Sound production in Scolytidae: Rivalry and premating stridulation of male Douglas-fir beetle

The previously reported attractant chirp of male Dendroctonus pseudotsugae changed to a distinctive long interrupted chirp after several minutes in the female gallery, and the same chirp occurred during fighting with another male. It was experimentally evoked by synthetic 3-methylcyclohex-2-en-1-one and frontalin, both known pheromone components of the male. A different, short, uninterrupted male chirp emitted shortly before copulation, was not evoked by these pheromones. It is suggested that chemostimulus allows or maintains dual contexts for the interrupted chirp. Observation of premating and mating behaviour showed some initial male aggressiveness with the female that subsided as gallery cooperation was established before mating and as the uninterrupted chirp replaced the interrupted chirp.     

Acoustic communication in the pine engraver bark beetle: do signals vary between behavioural contexts?

Acoustic communication is taxonomically widespread in bark beetles and is proposed to play an important role in a variety of social and defensive behavioural contexts. Yet our understanding of how signals vary between contexts is currently limited. The present study tests the hypothesis that acoustic signals vary between behavioural contexts in the female pine engraver beetle Ips pini (Say) (Coleoptera: Curculionidae: Scolytinae). Female Ips pini produce acoustic chirps using a vertex‐pronotal stridulatory organ. Randomly sampled chirps generated under three contexts (i.e. distress, predation and premating) are compared for their duration, number of pulses, interpulse intervals, pulse rate and amplitude envelope shapes. The results obtained show that, during premating events, chirps are significantly longer in duration and tend to have a higher proportion of descending amplitude envelopes than chirps occurring during distress and predation events. Chirps produced during distress and predation conditions are indistinguishable from one another. By contrast to the results from previous bark beetle studies, no support is found for categorizing chirps as ‘interrupted’ or ‘uninterrupted’ types based on temporal patterns. The functional significance of context‐dependent variation in chirp characteristics is discussed. Previous studies on acoustic communication in bark beetles are limited as a result of a general lack of objective sampling and measurement criteria for characterizing signals. Recommendations are outlined for future studies on the functions and evolution of acoustic communication in bark beetles.     

Female giant panda (Ailuropoda melanoleuca) chirps advertise the caller's fertile phase

Although female mammal vocal behaviour is known to advertise fertility, to date, no non-human mammal study has shown that the acoustic structure of female calls varies significantly around their fertile period. Here, we used a combination of hormone measurements and acoustic analyses to determine whether female giant panda chirps have the potential to signal the caller''s precise oestrous stage (fertile versus pre-fertile). We then used playback experiments to examine the response of male giant pandas to female chirps produced during fertile versus pre-fertile phases of the caller''s reproductive cycle. Our results show that the acoustic structure of female giant panda chirps differs between fertile and pre-fertile callers and that male giant pandas can perceive differences in female chirps that allow them to determine the exact timing of the female''s fertile phase. These findings indicate that male giant pandas could use vocal cues to preferentially associate and copulate with females at the optimum time for insemination and reveal the likely importance of female vocal signals for coordinating reproductive efforts in this critically endangered species.     

Synchrony during acoustic interactions in the bushcricket <Emphasis Type="Italic">Mecopoda</Emphasis> ‘Chirper’ (Tettigoniidae:Orthoptera) is generated by a combination of chirp-by-chirp resetting and change in intrinsic chirp rate

In several bushcricket species, individual males synchronise their chirps during acoustic interactions. Synchrony is imperfect with the chirps of one male leading or lagging the other by a few milliseconds. Imperfect synchrony is believed to have evolved in response to female preferences for leading chirps. We investigated the mechanism underlying synchrony in the bushcricket species Mecopoda ‘Chirper’ from Southern India using playback experiments and simulations of pairwise interactions. We also investigated whether intrinsic chirp period is a good predictor of leading probability during interactions between males. The mechanism underlying synchrony in this species differs from previously reported mechanisms in that it involves both a change in the oscillator’s intrinsic rate and resetting on a chirp-by-chirp basis. The form of the phase response curve differs from those of previously reported firefly and bushcricket species including the closely related Malaysian species Mecopoda elongata. Simulations exploring oscillator properties showed that the outcome of pairwise interactions was independent of initial phase and alternation was not possible. Solo intrinsic chirp period was a relatively good predictor of leading probability. However, changing the intrinsic period during interactions could enable males with longer periods to lead during acoustic interactions.     

