Filtering of temporal parameters of the calling song by cricket females of two closely related species: a behavioral analysis

The phonotactic response of cricket females was investigated on a locomotion compensator to determine the temporal parameters of the male's calling song which are important for species recognition. Two sympatric species (Teleogryllus commodus, T. oceanicus) that show different syllable periods in the chirp and trill parts of their calling songs were used. By their responses T. commodus females exhibited two temporal filters for syllable periods, which were tuned to the species-specific syllable periods occurring during chirp and trill. For song recognition both filters had to be activated and for both a minimum number of three to five consecutive syllable periods was necessary. In contrast, T. oceanicus females showed only one sharply tuned filter corresponding to the chirp part of the male's calling song. This filter was sufficient for calling song recognition. Syllable periods of the trill part also influenced calling song recognition, but these played only a minor role. Carrier frequency was also important for positive phonotaxis. Calling song recognition by T. commodus females is largely based on central nervous processing, while for T. oceanicus both peripheral frequency filtering and central temporal filtering is important. Accepted: 17 January 1997     

Temperature coupling in cricket acoustic communication

Summary Temperature effects on calling song production and recognition were investigated in the North American field cricket, Gryllus firmus. Temporal parameters of field-recorded G. firmus calling song are strongly affected by temperature. Chirp rate and syllable rate increase, by factors of 4 and 2, respectively, as linear functions of temperature over the range in which these animals sing in the field (12°–30 °C). Temperature affects syllable duration to a lesser extent, and does not influence calling song carrier frequency. Female phonotactic preference, measured on a spherical treadmill in the laboratory, also changes with temperature such that warmer females prefer songs with faster chirp and syllable rates. Best phonotaxis, measured as accuracy of orientation to the sound source, and highest walking velocity, occur in response to temperature-matched songs at 15°, 21°, and 30 °C. Experiments under semi-natural conditions in an outdoor arena revealed that females perform phonotaxis at temperatures as low as 13 °C. Taken together, the song and phonotaxis data demonstrate that this communication system is temperature coupled. A strategy is outlined by which temperature coupling may be exploited to test hypotheses about the organization of neural networks subserving song recognition.Abbreviations CP chirp period - SP syllable period - SD syllable duration     

Calling is an honest indicator of paternal genetic quality in poison frogs

Several competing hypotheses have been put forward to explain why females of many species mate preferentially with males possessing the most conspicuous signals (e.g., ornaments, displays, or songs). We performed a laboratory experiment using two species of poison frogs, Dendrobates leucomelas and Epipedobates tricolor, to test the hypothesis that male calling performance is an honest indicator of parental quality. Our analyses are based on data from behavioral observations of mating activities of captive-reared individuals (and their offspring) that were housed in terraria for four consecutive breeding seasons. Male mating success increased with male calling rate and chirp duration in both species, suggesting that females preferred males with more elaborate calls. Because calling performance improved with age in D. leucomelas, female poison frogs that prefer males with more elaborate calls in the wild may end up mating with older males that have already proven their ability to survive. Females that mated with good callers obtained higher quality offspring. Eggs fertilized by males with high calling rates and long chirp durations had higher hatching success and produced tadpoles that were more likely to metamorphose into surviving frogs. As a consequence, females that mated with males with high calling performance obtained more surviving offspring per egg, compared to females that mated with poor callers. Collectively, our findings comply with the notion that female poison frogs prefer to mate with good callers because calling performance is a reliable predictor of offspring quality. The possible influence of maternal allocation and reasons for the strong effect size compared to previous studies are discussed.     

