Acoustic communication in Okanagana rimosa (Say) (Homoptera: Cicadidae)

The cicada Okanagana rimosa (Say) has an acoustic communication system with three types of loud timbal sounds: (i) A calling song lasting several seconds to about 1 min which consists of a sequence of chirps at a repetition rate of 83 chirps per second. Each chirp of about 6 ms duration contains 4-5 pulses. The sound level of the calling song is 87-90 dB SPL at a distance of 15 cm. (ii) An amplitude modulated courtship song with increasing amplitude and repetition rate of chirps and pulses. (iii) A protest squawk with irregular chirp and pulse structure. The spectra of all three types are similar and show main energy peaks at 8-10 kHz. Only males sing, and calling song production is influenced by the songs of other males, resulting in an almost continuous sound in dense populations. In such populations, the calling songs overlap and the temporal structure of individual songs is obscured within the habitat. The calling song of the broadly sympatric, closely related species O. canadensis (Provander) is similar in frequency content, but distinct in the temporal pattern (24 chirps per second, 24 ms chirp duration, eight pulses per chirp) which is likely important for species separation in sympatric populations. The hearing threshold of the auditory nerve is similar for females and males of O. rimosa and most sensitive at 4-5 kHz. Experiments in the field show that female phonotaxis of O. rimosa depends on parameters of the calling song. Most females are attracted to calling song models with a 9 kHz carrier frequency (peak frequency of the calling song), but not to models with a 5 kHz carrier frequency (minimum hearing threshold). Phonotaxis depends on temporal parameters of the conspecific song, especially chirp repetition rate. Calling song production is influenced by environmental factors, and likelihood to sing increases with temperature and brightness of the sky. Correspondingly, females perform phonotaxis most often during sunny conditions with temperatures above 22 degrees C. Non-mated and mated females are attracted by the acoustic signals, and the percentage of mated females performing phonotaxis increases during the season.     

Why There is a One-way Crossability Between D. melanogaster and D. simulans?

In the Drosophila melanogaster complex, females D. melanogaster mate relatively easily with males Drosophila simulans but the reciprocal cross is rare. The species sexual isolation is mainly based on chemical and acoustic signal exchanges between partners. The male side of this communication is investigated in this paper in order to understand the asymmetry. In D. melanogaster the acoustic signature is highly significant, and is synergistically reinforced by the chemical signal. In D. simulans the importance of the two signaling channels seems to be reversed. This could explain why D. simulans males produce less precise interpulse interval (IPI) mean value in the courtship song, which can readily overlap those of D. melanogaster. As the males of the two species use the same chemical key, D. simulans males could be recognized by D. melanogaster females as a conspecific.     

Variation and consistency of female preferences for simulated courtship songs in Drosophila virilis

To study the variation and consistency of song preferences in Drosophila virilis females, we played them species-specific song and songs with modified sound pulses and/or interpulse intervals on 3 consecutive days. Species-specific song was played again on the fourth day. All playbacks were done without the presence of males. About 62% of the females indicated their readiness to mate by spreading their wings in at least one of the trials. The proportion of the females responding to species-specific song was about twice that of the females responding to modified songs. The majority of females responded to only one song type, which suggests that the females varied in their preferences and that their preference windows were rather narrow. The females were consistent in their responsiveness to species-specific song played on 2 days. If the female responded to normal song during the first trial, the probability of her responding to the same song during the second trial increased by about 32%. The number of songs required by the females before responding in the two subsequent trials was also correlated within the females (repeatability 0.328). Repeatability of female preferences for male sexual traits is expected both in the viability and Fisherian models of sexual selection. Copyright 1999 The Association for the Study of Animal Behaviour.     

Ultrasound production by genital stridulation in Syntonarcha iriastis (Lepidoptera: Pyralidae): long-distance signalling by male moths?

