Auditory pattern recognition and steering in the cricket Teleogryllus oceanicus

Male crickets Teleogryllus oceanicus (Le Guillou) produce a complex species‐specific calling song with phrases combining groups of single pulses (chirps) and groups of double‐pulses (trills) to attract females, which fly or walk towards singing males. In open‐loop trackball experiments, phonotactic steering responses to normal calling song phrases consisting of chirps and trills are strongest, suggesting that both components are necessary for maximal attractiveness. Sequences of just chirps or trills are less effective in eliciting phonotactic walking and steering. Split‐song paradigms are used to analyze the steering behaviour underlying orientation in more detail. The females' phonotactic steering reflects the alternating acoustic pattern of the split‐song paradigm. Analysis with high temporal resolution demonstrate, that even when the calling song is presented only from one side, the steering velocity and lateral deviation towards the song is modulated by steering events to single‐sound pulses. Therefore, pattern recognition, which integrates the structure of the song, appears not to be directly involved in the rapid steering response. This organization of phonotactic behaviour with a parallel processing of pattern recognition and steering is similar to other cricket species and may allow T. oceanicus females to steer transiently towards distorted song patterns as they occur in natural habitats.     

Acoustic communication in the trilling field cricket,Gryllus rubens (Orthoptera: Gryllidae)

Male field crickets produce calling songs that are used for attracting conspecific females for mating. Acoustic communication was studied in the trilling field cricket, Gryllus rubens.A trilled calling song consists of a continuous train of sound pulses, each of which was repeated at a stereotyped rate. Singing males were recorded at different temperatures. The pulse period of the calling song decreased with increasing temperature; there was less effect of temperature on pulse duration. Female phonotaxis was studied on a noncompensating, spherical treadmill. In two-stimulus (choice) playback experiments, females preferred the conspecific trilled song over the chirped calling song of a sympatric species, Gryllus fultoni.This preference persisted even when the song of G. fultoniwas 6 dB louder. Females also discriminated between synthetic trills having different pulse periods; females chose trills with the conspecific pulse period over trills having lower and higher pulse periods.     

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.     

Phonotaxis in flying crickets

The steering responses of three species of field crickets, Teleogryllus oceanicus, T. commodus, and Gryllus bimaculatus, were characterized during tethered flight using single tone-pulses (rather than model calling song) presented at carrier frequencies from 3-100 kHz. This range of frequencies encompasses the natural songs of crickets (4-20 kHz, Fig. 1) as well as the echolocation cries of insectivorous bats (12-100 kHz). The single-pulse stimulus paradigm was necessary to assess the aversive nature of high carrier frequencies without introducing complications due to the attractive properties of repeated pulse stimuli such as model calling songs. Unlike the natural calling song, single tone-pulses were not attractive and did not elicit positive phonotactic steering even when presented at the calling song carrier frequency (Figs. 2, 3, and 9). In addition to temporal pattern, phonotactic steering was sensitive to carrier frequency as well as sound intensity. Three discrete flight steering behaviors positive phonotaxis, negative phonotaxis and evasion, were elicited by appropriate combinations of frequency, temporal pattern and sound intensity (Fig. 12). Positive phonotactic steering required a model calling song temporal pattern, was tuned to 5 kHz and was restricted to frequencies below 9 kHz. Negative phonotactic steering, similar to the 'early warning' bat-avoidance behavior of moths, was produced by low intensity (55 dB SPL) tone-pulses at frequencies between 12 and 100 kHz (Figs. 2, 3, and 9). In contrast to model calling song, single tone-pulses of high intensity 5-10 kHz elicited negative phonotactic steering; low intensity ultrasound (20-100 kHz) produced only negative phonotactic steering, regardless of pulse repetition pattern. 'Evasive', side-to-side steering, similar to the 'last-chance' bat-evasion behavior of moths was produced in response to high intensity (greater than 90 dB) ultrasound (20-100 kHz). Since the demonstration of negative phonotactic steering did not require the use of a calling song temporal pattern, avoidance of ultrasound cannot be the result of systematic errors in localizing an inherently attractive stimulus when presented at high carrier frequencies. Unlike attraction to model calling song, the ultrasound-mediated steering responses were of short latency (25-35 ms) and were produced in an open loop manner (Fig. 4), both properties of escape behaviors.(ABSTRACT TRUNCATED AT 400 WORDS)     