Temperature coupling in cricket acoustic communication

Acoustic communication in Gryllus firmus is temperature-coupled: temperature induces parallel changes in male calling song temporal pattern, and in female preference for song. Temperature effects on song production and recognition networks were localized by selectively warming head or thorax or both head and thorax of intact crickets, then eliciting aggression song production (males) or phonotaxis to synthetic calling song (females). Because male song is produced by a thoracic central pattern generator (CPG), and because head ganglia are necessary for female song recognition, measurements of female phonotaxis under such conditions may be used to test the following competing hypotheses about organization of the song recognition network: 1. A set of neurons homologous to the male song CPG exist in the female, and are used as a template that determines preferred values of song temporal parameters for song pattern recognition (the common neural elements hypothesis), and 2. temporal pattern preference is determined entirely within the head ganglia. Neither selective warming of the head nor of the thorax was effective in changing female song preference, but simultaneous warming of head and thorax shifted preference toward a faster song in most preparations, as did warming the whole animal by raising ambient temperature. These results suggest that phonotactic preference for song temporal pattern is plurisegmentally determined in field crickets. Selective warming experiments during aggression song production in males revealed that syllable period is influenced but not completely determined by thoracic temperature; head temperature is irrelevant. The song CPG appears to receive some rate-setting information from outside the thoracic central nervous system.     

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.     

Mate recognition in the Aphis fabae complex: daily rhythm of release and specificity of sex pheromones

The two subspecies of the host-alternating black bean aphid, Aphis fabae Scopoli, used in this study have different secondary host plants but reproduce sexually in autumn on the same host plant, spindle, Euonymus europaea L. In both Aphis fabae fabae Scopoli and A. f. solanella Theobald the percentage of adult mating females (oviparae) calling (releasing sex pheromone) each day increased with age up to day 8. During the course of a day the mating females of A. f. solanella called most actively early in the morning and then showed a gradual decrease in calling activity. In contrast, few A. f. fabae called early in the day and were most active around midday, after which the incidence of calling decreased slightly, with 50% still calling in the evening. In both taxa the activity of the males in the absence of mating females paralleled the calling activity of their respective females. In an olfactometer, males showed a marked preference for the sex pheromone released by conspecific females over that of the other subspecies and did not respond to the odour of spindle. Thus, as in the other species of aphids there are specific mate recognition systems in A. f. fabae and A. f. solanella, which are likely to reduce the incidence of hybridisation.     

Alternative mating tactics in the Great Plains toad

Male Great Plains toads, Bufo cognatus, exhibit two mating tactics. At any time, most males give advertisement calls to attract females, while other males, ‘satellites’, remain silent and station themselves near callers in an attempt to intercept females. Females are usually capable of moving through choruses undetected by males. Those females detected by males can avoid being clasped by quickly darting away; but if clasped, can detach the male by inflating. Females initiated amplexus with callers; subsequently calling males mated with 92% of the females and satellites mated with the remaining 8%. Toads employing the satellite tactic associated with males that had longer calls and that were larger, although call duration was the stronger predictor. The percentage of males exhibiting satellite behaviour during a sampling period varied from 0 to 57%. Several hypotheses explaining satellite frequency were explored, including variation in male density, male quality, and operational sex ratio. Of the variables considered, only two aspects of male density (namely, the number of unmated males per m of shoreline and number of calling males per m shoreline) affected satellite frequency. High male density may cause at least some calling males to lose vocal conspicuousness, thereby favouring satellite behaviour. However, male density accounted for only 17% of the observed variation in satellite frequency. This, plus the fact that up to 74% of marked males switched between tactics, suggests that factors such as higher energetic demands of calling may affect satellite frequency.     

Intrasexual mounting in the beetle Diaprepes abbreviatus (L.).

The weevil Diaprepes abbreviatus shows three kinds of same-sex mountings: males mount other unpaired males, males mount males already engaged in copulation and females mount other females. Four hypotheses were evaluated in order to explain same-sex matings by males: (i) female mimicry by inferior males, (ii) dominance of larger males which affects the behaviour of small males, (iii) sperm transfer in which smaller males gain some reproductive success by 'hitchhiking' their sperm with the sperm of larger males, and (iv) poor sex recognition. Data from mate choice and sperm competition experiments rejected the female mimicry, dominance and sperm transfer hypotheses and supported the poor sex recognition hypothesis. We tested three hypotheses in order to explain female mounting behaviour: (i) females mimic male behaviour in order to reduce sexual harassment by males, (ii) females mount other females in order to appear larger and thereby attract more and larger males for mating, and (iii) female mimicry of males. The results of our mate choice experiments suggested that the female mimicry of males hypothesis best explains the observed female mounting behaviour. This result is also consistent with the poor sex recognition hypothesis which is the most likely explanation for male and female intrasexual mating behaviour in many insect species.     

