Chemosystematic Studies of the Triboliini (Coleoptera: Tenebrionidae): Phylogenetic Inferences from the Defensive Chemicals of Eight Tribolium spp., Palorus ratzeburgi (Wissmann), and Latheticus oryzae Waterhouse1

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Abstract
The organic chemicals in the defensive secretions of 10 species representing both genus groups in the tenebrionid tribe Triboliini (subfamily Tenebrioninae) have been identified and quantified. All taxa contain substantial quantities of 2-methyl- and 2-ethylbenzoquinone with approximately 10-fold less of the corresponding hydroquinones. These are the only components in Palorus ratzeburgi, the only species examined from the Palorus genus group. Within the Tribolium genus group, the defensive secretion of Latheticus oryzae also contains 15-, 16-, and 17-carbon olefins. Eleven nonquinoid chemicals occur in the eight species of Tribolium examined. Members of the confusum species group (Tribolium anaphe, Tribolium confusum, Tribolium destructor) contain the 15-, 16-, and 17- carbon olefins. Those in the castaneum species group (Tribolium castaneum, Tribolium freemani, Tribolium madens) contain these olefins and two aromatic ketones. Species in the brevicornis group (Tribolium brevicornis, Tribolium audax) contain the olefins, the same aromatic ketones, and two aromatic esters. No taxonomically significant differences were found between the secretions of males and females. A comparison of the defensive secretions from six populations of T. castaneum, four of T. confusum, and two of T. audax indicated only quantitative intraspecific variation. A new species-group placement of T. audax is suggested based on the available morphological, biochemical, and biogeographical evidence; and chemosystematic inferences regarding the phylogeny of Tribolium are presented.

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