Males of the pyralid moth, Syntonarcha iriastis Meyrick, perch on vegetation at the tops of trees and bushes and produce ultrasound while their wings are spread and while sclerites at the end of the abdomen are spread to expose the genitalia. Exposing the genitalia appears to engage the sound-producing mechanism; the male genitalia and eighth abdominal sternite of this species are greatly modified and include a file and scraper and possible resonating areas. Sounds produced are consistent between individuals and comprise pulses which are narrow in frequency range, the first pulse being at about 42 kHz and two following pulses at about 57 kHz. The signal is detectable with an ultrasound 'bat' detector from 20 m. Two tettigoniid species (Orthoptera) at the same site produced ultrasonic calls of similar frequencies at the same times as the moth; differences in time-amplitude patterns could be used by orienting moths to recognize conspecifics. Signalling by male S. iriastis is compared with that of other pyralid species in which females are attracted to signalling males. The behaviour of S. iriastis males differs from that of other pyralids in that they do not signal in groups or from a resource attractive to females, and do not possess glands known to produce a 'calling' pheromone. It is suggested, because of these differences, that sound production in this species does not function at close range, as argued for the wax moth Achroia grisella (Fabricius), but instead as a long-distance calling signal.     

Detection of short pure-tone stimuli in the noctuid ear: what are temporal integration and integration time all about?

The interception of a pheromone filament induces flying moths to surge briefly nearly straight upwind; in the absence of pheromone moths cease upwind progress and zigzag crosswind. We tested males of the almond moth, Cadra cautella (Lepidoptera, Pyralidae), in a low-turbulence wind tunnel in wind velocities of 20, 40 and 80 cm s⁻¹. A mechanical pulse generator was set to produce plumes either with same pheromone pulse frequency (pulse generation frequency of 2.9 Hz, interpulse distances from 7 cm to 28 cm) or plumes with same interpulse distance across the three wind velocities (interpulse distance of 14 ± 2 cm, pulse generation frequency of 1.7–5.0 Hz). In plumes of similar pulse frequency, the faster the speed of the wind the slower the ground speed of flight. However, in plumes of similar interpulse distance, ground speed remained relatively constant independent of the wind speed. A `realized' frequency of pulse interception for males flying along the various combinations of pulse frequencies and wind velocities was calculated using the males' average airspeed and the spatial distribution of pheromone pulses in the plume. Realized frequency of pulse interception ranged from 1.3- to 3.0-fold higher than the frequency of pulse generation. The flight tracks of males reflected the regime of realized pulse interception. These results suggest that upwind flight orientation of male C. cautella to pheromone in different wind velocities is determined by the flux of filament encounter. Accepted: 3 September 1997     

Courtship song rhythms in Drosophila yakuba

We studied the courtship songs of males from several strains of Drosophila yakuba. There was clear evidence for two types of pulse songs, generated by the male's wing vibration, which we called 'thud' song and 'clack' song. Clack song was generated by both wings, at distances further from the female than the thud component, which was produced by one-wing displays. The carrier frequency of clack song was considerably higher than that of thud, whereas the interpulse intervals (IPI) of thud song, which usually predominated, were shorter. Furthermore, thud song revealed a cycling of IPIs with a period of about 70 s, significantly longer than that of D. melanogaster or D. simulans. Copyright 1999 The Association for the Study of Animal Behaviour.     

The song structure and seasonal patterns of vocal behavior of male and female bellbirds (Anthornis melanura)

The bellbird (Anthornis melanura) is a honeyeater endemic to New Zealand, which uses song to defend breeding territories and/or food resources year round. Both sexes sing and the song structure and singing behavior have not yet been quantified. The number of song types, spectral structure, repertoire size, and singing behavior of male and female bellbirds was investigated for a large island population. Song types differed between the sexes with males singing a number of structurally distinct song types and females producing song types that overlapped in structure. Singing behavior also differed between the sexes; males often sung long series of songs while females sung each song at relatively long and variable intervals. Singing by both sexes occurred year round but the frequency of male and female singing bouts showed contrasting seasonal patterns. The frequency of female singing bouts increased as the breeding season progressed, whereas male singing bouts decreased. In contrast to almost all studied passerines, female bellbirds exhibited significant singing behavior and sung songs of complex structure and variety that parallel male song. These results provide a quantitative foundation for further research of song in bellbirds and in particular the function of female vocal behavior.     

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.     

Courtship Behavior of Zaprionus indianus (Gupta) (Diptera: Drosophilidae) from Populations Colonizing South America

We describe for the first time the sexual behavior and the courtship song of males of the African fly Zaprionus indianus (Gupta), a recent invader of South America. The male courtship song is formed by monocyclic pulses and the courtship behavior is simple when compared to that of species of Drosophila. Two interpulse interval (IPI) distributions were observed: pre-mounting and mounting. No significant difference was observed between the pre-mounting IPIs of males that descended from three geographical populations from South America. We also observed the songs produced by females and the homosexual behavior exhibited by males. A sequence of bursts is produced by females as a refusal signal against males, while males emit a characteristic song that identifies sex genus, which differs from the courtship song. The short courtship and mating latencies recorded reveal vigorous males and receptive females, respectively.     