Calling behaviour of adult female Helicoverpa armigera (Hübner) (Lep., Noctuidae) of overwintering generation and effects of mating

The calling behaviour of overwintering generation females of Helicoverpa armigera and the effects of mating were studied in the laboratory at 24 ± 1°C and under reversed light-dark cycle (16 h light : 8 h dark). Age had a significant influence on calling patterns. Based on calling age, mean number of calling bouts and total calling length of virgin females increased significantly, and mean onset time of calling advanced significantly from calling day 1 to subsequent calling days. Females of the overwintering generation exhibited more short bouts in calling, and some females that initiated calling on a previous day did not call on subsequent days. Mating had no effect on the overall patterns, but did affect calling behaviour. Mated females did not resume calling after mating during the same scotophase and, on the day following mating, mated females called less frequently and for a shorter duration, but thereafter increased to the same level of virgin females of the same calling age. Furthermore, as the moth aged, the percentage of mated females calling was lower than that of virgin females.     

Pupal Period Affects Calling Behavior of the Wheat Moth, Pseudaletia sequax (Lepidoptera: Noctuidae)

The effects of pupal period and age on calling behavior of virgin females of Pseudaletia sequax were determined. Calling behavior of groups of females of similar age of pupation was observed every 10 min for six calling days. Females were considered calling when they assumed a characteristic position: wings and abdomen elevated, displaying the ovipositor and presumably releasing pheromone. Calling behavior was discontinuous. Females with a shorter pupal period took more time to start calling than females with a longer pupal period and called for a longer period. On the first day of calling, females took more time to initiate calling, on average during the seventh hour of the scotophase, whereas on the second day of calling, most females initiated calling during the fifth hour of the scotophase. On subsequent days of calling, the average time to initiate calling changed to the fourth hour of the scotophase. The mean time of calling per day decreased significantly with the number of days since first calling, and the mean number of calling bouts increased with days since first calling.     

Acoustic mimicry and disruptive alternative calling tactics in an Australian bushcricket (Caedicia; Phaneropterinae; Tettigoniidae; Orthoptera): does mating influence male calling tactic?

Abstract.  Male calling and searching tactics are described for a duetting Australian bushcricket, Caedicia sp. 12 (Phaneropterinae; Tettigoniidae; Orthoptera). The repertoire of Caedicia sp. 12 consists of the calling song and, by nonduetting males, a series of calling tactics that include short-click calling, disruptive over-singing and a call mimicking the entire duet. Nonduetting males respond to the production of a duet by another male and a female with short-click calls that mimic the female call at the conclusion of a duet. By manipulating the male's mating history, it is found that this form of calling behaviour is more likely to occur within the male's 6-day postmating refractory period; the low cost tactic allows males to re-mate during spermatophore replenishment. Males also produce disruptive calls in response to a duet, where the male may over-sing the duetting male's signal or produce a call that appears to mimic the entire duet; the male produces a calling song followed by a short signal that has the same latency as the female's reply within a duet. Males also over-sing crucial elements of the duetting-male's song that are normally critical for the female's conspecific recognition. There is no evidence that females search for the duetting male partner, but males unable to enter a duet will search for the call of a responding female. Searching by males is more common when these males are producing disruptive calls. Alternative male calling tactics are discussed as a set of conditional strategies for securing unmated females.     

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.     

Effects of a host plant,Brassica juncea, on calling behaviour and egg maturation in the diamondback moth,Plutella xylostella

The influence of the presence of seedlings of a host plant, Oriental mustard,Brassica juncea (L.) Coss., on the calling behaviour and egg maturation in the diamondback moth,Plutella xylostella (L.), was investigated. In the presence of plants, females began calling at a younger age, began calling earlier in the night, and spent more time calling. Females which were kept with plants until after their first night of calling and then were isolated from plants, subsequently called less, at a level not significantly different from females which had never been exposed to plants. Of all females, both with and without plants, which were dissected at the end of the first night of calling (n=150), 93% contained mature eggs. Heavier females were significantly more likely to have mature eggs at adult emergence, to begin calling at a younger age and to spend more time calling on their first night of calling. The number of developed eggs in calling females was not significantly different in the presence or absence of host plants on the first night of calling. However, by the second night, females in the presence of plants contained significantly more eggs, indicating that the presence of plants accelerated egg maturation, both before and after the onset of calling.     

