1984] 
Frank — Fig wasps 
301 
across the intersegmentus, while maintaining their angle relative to 
the median sternal line. This movement of the coxae is accomp- 
lished by sliding the pronotum forward and down, which causes the 
prosternum to form about an 80° angle with the mesosternum. The 
coxae are now against the flattened medial region of the mesoster- 
num (Fig. 3). Next the coxae are brushed laterally over the meso- 
thoracic pockets (Figs. 9,10). The lateral membranous flap of the 
pockets (Fig. 4) may aid in scooping the pollen from the coxal 
pockets. The coxae are brought forward again when the pronotum 
is raised; thus the coxae are moved in a circular motion. The 
sequence is usually repeated 3-8 times, although the observed range 
was 2-34 (n=15). Sometimes two or three females will collect pollen 
from the same anther. If only a small amount of pollen is collected, 
the wasp will find another anther and repeat the pollen collecting 
sequence. 
After collecting pollen the wasps leave the syconium through the 
exit tunnel and quickly take to the air, perhaps because predaceous 
ants of the genus Pseudomyrmex are often waiting for escaping 
wasps. Those wasps that successfully find receptive syconia begin 
the cycle again. 
Discussion 
The use of behavioral characters in systematics has been reviewed 
and discussed several times (e.g., Lorenz 1950; Evans 1952; Mayr 
1958; Alexander 1962; Blair 1962; Atz 1970; Hinde 1970). Three 
examples are Lorenz’s (1941) phylogeny of ducks and geese (Anati- 
dae) based solely on behavioral characters, leading to Delacour and 
Mayr’s (1945) revision based on both morphology and behavior, 
Evans’ ( 1 952) work on the comparative behavior and systematics of 
spider wasps (Pompilidae), and van den Assem et al.’s (1982) subdi- 
vision of Melittobia (Eulophidae) based on mating behavior. Also, 
several workers have noted a reasonable congruence between 
inferred phylogenies based on behavior and those based on mor- 
phology (e.g., Speith 1952 for Drosophila; Crane 1952 for mantids; 
and Michener et al. 1978 for Apidae). 
The phylogeny of the Agaonidae is controversial (Wiebes 1982), 
and much work remains. Behavioral characters have not been used 
in fig wasp systematics, simply because there is not enough compar- 
ative information available. Fig wasps seem ideal for comparative 
studies of behavior, since so much of their behavior is extremely 
