Multimodal communication in a noisy environment: a case study of the Bornean rock frog Staurois parvus

High background noise is an impediment to signal detection and perception. We report the use of multiple solutions to improve signal perception in the acoustic and visual modality by the Bornean rock frog, Staurois parvus. We discovered that vocal communication was not impaired by continuous abiotic background noise characterised by fast-flowing water. Males modified amplitude, pitch, repetition rate and duration of notes within their advertisement call. The difference in sound pressure between advertisement calls and background noise at the call dominant frequency of 5578 Hz was 8 dB, a difference sufficient for receiver detection. In addition, males used several visual signals to communicate with conspecifics with foot flagging and foot flashing being the most common and conspicuous visual displays, followed by arm waving, upright posture, crouching, and an open-mouth display. We used acoustic playback experiments to test the efficacy-based alerting signal hypothesis of multimodal communication. In support of the alerting hypothesis, we found that acoustic signals and foot flagging are functionally linked with advertisement calling preceding foot flagging. We conclude that S. parvus has solved the problem of continuous broadband low-frequency noise by both modifying its advertisement call in multiple ways and by using numerous visual signals. This is the first example of a frog using multiple acoustic and visual solutions to communicate in an environment characterised by continuous noise.     

Integration time for short broad band clicks in echolocating FM-bats (Eptesicus fuscus)

Vespertilionid FM-bats (four Eptesicus fuscus and one Vespertilio murinus) were trained in an electronic phantom target simulator to detect synthetic echoes consisting of either one or two clicks. The threshold sound pressure for single clicks was around 47 dB peSPL for all five bats corresponding to a threshold energy of -95 dB re 1 Pa² * s. By varying the interclick interval, T, for double clicks it was shown that the threshold intensity was around — 3 dB relative to the threshold for single clicks at T up to 2.4 ms, indicating perfect power summation of both clicks. A threshold shift of -13.5 dB for a 1 ms train of 20 clicks (0.05 ms interclick interval) confirmed that the bats integrated the power of the stimuli. At T longer than around 2.5 ms the threshold for double clicks was the same as for single clicks. Thus, the bats performed like perfect energy detectors with an integration time of approximately 2.4 ms. This integration time is an order of magnitude shorter than that reported for bats listening passively for pure tones. In our setup the bats emitted sonar signals with durations of 2–3 ms. Hence, the results may indicate that while echolocating the bats integration time is adapted to the duration of the sonar emissions.Abbreviations AGC automatic gain control - FM frequency modulated - peSPL peak equivalent sound pressure level - rms root mean square - SD standard deviation - SE standard error of mean - T interclick interval     

Confusion of Phonotaxis by Masking Sounds in the Bushcricket Conocephalus brevipennis (Tettigoniidae: Conocephalinae)

Female Conocephalus brevipennis, executing phonotaxis to the call of a conspecific male broadcast from a speaker, were subjected to varying levels of background sound so as to progressively mask the attracting signal. The background consisted of either random noise or the call of a sympatric congener, C. nigropleurum and was delivered from above the orienting female. Disruption of phonotaxis was assayed by monitoring the angular dispersion of the female's exit path from the arena, relative to the position of the conspecific call. Random noise significantly interfered with phonotaxis when it was within 2 dB of the conspecific call intensity, both signals measured at the centre of the arena. The equivalent effect of the call C. nigropleurum was 5 dB above conspecific call intensity. The implication of signal loss within background noise is discussed as are the possible interactive behaviours of competing sympatric syntopic species.     

