Chemical polymorphism in defense secretions during ontogenetic development of the millipede Niponia nodulosa

Abstract

A mixture of defense compounds (benzaldehyde, benzoyl cyanide, benzoic acid, mandelonitrile, and mandelonitrile benzoate), found commonly in cyanogenic polydesmid millipedes, was identified in the non-cyanogenic millipede Niponia nodulosa. These compounds were major components in 1st-4th instars, but were absent in older instars and adults. Extracts of older instars and adults contained 1-octen-3-ol, 2-methyl-2-bornene, E-2-octen-1-ol, 2-methyl-isoborneol, and geosmin; these compounds were minor components in 1st-4th instars. This ontogenetic allomone shift may be explained by the high cost of biosynthesis of polydesmid compounds from L-phenylalanine being offset by their potency in protecting the insect during fragile and sensitive growth stages. However, as the cuticle hardens in older juveniles (5th, 6th, 7th instars) and adults, this allows for a switch in defense to using less effective and less costly volatile organic compounds (presumably microbial in origin) that are ubiquitous in the millipede's habitat or are produced by symbiotic microbes and may be readily available through food intake or aspiration.