1983] 
Ross & Visscher — Polygynous Vespula 
187 
queen brood (MacDonald & Matthews, 1981; Spradbery, 1971; 
MacDonald et al., 1974). Perhaps pressure to find empty cells 
resulted in workers ovipositing in large cells. 
The presence of many laying workers in large, diffuse nest struc- 
tures is not unexpected if queens exert reproductive control via the 
dissemination of volatile or trophic pheromones (Ikan et al., 1969; 
Landolt et al., 1977). The percentages of laying workers and super- 
numerary eggs we found were in close agreement to those reported 
for a queenless nest of Vespa simillima Smith (Yamane, 1974). Lay- 
ing workers in annual vespine colonies may be common during the 
phase of colony decline, also suggesting diminished queen control 
(R. W. Matthews, personal communication; Montagner, 1966; Akre 
et al., 1982). These workers appear to occupy regions of the nest not 
frequented by the queen (Edwards, 1980). 
Vespine workers may indeed represent “hopeful reproductives” 
(West Eberhard, 1978; Lin & Michener, 1972): while many workers 
never lay eggs, a significant proportion of them do and all can be 
regarded as having some probability of directly contributing genes 
to subsequent generations. This factor has not been adequately con- 
sidered by theories attempting to explain the origin of eusociality by 
reference to a polarized view of reproductive castes (Hamilton, 
1964a, b; Alexander, 1974; Spradbery, 1973a). 
Our discovery of an overwintered, polygynous colony of V. macu- 
lifrons completes the series of free-living North American species in 
the subgenus Paravespula with this life history. No members of the 
subgenus Vespula or the aerial-nesting genus Dolichovespula have 
been reported to exhibit this atypical colony cycle (Akre & Reed, 
1981). Perennial, polygynous colonies have been reported for V. 
squamosa (Tissot & Robinson, 1954; Ross & Matthews, 1982), 
whose affinities with other Vespula species are unclear (Akre et al., 
1980; see also Archer, 1981). 
The ability to retain colony social cohesion through two develop- 
mental seasons and to tolerate the existence of multiple functional 
queens points to great ecological and behavioral plasticity in the 
subgenus Paravespula. Members of this group differ from other 
temperate vespines in additional biological attributes including: (1) 
delay of reproductive production until fall or early winter and con- 
sequent increased duration of colony life span, (2) development of 
populous colonies and large nests, (3) ability to successfully colonize 
