Defensive Allomones Function as Aggregation Pheromones in Diapausing Ladybird Beetles, Hippodamia convergens

Abstract

Identification of the stimuli responsible for the formation of an aggregation can be used to distinguish between social and non-social aggregations and help in the process of identifying the adaptive benefits of the gregarious behavior. The convergent ladybird beetle, Hippodamia convergens, forms dense aggregations during winter diapause. The mechanisms of conspecific attraction and hibernacula site selection of H. convergens are not well understood. In laboratory and field bioassays, we evaluated the role of three defensive compounds in the formation of H. convergens aggregations. Diapausing H. convergens aggregated within the section of an arena exposed to alkylmethoxypyrazines. 2-Isobutyl-3-methoxypyrazine (IBMP) caused the strongest aggregative effect. Beetles also aggregated to some doses of 2-sec-butyl-3-methoxypyrazine, but appeared to be repelled at higher doses. A third constituent, 2-isopropyl-3-methoxypyrazine, generally had little effect on the distribution of beetles, although the highest dose tested was repellent. Beetles also aggregated to a blend of these alkylmethoxypyrazines at their natural ratio. During fall migration to overwintering sites, more beetles aggregated in artificial hibernacula baited with IBMP, confirming its function as an aggregation pheromone. These three pyrazines also function as warning odors that, in conjunction with other aposematic displays (contrasting red and black coloration, gregarious behavior, reflex bleeding), contribute to the multi-modal, anti-predatory defense of coccinellid beetles and some other arthropods. Confirmation of the role of some alkylmethoxypyrazines in coccinellid aggregations suggests that these defensive allomones have been co-opted for intraspecific communication.