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Structure

Two main cell types constitute the defensive osmeterium gland of Papilio larvae. Ellipsoid gland cells have an extensively infolded basal plasma membrane, abundant ribosomes and whorls of smooth endoplasmic reticulum. The apical plasma membrane bears long microvilli extending into a mass of granular material containing electron-lucid cavities. Tangential slits occur in the epicuticle. Tubular arm cells contain heterogeneous, electron-dense inclusions, extensively-branched nuclei and large mitochondria sometimes distended with electron-dense material. The apical plasma membrane bears short microvilli. The inner, dense epicuticle forms a complex ramifying system. The two-phase defensive fluid consists mainly of water, 2-methyl propionic acid, and 2-methyl butyric acid.

Osmetrial secretion

A survey of the present results and others described previously on osmeterial secretion makes it possible to classify the Papilionid species into at least two large categories in terms of the chemical property of osmeterial secretion. One is a group in which the chemical constitution of osmeterial secretion of the last larval instar markedly differs in quality from those of the younger larvae. The results derived from Papilio protenor (HONDA, 1980), P. demodocus (BURGER et al., 1978) and other Papilio species (P. helenus, P. machaon, P. memnon, P. bianor, P. maccki, P. xuthus, etc., unpublished work) may assign the genus Papilio (tribe Papilionini) to this group, which can be termed 'heterogeneous type'. The genera Luehdorfia (tribe Zerynthiini), Graphium (tribe Graphiini) and Atrophaneura (tribe Troidini) apparently belong to the other group in which no qualitative change of osmeterial secretion occurs at the last larval ecdysis. This group can be designated as 'homogeneous type', which is further subdivisible into three types. The reciprocal relation and classification among the four genera on the basis of chemical nature of larval osmeterial secretions are summarized in Fig.