Sound production in Scolytidae: Female sonic stimulus of male pheromone release in two Dendroctonus beetles

The hypothesis that female sonic stimulus may evoke male pheromone release in a behavioural interaction analogous to the known male sonic stimulus of female pheromone release, was confirmed in Dendroctonus pseudotsugae, and also in D. brevicomis. In both species known male-produced substances collected from males stimulated by recorded female stridulation were identified by coupled gas chromatography/mass spectrometry. In a second test with D. pseudotsugae, male pheromone release during recorded female stridulation was evident in the change of male stridulation from the simple attractant chirp to the interrupted chirp, which is known to result from a medium concentration of 3,2-MCH. Also, the D. pseudotsugae male attractant chirp was synthesised with an electronic pulse generator and used to evoke pheromone release. It is concluded that the antiaggregative pheromone of this species is released by each sex at the sonic stimulus of the other sex.     

Acoustic characterization of ultrasonic vocalizations by a nocturnal primate Tarsius syrichta

This preliminary study characterizes the ultrasonic vocalizations produced by Philippine tarsiers, Tarsius syrichta. Data were collected at the Philippine Tarsier Foundation Sanctuary in Corella, Bohol, Philippines, from July through October 2010. Recordings were made on a Wildlife Acoustics Ultrasonic Song Meter 2 BAT from 29 wild, free-living adult resident T. syrichta (23 females and six males). A total of 10,309 USVs were recorded. These vocalizations fell into three main categories: chirps, twitters, and whistles. Chirps were the most frequent, followed by twitters and whistles. Whereas chirps and twitters were emitted by both male and female Philippine tarsiers, whistles were only emitted by adult males. Given that vocalizations reported in this study were exclusively recorded during capture and handling, it is very likely that these vocalizations function as distress calls. However, as the long whistle was only given by adult males who were captured at the same time as the female or the group’s infant, the function of the long whistle might be slightly different than the function of the other relatively lower-frequency USVs.     

First Evidence of Vibrational Communication in Homotomidae (Psylloidea) and Comparison of Substrate-Borne Signals of Two Allied Species of the Genus <Emphasis Type="Italic">Macrohomotoma</Emphasis> Kuwayama

Substrate-borne signals are widely used in Hemiptera and are known to be utilized in mate searching and recognition. Within this Order, the superfamily Psylloidea is a diverse taxon which uses this type of signal modality during mating behavior between the two sexes. This study describes and compares the previously unreported vibrational communication of two closely related species of Macrohomotoma (Homotomidae). Both genders of these two species, Macrohomotoma gladiata Kuwayama 1908 and Macrohomotoma robusta Yang 1984, emit vibrational signals and establish duets during mating. The structure of male calling consists of two chirps while the female response is a single chirp. Males may sometimes follow the female response by emitting a single chirp that sounds similar to the female response with respect to the chirp duration and dominant frequency. This behavior is novel among Psylloidea and its potential function is discussed. Specific comparison of signal characteristics has revealed that the two species of Macrohomotoma are clearly distinguishable from each other which opens the possibility of acoustic signals being used for species delineation.     

(R)-Desmolactone Is a Sex Pheromone or Sex Attractant for the Endangered Valley Elderberry Longhorn Beetle Desmocerus californicus dimorphus and Several Congeners (Cerambycidae: Lepturinae)

We report here that (4R,9Z)-hexadec-9-en-4-olide [(R)-desmolactone] is a sex attractant or sex pheromone for multiple species and subspecies in the cerambycid genus Desmocerus. This compound was previously identified as a female-produced sex attractant pheromone of Desmocerus californicus californicus. Headspace volatiles from female Desmocerus aureipennis aureipennis contained (R)-desmolactone, and the antennae of adult males of two species responded strongly to synthetic (R)-desmolactone in coupled gas chromatography-electroantennogram analyses. In field bioassays in California, Oregon, and British Columbia, traps baited with synthetic (R)-desmolactone captured males of several Desmocerus species and subspecies. Only male beetles were captured, indicating that this compound acts as a sex-specific attractant, rather than as a signal for aggregation. In targeted field bioassays, males of the US federally threatened subspecies Desmocerus californicus dimorphus responded to the synthetic attractant in a dose dependent manner. Our results represent the first example of a “generic” sex pheromone used by multiple species in the subfamily Lepturinae, and demonstrate that pheromone-baited traps may be a sensitive and efficient method of monitoring the threatened species Desmocerus californicus dimorphus, commonly known as the valley elderberry longhorn beetle.     

