The acoustic behaviour of the bushcricket Tettigonia cantans I. Behavioural responses to sound and vibration

The responses of $\text{Tettigonia cantans}$ to species-specific song and substrate conducted vibration have been tested under three experimental regimes.When directional cues are adequate the sound source can be located by audition alone; where such cues fail to provide sufficient information then vibration signals become of value. In two-choice experiments vibrating substrates are preferred by both males and females moving towards the sound source. In multiple-choice experiments Tettigoniids may either detect and follow vibration gradients or the direction of the bending waves coming from the vibration source to orientate towards a conspecific.     

Neuronal basis for spectral song discrimination in the bushcricket Tettigonia cantans

Previous experiments demonstrated that in choice situations female Tettigonia cantans prefer signals with the conspecific spectrum over signals with the spectrum of the sibling species T. viridissima, whose signals lack the T. cantans-typical 8-kHz component but are otherwise identical in spectral composition. The underlying neuronal mechanisms were investigated using whole-nerve recordings of the tympanal nerve. Hearing thresholds did not differ significantly between the species in the range between 7 and 25 kHz. The responses of the tympanic nerve at 30 dB above threshold did, however, differ significantly between the species. While in T. viridissima response amplitudes did not differ between 7 and 12 kHz, in T. cantans responses at 8 kHz were significantly larger than at 10 and 12 kHz. Since possible influences of axonal diameters on response magnitudes were excluded, these results indicate that in T. cantans more receptor cells are tuned to 8 kHz than to 10–12 kHz, while in T. viridissima tuning of receptor cells is more evenly distributed. The higher response magnitude in T. cantans at 8 kHz than at 10–12 kHz is equivalent to an amplitude difference of 3.8–7 dB, which agrees well with previous behavioural estimates. Accepted: 13 January 1999     

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.     

Influence of vibratory signals on the phonotaxis of the gryllid Gryllus bimaculatus DeGeer (Ensifera: Gryllidae)

Summary Biophysical measurements exhibit that males of the ground-living gryllid Gryllus bimaculatus DeGeer (Ensifera: Gryllidae) produce vibratory signals while stridulating. Transmitted via the ground, these signals are perceiveable for conspecifics at a distance of a few dm. Experiments on the phonotactic behaviour of females of this species reveal that vibratory signals do influence this phonotactic behaviour pattern when no other directional cue is available. The significance of vibratory signals for the orientation of the gryllid in the biotope is discussed.     

The effect of vibratory disturbance on sexual behaviour of the southern green stink bug Nezara viridula (Heteroptera, Pentatomidae)

The effect of vibratory disturbance on sexual behaviour and substrate-borne sound communication of the southern green stink bug, Nezara viridula L. was studied. Disturbance signals do not change the time N. viridula males need to locate the source of vibratory signals, but decrease the number of males responding with the calling and courtship song to calling females. Female N. viridula proceed calling during stimulation with disturbance signals but some of them change the song rhythm by skipping one or more signal intervals or emitting the repelling signals. The number of females which change the dominant frequency of the calling song decreases proportionally with increasing differences between the dominant frequency of the disturbance signals and the emitted female calling song. Variation of the song dominant frequency probably serves females to avoid interference by increasing the signal to noise ratio. Signal duration and repetition rate do not change significantly when the female is stimulated with the disturbance signals. This indicates that frequency shift by calling females is the main strategy for reducing interference by competitive signalers in N. viridula vibrational communication.     

Range of corixid sound signals in the biotope

ABSTRACT. The propagation of sound in the frequency band (2–12 kHz) used by the Corixidae was measured in two shallow natural ponds. At distances of more than 1 m from the shore the water was at least O.4 m deep. The first pond was eutrophic and contained no plants. The spread of sound into the open water obeyed approximately the geometric attenuation of the sound pressure level (SPL); a loss of 6dB for each doubling of the distance from the point sound source. Near the shore the attenuation was considerably greater, especially for low frequencies; for a 2 kHz signal the damping of the SPL was c. 40–50 dB/m.
The second pond had dense plant growth, and the sound attenuation depended strongly on the photosynthetic activity of the waterplants. Measurements in winter, with an overcast sky, revealed only a slight damping effect of the plants for a 10kHz test signal. During intense sunlight in summer, however, in addition to the geometric attenuation the damping effect of the plants over a distance of O.5 m was 50 dB for a 2 kHz signal and 80 dB for 10 kHz. This effect was due to gas bubbles produced during intense photosynthesis.
Song A of Corixa dentipes Thms. (Heteroptera) males elicits usually a response by male conspecifics. The threshold SPL for this response was measured to be c. 40 dB lower than the SPL at a distance of O.1 m from a stridulating animal. From the measurements of sound propagation it follows, therefore, that the effective range of Song A in the most favourable case is at least 10m, though in a pond overgrown with plants it can be less than O.4 m.     

