Comparison of disturbance stridulations in five species of triatominae bugs
Introduction
Several insect species produced different vibratory signals that are emitted in different behavioural contexts (Michelsen et al., 1982, Markl, 1983). For example, the bug Phymata crassipes (Heteroptera: Phymatidae) performs precopulatory, locomotory and alarm vibratory signals (Gogala and Cokl, 1983). In Phrydiuchus tau (Coleoptera: Curculionidae), the occurrence of two kinds of stridulations performed in both disturbance and sexual contexts was demonstrated. In one case, both males and females of this species stridulated when they were disturbed and, in the other, only males stridulated when they approach towards the female and during copula (Wilson et al., 1993). Triatomine bugs posses a prosternal stridulatory organ which consists of a cuticular, longitudinal groove composed of a number of transverse ridges (Lent and Wygodzinsky, 1979, Di Luciano, 1981). Stridulation is produced by these insects when the tip of the proboscis is rubbed against the prosternal groove. Like other insects, the triatomine bugs Triatoma infestans and Rhodnius prolixus have been observed to stridulate, in at least, two different behavioural contexts. Females of these species stridulate either to reject copulatory attempts performed by males, or males and females both stridulate when they are disturbed or handled (Moore, 1961, Schofield, 1977, Manrique and Lazzari, 1994, Roces and Manrique, 1996, Manrique and Schilman, 2000). The hypothesis that the stridulations emitted in response to disturbance are defensive responses to predators' attack had been proposed for wasps and beetles species (Bauer, 1976, Masters, 1979) and for Reduviidae (Leston, 1957). Taking into account this hypothesis, we would expect that the disturbance signal might be generalised for all triatomines. Thus, a comparative analysis of these vibratory signals among different species becomes important.
In this study, the temporal structure and frequency spectra of stridulations performed by mechanically-disturbed T. infestans, T. sordida, T. guasayana, R. prolixus and Dipetalogaster maxima were characterised and compared. The morphology of the stridulatory groove of these species was also compared.
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Insects
Adult males and females of T. infestans, T. sordida, T. guasayana, R. prolixus and D. maxima, were used. Insects were reared at the Servicio Nacional de Chagas, Santa Marı́a de Punilla, Córdoba, Argentina, or in our laboratory. One week before the assays, insects were fed ad libitum with heparinised bovine blood using an artificial feeder (Núñez and Lazzari, 1990).
Experimental design and procedure
To generate disturbance stridulations each bug was clasped with forceps by its hindleg and then, its dorsal thorax was smoothly
Results
The temporal pattern of stridulations, as well as the frequency spectra of the five triatomine species, are shown in Fig. 1, Fig. 2 and Table 1. As no signal differences were found between sexes, they were analysed together. In the cases presented here, stridulations consisted of long series of repetitive syllables, each one composed of two chirps, a short and a long one. The exception was R. prolixus, which produced a series of short chirps and then a long chirp. In addition, in all species
Discussion
Triatomine species were able to produce at least two different vibratory signals with different meanings. On the one hand, the sexual signal of triatomines has an intraspecific role in communication between non-receptive females that stridulate to reject the copulatory attempts performed by males (Manrique and Lazzari, 1994, Roces and Manrique, 1996, Manrique and Schilman, 2000). This signal must match the perception capacities of the insects and we would expect structure specificity. On the
Acknowledgements
The authors are indebted to F. Roces for critically reading the early version of the manuscript, to all Insect Physiology Laboratory staff for fruitful discussions, to Delmi Canale (Servicio Nacional de Chagas, Argentina) for the provision of much of the insects used in the study, to M. Lorenzo for body length measurements of D. maxima and to two anonymous reviewers for very helpful comments. Data analysis was performed thanks to F. Roces from the University of Würzburg, Germany. This
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