significant morphological overlap with other song species. Larval morphology and ecophysiology also fail to distinguish C. mediterranea from other song species. Based on unique song phenotype, however, we conclude that C. mediterranea is a valid biological species, which can also be recognized in museum collections when a suite of physical attributes is used.     

A NEW SOUND TRAP,AND OBSERVATIONS ON THE MIGRATORY AND ACOUSTIC BEHAVIOUR OF THE FIELD CRICKET TELEOGRYLLUS COMMODUS (WALKER)

ABSTRACT

A new sound trap for capturing male and female crickets attracted to the calling song of the male is described. Major advantages of the trap are its portability and effectiveness. It was designed to operate in conjunction with a previously described cricket-song generator. The trap is compared with other designs and the acoustic and migratory behaviour of crickets revealed by sound trapping are discussed.     

SONG MIMICRY AND SPECIES STATUS OF THE GREEN WIDOWFINCH VIDUA CODRINGTONI

Summary

Payne, R. B., Payne, L. L. &; Nhlane, M. E. D. 1992. Song mimicry and species status of the Green Widowfinch Vidua codringtoni. Ostrich 63:86-97.

The Green Widowfinch Vidua codringtoni mimics the songs of the Redthroated Twinspot Hypargos niveoguttatus, its apparent foster species, in Zimbabwe, Zambia and Mala?i. Of 38 male Green Widowfinches, 37 mimicked the twinspot; one mimicked a firefinch and not the twinspot. A wild-captured juvenile developed mimicry of twinspot song after six months and retained its mimicry through the next year. Male Green Widowfinches have glossy breeding plumage (green to blue), black wings, white bill and bright orange feet, a colour combination unlike other widowfinches in their range in southern Africa. Females are marginally distinguishable from other widowfinch species. Green Widowfinches occur together locally with Black V. funerea nigerrima, Purple V. purpurascens and Steelblue V. chalybeata Widowfinches and do not interbreed with them. The four species of widowfinches in southern Africa each have distributional ranges within the limits of their foster species.     

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.     

ACOUSTIC COMMUNICATION DISTANCES OF A GOMPHOCERINE GRASSHOPPER

ABSTRACT

I studied in the laboratory and in the field the communication distance of the gomphocerine grasshopper Chorthippus biguttulus, which lives close to the ground in stony, short grass habitats. Using sound measurements and behavioural tests, I examined the frequency dependent attenuation and temporal degradation of the song pattern. Sound measurements (2–40 kHz) in the field revealed that excess attenuation increases with increasing frequency. Close to the ground, excess attenuation reaches 18 dB/m. On the basis of (i) excess attenuations, (ii) tympanic receptor threshold curves and (iii) song spectra, perception distances of female tympanic receptors for male songs and vice versa were calculated. Behavioural tests revealed that, especially in short and stony grass vegetation, the typical habitat of Ch. biguttulus, the temporal song pattern is strongly degraded. Thus, in the laboratory and in tall grass vegetation, sound attenuation was found to limit the communication distance. In contrast, in short grass vegetation, degradation of the temporal pattern was found to limit the communication distance of males listening to female songs. I argue that the exploitation of fast amplitude modulations for song recognition restricts the acoustic communication of grasshoppers to short distances (1–2.2 m). In this respect it seems adaptive to stridulate at low intensities.     

Syringeal specialization of frequency control during song production in the Bengalese finch (Lonchura striata domestica)

Background

Singing in songbirds is a complex, learned behavior which shares many parallels with human speech. The avian vocal organ (syrinx) has two potential sound sources, and each sound generator is under unilateral, ipsilateral neural control. Different songbird species vary in their use of bilateral or unilateral phonation (lateralized sound production) and rapid switching between left and right sound generation (interhemispheric switching of motor control). Bengalese finches (Lonchura striata domestica) have received considerable attention, because they rapidly modify their song in response to manipulations of auditory feedback. However, how the left and right sides of the syrinx contribute to acoustic control of song has not been studied.

Methodology

Three manipulations of lateralized syringeal control of sound production were conducted. First, unilateral syringeal muscular control was eliminated by resection of the left or right tracheosyringeal portion of the hypoglossal nerve, which provides neuromuscular innervation of the syrinx. Spectral and temporal features of song were compared before and after lateralized nerve injury. In a second experiment, either the left or right sound source was devoiced to confirm the role of each sound generator in the control of acoustic phonology. Third, air pressure was recorded before and after unilateral denervation to enable quantification of acoustic change within individual syllables following lateralized nerve resection.

