Satellite Nests, Early Males, and Plasticity of Reproductive Behavior in a Paper Wasp

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Abstract
We studied 298 primary nests of a single population of the paper wasp Polistes fuscatus variatus over a period of 2 yr. This population demonstrated a remarkable amount of reproductive behavioral plasticity. At least 14% of all nests produced a few males early in the season, before the emergence of the majority of reproductives. At least 17% of all nests were part of polydomous colonies that consisted of 2-9 separate nests with 2-9 foundresses. During the first year of the study, 10 satellite nests were founded, some of which contained mated, spring-produced females and, in late summer, produced male and female offspring. Early males apparently mate with early-season reproductive females that supercede orphaned nests or oviposit on satellites. The small number of males produced may represent a trade-off between their early-season reproductive value and their cost later in the season. The cost of an early male is the reproductive value of males and females that could have been produced in late summer if resources had been invested in workers. Satellite nests were closely associated with polydomous colonies. This suggests that a reduction in the ability of alpha foundresses to maintain their reproductive dominance of polydomous colonies leads to the expression of alternative reproductive behavior. Functional polygyny results in reduced genetic relationships between adults and reproductive offspring and should favor behavior that increases individual reproduction. Satellite nests were functionally like additional nests in polydomous colonies; however, alpha foundresses (queens) were never observed on them. Instead, spring-produced, female offspring appear to have been the sole egg layers on satellite nests. Other unmated, nonreproductive (presumably worker) offspring incorporated satellite nests into the set of nests they worked. Our results demonstrate the difficulty of testing kin-selection theory. Recent studies of kin recognition in the social Hymenoptera show that estimates of the genetic relationships of specific interacting individuals are needed because some important interactions among nest mates are deterministic and biased according to relatedness. In addition, our demonstration of the large repertoire of alternative reproductive options available reaffirms the need for more-comprehensive studies of reproductive biology before assessing the relative roles of genetics and ecology in the evolution of insect societies.