Pheromone Release Behaviour in Females of Phyllonorycter junoniella (Z.) (Lepidoptera, Gracillariidae) Under Constant and Cycling Temperatures

The calling activity of virgin Phyllonorycter junoniella (Z.) females under a cycling thermal regime differed from that under constant temperature in the following ways: the percentage of females calling at the morning activity peak was increased; the morning period of calling activity was prolonged; the total period of calling activity of an individual female was increased; an extra peak of activity occurred at the end of the photophase, and females changed the calling pattern during the active period. We assume that the occurrence of two calling peaks and the extension of the calling period might be adaptive for a Ph. junoniella species with a sex ratio strongly shifted towards females (8:1 females:males), as it could lead to an increased proportion of females mated.     

Effects of temperature and light on calling in the tiger moth Holomelina lamae (Freeman) (Lepidoptera: Arctiidae)

ABSTRACT. In Holomelina lamae Freeman daily eclosion of adults is gated, with males emerging before females. By advancing the onset of photophase and by delaying the onset of scotophase, it was demonstrated that lights-on acts as the main phase-setting cue for calling. Few females call on the day they eclose. Calling is initiated c. 9 h after the onset of photophase in 2-day-old females, and shifts to earlier times in older females. The duration of calling also increases with age. That calling is controlled by an endogenous circadian clock is indicated by its persistence in continuous light (LL) and dark (DD). In LL calling is dampened rapidly, but a single scotophase re-entrains the rhythm. Decreases in temperature advance the onset of calling and the mean hour of calling, while increases in temperature delay both. However, the magnitudes of such phase-shifts depended upon hour of the photoperiod. Moreover, cooling and heating appears to exert both transient and long-term effects on the calling rhythm. An 8 h period at a reduced temperature in LL induces calling in females whose calling is dampened, and entrains the calling rhythm. Females maintained in DD from second instar larvae to the adult stage exhibit a circadian calling rhythm set by eclosion.     

The function of Barbary macaque copulation calls.

In a wide variety of animal species, females produce vocalizations specific to mating contexts. It has been proposed that these copulation calls function to incite males to compete for access to the calling female. Two separate advantages of inciting male-male competition in this way have been put forward. The first suggests that as a result of calling, females are only mated by the highest ranking male in the vicinity (indirect mate choice hypothesis). The second proposes that copulation calling results in a female being mated by many males, thus promoting competition at the level of sperm (sperm competition hypothesis). In this paper, I give results from the first experimental study to test these hypotheses. Playback was used to examine the function of copulation calls of female Barbary macaques (Macaca sylvanus) in Gibraltar. Although rank did not affect lone males'' likelihood of approaching copulation calls, when playbacks were given to pairs of males only the higher ranking individual approached. Moreover, females were mated significantly sooner after playback of their copulation call than after playback of a control stimulus. These results suggest that the copulation calls of female Barbary macaques play a key role in affecting patterns of male reproductive behaviour, not only providing an indirect mechanism of female choice, but also promoting sperm competition by reducing the interval between copulations. Potential fitness benefits of inciting male-male competition at these two levels are discussed.     

Age-dependent pattern of calling behavior in <Emphasis Type="Italic">Atheloca subrufella</Emphasis> (Hulst) (Lepidoptera: Phycitidae)

Changes in the signaling effort of virgin females are known to occur until their first mating. Young females tend to increase the calling effort (number and time duration of bouts) as they age. In this context, the calling behavior of virgin females of Atheloca subrufella, an important pest of coconut palms (Cocos nucifera), was studied under laboratory conditions. Thirty four virgin females were observed during 16 scotophases (25 ± 2 °C, 75 ± 5 % RH and 10 h scotophase), until they stopped calling in the following photophase. The calling position, pattern and frequency were evaluated. Females of A. subrufella showed only one calling position, and calling peaked between the second and fifth scotophases. Most females did not start calling immediately after emergence. Neither the individual duration of calling nor the number of calling bouts changed over time. However, the time that calling started was earlier in older females, probably to avoid competition with younger females and increase the probability of mating. These data support the hypothesis that there is an age-dependent pattern of pheromone emission in virgin females of A. subrufella.     