When are good genes good? Variable outcomes of female choice in wax moths

Female lesser wax moths (Achroia grisella) choose males based on characters of their ultrasonic advertisement signals. Because a female''s opportunity to obtain increased somatic benefits by mating with a particular male is limited, we investigated whether females obtain genetic benefits for their offspring via mate choice. Controlled breeding experiments conducted under favourable food and temperature conditions showed that developmental characters are heritable, that sire attractiveness and offspring survivorship are unrelated, but that females mating with attractive signallers produce offspring who mature faster than the offspring of females mating with non-attractive signallers. However, under some unfavourable food or temperature conditions, it is the offspring of females mating with non-attractive males who mature faster; these offspring are heavier as well. Thus, the relationship between male attractiveness and offspring development is not environmentally robust, and support for a good genes model of mate choice in A. grisella is dependent on conditions. These findings suggest genotype–environment interactions and emphasize the necessity of testing sexual selection models under a range of natural environments.     

Zebra Finch Mates Use Their Forebrain Song System in Unlearned Call Communication

Unlearned calls are produced by all birds whereas learned songs are only found in three avian taxa, most notably in songbirds. The neural basis for song learning and production is formed by interconnected song nuclei: the song control system. In addition to song, zebra finches produce large numbers of soft, unlearned calls, among which “stack” calls are uttered frequently. To determine unequivocally the calls produced by each member of a group, we mounted miniature wireless microphones on each zebra finch. We find that group living paired males and females communicate using bilateral stack calling. To investigate the role of the song control system in call-based male female communication, we recorded the electrical activity in a premotor nucleus of the song control system in freely behaving male birds. The unique combination of acoustic monitoring together with wireless brain recording of individual zebra finches in groups shows that the neuronal activity of the song system correlates with the production of unlearned stack calls. The results suggest that the song system evolved from a brain circuit controlling simple unlearned calls to a system capable of producing acoustically rich, learned vocalizations.     

Multivariate sexual selection on male song structure in wild populations of sagebrush crickets,Cyphoderris strepitans (Orthoptera: Haglidae)

While a number of studies have measured multivariate sexual selection acting on sexual signals in wild populations, few have confirmed these findings with experimental manipulation. Sagebrush crickets are ideally suited to such investigations because mating imposes an unambiguous phenotypic marker on males arising from nuptial feeding by females. We quantified sexual selection operating on male song by recording songs of virgin and mated males captured from three wild populations. To determine the extent to which selection on male song is influenced by female preference, we conducted a companion study in which we synthesized male songs and broadcast them to females in choice trials. Multivariate selection analysis revealed a saddle‐shaped fitness surface, the highest peak of which corresponded to longer train and pulse durations, and longer intertrain intervals. Longer trains and pulses likely promote greater mate attraction, but selection for longer intertrain durations suggests that energetic constraints may necessitate “time outs”. Playback trials confirmed the selection for longer train and pulse durations, and revealed significant stabilizing selection on dominant frequency, suggesting that the female auditory system is tightly tuned to the species‐specific call frequency. Collectively, our results revealed a complex pattern of multivariate nonlinear selection characterized primarily by strong stabilizing and disruptive selection on male song traits.     

ABSOLUTE VERSUS RELATIVE MEASUREMENTS OF SEXUAL SELECTION: ASSESSING THE CONTRIBUTIONS OF ULTRASONIC SIGNAL CHARACTERS TO MATE ATTRACTION IN LESSER WAX MOTHS,ACHROIA GRISELLA (LEPIDOPTERA: PYRALIDAE)

