A Comunicação Entre Insetos Através de VibraçõesRESUMO -A comunicação através de vibrações originárias do substrato tem sido reconhecida há muito tempo, mas tem recebido muito menos atenção que a comunicação através do som propagado pelo ar. Recentemente, entretanto, tem se tornado cada vez mais claro o papel crucial que sinais vibratórios desempenham na comunicação em muitos grupos de insetos, o que motivou essa revisão suscinta. Sinais vibracionais estão relacionados ao comportamento sexual, de alarme e de defesa, e são em geral usados para mediar ações coordenadas em grupo e interações sociais complexas. Para insetos pequenos esses sinais representam um custo energético mínimo, com alcance máximo, e que não são percebidos facilmente por predadores e por parasitóides em potencial. Sinais originários dos substrato são produzidos por diversos métodos e detectados por receptores presentes nas seis pernas. O comportamento de corte do percevejo verde, Nezara viridula (L.) (Hemiptera: Pentatomidae) é analisado como um modelo representativo na ilustração de alguns mecanismos importantes na comunicação vibracional em insetos. Sinais vibratórios são específicos das espécies e de sexos, os quais são produzidos durante a corte e que são adequados para serem transmitidos pelas plantas, contendo informações relevantes sobre a espécie e o sexo do organismo emissor, bem como informações de onde localizar o parceiro. O papel de sinais originados do substrato, os quais são únicos para cada espécie, faz com que os estudos em sinais vibracionais sejam uma importante ferramenta para resolver problemas taxonômicos. PALAVRAS-CHAVE: Comunicação vibracional, vibração do substrato, comportamento, Nezara viridulaABSTRACT -Communication through substrate-borne vibrations has for long been recognized but in comparison with air-borne sound it has received very little attention. However, in recent years it has become increasingly clear that vibrational signals play a crucial role in communication in many insect groups and we provide a short overview. Vibrational signals are related to sexual behavior, alarm and defensive behavior and are often used to mediate coordinated group actions and complex social interactions. For small insects they are probably the least costly and most far-reaching signals for intraspecific communication and also not easily perceived by a potential predator or parasitoid. Substrateborne signals are produced by diversed methods and detected by sensitive receptors in all six legs. The courtship behavior of the southern green stink bug Nezara viridula (L.) (Hemiptera: Pentatomidae) is taken as a representative model in illustrating some principal mechanisms of vibrational communication in insects. Species and sex specific vibrational signals produced during the courtship are well suited for propagation through plants and to transmit the relevant information about the species and sex of the sender as well as provide the directional cue for locating the mate. The role of substrate-borne signals as a part of the specifi...
Vibratory signals of plant-dwelling insects, such as land bugs of the families Cydnidae and Pentatomidae, are produced mainly by stridulation and/or vibration of some body part. Signals emitted by the vibratory mechanisms have low-frequency characteristics with a relatively narrow frequency peak dominant around 100 Hz and differently expressed frequency modulation and higher harmonics. Such spectral characteristics are well tuned to the transmission properties of plants, and the low attenuation enables long-range communication on the same plant under standing wave conditions. Frequencies of stridulatory signals extend up to 10 kHz. In some groups, vibratory and stridulatory mechanisms may be used simultaneously to produce broadband signals. The subgenual organ, joint chordotonal organs, campaniform sensilla and mechanoreceptors, such as the Johnston's organ in antennae, are used to detect these vibratory signals. Species-specific songs facilitate mate location and recognition, and less species-specific signals provide information about enemies or rival mates.
High-frequency airborne sound is quickly absorbed in vegetation, and outside the plant shelter such signals attract predators. Furthermore, small insects cannot emit efficiently low-frequency airborne sound when their diameter is smaller than a third of the radiated wavelength. For plant-dwelling insects, substrate-borne sound signaling remains the best solution for communication in dense vegetation typical for the tropics. Communication range in such an environment depends on tuning of vibratory signal properties with mechanical characteristics of plants and sensory abilities to extract the information from the environmental noise. Nezara viridula and other until now investigated species of the stink bug family Pentatomidae represent the model for solutions optimizing long-range communication through green plants. They communicate with vibratory signals of the dominant frequency around 100 Hz, which travel through plants with low attenuation, creating standing wave conditions in the plant’s rod-like structures. Green plants act as low-pass filter and their resonant peaks fit well with the spectral peaks of stink bug vibratory emissions. The species’ leg sensory organs with the underlying neural network are sensitive enough to enable communication through a plant on a distance of several meters. Comparative data on substrate-borne sound communication in burrower (Cydnidae) and predatory (Asopinae) bugs will be discussed.
Scaphoideus titanus Ball (Hemiptera: Cicadellidae: Deltocephalinae) is the vector of the grapevine disease Flavescence dorée. In S. titanus the male–female duet (MFD), based on species‐specific vibrational signals, is essential for successful copulation. The female reply within a duet is a single pulse that is coupled with the male pulse with constant latency. It has been shown that a rival male can interrupt an existing duet by emitting disruptive noise signals. We tested whether the reproductive behaviour of S. titanus can be disrupted by the playback of intra‐specific and synthesized vibrational signals. Tested males responded to the playback of an MFD with typical rivalry behaviour. Such behaviour includes silent search for a duetting female (satellite behaviour) and/or emission of disruptive signals. These signals were emitted either after exchange of male–female pulses or after two male pulses coupled by latency corresponding to the female response window. The onset of male disruptive signals overlapped with a female pulse. We suggest that the intruder’s disruptive signals can mask the female reply and confuse courting males. Playback of disruptive vibrational signals reduced the level of male calling and interrupted an established MFD that consequently resulted in a significantly reduced number of copulations. These results indicate that the vibrational communication channel is open to interference either from abiotic environmental noise or from signals produced by sexual competitors or heterospecifics. The present study also suggests that a detailed understanding of leafhopper behaviour is essential for trying new approaches in the development of more environmentally friendly control practices.
Nezara viridula (L.) (Pentatomidae: Heteroptera) from Brazil, Florida, Italy and Slovenia, communicate by vibratory songs associated with long-range calling and close-range courting, rivalry and repelling. Each song is composed of spectrally and temporally different units. Spectrally different pulses of duration less than 300 ms are present in the male calling song. The female calling song is characterized by pulse trains composed of pulses shorter than 150 ms and pulse trains composed of a longer (> 700 ms) and shorter (< 250 ms) pulse. Shorter and longer pulses have different spectral characteristics. The male and female courtship songs are characterized by fusion of shorter (< 150 ms) pulses into a pulse train usually followed by a shorter (< 200 ms) postpulse in the case of the male courtship song. The female repelling song is a several seconds long vibration of irregular temporal structure. The short (< 400 ms) male rival song pulses are frequency modulated. The dominant frequency peaks of the songs investigated lie between 70 and 130 Hz. The dominant frequency and the microstructure of song spectra show no population speci®city. The average duration varies more in calling than in courtship songs. The repetition time varies extensively in songs of different populations. Normal communication followed by copulation was observed between mates from Slovenia and Brazil and between mates from Florida and Italy. The potential role of different temporal and spectral parameters for species recognition and mate location is discussed in view of the expected distortion of the characteristic signal structure during transmission through plants.
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