Rivalry between Stink Bug Females in a Vibrational Communication Network

In the field, male pheromone attracts stink bugs to meet on the same plant and triggers females to call a male by the emission of the calling song. As first among Pentatomidae we describe female rivalry in Chinavia impicticornis, C. ubica and Euschistus heros. Rivalry starts in C. impicticornis by synchronized exchange of the first type of the female calling song pulse trains and proceeds by one of them either to change pulse trains from the first to the second type or to produce readily repeated single pulses. Both reactions either inhibit calling of the rival female or trigger her to respond by alternation with the second type of the calling song pulse trains. Female rivalry in C. ubica differs by the emission of the rival song that replaces alternation with the second type of the calling song typical for C. impicticornis. E. heros females synchronize pulses of the calling song duets and induce emission of the female rival song by one of them that partly inhibits singing of the other. These competitive interactions in Chinavia species reduce the proportion of couples when compared with single couples on a plant. Contrary to both Chinavia species, E. heros female rivalry does not inhibit male response, male signals overlap female emissions and create complex vibrations with modified amplitude modulation pattern caused by interference.     

Selective attention in a synchronising bushcricket: physiology,behaviour and ecology

Synchronising bushcricket males achieve synchrony by delaying their chirps in response to calling neighbours. In multi-male choruses, males that delay chirps in response to all their neighbours would remain silent most of the time and be unable to attract mates. This problem could be overcome if the afferent auditory system exhibited selective attention, and thus a male interacted only with a subset of neighbours. We investigated whether individuals of the bushcricket genus Mecopoda restricted their attention to louder chirps neurophysiologically, behaviourally and through spacing. We found that louder leading chirps were preferentially represented in the omega neuron but the representation of softer following chirps was not completely abolished. Following chirps that were 20 dB louder than leading chirps were better represented than leading chirps. During acoustic interactions, males synchronised with leading chirps even when the following chirps were 20 dB louder. Males did not restrict their attention to louder chirps during interactions but were affected by all chirps above a particular threshold. In the field, we found that males on average had only one or two neighbours whose calls were above this threshold. Selective attention is thus achieved in this bushcricket through spacing rather than neurophysiological filtering of softer signals.     

Anatomy of the stridulation apparatus of the beech leaf‐mining weevil and characterization of,and behavioral responses to,stridulation sounds

We investigated auditory signals and morphology of the stridulatory apparatus of the European beech leaf‐mining weevil, Orchestes fagi L. (Coleoptera: Curculionidae), an invasive herbivore now established in Nova Scotia, Canada, to determine their potential for enhancing survey tools to monitor the spread of the species in Canada. We recorded and described sounds produced by adult O. fagi, analyzed the morphology of the stridulatory mechanism for intersexual differences and asymmetry, and examined behavioral responses elicited in conspecifics by playback of stridulation recordings. Adult O. fagi produced sounds under three conditions: male in distress, female in distress, and male in the presence of female. Female distress chirps lasted significantly longer than male distress chirps and male chirps in the presence of females, but peak frequencies and mean number of chirps per s did not differ significantly among the three groups. Morphology of the stridulation structures in male and female O. fagi was compared using scanning electron microscopy. Orchestes fagi have an elytro‐tergal file‐ and scraper‐type sound production apparatus, through which sound is produced upon anterior motion of the abdomen. Female O. fagi have a ‘pars stridens’ that is longer and has more ridges than males. Width and number of ridges per length of pars stridens did not differ between the sexes. Evidence of asymmetry was found in male pars stridens, with the right side being longer than the left. Playback of recorded sounds to adult weevils suggests female O. fagi were repelled by sounds produced by distressed males.     

