104 
Psyche 
[March 
Subordinate females check the cells periodically but do not deposit 
eggs in them. They feed the larvae material brought in by them or 
by other cofoundresses. They also “smear” to some degree, although 
most of this is done by the queen (Hermann and Dirks, 1974). 
It is obvious at this stage that the subordinate cofoundresses be- 
come workers (beta individuals, Pardi, 1948), in spite of the fertile 
condition of their spermatheca. They are subordinate to a single 
dominant queen (alpha individual). The establishment and main- 
tenance of a well defined dominance hierarchy takes as long as two 
and a half months, until the first emerging workers arrive. 
The emergence of workers further complicates the dominance 
hierarchy since workers begin competing with the subordinate co- 
foundresses for dominance and the workers evidently put the cofound- 
resses under extensive pressure at this time (about June 16, Table 
1 ) ; with the appearance of the first workers, the subordinate co- 
foundresses leave the nest, never to return, leaving only the dominant 
queen and workers (new beta individuals) on the nest. Workers 
begin to “tail wag” to some degree and “smear,” behavior normally 
demonstrated by the queen or cofoundresses. The leaving of the 
subordinate cofoundresses is a strong indication of their extremely 
subordinate position in the colony. It appears that once a cofound- 
ress becomes subordinate it will be dominant to no wasp, except those 
already subordinate to her, including a worker. The queen (domi- 
nant cofoundress) usually remains on the nest through August but 
leaves the nest permanently long before cold weather. According to 
Pardi (1948), the queen (representing the alpha individual on the 
nest during most of the nest life) shows a lower dominance frequency 
upon growing old but maintains supremacy on the nest until later in 
the season. Rau (1939) indicated that his observations of P. annu- 
laris colonies likewise revealed first the arrival of workers and a sub- 
sequent departure of subordinate cofoundresses. 
Pardi (1948) reported that after the eclosion of P. gallicus work- 
ers and the elimination of auxiliary females (if the queen dies), a 
worker that is in the beta position on the nest begins to dominate its 
siblings and soon begins to deposit eggs. Rau (1931) also mentions 
egg deposition by workers when the queen is lost. Pardi does not 
mention the pressures of the dominance hierarchy as being put on 
colony cofoundresses, thus resulting in their permanent departure 
from the nest. 
The build up and atrophy of colony life encompasses approximately 
five months. This is much the same as the cycle found in colonies of 
P. canadensis (Eberhard, 1969). Colony enlargement ceases when 