Evoked potentials of the auditory cortex of the porpoise,Phocoena phocoena

Summary Evoked potential (EP) recordings in the auditory cortex of the porpoise,Phocoena phocoena, were used to obtain data characterizing the auditory perception of this dolphin. The frequency threshold curves showed that the lowest EP thresholds were within 120–130 kHz. An additional sensitivity peak was observed between 20 and 30 kHz. The minimal EP threshold to noise burst was 3·10^–4–10^/s-3 Pa. The threshold for response to modulations in sound intensity was below 0.5 dB and about 0.1% for frequency modulations. Special attention was paid to the dependence of the auditory cortex EP on the temporal parameters of the acoustic stimuli: sound burst duration, rise time, and repetition rate. The data indicate that the porpoise auditory cortex is adapted to detect ultrasonic, brief, fast rising, and closely spaced sounds like echolocating clicks.Abbreviation EP evoked potential     

Energy detection and temporal integration in the noctuid A1 auditory receptor

The temporal integration of the A1 auditory receptor of two species of noctuid moths (Lepidoptera, Noctuidae) was investigated. Tympanal nerve spikes were recorded while stimulating the ear with broad band clicks. Thresholds were measured for single clicks, pairs of clicks with a separation of 1–20 ms, and trains of up to 8 clicks at separations of 1–2 ms. The average threshold for single clicks was 52.9 dB peSPL (SD 1.7 dB, n = 40) for Noctua pronuba and 50.1 dB peSPL (SD 4.0 dB, n = 27) for Spodoptera littoralis. The thresholds for double clicks with a 1 ms separation were lower than the thresholds for single clicks. The difference decreased as the separation between the clicks was increased. The results were fully consistent with an energy detector model (a leaky integrator with an exponential decay) with a time constant of about 4 ms.The results are compared to previously published results with pure tone intensity/duration trading. A common underlying mechanism is suggested, based on the passive electric properties of the receptor cell membrane.It is suggested, that the time constant revealed in the present study characterizes auditory receptors in general, and is related to the short time constants in vertebrate audition.Abbreviations peSPL peak equivalent sound pressure level - SD standard deviation - time constant     

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.     

Sound radiation in a cicada: the role of different structures

Sound radiation was studied in males of Tympanistalna gastrica St»l during a spontaneous song with the characteristics of the conspecific calling song, which was elicited as an after effect of brain stimulation. The song contains two different kinds of sound pulses: 1) loud clicks and 2) soft pulses, presenting different spectra.The timbals, abdomen, tympana, folded membranes and opercula were tested as potential radiators of the song. The experiments included: 1) probe microphone measurements of the spectra of loud clicks and soft pulses in several positions around the animal and close to the body surface; 2) measurements of the spectra before and after covering with vaseline different structures that might be relevant to the radiation of the song, and manipulations of the size and shape of the abdominal and thoracic portions of the tracheal air sac; 3) laser vibrometry measurements in different parts of the body, both during singing and external sound stimulation.The data obtained demonstrate that several structures contribute differently to the radiation of clicks and soft pulses: 1) The timbals are the main radiators at frequencies around the dominant spectral peak, 10–11 kHz in clicks and 12–13 kHz in soft pulses; 2) The tympana are important in radiation of frequencies below and above the timbal peak, especially during the generation of soft pulses; 3) The abdomen is more activated during the generation of clicks, and is more important in the radiation of low frequencies around 5 kHz.Manipulations of the body cavities showed that neither the thoracic nor the abdominal portions of the air sac are critical for the song tuning. The large abdominal cavity do not seem to work as a Helmholtz resonator. We found no evidence that resonances inside this cavity should play an important role in enhancing sound radiation in T. gastrica.     

Dynamic properties of the advertisement calls of gray tree frogs: patterns of variability and female choice