When females choose a mate among a group of signaling males concentrated in a small area, a male's mating success is often determined not only by his absolute attractiveness but by the attractiveness of his neighbors as well. Multivariate analyses of sexual selection measurements based on absolute values of predictor variables are then misleading, because such analyses assume that the fitness of a given individual is not influenced by others. We addressed this problem of relative fitness in sexual selection by developing two adjustments of the predictor variables in the multivariate analyses by including group means in addition to absolute values and by using relative values, deviations from group means. In the lesser wax moth, Achroia grisella (Lepidoptera: Pyralidae), aggregated males produce ultrasonic signals attractive to females in the vicinity of honey bee colonies. Playback experiments showed that females prefer signals whose pulses are louder, longer, delivered at a faster rate, and include lengthy silent gaps within pulse pairs (long asynchrony intervals). To measure sexual selection on ultrasonic signals, attractiveness was assessed by observing the number of females orienting toward given males in four-choice trials. The various signal characters of the males were computed from ultrasound recordings. Multivariate regression and nonparametric analyses revealed that peak amplitude, asynchrony interval, and pulse rate were the direct targets of selection. Nonlinear stabilizing and correlational selection were also detected. Comparisons of results from multivariate analyses based on absolute values and on the two relative adjustments indicated that the quality of a four-male group did not affect the strength or direction of linear selection but changed the strength of nonlinear selection. Both relative adjustments improved the prediction of male attractiveness. Three-dimensional surface plots generated by the nonparametric regression analyses showed that attractiveness increased monotonically with any combination of the signal characters. This graphical technique also showed that female choice in A. grisella was based on a relative, as opposed to a threshold, decision rule. Overall, male attractiveness for the population remained consistent between testing days. However, attractiveness values for some individual males changed between successive testing days. In these cases, the change in attractiveness was related only to the change in peak amplitude.     

Double meaning of courtship song in a moth

Males use courtship signals to inform a conspecific female of their presence and/or quality, or, alternatively, to ‘cheat’ females by imitating the cues of a prey or predator. These signals have the single function of advertising for mating. Here, we show the dual functions of the courtship song in the yellow peach moth, Conogethes punctiferalis, whose males generate a series of short pulses and a subsequent long pulse in a song bout. Repulsive short pulses mimic the echolocation calls of sympatric horseshoe bats and disrupt the approach of male rivals to a female. The attractive long pulse does not mimic bat calls and specifically induces mate acceptance in the female, who raises her wings to facilitate copulation. These results demonstrate that moths can evolve both attractive acoustic signals and repulsive ones from cues that were originally used to identify predators and non-predators, because the bat-like sounds disrupt rivals, and also support a hypothesis of signal evolution via receiver bias in moth acoustic communication that was driven by the initial evolution of hearing to perceive echolocating bat predators.     

Co-Occurrence of Preference Functions and Acceptance Thresholds in Female Choice: Mate Discrimination in the Lesser Wax Moth

Basic economic models adapted from foraging theory predict that decisions in mate choice may be determined either by ‘best‐of‐n’ preference functions or by sequential rules incorporating acceptance thresholds. However, in some species, more complex determinations that incorporate versions of both protocols are found. To understand the functions of co‐occurring protocols, we studied mating decisions in the lesser wax moth, Achroia grisella (Lepidoptera: Pyralidae), an acoustic species in which females prefer males, the advertisement songs of which are delivered at relatively high ‘pulse‐pair’ rates. In addition to this preference, A. grisella females avoid mating with a male, the song of which does not exceed a minimum pulse‐pair rate, and they hold to this criterion even when no other singing males are present and regardless of song amplitude. Thus, mating decisions are not simply based on acoustic power (pulse‐pair rate × amplitude). We recorded male songs and female responses in an A. grisella population and found that male pulse‐pair rates showed a median of 87/s and ranged from 50 to 115/s, while female acceptance thresholds for male song showed a median of 60/s and ranged from 30 to 105/s. The distributions of thresholds were approximately normal and were not significantly skewed toward the right. Male song rates declined slightly with age, but female thresholds remained stable over the adult lifespan. Both the male and female traits showed significant repeatability within individuals. Whereas phylogenetic inference indicates that hearing in pyralid moths originated as a means of avoiding predation by insectivorous bats, the specific distribution of female acceptance thresholds suggests that currently this protocol does not primarily function to preclude inappropriate, and potentially lethal, responses to bat echolocations: pulse rates in the searching‐phase echolocations used by either aerial‐hawking or substrate‐gleaning bats mostly range from 10 to 20/s, and the lack of positive skew in the distribution of thresholds indicates an absence of directional selection from the left. Rather, we infer that thresholds augment preference functions in A. grisella by precluding mating with males which are markedly inferior in a critical song character. In general, co‐occurring protocols may be important where population density fluctuates markedly, as preference functions may be ineffective in preventing mating with inferior males when density is low.     