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     

Predatory bug <Emphasis Type="Italic">Picromerus bidens</Emphasis> communicates at different frequency levels

The Asopinae (Heteroptera: Pentatomidae) are a subfamily of stinkbugs with predaceous feeding habits and poorly understood communication systems. In this study we recorded vibratory signals emitted by Picromerus bidens L. on a non-resonant substrate and investigated their frequency characteristics. Males and females produced signals by vibration of the abdomen and tremulation. The female and male songs produced by abdominal vibrations showed gender-specific time structure. There were no differences in the temporal patterns of male or female tremulatory signals. The signals produced by abdominal vibrations were emitted below 600 Hz whereas tremulatory signals had frequency ranges extending up to 4 kHz. Spectra of male vibratory signals produced by abdominal vibrations contained different peaks, each of which may be dominant within the same song sequence. Males alternated with each other during production of rivalry signals, using different dominant frequency levels. We show that the vibratory song repertoire of P. bidens is broader than those of other predatory stinkbugs that have been investigated. The emission of vibrational signals with different dominant frequencies but the same production mechanism has not yet been described in heteropteran insects, and may facilitate location of individual sources of vibration within a group.     

Context Specific Signaling with Different Frequencies - Directed to Different Receivers? A Case Study in <Emphasis Type="Italic">Gonatoxia</Emphasis> Katydids (Orthoptera<Emphasis Type="Italic">,</Emphasis> Phaneropteridae)

In katydids (Orthoptera: Tettigonioidea) of the subfamily Phaneropterinae females ready to mate initiate a duet, announcing her position to the male singer, but also potentially to eavesdropping rivals. In many species the male seems to defend the communication by adding self-produced imitations of a female response. If these signals occur within the male sensory time-window after the female song, they can disturb the orientation of rivals. In two species of the genus Gonatoxia, males and females use short, relatively narrow-banded sounds (width 2–7 kHz 10 dB below peak). Male song and female response, however, differ considerably in peak frequency. In G. maculata, the peak frequency of the last part of the male song (13 kHz) is between that of the first part (15 kHz) and the female response (9 kHz), in G. helleri the last part (9 kHz; assumed imitation) and the female song are identical in peak frequency and by a factor two lower than the first part (19 kHz). The male stridulatory file of this species is correspondingly modified and differs from all other members of the genus. The imitation of spectral properties of the female response is not known from any other katydid.     

The role of signals of different modalities in initiating vibratory communication in Nezara viridula

Signals of different modalities are involved in the behaviour of the green stink bug, Nezara viridula (L.) (Pentatomidae, Heteroptera). Long range attraction is mediated by male pheromones, resulting in aggregation of bugs on the same plant where vibratory signals, vision and various chemical signals become important. Both males and females sing spontaneously. When both are on the plant, males start vibratory communication as often as females. Females induce the exchange of vibratory signals spontaneously or triggered by the male pheromone while males initiate the duet either spontaneously or after seeing the female. Males and females sing spontaneously and respond to signals of different modalities more often in the daylight than in the dark. Long lasting autonomous emission of the female calling song is present when triggered by the male pheromone and males respond to female calling predominantly by the emission of the courtship song.     