We assessed the potential for several acoustic properties ofthe advertisement calls of male gray tree frogs to affect relativemating success by relating patterns of variation in these propertiesto minimum differences required to elicit female choice. Dynamicproperties (pulse number, PN; call rate, CR; and duty cycle,DC, the ratio of call duration to call period) varied much morewithin bouts of calling than a static property (dominant frequency,DF) but nevertheless exhibited significant between male variationin three of four breeding seasons. Many multiply recorded malesconsistently produced calls with values substantially aboveor below mean values of males recorded on the same nights. Nightlyranges of variation in PN and CR were often greater than theminimum differences required to elicit female preferences inthe laboratory. In most experiments, females chose high-PN orfast-CR calls over low-PN or slow-CR alternatives, respectively,even if the preferred stimuli were farther away or 6-10 dB lowerin sound pressure level (SPL), provided that differences inPN or CR were 100%. Consistent with these results, females didnot always choose the closer of two calling males in the field.Nightly ranges of variation in DF rarely equaled the minimumdifference required to elicit SPL independent preferences. Femalespreferred a stimulus of high-PN and slow-CR over an alternativeof low-PN or fast-CR with the same acoustic on-time; in twoexperiments, females chose calls of high-PN over low-PN alternativeseven though the playback of the high-PN call was interruptedand the low-PN call was broadcast continuously. Thus, femalepreferences were not merely based on the total time of acousticstimulation. Responses of females tested twice in the same experimentsuggest that phenotypic variation in preference was limitedin our study populations.     

A Cervid Vocal Fold Model Suggests Greater Glottal Efficiency in Calling at High Frequencies

Male Rocky Mountain elk (Cervus elaphus nelsoni) produce loud and high fundamental frequency bugles during the mating season, in contrast to the male European Red Deer (Cervus elaphus scoticus) who produces loud and low fundamental frequency roaring calls. A critical step in understanding vocal communication is to relate sound complexity to anatomy and physiology in a causal manner. Experimentation at the sound source, often difficult in vivo in mammals, is simulated here by a finite element model of the larynx and a wave propagation model of the vocal tract, both based on the morphology and biomechanics of the elk. The model can produce a wide range of fundamental frequencies. Low fundamental frequencies require low vocal fold strain, but large lung pressure and large glottal flow if sound intensity level is to exceed 70 dB at 10 m distance. A high-frequency bugle requires both large muscular effort (to strain the vocal ligament) and high lung pressure (to overcome phonation threshold pressure), but at least 10 dB more intensity level can be achieved. Glottal efficiency, the ration of radiated sound power to aerodynamic power at the glottis, is higher in elk, suggesting an advantage of high-pitched signaling. This advantage is based on two aspects; first, the lower airflow required for aerodynamic power and, second, an acoustic radiation advantage at higher frequencies. Both signal types are used by the respective males during the mating season and probably serve as honest signals. The two signal types relate differently to physical qualities of the sender. The low-frequency sound (Red Deer call) relates to overall body size via a strong relationship between acoustic parameters and the size of vocal organs and body size. The high-frequency bugle may signal muscular strength and endurance, via a ‘vocalizing at the edge’ mechanism, for which efficiency is critical.     

Communication in a noisy environment: short-term acoustic adjustments and the underlying acoustic niche of a Neotropical stream-breeding frog

Acoustically active animals may show long- and short-term adaptations in acoustic traits for coping with ambient noise. Given the key role of calls in anurans’ life history, long- and short-term adaptations are expected in species inhabiting noisy habitats. However, to disentangle such adaptations is a difficult task, incipiently addressed for Neotropical frogs. We investigated if males of a stream-breeding frog (Crossodactylus schmidti) adjust call traits according to the background noise, and if the signal-to-noise ratio (SNR) varies between call harmonics and along call notes. We measured sound pressure levels of calls and noise in the field and used a fine-scale acoustic analysis to describe the signal and noise structure and test for noise-related call adjustments. The multi-note harmonic call of C. schmidti greatly varied in the spectral structure, including a trend for increasing note amplitude along the call, a wide frequency bandwidth of the 2nd harmonic, a minor call frequency modulation due to a trend for increasing note frequency within the same harmonic, and a major call frequency modulation due to the variable location of the dominant harmonic along the call. Calls had significantly higher frequencies than the noise at the range of the 1st and the 2nd call harmonics, and significantly louder sound pressure than the noise at the range of all harmonics. Males emitted the majority of call notes showing positive SNR, and though males also emitted some notes with negative SNR, when a given harmonic was negative the other harmonics in the same note did not tend to be SNR-negative. Our results indicate that male C. schmidti show short-term acoustic adjustments that make the advertisement call effective for coping with the interference of the stream-generated noise. We suggest that the call spectral plasticity serves for coping with temporary changes in the background noise, whilst we also discuss the possibility that the redundant, harmonic-structured call may have evolved to diminish masking interference on the acoustic signal by the background noise. This is the first study to uncouple noise-related acoustic adjustments and putative long-term acoustic adaptations for a Hylodidae, providing insights on behavioral plasticity and signal evolution of stream-breeding frogs.     

