Figure 1. Colony cycle of an annual Polistes paper wasp, exemplified by P. metricus.

Foundress Phase. The annual cycle begins (arrow) when a female emerges from quiescence with undeveloped ovaries. Feeding at flowers enables ovary development, which precedes nest founding. Oviposition occurs immediately upon initiation of each nest cell by the now-maternal foundress (crown icon) that performs all maternal behaviors in addition to oviposition, including foraging, feeding larvae, and nest construction. Worker Phase. The first female offspring to emerge in early summer are workers (hard hat icon) that forage, feed larvae, and construct the expanding nest. The foundress transitions into a true social queen that limits her activities to oviposition and feeding larvae using foods brought to the nest by workers. Reproductive Phase. Female offspring emerging in mid-summer are non-working future foundresses called gynes (tiara icon). Males, not shown, emerge synchronously with gynes. Decline Phase. In late summer the queen and workers die. Gynes and males depart the nest, which now is empty and unattended. Quiescence Phase. Gynes and males mate and feed at fall flowers. With the decline in flower availability and onset of cooler weather, males die, and gynes enter a torpor-like quiescence in sheltered concealment. Workers live only from their emergence until the end of the reproductive phase. A gyne from the reproductive phase of one year has the potential to become a foundress and then queen the following year, thus the tiara to crown transition can represent single individuals that pass from gyne to foundress to queen. Larval treatments and sampling reported in this experiment were conducted in the Reproductive Phase of the colony cycle.