Variation and Repeatability of Calling Behavior in Crickets Subject to a Phonotactic Parasitoid Fly

Male Teleogryllus oceanicus (Orthoptera: Gryllidae) produce a conspicuous calling song to attract females. In some populations, the song also attracts the phonotactic parasitoid fly Ormia ochracea (Diptera: Tachinidae). I examined the factors affecting calling song by characterizing the calling behavior of caged crickets from an area where the fly occurs. Calling activity (proportion of time spent calling) was repeatable and a significant predictor of female attraction. However, calling activity in the parasitized population was lower than in an unparasitized Moorea population (Orsak, 1988), suggesting a compromise between high activity to attract females and low activity to avoid flies. Calling activity peaked simultaneously with fly searching, so crickets did not shift to calling when the fly is less active. Males harboring larvae did not call less than unparasitized males; however, a more controlled study of the effects of parasitization on calling behavior is needed to evaluate this result. The results are discussed in the context of other studies of the evolutionary consequences of sexual and natural selection on cricket calling behavior.     

The effects of injecting Juvenile Hormone III into the prothoracic ganglion on phonotaxis by female crickets Gryllus bimaculatus

Nanoinjection of Juvenile Hormone III (JH III) into the prothoracic ganglion causes virgin female crickets Gryllus bimaculatus De Geer to become more phonotactically selective for the syllable periods (SPs) of model calling songs. Females responding to all, or almost all, of the SPs presented before JH III injection significantly narrow their responses to a range of SPs that is usually centered on the SPs included in the conspecific males' calling song. Control injections of acetone (i.e. the solvent for JH III) into the prothoracic ganglion do not significantly change the recipient females' phonotactic responses. Injection of JH III into the metathoracic ganglion also has no effect the females' phonotactic choices.     

Nanoinjection of neurotransmitters into the prothoracic ganglion of female cricket Acheta domesticus changes phonotactic selectivity

The phonotactic response by female crickets is influenced by Juvenile Hormone III, which affects selectivity to the syllable period of the calling song. This pathway is influenced by an inhibitory input in the prothoracic ganglion, possibly chloride-mediated inhibition. In order to identify potential neurotransmitters involved in such pathway, we performed nanoinjection of five neurotransmitters into the prothoracic ganglion of virgin female Acheta domesticus. Phonotaxis for these females was evaluated before and after injections. All five neurotransmitters that were nanoinjected are known to bind to chloride channels. Nanoinjection of histamine significantly decreased phonotactic selectivity to the syllable period of the calling song while glycine, gamma aminobutyric acid, serotonin and saline controls did not. Octopamine significantly decreased phonotactic responses overall. The effect of histamine was tested further by nanoinjecting the antihistamine pyrilamine into the prothoracic ganglion of older unselected females, which resulted in increased phonotactic selectivity.     

Phonotactic specificity of the cricketTeleogryllus oceanicus: intensity-dependent selectivity for temporal parameters of the stimulus

Summary We have investigated the effects of alterations of several temporal parameters of auditory stimuli, as well as of stimulus intensity changes, on the attractiveness of these stimuli to femaleTeleogryllus oceanicus, as measured by monitoring sound-elicited flight steering responses. AlthoughT. oceanicus has a rhythmically complex calling song, females are attracted by a simpler model consisting of regularly repeating sound pulses. We have found that the two major temporal features of this model, sound pulse duration and pulse repetition rate, are both important for eliciting phonotactic steering responses.Stimuli with altered temporal features had intensity thresholds indistinguishable from the control stimulus (Fig. 3). The majority of crickets, however, ceased to respond to the altered stimuli when the stimulus intensity was sufficiently increased (Figs. 4–7). In some cases, intensity increases resulted in a reversal of the steering response from positive to negative (Fig. 10). Effects of altered temporal parameters were also apparent at lower stimulus intensities, where the amplitudes of steering responses to stimuli with altered parameters were smaller than those in response to the control stimulus (Figs. 8, 9).We considered the possibility that the cessation of responsiveness to stimuli with altered temporal features was due to a temporal pattern-specific diminution of binaural cues for sound localization at high intensities. Experiments performed with unilaterally deafened crickets (Fig. 11) led us to conclude that this was not the case, and that our findings instead reflect the properties of the song recognition mechanism.Abbreviations UIL upper intensity limit - RAF ratio of abdominal flexion     