Species differences of male loud calls and their perception in sulawesi macaques

Playback experiments were conducted to investigate interspecific discrimination of male loud calls in Sulawesi macaques. Loud calls of four macaque species living in Sulawesi (Macaca tonkeana, M. maurus, M. hecki, andM. nigrescens) and a control stimulus (an 8-sec frequency modulated sound) were played back to semi-free-ranging Tonkean macaques (M. tonkeana). A preliminary acoustic analysis indicated that the calls of these four species differ in some spectral and temporal features. In the playback experiments, Tonkean macaques responded in a similar manner to conspecific calls and calls of two other species,M. maurus andM. hecki. In contrast, animals responded more weakly to the call ofM. nigrescens and the control stimulus. Males responded more strongly than females to all stimuli, while females appeared to be more discriminating for species differences than males. Analyses on the acoustic features of loud calls suggested that high frequency, wide frequency range, and repetition of sound units at a high rate elicit quick responses from animals.     

Acoustic courtship signals in the Caribbean fruit fly, Anastrepha suspensa (Loew)

Male Caribbean fruit flies, Anastrepha suspensa (Loew) produce two sounds in sexual contexts, calling songs and precopulatory songs. Calling song occurs during pheromone release from territories within leks and consists of repeated bursts of sound (pulse trains). Virgin female A. suspensa became more active in the presence of recorded calling songs. Activity during the broadcast of a heterospecific song did not differ from movement during periods of silence. A conspecific song typical of smaller males, i.e. conspicuous for its long periods between pulse trains, also failed to elicit more activity by virgin females than silence. Mated females were most active during silences. Unmated males had no obvious reaction to sound. Calling songs are apparently sexually important communications which females discriminate among and may use as cues for locating and/or choosing between mates. Precopulatory song is produced by mounted males just before and during the early stages of copulation. Males that did not produce such songs remained coupled for shorter periods, perhaps passing fewer sperm. Wingless (muted) males were more likely to complete aedeagal insertion if a recorded precopulatory song was broadcast. Calling song played at the same level (90 dB) had no significant effect on the acceptance of males, nor did precopulatory song at a lower SPL (52dB). Precopulatory song may be used to display male vigour to choosing females.     

Interactions Between Swarming Chironomus annularius (Diptera: Chironomidae) Males: Role of Acoustic Behavior

Males of swarming species of chironomids use their auditory system (Johnston's organs) to recognize a female within swarm and do not respond to male flight tones. However, in some cases the male–male interactions were observed at a high frequency. The role of acoustic behavior in this phenomenon in C. annularius was studied. The results showed that male Johnston's organs were sensitive to male flight tones from a distance of about 1–1.5 cm. The carrier frequencies of these sounds negatively correlated with male body size. Thus we would expect that male–male interactions will occur mainly between large males. Nonetheless, the analysis of caught pairs revealed that in both male–male and female–male interactions small males had an advantage. The ability of males to perceive the male flight tones is discussed with respect to swarming behavior and mating success.     

