display a unique, behavioral phenotype demonstrated by only a small percentage of workers. These individuals leave their natal nest to overwinter to found new colonies in the spring. Compared to nonhelpers, these helpers exhibit higher overall levels of activity, demonstrating higher frequencies of behaviors. Specifically, they gave more trophallaxis, attacked more, and received more ritualized dominance behavior (RDB). Nonhelpers received more trophallaxis and performed more RDB. The survival rate of helpers is around 14%, while the survival rate of nonhelpers is around 59%. These behavioral divisions are not permanent. For instance, if an alpha female is removed from a nest, then another female, usually the second-most dominant, beta female, assumes the role and behavioral profile of the removed dominant. Indeed, individuals alternate between different profiles of behavior within their own dominance rank position. When larvae were artificially removed, the frequency of worker reproduction increased. Therefore, workers lay eggs when they perceive a decline in queen power, as demonstrated by artificially empty cells. The interactions of females in the nest can influence which daughters become workers and which become gynes. Despite some minor physiological differences (primarily in the fat body), gyne-destined females produced late in the colony cycle can be induced to become workers if placed on nests that are at an earlier stage of colony development, and the reverse is also true. This indicates a significant degree of flexibility in the caste system of this species. Female dominance Typically, the alpha female dominates all other individuals of a colony, laying the majority of eggs and partaking in differential oophagy. The alpha female devotes much of her time to social interaction, in comparison to subordinates that are much more involved in foraging and brood care. The clypeus - the yellow region above the mouth, in other words the face - is extremely variable in number, size, and shape of black spots, and this variation correlates to dominance; more dominant individuals have more black spots. The arrival at the nest correlates with the dominance hierarchy. Therefore, individuals that join the nest later are seldom dominant. P. dominula shows distinct behavioral differences in response to face marking. Researchers used paint to alter the number of facial spots on two wasps of the same size after killing them. They put these dead wasps as guards in front of food sources and introduced a third wasp to see where this wasp would go. The third