Significance

These experiments demonstrate that the left sound source produces louder, higher frequency, lower entropy sounds, and the right sound generator produces lower amplitude, lower frequency, higher entropy sounds. The bilateral division of labor is complex and the frequency specialization is the opposite pattern observed in most songbirds. Further, there is evidence for rapid interhemispheric switching during song production. Lateralized control of song production in Bengalese finches may enhance acoustic complexity of song and facilitate the rapid modification of sound production following manipulations of auditory feedback.     

EVIDENCE FOR A HYDRODYNAMIC MECHANISM OF SOUND PRODUCTION BY COURTING MALES OF THE NOTCHTONGUE GOBY,BATHYGOBIUS CURACAO (METZELAAR)

ABSTRACT

Male gobies of the genus Bathygobius are soniferous during courtship. The mechanism by which the sounds are produced is, however, unknown. Early studies on sound production by males of B. soporator suggested that these sounds are hydrodynamic in nature, being produced by the forcible ejection of water through the gill opening. The mechanism of sound production by the closely related species B. curacao was investigated and three lines of evidence are presented which support the hydrodynamic hypothesis. First, similarities between the sounds produced by courting males and by ejecting water through a pipette demonstrated that hydrodynamic forces can readily produce such sounds. Second, the behavioural motor patterns occurring during sound production are consistent with the hypothesis that water is being ejected through the gill openings. Finally, morphological examination revealed an apparent lack of specialised features associated with sound production, effectively eliminating stridulatory and swim bladder mechanisms. These results represent the strongest evidence to date in support of any proposed mechanism of sound production in the gobiids.     

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.     

INDIVIDUAL AND SPECIES-SPECIFIC SONG PATTERNS OF ROCK AND MEADOW PIPITS: PHYSICAL CHARACTERISTICS AND EXPERIMENTS

ABSTRACT

The songs of Anthus spinoletta littoralis and A. pratensis, in sympatric populations in SW Sweden were recorded, analysed in sound spectrographs and tested in the field. The purpose was to investigate: a) if the strophes are species specifically and individually distinct and, if so, whether pipits are able to make such distinctions; b) if the species-specific and individual-specific patterns are concentrated in different sections of the song and, if so, whether the species-specific section transmits individual-specific information and the individual-specific section transmits species-specific information as well.

The basic structure of the syllables in the terminating section of the strophe was found to be intraspecifically stereotyped, but varied between the two species. Replay experiments showed that territorial males could discriminate between con- and heterospecific strophes and this ability persisted when the terminating section of the strophe was replayed alone. The terminating part of the strophe did not seem to transmit individual-specific information to conspecific males. On the other hand, the basic structure of the syllables in the first section of the strophe was intraindividually stereotyped but varied between most conspecific males. Playback experiments showed that a territorial male could discriminate his neighbours' song patterns and this ability remained intact when the introductory phrase of the strophe was replayed alone. Both song pattern and position were necessary properties of the identifying signal in these pipits. But the introductory phrase of a total stranger did not seem to transmit unambiguous species-specific information to conspecific males.     

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.     

SOUND PRODUCTION BY THE LUSITANIAN TOAD FISH,HALOBATRACHUS DIDACTYLUS

ABSTRACT

Several batrachoidids have been known to produce sounds associated with courtship and agonistic interactions, and their repertoires have been studied acoustically and behaviourally. In contrast, sound production of the Lusitanian toadfish Halobatrachus didactylus, although often noted, has not been acoustically studied.

This sedentary predator of Northeastern Atlantic coastal waters is usually found in sandy and muddy substrates, under rocks or crevices. Sound recordings were made in Ria Formosa, a lagoon complex in southern Portugal. The sound producing apparatus was studied in adult individuals of both sexes captured by local fishermen.

It is shown that this species produces acoustic emissions similar to other batrachoidids. It produces a long, rhythmical, tonal sound, often in choruses, which is comparable to the boatwhistle or hum signals of Opsanus and Porichthys, and a complex of signals that were classified as grunts, croaks, double croaks and mixed calls (‘grunt-croak’). As in other toadfishes, H. didactylus presents sonic muscles connected to a bi-lobed swimbladder. Asynchronous contractions of the sonic muscles were detected when massaging the ventral surface of the fish.     

SOUND PRODUCTION IN THE LARVAE OF GEOTRUPES SPINIGER (MARSHAM) (COLEOPTERA: GEOTRUPIDAE)

ABSTRACT

This paper focuses on sound production in the larvae of Geotrupes spiniger (Marsham) (Coleoptera: Geotrupidae). A single stridulatory apparatus was identified and then analysed from a structural point of view. Sound was studied using spectography and results included a quantitative analysis with characterizing physical parameters. Results highlight the need to study stridulation in Geotrupidae more in detail, to improve our understanding of its evolutionary and biological meaning.