1987] 
Letourneau & Choe — Wasps and ants 
85 
nymphs also produced honeydew, but presented droplets on the 
dorsal surface about 2 mm from the tip of the abdomen. It appeared 
that nymphs did not have fully developed tubes. 
Aetalion reticulatum appeared to discriminate between attend- 
ants. The mean number of droplets per feeding bout was signifi- 
cantly higher (Kruskal- Wallis test: p < 0.001) when P.fraternus was 
soliciting than when either C. abdominalis, the major nocturnal 
tender, or Z. porrasi were tending (Table 1). The size of droplets 
from the same adults was consistently larger (about twice in diame- 
ter) when P.fraternus was tending. Duration of feeding bouts, how- 
ever, did not differ (Table 1). In the presence of P. fraternus, 
aetalionids commonly raised a pair of legs nearest an approaching 
ant, kicked at it, and prevented it from collecting honeydew. Thus 
A. reticulatum appeared to play an active role in selecting an 
attendant. 
Discussion 
Associations of Hymenoptera with Homoptera cover the entire 
range of interaction categories including predation (e.g., Evans 
1968, Hook 1981), parasitism, mutualism, commensalism, and 
behavioral combinations such as mutualism-predation (Pontin 
1958) and parasitism-predation (Gerling 1966). It is well known that 
wasps and bees, like ants, depend upon homopterans as sources of 
carbohydrates (Evans and West Eberhard 1970, Spradbery 1973); 
however, the behaviors associated with honeydew collection in 
wasps and bees have not been reviewed. 
Our literature survey and field study revealed that bees and wasps 
exhibit nearly as great a range of interactions as do ant associates. 
Honeydew foraging of bees and wasps involves various types of 
behavior ranging from little or no contact with Homoptera to dis- 
plays of active herding and guarding (Table 2). Unlike some ants 
(e.g., Maschwitz and Hanel 1985), however, no bees or wasps have 
been observed to move homopterans to favorable feeding sites. At 
least eight species of Trigona bees and ten species of polistine wasps 
are now known to tend and collect honeydew from various Homop- 
tera. It is interesting to note that all hymenopteran species known to 
collect honeydew directly from Homoptera are meliponine bees and 
polistine wasps. Other social hymenopterans such as honey bees, 
bumble bees, and vespine wasps have been observed only to obtain 
