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[Vol. 90 
ments and determining their sex (n = 528). We counted and sexed all 
pupae and pharate adults in large cells. 
To estimate the sex ratio of the adult population of the nest we 
weighed the two sub-populations of workers and males retrieved 
from the nest interior and the entrance traps. We then weighed, 
counted, and sexed 4 and 5 samples, respectively, of these popula- 
tions. The mean number of wasps/ g and the mean sex ratios of these 
samples were extrapolated to the population and sex ratio of the 
colony adults as a whole. 
Reproductive status of queens and workers 
We dissected all queens in the nest and classified their reproduc- 
tive status as follows: “Developed” ovaries had oocytes greater than 
2 mm in length, “undeveloped” ovaries had oocytes less than 1 mm 
in length. “Inseminated” queens had spermatozoa visible microscop- 
ically in a squash mount of their spermathecae, while “not insemi- 
nated” queens had none. 
To estimate the reproductive status of workers in the colony, we 
dissected a random sample of 100 workers and examined their ova- 
ries. We classified them as “undeveloped” [ovarian index less than 1 
(Cumber, 1949), oocytes not developed], “moderately developed” 
(ovarian index 1-2, maximum oocyte length 0.9 mm), or “well devel- 
oped” (ovarian index 3-12, oocyte length greater than 1.0 mm). 
Results 
Nest structure and brood composition 
Estimated cell number and brood composition of the nest are 
given in Table 1 (see also Fig. 1). From comb measurements the 
volume of the nest cavity was estimated to be ca. 80 liters. The nest 
structure was roughly ellipsoid, 71 cm by 45 cm. Two nest entrances 
were in use at the time of discovery; flight activity from each was 95 
± 14 and 15 ± 6 wasps/ min returning at midday on 1 1 March (here 
and throughout this paper numbers in this form denote mean ± 1 
SD, except as noted). The combs were present in at least 15 layers. 
Total comb area was 15,652 cm^; the combs contained an estimated 
100,120 cells. 
Small (worker-size) cells comprised 99.3% of all cells in the nest 
(4.0 ± 0.20 mm in diameter between parallel sides, range 3. 7-4. 7 
mm, n = 44). Of the small cells, 16.7% contained eggs; 41.9% of 