Soft Song in Song Sparrows: Acoustic Structure and Implications for Signal Function

In many species of songbirds, males sometimes produce songs at distinctly lower amplitude than in normal singing. Depending on the species, these 'soft songs' may be sung in the context of female courtship, male–male aggression, or both. In song sparrows, males produce soft songs during aggressive interactions with other males, and the amount of soft song produced is the only singing behavior that can be used to reliably predict a subsequent attack by the singer. Although soft song is clearly an important signal in this species, little is known about the acoustic structure of soft song or about how that structure compares to the structure of normal 'broadcast song'. We recorded a large sample of soft songs and broadcast songs from 10 male song sparrows, and measured song amplitudes in the field while controlling the subject's distance to a calibrated microphone. We show that song sparrow males produce songs over a wide range of amplitudes, with soft songs in the range of 55–77 dB sound pressure level and broadcast songs in the range of 78–85 dB. We present evidence for two types of soft song: 'crystallized' soft songs that are broadcast repertoire song types sung at low amplitude, and 'warbled' soft songs that are not found in the broadcast repertoire. Although highly variable, warbled soft songs produced by individual birds could be grouped into song types based on spectrographic similarity. To our knowledge, a distinct repertoire of soft song types has not been previously reported for any songbird.     

Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow,Acoustically Complex Habitat

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPa_pp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPa_pp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.     

Individually Specific Call Feature Is Not Used to Neighbour-Stranger Discrimination: The Corncrake Case

In various contexts, animals rely on acoustic signals to differentiate between conspecifics. Currently, studies examining vocal signatures use two main approaches. In the first approach, researchers search for acoustic characteristics that have the potential to be individual specific. This approach yields information on variation in signal parameters both within and between individuals and generates practical tools that can be used in population monitoring. In the second approach, playback experiments with natural calls are conducted to discern whether animals are capable of discriminating among the vocal signatures of different individuals. However, both approaches do not reveal the exact signal characteristics that are being used in the discrimination process. In this study, we tested whether an individual-specific call characteristic – namely the length of the intervals between successive maximal amplitude peaks within syllables (PPD) – is crucial in neighbour-stranger discrimination by males of the nocturnal and highly secretive bird species, the corncrake (Crex crex). We conducted paired playback experiments in which corncrakes (n = 47) were exposed to artificial calls with PPD characteristics of neighbour and stranger birds. These artificial calls differed only in PPD structure. The calls were broadcast from a speaker, and we recorded the birds'' behavioural responses. Although corncrakes have previously been experimentally shown to discriminate between neighbours and strangers, we found no difference in the responses to the artificial calls representing neighbours versus strangers. This finding demonstrates that even if vocal signatures are individual specific within a species, it does not automatically mean that said signatures are being crucial in discrimination among individuals. At the same time, the birds'' aggressive responses to the artificial calls indicated that the information transmitted by PPDs is important in species-specific call recognition and may be used by males and/or females to evaluate sender quality, similarly like sound frequency in some insect species.     

Arginine Vasotocin Injection Increases Probability of Calling in Cricket Frogs, but Causes Call Changes Characteristic of Less Aggressive Males

Male cricket frogs,Acris crepitanscommunicate to males and females using advertisement calls, which are arranged into call groups. Calls at the middle and end, but not beginning of the call group, are modified in response to male–male aggressive interactions. We found in this field study of male cricket frogs in natural breeding choruses that the peptide hormone arginine vasotocin (AVT) not only increased the probability that males called after injections, but also caused modifications in middle and end calls to produce calls characteristic of less aggressive males. Moreover, AVT-injected males showed significantly greater increases in call dominant frequency than saline-injected males, again, a characteristic of less aggressive males. Cricket frog calls are used to both repel males and attract females, thus call changes may relate to male–male and/or male–female interactions. Saline-injected males also demonstrated significant changes in several call traits, including changes that occurred in the beginning and middle calls of the call groups, but not the end calls. AVT appeared to block some call changes produced through handling. These data suggest that AVT can influence acoustic communication in frogs in several ways, including effects on call characteristics and dominant frequency, as well as potentially blocking some handling effects.     