Phonotactic Behavior of Male Field Crickets (Gryllus texensis) in Response to Acoustic Calls From Conspecific Males

We studied male phonotactic behaviors elicited by acoustic cues that simulate conspecific male songs in the field cricket, Gryllus texensis. Males exhibited significant positive phonotaxis in response to the simulated song stimuli, but showed no such response to atypical song stimuli. We found no significant relationship between males’ own calling behavior and their phonotactic responses to the stimuli. Analyses indicated that larger males exhibited greater phonotactic responses, which may indicate a greater tendency to engage in aggressive interactions if size is an indicator of fighting ability. Male phonotactic responses were significantly weaker than those exhibited by females, and adult males did not exhibit stronger responses with increasing age as has been documented for females. Observed sex differences in the strengths of phonotactic responses may reflect differences in the fitness-payoffs of responding. That is, females are under strong selection pressure to respond to male songs and subsequently mate. In contrast, males responding to acoustic signals from other males need not precisely locate the signaler but would likely move to areas where females are likely to be found. Alternatively, males might benefit from avoiding areas with calling males and establishing their own calling stations away from competing males.     

The effect of ultrasound on the attractiveness of acoustic mating signals

Abstract. Previous laboratory studies ( Nolen & Hoy 1986b ) have shown that the phonotactic responses of flying crickets are influenced by the relative intensities of attractive (mating signal) and repulsive (predator) stimuli. At the functional level, these results suggest that predator cues (ultrasound) can change the attractiveness of a calling song. Using extracellular recordings from cervical connectives it was shown that, like other field crickets, Gryllus rubens (south-eastern field cricket) is sensitive to ultrasound. This ultrasonic sensitivity has probably evolved in response to predation pressure from echolocating bats. Using acoustic playback under field conditions, it was tested whether the relative attractiveness of two male calling songs was influenced by the simultaneous broadcast of ultrasound. A simulated male calling song of G. rubens was broadcast at two different intensities (109 and 103 dB) from two sound traps that caught flying crickets attracted to the songs. Simulated bat ultrasound was broadcast simultaneously with the high-intensity calling song (109 dB) and the relative catch in each of the two traps was measured. The intensity of the ultrasound was varied on different nights. The relative attractiveness of the high-intensity sound trap decreased significantly as the intensity of the ultrasound broadcast with it was increased. For the lowest of the ultrasound broadcast levels, the relative attractiveness did not differ from that expected for two calling songs broadcast without ultrasound. Thus, increased levels of simulated predation risk decreased the attractiveness of the calling song associated with it. These are the first field experiments to show that predation risk in the form of simulated bat ultrasound influences the phonotactic behaviour of flying crickets.     

蟋蟀对召唤声识别的趋声性定位行为

为了探究蟋蟀种内个体间的声音通讯原理,本研究通过回放利用Cool-edit2000声音采集软件采集的双斑蟋Gryllus bimachlatus召唤声,分别对雌、雄蟋蟀识别召唤声的趋声性定位行为进行了观测研究。实验结果表明:雌、雄双斑蟋对回放的召唤声均有较强的识别反应,但雌蟋蟀的趋声定位性较强,能在较短的时间内积极寻找到声源,路径成"s"型;雄蟋蟀反应相对较弱,路径较长,找到声源时间比雌蟋蟀长。本研究结果揭示召唤声在蟋蟀种内通讯中不仅对个体间的联系有作用,对异性求偶也有重要作用。     

