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Investigations were made on the pygidial defensive components and gland morphologies of 132 species in 60 genera of carabid beetles. Components so far detected may be divided into six groups, : (1) formic aicd, (2) methacrylic, tiglic and/or ethacyrlic acids, (3) other acids, such as n-butyric, iso-butyric, iso-valeric, iso-crotonic, angelic and several other acids (4) m-cresol, (5) benzoquinones with one chamber, (6) benzoquinones with two chambers. Gland types so far observed may be divided into three types : (1) spherical lobes which produce fatty acids, (2) elongated lobes which produce m-cresol, (3) long thick lobes which produce benzoquinones. There were also three different kinds of fatty-acid-producing glands at the points of entry of the collecting canals to the resevoris. The same genera, or at least the same subgenera, shared similar components and gland types.
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Kanehisa et al. (1977) studied this question.
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