In the evolution of caste-based societies in Hymenoptera, the classical insect hormones juvenile hormone (JH) and ecdysteroids were co-opted into new functions. Social wasps, which show all levels of sociality and lifestyles, are an ideal group in which to study such functional changes. Virtually all studies on the physiological mechanisms underlying reproductive division of labor and caste functions in wasps have been done on independent-founding paper wasps, and the majority of these studies have focused on species specially adapted for overwintering. The relatively little-studied tropical swarm-founding wasps of the Epiponini (Vespidae) are a diverse group of permanently social wasps, with some species maintaining caste flexibility well into the adult phase. We investigated the behavior, reproductive status, JH and ecdysteroid titers in hemolymph, ecdysteroid content of the ovary and cuticular hydrocarbon (CHC) profiles in the caste-monomorphic, epiponine wasp Polybia micans Ducke. We found that the JH titer was not elevated in competing queens from established multiple-queen nests, but increased in lone queens that lack direct competition. In queenless colonies, JH titer rose transiently in young potential reproductives upon challenge by nestmates, suggesting that JH may prime the ovaries for further development. Ovarian ecdysteroids were very low in workers but higher and correlated with the number of vitellogenic oocytes in the queens. Hemolymph ecdysteroid levels were low and variable in both workers and queens. Profiles of P. micans CHCs reflected caste, age and reproductive status, but were not tightly linked to either hormone. These findings show a significant divergence in hormone function in swarm-founding wasps compared with independently founding ones.
KEY WORDS: Challenge hypothesis, Cuticular hydrocarbons, Ecdysteroids, Epiponini, Juvenile hormone, Polistes
INTRODUCTIONEusocial insects within the order Hymenoptera (bees, wasps and ants) are characterized by the production of distinct phenotypes (caste polyphenism) in the female sex: the queen, the worker and often additional physiologically or morphologically specialized types within the worker caste (Wilson, 1971;Hunt, 2006;Hunt et al., 2007;Robinson, 2009;Tibbetts and Izzo, 2009;Shukla et al., 2013). In most caste-based societies the development of such specialized phenotypes is induced or at least biased in a pre-imaginal stage, where qualitative or quantitative differences in diet, pheromonal signals, mechanical stimuli or other environmental cues trigger endogenous signaling cascades toward alternative developmental trajectories (Wilson, 1971; Chavarría-Pizarro and West-Eberhard, 2010;Suryanarayanan et al., 2011;Hartfelder and Emlen, 2012;Penick and Liebig, 2012). As with most polyphenisms in insects, juvenile hormone (JH) and ecdysteroids were co-opted to orchestrate distinct ontogenetic pathways on top of their wellconserved role in regulating the episodic, molt-based growth events in immature animals (Hartfelder and Emlen, 2012)....