1983] 
Strassmann & Orgren — Polistes 
245 
TABLE 4. ANOVA of the effects of date and location in nest on pupa development. 
Main Effects 
Location in nest Date 
Interaction 
Location in nest 
X date 
Small nest 
Sum of squares 
7 
33 
3 
F 
1.1 
11 
1.1 
df 
2 
1 
1 
% of variance explained 
8 
39 
4 
Medium nest 
Sum of squares 
21 
333 
180 
F 
0.4 
4.8** 
1.0 
df 
3 
4 
10 
% of variance explained 
8 
5 
Large nest 
Sum of squares 
57 
278 
105 
F 
1.4 
5.0 
0.8 
df 
3 
4 
9 
% of variance explained 
2 
7 
3 
df= degrees of freedom, *p<0.05, ♦♦p<0.01, ***p <0.001 
tages to flexibility in development time of larvae which allow more 
time for development when food is limiting, either because there are 
fewer females to harvest it, or because of a general scarcity of prey in 
the environment. 
Data on development times were found in the literature for 4 
populations of P. fuscatus, and 1 each of P. hunteri, P. annularis, P. 
gallicus and P. exclamans (Table 6). Development times varied from 
10 to 25 days for eggs, from 15 to 25 days for larvae and from 13 to 
22 days for pupae (Table 6). P. exclamans in this study falls towards 
the faster end of this range, particularly for pupa development time. 
All reports of pupa development times averaged over 18 days except 
Rabb’s study of P. exclamans in N. Carolina. P. exclamans was 
studied in the most southern climate, so it is possible that the differ- 
ences are due to temperature. There is a trend in P. fuscatus towards 
shorter pupa development times in more southern populations. P. 
exclamans and P. annularis are the only two members of the subge- 
nus Aphanilopterus represented here. It is possible that Aphanilop- 
terus, which generally has larger nest sizes and more adults tending 