Discrimination of calling song models by the cricket,Teleogryllus oceanicus: the influence of sound direction on neural encoding of the stimulus temporal pattern and on phonotactic behavior

Summary Recordings were made from an identified auditory neuron, the omega neuron, in the cricketTeleogryllus oceanicus. Models of the conspecific calling song and of the song of another species were presented either singly or simultaneously, and the degree to which the temporal pattern of the conspecific model was encoded in the neuron's spike train was determined. When a single stimulus was presented alone, its temporal pattern was faithfully reflected by the cells's spiking activity, no matter what the azimuth of the broadcasting loudspeaker (Fig. 3). When two stimuli were presented simultaneously from opposite sides, encoding of the pattern ipsilateral to the recorded neuron was interfered with only slightly by the contralateral pattern, as long as the two loudspeakers were sufficiently separated (Figs. 2, 3, 4). When the loudspeakers were each 15° from the cricket's midline, however, the encoding of the temporal pattern of the ipsilateral song model was severely disrupted (Figs. 3, 4). Bilateral interactions are important in determining the response level of the neuron, but do not appear to contribute to the direction-selective encoding of the stimulus temporal pattern (Figs. 5, 6).Phonotactic steering movements of tethered, flying crickets were recorded under stimulus conditions similar to those used in the neurophysiological tests. Under one-stimulus conditions, crickets attempted to turn towards the conspecific model for all tested speaker locations. The heterospecific model elicited reliable steering behavior when it was broadcast from azimuths of 90° and 60°, but often failed to elicit consistent responses when the speaker was positioned closer to the cricket's midline (Figs. 7, 8A and 8B). Responses to the heterospecific pattern were smaller in amplitude than those to the conspecific song model (Figs. 7, 8B). Under two-stimulus conditions, the conspecific model was consistently preferred over the heterospecific song for all tested speaker locations in half the tested crickets. In the remaining animals, preference for the conspecific pattern was only evident for the larger loudspeaker azimuths (Figs. 7, 8C).These results demonstrate that simultaneouslypresented stimuli can be represented separately in the nervous system as a consequence of auditory directionality. It is postulated that the cricket's ability to choose between these stimuli may result from the interactions between two bilaterallypaired song recognizers, each of which may be driven primarily by sound stimuli from one side.     

Sound localization in intact and one-eared crickets

Summary Intact and one-eared crickets,Gryllus bimaculatus, were tested for phonotactic behavior in a closed-loop and an open-loop situation and for related physiological characteristics of an identified auditory neuron pair, the left and the right AN2.Intact animals that performed phonotaxis in the closed-loop condition showed intended turning tendencies in the open-loop condition that correlated with the directional characteristics of their AN2s (Figs. 1–3).Animals in which one foreleg had been amputated during postembryonic development (one-eared regenerates) were classified according to their phonotactic performance as tracking or unoriented animals. The ability of one-eared regenerates to track a sound source was closely correlated with the direction of turning tendencies in the open-loop behavior and to specific features of their AN2 pair (Figs. 4–6).Some animals with one foreleg amputated as adults (one-eared amputees) perform stable phonotactic walking. Their open-loop behavior, however, is different from that of the tracking one- eared regenerates (Fig. 7).One-eared amputees showed stable phonotactic walking when calling song was presented from above and the sound intensity was varied according to the actual walking angle of the animal. The only orientational cue under this condition is the difference of sound intensity at different walking directions (Figs. 8–11).Different mechanisms are discussed for sound localization in one-eared regenerates and one- eared amputees based on turning tendencies which depend on the instantaneous stimulus intensity or on the intensity change between successive stimuli.     

Field Cricket Spacing,and the Phonotaxis of Crickets and Parasitoid Flies to Clumped and Isolated Cricket Songs

Nearest neighbor analyses of the field crickets Gryllus integer, G. veletis, and Teleogryllus oceanicus demonstrated that calling ♂♂ were aggregated. Broadcasts of conspecific song to calling ♂♂ indicated that attraction of neighboring ♂♂ maintained inter-male distances. Broadcasts of G. integer song through aggregated and isolated loudspeakers showed that the total number of crickets and parasitoid flies, Euphasiopteryx ochracea, attracted to aggregated loudspeakers was greater than that to an isolate. The average number of attracted crickets and flies in an aggregation was comparable to the isolated total.     

