1975] 
Herjnann & Dirks — Polistes ananularis 
99 
Although they form aggregations, each adult wasp is an individual 
which functions almost independently from other wasps in the aggre- 
gation (i.e., no individual forms a “nucleus” as with the queen 
honey bee). This independence remains to some extent throughout 
colony life, although each wasp on a nest influences and is influenced 
by the other co-foundresses of the colony. 
In addition, the establishment of dominance hierarchies begins 
during the pre-hibernation period in late fall ( at the end of the 
nesting period) and continues through the post-hibernation period of 
early spring (Hermann and Dirks, 1975). These hierarchies later 
are instrumental in determining nest duties (division of labor among 
colony cofoundresses). Dominant females remain at the “parent site” 
and because of this struggle for dominance some of the cofoundresses 
are forced to disseminate. Each dominance hierarchy has, within its 
own structure, characteristic behavioral patterns that are present 
regardless of colony size. 
Dominance Hierarchies 
Each colony of these pleometrotic females commences under the 
pressure of dominance establishment. Such establishment often in- 
volves up to eight or ten females (cofoundresses) which are all 
making an attempt to become the dominant queen of the colony 
(alpha individual of Pardi, 1948). Although single Polistes cana- 
densis females have been reported by Eberhard (1969) to most often 
initiate a nest alone, similar to the fashion demonstrated by P. fus- 
catus, Rau (1933) has described colony founding by several P. cana- 
densis nestmates (siblings) that were chased from their original nest. 
The females of P. annularis , a close relative of P. canadensis , readily 
initiate their nests with a number of females. However, like with 
P. canadensis , additional females often join the P. annularis nest 
after initiation (Rau, 1940). 
A colony usually benefits by having a number of cofoundresses 
rather than a single queen, but maximum efficiency is not achieved 
because of dominance struggles. As an example, by April 15, 1971,, 
nest #18 occupied by a single queen had 6 cells. On the same date 
nests #16 and #17 with multiple queens (2 and 5 respectively) 
had 11 and 13 cells respectively; the number of cells, however, is 
not in proportion to the number of cofoundresses present. If the 
number of cofoundresses could be directly correlated with the cells 
present in a nest, we would expect nest #16 to have 12 cells on 
April 15 and nest #17 should have 18 on the same date. In addition, 