Relative Amplitude of Courtship Song Chirp and Trill Components Does Not Alter Female Teleogryllus oceanicus Mating Behavior

Low‐amplitude acoustic signals intended for short‐range communication, often called soft songs, remain poorly studied, especially among acoustically communicating invertebrates. Some insects do employ low‐amplitude acoustic signals, but it remains unclear what the specific function, if any, is of quietness per se. Male Teleogryllus oceanicus, or Pacific field crickets, produce a two component, short‐distance courtship song consisting of a high‐amplitude series of chirps followed by a lower‐amplitude trill. We investigated whether female T. oceanicus prefer to mate with males that sing courtship songs containing trill components that are equally as loud as (?0 dB) or quieter than (?3 dB and ?10 dB) the loudest chirp (90 dB). We found no evidence that modifying trill amplitude affects female mate preference. We did, however, find that previously unmated females were faster to mount males than were females that had mated once before. Previous mating status showed no significant interaction with trill amplitude. What, if any, function of low‐amplitude components of field cricket courtship song remains to be determined.     

Linyphid spider courtship: Releaser and attractant functions of a contact sex pheromone

Experiments with male bowl-and-doily spiders (Frontinella communis) on methanol-washed and unwashed female webs indicate that the courtship of males is chemically released. Parallel experiments on the webs of females of another linyphiid species and on immature F. communis webs indicate that the releaser pheromone is species specific and that it is produced only by mature females. The pheromone that releases male courtship also functions as an attractant, changing the male's orientation from negatively to positively geotactic. The presence or absence of a female on the test web has only a slight effect on the male's behaviour.     

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.     

Model of alternating and aggressive communication with the example of katydid chirping.

Alternating and aggressive communication patterns are present in many species. A theory is developed that explains these two modes of communication. The theory covers both central neural control as well as phonic interaction between two partners. A concrete example, chirping in the katydid Pterophylla camellifolia (Fab.), is shown in detail. Upon receiving acoustical stimuli from the partner a male katydid generates a characteristic response function, with three parts, which regulates solo-overlapping chirps, partially delayed chirps and alternating chirps. Each partner is considered as an element in a closed feedback loop, and described through its response period versus stimulus period curves. It is shown that in alternating mode the communication loop converges towards a stable operating point, while in aggressive mode stability is never achieved. Based on the theory, computer models have been designed simulating both deterministic and random components of the communication signal. Computer simulated chirping sequences are in excellent agreement with field measured data.     

Plant volatiles and density-dependent conspecific female odors are used by Asian citrus psyllid to evaluate host suitability on a spatial scale

We investigated how chemical cues derived from female Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae) and their host plants affect host acceptance choices by conspecifics. In four-choice cage and two-choice olfactometer assays, female psyllids avoided conspecific female cues in a density-dependent manner. However, odors from citrus plants actively damaged by psyllid feeding were attractive to conspecific females. When odors from feeding-damaged plants were presented simultaneously with odors from female D. citri, attraction of female conspecifics was no longer observed as compared with a clean air control in olfactometer assays. In subsequent experiments, D. citri females were released within arenas that contained actively feeding-damaged or non-damaged (control) citrus plants, each with previously psyllid-infested and uninfested young leaves. D. citri development is linked to the presence of these newly emerging leaves which is the only site of nymphal development. Female D. citri were initially attracted by the actively damaged plants as compared with non-damaged controls. After acceptance of plants that were actively damaged by feeding, D. citri females preferentially chose and settled on uninfested young leaves as compared with previously infested young leaves. A herbivore-induced plant volatile attractant and a female-specific odor repellent appear to be complementary foraging cues providing psyllids with information at two spatial scales: (1) the whole plant level for choosing a plant potentially harboring male conspecifics for mating, and (2) the within plant level to reduce intra-conspecific competition by identifying previously exploited resources.     