Cricket phonotaxis: localization depends on recognition of the calling song pattern

Summary Calling song with a carrier frequency of 5 kHz evokes positive phonotaxis in female crickets,Gryllus bimaculatus, when presented at an azimuth. In contrast, a continuous tone of 4.7 kHz in the same position when paired with calling song from above leads to negative phonotaxis. Under open-loop conditions, when a tethered animal runs on a paired tread wheel, characteristic curves are produced with the stable equilibrium point towards or away from the stimulus, respectively (Fig. 3).In order to understand this sign reversal at the neuronal level, directional characteristics of the ascending acoustic inter neurons AN1 and AN2 were measured using extracellular recordings from the cervical connectives.Taking the mean spike rate of the interneurons as a measure for their excitation, the function relating response magnitude to stimulus direction for calling song corresponds well to the behavioural characteristic curve (Fig. 5). The response function obtained using a continuous tone with simultaneous presentation of calling song from above is similar (Fig. 5) and hence does not correspond to the inverse behavioural characteristic curve.However, consideration of the extent to which the temporal parameters of the calling song (syllables and chirps) are reflected in the neuronal response (amplitudes of the Fourier components) leads to characteristic curves for AN1 and AN2 which are in good agreement with the behaviour for stimulation with calling song as well as for the continuous tone experiment (Fig. 8). In addition, the neural response curves correspond to the behaviour in showing smaller amplitudes when a continuous tone rather than the calling song is presented on the horizon (Fig. 8).From these data we conclude that the activity in interneurons AN1 and AN2 does not directly guide orientation in mating behaviour but first is filtered by a mechanism tuned to the frequency of syllables and/or chirps. According to this hypothesis recognition of conspecific song and localization proceed sequentially inGryllus.     

Phonotaxis in flying crickets

The effects of two-tone stimuli on the high frequency bat-avoidance steering behavior of flying crickets (Teleogryllus oceanicus) were studied during tethered flight. Similarly, the effects of two-tone stimuli on the ultrasound sensitive auditory interneuron, Int-1, which elicits this behavior, were studied using intracellular staining and recording techniques. When a low frequency tone (3-8 kHz) was presented simultaneously with an aversive high frequency tone (in a two-tone stimulus paradigm), the high frequency avoidance steering behavior was suppressed. Suppression was optimal when the low frequency tone was between 4 and 5 kHz and about 10-15 dB louder than the high frequency tone (Figs. 2, 3). Best suppression occurred when the low frequency tone-pulse just preceded or overlapped the high frequency tone-pulse, indicating that the suppressive effects of 5 kHz could last for up to 70 ms (Fig. 4). The threshold for avoidance of the bat-like stimulus was elevated when model bat biosonar (30 kHz) was presented while the animal was performing positive phonotaxis toward 5 kHz model calling song, but only if the calling song intensity was relatively high (greater than 70-80 dB SPL) (Fig. 1). However, avoidance steering could always be elicited as long as the calling song was not more than 10 dB louder than the ultrasound (Fig. 1). This suppressive effect did not require performance of positive phonotaxis to the calling song (Fig. 2) and was probably due to the persistence of the suppressive effects of the 5 kHz model calling song (Fig. 4). The requirement for relatively high intensities of calling song suggest that the suppression of bat-avoidance by the calling song is not likely to be of great significance in nature. The high frequency harmonics of the male cricket's natural calling song overlap the lower frequency range used by insectivorous bats (10-20 kHz) and are loud enough to elicit avoidance behavior in a flying female as she closely approaches a singing male (Fig. 5). The high frequency 'harmonics' of a model calling song were aversive even if presented with a normally attractive temporal pattern (pulse repetition rate of 16 pps) (Fig. 6A). When the 5 kHz 'fundamental' was added to one of the high frequency 'harmonics', in a two-tone stimulus paradigm, this complex model calling song was attractive; the high frequency 'harmonic' no longer elicited the avoidance behavior (Fig. 6) and the animals steered toward the model CS. Thus, addition of 5 kHz to a high frequency harmonic of the calling song 'masked' the aversive nature of this stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)     

