02057nas a2200157 4500008004100000022001300041245013800054210006900192260001600261300001400277490000700291520149200298100002301790700002101813856006501834 1995 eng d a0003347200aVariation and repeatability of male agonistic hiss characteristics and their relationship to social rank in Gromphadorhina portentosa0 aVariation and repeatability of male agonistic hiss characteristi cJan-01-1995 a719 - 7290 v503 a

Male Madagascar hissing cockroaches, Gromphadorhina portentosa, produce a variety of sounds or ‘hisses’ during social interactions. The agonistic signals produced during male-male competition were examined, specifically the intra- and inter-individual variation of agonistic hisses, to determine if these sounds could be reliable indicators of male rank or male size. The relationship between characteristics of the agonistic hiss and social rank were also examined. Among males, agonistic hisses ranged from sounds without clear frequency bands to those with frequency modulation. Agonistic hisses varied in dominant frequency, which was measured at the beginning, middle, and end of the hiss, duration, and hiss rate (number per min). With males, the characteristics of a hiss were repeatable within and between days. Dominant males hissed significantly more than middle-ranking or subordinate males. While dominant, middle-ranking and subordinate males differed in the rate of hissing, there were no significant differences in the duration or dominant frequency of their hisses. Finally, the relationship between male weight and features of agonistic hisses was examined. Both duration and frequency were significantly correlated with male weight. Larger males had longer, lower frequency hisses than smaller males. The possible function of agonistic hissing during male-male interactions and the role of hisses in discrimination between individuals is discussed.

1 aClark, Deborah, C.1 aMoore, Allen, J. uhttp://linkinghub.elsevier.com/retrieve/pii/000334729580132402180nas a2200157 4500008004100000022001400041245010400055210006900159260001600228300001400244490000600258520166800264100002501932700001701957856004801974 1980 eng d a0340-544300aSound production in the cockroach, Gromphadorhina portentosa: evidence for communication by hissing0 aSound production in the cockroach Gromphadorhina portentosa evid cJan-03-1980 a305 - 3140 v63 a

1. Sound production of the giant Madagascar cockroach, Gromphadorhina portentosa, was examined by behavioral and acoustical methods in order to determine the functions of the hisses produced by this species.

2. Gromphadorhina is able to produce audible hisses from a pair of modified spiracles. Adult males hiss in three social contexts: during aggressive encounters, during courtship (when two types of hisses are discernable), and during copulation. Adults and nymphs of both sexes also hiss when disturbed (Figs 2 and 3).

3. There are reliable differences among hisses emitted in these social contexts which depend on several features: the shape of the amplitude envelope (Fig. 4), the relative loudness (Table 1), and the temporal characteristics both of single hisses and of hiss trains (Table 1 and Fig. 4).

4. In both courtship and aggression, hissing accompanies characteristic, stereotyped behavior patterns (Figs. 5 and 7); during aggressive encounters between males, hissing is predictive of winning (Fig. 6).

5. Males which have been silenced by occlusion of the specialized spiracles carry on apparently normal courtship, but they are unsuccessful in copulating due to a lack of receptive behavior by the female (Fig. 8).

6. Playback of recorded courtship hisses during courtship of females by silenced males leads to receptive behavior by females, and to normal rates of copulation (Fig. 8).

7. Our evidence supports the hypothesis that G. portentosa has evolved a system of communication in which hisses serve as auditory social signals.

1 aNelson, Margaret, C.1 aFraser, Jean uhttp://link.springer.com/10.1007/BF0029277302273nas a2200145 4500008004100000022001400041245009500055210006900150260001600219300001200235490000800247520179900255100002502054856004802079 1979 eng d a0340-759400aSound production in the cockroach,Gromphadorhina portentosa: The sound-producing apparatus0 aSound production in the cockroachGromphadorhina portentosa The s cJan-01-1979 a27 - 380 v1323 a

1. The giant Madagascar cockroach,Gromphadorhina portentosa, hisses by expelling air from a pair of specialized abdominal spiracles. The anatomy and innervation of serially homologous respiratory and sound-producing spiracles were compared in order to determine the evolutionary steps by which a new behavior has developed.