Sexual differences in cricket calling song recognition

Summary Phonotactic behavior was studied in male crickets,Teleogryllus oceanicus. Tethered flying males were presented with electronically synthesized calling song models in a two-choice phonotaxis assay, and their song preferences were determined and compared with previous findings for females.Males are poorer at discriminating between songs than females; they do not display choice behavior as frequently as females, and the choices they do make are not as consistent as those of females (Figs. 3, 4). T. oceanicus calling song is composed of rhythmically different chirp and trill sections. The selectivity of males for these two components differs from that of females. Females prefer chirp to trill, but the opposite is true for males (Fig. 5B-F). Males are similar to females in that they prefer either a conspecific song model or its separate components to a heterospecific model (Fig. 5A, G, H).Behavioral and neural implications of these findings are discussed.     

Modelling female mating success during mass trapping and natural competitive attraction of searching males or females

Simulation models of insects encountering sex pheromone with or without mass trapping in which the searching sex is either male (moths and many insect species) or female (some true bugs, beetles, and flies) were developed. The searching sex moved as a correlated random walk, while the opposite sex remained stationary (calling) and released an attractive sex pheromone. The searching sex was caught when encountering a pheromone‐baited trap, and females mated when encountering a male. An encounter with pheromone was defined by the searcher's interception of a circle termed the effective attraction radius (EAR_c). Parameters of movement (speed and duration), initial numbers of calling sex and searching sex, number of traps, area, and EAR_c of traps and calling sex were varied individually to evaluate effects on the percentage of females mating. In the natural condition without traps, female mating success in both models was identical. Increasing the EAR_c of the calling sex caused diminishing increases in female mating success, suggesting that evolution of larger pheromone release and EAR_c is limited by increasing costs (production/sensitivity) relative to diminishing increases and benefits of mating encounters. With mass trapping, increasing the EAR_c of traps or density of traps caused similar declines in female mating in both models, but the female‐searching model predicted slightly lower mating success than the male‐searching model. Increasing the EAR_c of calling insects or the initial density of insects caused similar increases in female mating in both models, but again the female‐searching model had slightly lower mating success than the male‐searching model. The models have implications for mating lek formation and for understanding the variables affecting the success of mass trapping programs for insect pests with either male or female sex pheromones.     

The effect of photoperiod on the calling behaviour of virgin females of the true armyworm,Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae)

The effect of photoperiod on the calling behaviour of Pseudaletia unipuncta virgin females was examined under five different photoperiodic regimes at 25°C, 65% r.h. The age at which females called for the first time following emergence varied with photoperiod; generally calling was later under long-scotophase conditions. However under a 6 h scotophase there was also a delay in calling and >63% of the females tested never called. There was a considerable variability in the daily calling patterns between the different photoperiods, and the mean onset time of calling was not constant with either “lights on” or “light off”. There was however a constancy of the mean onset time of calling relative to the mid-point of both the photo- and scotophase at all photoperiods tested, indicating that females could measure the absolute duration of either the photo- or scotophase. Transferring females from one photoperiodic condition to another once calling had been initiated, determined that it was the “lights off” signal that P. unipuncta females used to phase set the clock governing circadian calling behaviour.Females subjected to a decrease of 4 or 6 h in the length of the scotophase following the initiation of calling required several days to adjust to the new photoperiodic regime and a high proportion of females did not call during the night following the transfer. A 4 or 6 h increase in the scotophase did not inhibit calling on the night following the transfer but females still required several days to adjust completely. However, females experiencing a 2 h increase or decrease in the duration of the scotophase were able to maintain normal calling behaviour. The results of these experiments are discussed in relation to the seasonal biology of the true armyworm and the hypothesis that this is a migrant species.