Song recognition by temporal cues in a group of closely related bushcricket species (genus Tettigonia)

Female phonotaxis of Tettigonia viridissima and T. caudata was investigated on a walking compensator to determine the temporal parameters of the male song used for song recognition, and to compare them with the previously described pulse rate filtering of T. cantans. The T. cantans song is continuous with a ≈30-Hz pulse rate. The T. caudata song has a higher pulse rate (≈40 Hz) and duty cycle than T. cantans and a distinct verse structure. The T. viridissima song is continuous with a double-pulse pattern. While the pulse rate is essential for song recognition in T. cantans, neither pulse rate not verse structure were essential for song recognition in T. caudata: females responded to signals above a minimum duty cycle. T. viridissima females did not require the double-pulse structure, but a single long pulse, equivalent to the duration of the double pulses and interval between them, was effective. Song attractiveness was limited by a minimum duration of the merged double pulse, and by minimum and maximum duration of the interval between them. Pulse rate recognition had little if any importance in either of the species investigated. Thus, the three congeners use different mechanisms for temporal song recognition. Accepted: 18 June 1998     

Transmission and perception of acoustic signals in the desert clicker,Ligurotettix coquilletti (Orthoptera: Acrididae)

Males of the desert clicker, Ligurotettix coquilletti (Acrididae: Orthoptera) defend a femalerequired resource, the creosote bush Larrea tridentata, in desert habitats of the southwestern United States. Males signal acoustically to each other as well as to searching females. The call is produced by tegminal/femoral stridulation where one or both legs are used for sound production. Sound pressure levels, measured laterally, are influenced by the intervening tegmen between the stridulating leg and the microphone. Differences in measured sound pressure levels between sides can vary up to 7 dB. When clicks are produced multiply,these multiple clicks may be 4 dB louder than single clicks. We examine the structure of the call and the effective broadcast area of single males by monitoring acoustic ascending neurons of the ventral nerve cord in the neck. By taking the neurophysiological preparation into the field, we were able to map the broadcast area of isolated males and also of males calling within aggregations. The distance over which the signal of isolated males could be detected was 8–14 m, whereas neural representation of the calls of males within aggregation were detectable within 4–6 m. The sound spectrum of the song, although having a major lower-frequency component around 10 kHz, has extensive power in the ultrasonic range. The tuning characteristics of the ascending auditory neuron matched the overall structure of the male call. The importance of the acoustic cue, as compared to visual cues, is discussed in relation to female attraction.     

Auditory behaviour of a parasitoid fly (Emblemasoma auditrix, Sarcophagidae, Diptera)

Females of the parasitoid fly Emblemasoma auditrix find their host cicada (Okanagana rimosa) by its acoustic signals. In laboratory experiments, fly phonotaxis had a mean threshold of about 66 dB SPL when tested with the cicada calling song. Flies exhibited a frequency dependent phonotaxis when testing to song models with different carrier frequencies (pulses of 6 ms duration and a repetition rate of 80 pulses s(-1)). However, the phonotactic threshold was rather broadly tuned in the range from 5 kHz to 11 kHz. Phonotaxis was also dependent on the temporal parameters of the song models: repetition rates of 60 pulses s(-1) and 80 pulses s and pulse durations of 5-7 ms resulted in the highest percentages of phonotaxis performing animals coupled with the lowest threshold values. Thus, parasitoid phonotaxis is adapted especially to the temporal parameters of the calling song of the host. Choice experiments revealed a preference of a song model with 9 kHz carrier frequency (peak energy of the host song) compared with 5 kHz carrier frequency (electrophysiologically determined best hearing frequency). However, this preference changed with the relative sound pressure level of both signals. When presented simultaneously, E. auditrix preferred 5-kHz signals, if they were 5 dB SPL louder than the 9-kHz signal.     

THE ULTRASONIC SONG OF THE MOTH AMYNA NATALIS (LEPIDOPTERA: NOCTUIDAE: ACONTIINAE)

ABSTRACT

In Malaysia, males of the noctuid moth Amyna natalis were observed producing a continuous ultrasonic song of high intensity (about 102 dB SPL measured at a distance of 10 cm). The frequency spectrum of the sound impulses had its peak between 60 and 80 kHz. During song production the animals were perching on plants and moving their wings up and down quickly. Simultaneously, by twisting the wings it seems likely that a male-specific “bubble” in the forewing functions as a tymbal, resulting in sound production.     