2. The trachea leading to the sound-producing (fourth) spiracle shows a constriction proximally; distally it is greatly elongated with a conical bore (Fig. 2). These features, which are lacking in other spiracles, are sufficient to account for the character of the sound (Fig. 10).

3. The motoneurons innervating both types of spiracles were located by axonal diffusion of cobalt, and their morphology was determined in wholemounted ganglia. The number, ganglionic locations, and in some cases branching patterns of motoneurons serving the sound-producing and respiratory spiracles were essentially identical (Figs. 4, 5, 6).

4. Physiological activity was recorded along spiracle nerves and within spiracle muscle fibers; four units were identified for each spiracle, agreeing with the number of cells located anatomically. These included, for each abdominal spiracle, an opener exciter motoneuron, two closer exciter motoneurons, and one closer inhibitor motoneuron (Figs. 7, 8).

5. During normal respiration the output of these 4 units had similar phase relationships in all abdominal spiracles which were examined; lower firing rates in the motoneurons innervating the hissing spiracles rendered these nonfunctional during normal respiration (Fig. 9).

6. The findings are consistent with conservation of motor innervation and of central pattern generators during evolution.

1 aNelson, Margaret, C. uhttp://link.springer.com/10.1007/BF0061772901354nas a2200157 4500008004100000022001300041245009000054210006900144260001600213300001400229490000700243520083800250100001701088700002501105856006601130 1984 eng d a0003347200aCommunication in the courtship of a madagascan hissing cockroach. I. Normal courtship0 aCommunication in the courtship of a madagascan hissing cockroach cJan-02-1984 a194 - 2030 v323 a

he hissing Madagascar cockroach, Gromphadorhina portentosa, has a prolonged and complex courtship involving signals in several sensory modalities. Courtship was described for 13 pairs of cockroaches and the frequencies and sequencing of 16 behavioural units were analysed. Particular attention was paid to the function and interactions of acoustic, chemical and tactile components. The results indicate that posturing and sound production by males, and antennation by both males and females, are important in courtship. They also suggest that courtship in G. portentosa, rather than depending on a rigid sequence of behaviour determined by a series of discrete releasers, is quite flexible, using ‘behavioural monologues’ by both sexes as a means of achieving transitions from one stage of courtship to the next.

1 aFraser, Jean1 aNelson, Margaret, C. uhttp://linkinghub.elsevier.com/retrieve/pii/S000334728480337102143nas a2200157 4500008004100000022001400041245014700055210006900202260001600271300001400287490000800301520154700309100002301856700002101879856008501900 1995 eng d a0005-795900aSocial Communication in the Madagascar Hissing Cockroach: Features of Male Courtship Hisses and a Comparison of Courtship and Agonistic Hisses0 aSocial Communication in the Madagascar Hissing Cockroach Feature cJan-01-1995 a401 - 4170 v1323 a

Male Madagascar hissing cockroaches (Gromphadorhina portentosa) produce acoustic sounds or 'hisses' during courtship interactions with females. G. portentosa males also hiss during agonistic interactions with other males. In this study we documented intra- and inter- individual variation in male courtship hisses, addressed the possible role of hissing as an indicator of male rank, size or mating success, and determined if individual males produce different hisses during courtship and agonistic interactions. We found inter-individual variation in courtship hisses in dominant frequency and duration. Within males, these characteristics of the courtship hiss were repeatable within a day. Characteristics of courtship hisses and mating success were not related to male rank. However, courtship hisses may convey information about the size of the male: larger males had longer, lower frequency hisses than smaller males. There was a trend for males that mated to be larger than males that did not mate. However, characteristics of the' courtship hiss were not significantly related to mating success after controlling for differences in male weight. Finally, we found that the dominant frequency and duration of courtship and agonistic hisses of males did not differ suggesting that males are using similar signals during both courtship and agonistic interactions. We interpret these results in light of the possible functions of hissing, male-male competition, and female discrimination during courtship interactions.

1 aClark, Deborah, C.1 aMoore, Allen, J. uhttp://booksandjournals.brillonline.com/content/journals/10.1163/156853995x00630