The Steppengrille (Gryllus spec./assimilis): Selective Filters and Signal Mismatch on Two Time Scales

In Europe, several species of crickets are available commercially as pet food. Here we investigated the calling song and phonotactic selectivity for sound patterns on the short and long time scales for one such a cricket, Gryllus spec., available as "Gryllus assimilis", the Steppengrille, originally from Ecuador. The calling song consisted of short chirps (2-3 pulses, carrier frequency: 5.0 kHz) emitted with a pulse period of 30.2 ms and chirp rate of 0.43 per second. Females exhibited high selectivity on both time scales. The preference for pulse period peaked at 33 ms which was higher then the pulse period produced by males. Two consecutive pulses per chirp at the correct pulse period were already sufficient for positive phonotaxis. The preference for the chirp pattern was limited by selectivity for small chirp duty cycles and for chirp periods between 200 ms and 500 ms. The long chirp period of the songs of males was unattractive to females. On both time scales a mismatch between the song signal of the males and the preference of females was observed. The variability of song parameters as quantified by the coefficient of variation was below 50% for all temporal measures. Hence, there was not a strong indication for directional selection on song parameters by females which could account for the observed mismatch. The divergence of the chirp period and female preference may originate from a founder effect, when the Steppengrille was cultured. Alternatively the mismatch was a result of selection pressures exerted by commercial breeders on low singing activity, to satisfy customers with softly singing crickets. In the latter case the prominent divergence between male song and female preference was the result of domestication and may serve as an example of rapid evolution of song traits in acoustic communication systems.     

Seasonal Variation in Female Response to Male Calling Song in the Field Cricket, Gryllus rubens

Conspicuous traits that make males attractive to females may make them vulnerable to predators. Females that approach conspicuous males may increase their risk of predation. This means that selection for reduced male conspicuousness in the presence of predators may be due to sexual selection resulting from altered female behavior in the face of increased predator risk. We examine this hypothesis in the field cricket, Gryllus rubens, in which male calling song attracts both conspecific females for mating and parasitoid flies (Ormia ochracea) which kill their hosts within a week. Female crickets are also parasitized by these flies as a result of associating with calling males. In northern Florida crickets that emerge in the spring are not subject to fly parasitism whereas autumn crickets encounter large numbers of flies. We predicted that autumn females should be less attracted to male song than spring females. We tested female response to male calls in a rectangular arena in which male calling song was broadcast from a speaker. Spring females readily approached the speaker but autumn females were less likely to approach and remain in the vicinity of the speaker. These results emphasize the importance of considering how risk affects the evolution of conspicuous male behavior both directly through its effect on the male and indirectly through its effect on female responses to males.     

Is it the Song or the Singers? Acoustic and Social Experiences Shape Adult Reproductive Tactics and Condition

When sexual signals are perceived during growth and development they can provide information regarding the social conditions likely to be encountered as an adult. Perception of cues related to the presence and density of future mates and potential competitors can result in altered adult phenotypes. Previous studies have shown that adult male Teleogryllus oceanicus field crickets from a Kauai, Hawaii population reared alone and without hearing conspecific song are more phonotactic than those reared with song. These naïve males also reduce investment in body size and immunity. Here we examined whether another source of population density information, the presence of other males, affects behavior, size, and immunity. Specifically, we examined satellite behavior as evidenced by strength of phonotaxis, body condition, and immune response in males reared singly and in groups in the presence and absence of conspecific song. Body condition did not vary with rearing density, and immune response did not vary with either acoustic environment or rearing density. Interestingly, group-housed males were more phonotactic than singly-housed males. This pattern was largely driven by the low levels of phonotaxis exhibited by males that were singly-housed in the presence of conspecific song. These findings suggest that males respond to social cues in addition to conspecific song, but that these cues do not necessarily provide concordant information.     

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.