Male Courtship Song Frequency as an Indicator of Male Mating Success in Drosophila montana

Drosophila montana females have been found to prefer overwintered males with short and dense (high-frequency) sound pulses in the wild. In the present study males producing high-frequency song succeeded in their courtship more often than males producing low-frequency song in mate-choice experiments. Male mating success correlated with the carrier frequency of the male song recorded after, but not before, an artificial winter (flies kept 6 months at 4°C). The finding that female preference is based on a male song trait changing considerably during overwintering suggests that this trait may reflect the viability and condition of the males during the mating season of the flies in spring.     

Behavioural context regulates dual function of ultrasonic hearing in lesser waxmoths: bat avoidance and pair formation

Abstract.  Ultrasonic hearing in moths evolved to detect bat echolocation cries and has been co-opted into the mating system of some species. Some moths evade or approach ultrasonic stimuli that overlap in their spectral and temporal features. The present study addressed the evaluation of ultrasonic stimuli by lesser waxmoths, a species in which ultrasonic hearing functions in bat evasion and pair formation. Playback experiments show that the role of different stimulus features varies with behavioural context. Flying moths have a single evasive response, which is performed mainly on the basis of pulse length and stimulus power: they dive upon hearing loud pulses ≥ 1 ms in length, and respond more strongly to high-power stimuli above that pulse length. Female moths walking on the ground respond with positive phonotaxis mainly on the basis of pulse rate by orienting toward pulses delivered at rates ≥ 30 pulses s⁻¹. Consequently, moths on the ground approach stimuli that elicit evasion when detected in flight. However, the different responses given to stimulus features by flying and walking moths match the distinguishing features of bat cries and conspecific males in both contexts. Stimulus evaluation, modified by behavioural context, gives lesser waxmoths the flexibility to show appropriate evasion or approach responses to stimuli that overlap in their features.     

Energetic cost of sexual attractiveness: ultrasonic advertisement in wax moths

Pair formation in the lesser wax moth,Achroia grisella(Lepidoptera: Pyralidae), is initiated by male ultrasonic signals that attract receptive females. Individual males vary in attractiveness to females, and the most attractive males are distinguished by exaggeration of three signal characters: pulse rate, peak amplitude and asynchrony interval (temporal separation between pulses generated by movements of the left and right wings during a given wing upstroke or downstroke). Using flow-through respirometry, we measured the resting and signalling metabolic rates of males whose relative attractiveness was known. Acoustic recordings and metabolic measurements were made simultaneously, and we calculated net metabolic rates and factorial metabolic scopes as measures for the energetic cost of signalling. On average, attractive males had higher net metabolic rates and factorial metabolic scopes than unattractive ones, but many unattractive males also had high values. Thus, high expenditure of energy on signalling is necessary but not sufficient for attractiveness. This may result because only one of the three signal characters critical for female preference, pulse rate, is correlated with energy expenditure. Although the results are consistent with the good genes model of sexual selection, they do not conflict with other indirect or direct mechanisms of female choice.     

Ultrasonic courtship songs of male Asian corn borer moths assist copulation attempts by making the females motionless

Males of the Asian corn borer moth Ostrinia furnacalis (Guenée) produce an ultrasonic courtship song of extremely low‐intensity during copulation attempts. The song has been shown to significantly increase the mating success of the male; however, the mode of action of the sound in courtship remains to be resolved. Behavioural experiments using pairs with deafened females or muted males show that, without the aid of the sound, 63% of males eventually succeed in mating after several copulation attempts, whereas the remainder (37%) make repeated attempts in vain until interrupted by the escape of the female. Because few (2%) males fail to copulate when females hear the courtship song, it is evident that the song has an effect on females, promoting the success of copulation attempts. In support of this view, males produce louder songs if the first copulation attempt fails, suggesting that the males increase their sound levels to achieve successful copulation. It is suggested that the ultrasonic songs of the male render the females motionless, which is the same response as that to ultrasonic bat calls. Because even slight movements by the female can interfere with the attempt of the male to copulate, it is likely that, by making her motionless, the success rate of a single copulation attempt is increased greatly.