Acoustic Preference of Frog‐Biting Midges (Corethrella spp) Attacking Túngara Frogs in their Natural Habitat

In many animals, males aggregate to produce mating signals that attract conspecific females. These leks, however, also attract eavesdropping predators and parasites lured by the mating signal. This study investigates the acoustic preferences of eavesdroppers attracted to natural choruses in a Neotropical frog, the túngara frog (Engystomops pustulosus). In particular, we examined the responses of frog‐biting midges to natural variation in call properties and signaling rates of males in the chorus. These midges use the mating calls of the frogs to localize them and obtain a blood meal. Although it is known that the midges prefer complex over simple túngara frog calls, it is unclear how these eavesdroppers respond to natural call variation when confronted with multiple males in a chorus. We investigated the acoustic preference of the midges using calling frogs in their natural environment and thus accounted for natural variation in their call properties. We performed field recordings using a sound imaging system to quantify the temporal call properties of males in small choruses. During these recordings, we also collected frog‐biting midges attacking calling males. Our results revealed that, in a given chorus, male frogs calling at higher rates and with higher call complexity attracted a larger number of frog‐biting midges. Call rate was particularly important at increasing the number of midges attracted when males produced calls of lower complexity. Similarly, call complexity increased attractiveness to the midges especially when males produced calls at a low repetition rate. Given that female túngara frogs prefer calls produced at higher repetition rates and higher complexity, this study highlights the challenge faced by signalers when increasing attractiveness of the signal to their intended receivers.     

Acoustic communication in spring peepers

Summary In the previous study (Wilczynski et al. 1984) we found that neurons in the auditory nerve of female spring peepers (Hyla crucifer) are tuned to frequencies in the male advertisement call, whereas auditory units in male peepers are mismatched in spectral sensitivity to their call. We investigated, in the present report, behavioral consequences of this sexual dimorphism in auditory sensitivity. Call amplitude, rate of call attenuation with environmental transmission, and the amplitude of ambient noise were measured and used to compute the active space of this signal for males and females. The effect of calling height upon active space was considered. Measurements of active space were compared with intermale distances within breeding choruses.Results indicate that active space of the advertisement call for females is as much as 6 times greater than that for males, and varies directly with the height above ground from which males call. Observed maximum intermale distances correspond closely to the active space of the call for this sex. This suggests that males space themselves so that the amplitude of a neighbor's calls approximates their auditory neural threshold to call frequencies. By this proximal mechanism, peepers maximize intermale distance but ensure that they remain within a chorus.Abbreviation dB SPL decibels sound pressure levelre: 20 Pa     

Reproductive isolation inProkelisia planthoppers (Homoptera: Delphacidae): Acoustic differentiation and hybridization failure

Males and females of Prokelisia marginata (Van Duzee) and Prokelisia dolusWilson communicate through substrate-transmitted vibrations. The acoustic signals (attraction and courtship calls) of these planthoppers are effective in mate location, attraction, and mate choice. Attraction calls are structurally distinct for both species and differ in pulse type, pulse repetition rate, and pulse duration. Using playback of prerecorded calls, individuals discriminated between conspecific and heterospecific signals. Depending on the sex and species, response calls were produced three to eight times more frequently to conspecifics than to heterospecifics. However, acoustic signals alone did not explain reproductive isolation and hybridization failure in these two congeners. Some heterospecific pairs called, courted, and attempted to join genitalia, but no connections were successful and no progeny were produced. Thus, acoustic behavior is not a guaranteed premating isolating mechanism in no-choice situations. Other courtship behaviors and possibly morphological differences in genitalia also contributed to their isolation. Females displayed a variety of rejection behaviors to conspecific and heterospecific males, suggesting that sexual selection (female choice), in addition to species recognition, may be an important force in the evolution of the acoustic signals of planthoppers. Although signal structure was not dependent on wing form (planthoppers exhibit wing dimorphism), the age when females first began to call was related to wing form. Brachypterous (flightless) females of both species began calling early in adult life (day 2), whereas macropterous (migratory) females began calling later in adult life (day 6). This pattern is consistent with the oogenesis-flight syndrome, in which reproductive maturity is delayed until after migration occurs.