Frequency as a releaser in the courtship song of two crickets,Gryllus bimaculatus (de Geer) and Teleogryllus oceanicus: a neuroethological analysis

1. The courtship behavior of male field crickets, Gryllus bimaculatus (De Geer) and Teleogryllus oceanicus, is a complex, multimodal behavioral act that involves acoustic signals (a courtship song; Fig. 1A,B). The dominant frequency is 4.5 kHz for T. oceanicus song (Fig. 1A) and 13.5 kHz for G. bimaculatus (Fig. IB).
2. When courting males are deprived of their courtship song by wing amputation, their courtship success declines markedly but is restored when courting is accompanied by tape-recordings of their courtship songs or a synthetic courtship song with only the dominant frequency of the natural song; other naturally occurring frequency components are ineffective for restoring mating success (Figs. 4, 5).
3. It has been suggested that an identified auditory interneuron, AN2, plays a critical role in courtship success. Chronic recordings of AN2 in an intact, tethered female show that AN2's response to the natural courtship song and synthesized songs at 4.5 and 13.5 kHz is similar in T. oceanicus. By contrast, in G. bimaculatus, AN2's response to the natural courtship song and synthesized song at 13.5 kHz, but not at 4.5 kHz, is similar (Figs. 2,3).
4. In behavioral experiments, playback of a 30 kHz synthetic courtship song in G. bimaculatus does not restore courtship success, yet this same stimulus elicits as strong a response from AN2 as does the normal courtship song (Fig. 6). Thus, contrary to earlier work by others, we conclude AN2 is not, by itself, a critical neural link in the courtship behavior of these two species of crickets.

     

Commensal auditory communication in two species of Neoconocephalus (Orthoptera)

Auditory communication in Neoconcephalus ensiger and Neoconocephalus robustus, two stridulating tettigoniids that inhabit the same geographical area, was examined to determine how these two species interact. Computer and electrophysiological techniques were used to analyze sound production and reception. Although similar in appearance, the males of these species produce easily distinguishable acoustic communication signals (call songs) that serve to attract conspecific females. Both wave form and spectral analyses were done on selected segments of the species-specific call songs by using discrete Fourier transform techniques. Sound production and reception capacities were measured by octave-band analyzers and extracellular electrode techniques. The results show that each of these species has a distinct, primary carrier frequency band. Secondly, the log magnitude spectra revealed a significant secondary component in the call song of N. robustus which corresponds to the primary energy band in the call song of N. ensiger. This overlap in acoustic signals and other findings suggest that males of N. ensiger might use the louder stridulation of the more metabolically active N. robustus to attract conspecific females over great distances. N. ensiger males have a sound production capacity like N. robustus, but conserve energy for soft, near-field signaling. The audiogram representing sensitivity of the tympano-receptor in the foreleg showed sound reception to be matched precisely with sound production curves.     

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.     

Directional hearing of a cicada: biophysical aspects

The directional hearing of male and female cicadas of the species Tympanistalna gastrica was investigated by means of laser vibrometry. The results show that the tympanic organs act as pressure difference receivers. This mechanism can produce left-right differences of more than 10 dB. The main acoustic inputs to the inner surfaces of the ears are the tympana, in males supplemented by the timbals, and by the third spiracles in females. In addition the hollow abdomen of males seems to play a minor role. Tympanic membrane input is the source of left-right differences in the tympanic vibration velocity at frequencies below 9 kHz in males and below 15–18 kHz in females. The input via the (contralateral) timbal in males is responsible for a null in vibration velocity appearing between 12 and 14 kHz when the sound is coming from the contralateral direction. The highest energy components of the calling song are found in this frequency range. The mechanical sensitivity of the ears depends upon the sex. At low frequencies males are about 10 dB more sensitive than females.     

Identification of mellein in the mandibular gland secretions of carpenter ants

The mandibular gland secretions of males of the ants $\text{Camponotus herculeanus, C. ligniperda}$, and $\text{C. pennsylvanicus}$ contain three major volatile substances. These compounds have been identified as 3,4-dihydro-8-hydroxy-3-methylisocoumarin (mellein or ochracin), methyl 6-methylsalicylate, and 10-methyldodecanoic acid. Mellein has not been isolated previously from animal sources.