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Psyche 
[Vol. 90 
Several to 50 queens are typically present in perennial Vespula 
colonies, although Spradbery (1973b) found up to 1000 in perennial 
nests of V. germanica in Tasmania. On the other hand, Thomas 
(1960) reported only a single queen in each of the perennial V. 
germanica nests he studied from New Zealand. Six of the 23 queens 
we found in the study colony were inseminated and possessed well 
developed ovaries; these were probably functional queens. The large 
number of worker brood we found corroborates the evidence for 
several egg laying queens. An additional two queens were probably 
former reproductives. The remaining queens could have emerged 
recently, as queens at all stages of development were present in the 
colony. Newly emerged queens in perennial colonies may mate in 
the nest with sibs (Ross, 1983; R. E. Wagner, personal communica- 
tion) or embark on mating flights and return to the parental nest as 
newly recruited reproductives (Spradbery, 1973a, b). Thus, func- 
tional queens in polygynous Vespula colonies are typically regarded 
as being daughters of the original foundress (Spradbery, 1973b; 
Edwards, 1980; but see Ross & Matthews, 1982). 
The study colony contained fewer large cells than do average 
annual V. maculifrons colonies, indicating that fewer than normal 
queens had been reared during the first developmental season. The 
presence of large numbers of male and queen brood and adults 
indicates that the colony had been rearing reproductives throughout 
the winter [as is typical for other perennial Vespula (Edwards, 
1980)], and a newly initiated queen comb with eggs suggests that 
queens would have continued to have been reared into the spring. 
Presumably the number of queens produced over two seasons 
would far exceed the productivity of an annual colony. 
Over one-half of the capped brood sampled in small cells were 
males. The occurrence of so many male brood in the spring, the 
great number of supernumerary eggs, and the positions of eggs high 
on the cell walls suggest the likelihood of laying workers (R. W. 
Matthews, personal communication; Akre et al., 1982). Dissections 
confirmed that at least 14% of the workers possessed well developed 
ovaries and were probably ovipositing. 
Greater than 30% of the brood being reared in large cells at the 
time of colony collection were males. This represents a considerably 
larger figure than has been previously reported for annual Para- 
vespula colonies, in which large cells contain almost exclusively 