Understanding the Long-Lasting Attraction of Malaria Mosquitoes to Odor Baits

The use of odor baits for surveillance and control of malaria mosquitoes requires robust dispensing tools. In this study, the residual activity of a synthetic mosquito attractant blend dispensed from nylon or low density polyethylene (LDPE) sachets was evaluated at weekly intervals for one year without re-impregnation. The potential role of bacteria in modulating the attraction of mosquitoes to odor-treated nylon that had been used repeatedly over the one year study period, without re-impregnation, was also investigated. Significantly higher proportions of female Anopheles gambiae sensu stricto mosquitoes were consistently attracted to treated nylon strips than the other treatments, up to one year post-treatment. Additional volatile organic compounds and various bacterial populations were found on the treated nylon strips after one year of repeated use. The most abundant bacteria were Bacillus thuringiensis and Acinetobacter baumannii. Autoclaving of treated nylon strips prior to exposure had no effect on trap collections of laboratory-reared female An. Gambiae (P = 0.17) or wild female An. Gambiae sensu lato (P = 0.26) and Mansonia spp. (P = 0.17) mosquitoes. Trap catches of wild female An. Funestus (P < 0.001) and other anophelines (P < 0.007) were higher when treated strips had been autoclaved prior to deployment as opposed to when the treated nylon strips were not autoclaved. By contrast, wild female Culex mosquitoes were more strongly attracted to non-autoclaved compared to autoclaved treated nylon strips (P < 0.042). This study demonstrates the feasibility of using odor baits for sampling and surveillance of malaria as well as other mosquito vectors over prolonged periods of time. Preliminary evidence points towards the potential role of bacteria in sustaining prolonged use of nylon material for dispensing synthetic attractant odorants for host-seeking malaria and other mosquito vectors but further investigations are required.     

Sound production and mating in a waterboatman, Palmacorixa nana (Heteroptera: Corixidae)

Both male and female Palmacorixa nana stridulate by rubbing fields of pegs on the fore femora over thickened flanges on the maxillary plates. I sugges that the head is the sound-radiating structure and is driven into oscillation when its edges are plucked by rows of femoral pegs. P. nana males produce spontaneous, courtship, mounting and copulatory signals. The latter tow signals are produced by a new mechanism which constitutes only the third record of inter-individual stridulation in insects. The known acoustic behaviour of the female is limited to a signal indicating readiness to mate. Mating behaviour was observed and copulatory success found not to rely on a complete exchange of acoustic signals. Comparison of data from several sources indicates that corixids capable of singing can either have an obligatory or optional exchange of signals.     

Temporal Analysis of Image-Rivalry Suppression

During binocular rivalry, perception alternates between two different images presented one to each eye. At any moment, one image is visible, dominant, while the other is invisible, suppressed. Alternations in perception during rivalry could involve competition between eyes, eye-rivalry, or between images, image-rivalry, or both. We measured response criteria, sensitivities, and thresholds to brief contrast increments to one of the rival stimuli in conventional rivalry displays and in a display in which the rival stimuli swapped between the eyes every 333 ms–swap rivalry–that necessarily involves image rivalry. We compared the sensitivity and threshold measures in dominance and suppression to assess the strength of suppression. We found that response criteria are essentially the same during dominance and suppression for the two sorts of rivalry. Critically, we found that swap-rivalry suppression is weak after a swap and strengthens throughout the swap interval. We propose that image rivalry is responsible for weak initial suppression immediately after a swap and that eye rivalry is responsible for the stronger suppression that comes later.     

Vocalizations of Belding's ground squirrels (Spermophilus beldingi)

Vocalizations of Belding's ground squirrels (Spermophilus beldingi) were recorded during the summer of 1982 near Tioga Pass in the central Sierra Nevada of California. Sonagrams were made and call parameters were measured. Discriminant function analyses revealed that multiple-note calls (‘trills’) differed acoustically depending on whether they occurred in response to a predator, or were given by males following copulations. The post-copulatory trills of males were individually distinctive. Among anti-predator trills there was no evidence of predator-specificity within the narrow range of predators tested: trills given to two species of (stuffed) weasels (Mustela), to dogs and to humans were statistically indistinguishable. Sonagrams of trills occurring in agonistic contexts suggest that a third general category of trills may exist, but agonistic trills were more variable than either anti-predator or post-copulatory trills. The ground squirrels also gave single-note calls in the three contexts described above, either repetitively (‘chirps’) or singly (‘whistles’). Neither chirps nor whistles encoded any obvious situation-specific information, except that whistles were typically associated with rapidly-moving predators, usually raptors. The post-copulatory chirps of males were individually distinctive. Vocalizations of Belding's ground squirrels may not vary among contexts as much as do the analogous calls of California ground squirrels (S. beecheyi).