1977] Leuthold & Bruinsma — Behavior in Hodotermes 
111 
The swarming referred to in this paper occurred on 9.4.1976 
around 16.30 h (sunshine). After a short dispersal flight of less 
than 100 m the alates landed individually on the ground, shed their 
wings and started running about. [Behavioral details of this se- 
quence have been described by Hewitt and Nel, 1969.] The MALES 
rambled with their abdomens raised in “calling” position and their 
huge sternal glands exposed (Fig. 1) in search of a digging ground. 
If they found a suitable place, they started excavating a hole by 
flicking dust particles out with their legs and often carrying out 
soil bits with their mandibles. They held their abdomens perma- 
nently in “calling” posture as long as they were unpaired. The un- 
paired FEMALES, on the other hand, repeatedly climbed on ele- 
vated structures, such as grass stems, apparently for “sniffing” for 
the male’s scent. They clearly perceived the presence of a male 
from a distance of at least 2.5 m up-wind. They obviously became 
excited and ran slightly zig-zag towards the calling male, without 
ever failing to reach the goal (Fig. 2). The joining of a female with 
a digging male was analyzed from two film sequences: the female 
touched the male’s abdomen with her antennae. On this stimulus 
the male turned around through 180° and palpated the female’s 
abdomen intensively. With that stimulus, apparently, the male 
terminated calling behavior by lowering the abdomen and retract- 
ing the sternal gland. Henceforth, no other female was attracted, 
not even from as close as a few cm. The pair met either during the 
male’s exploratory run or on the male’s digging site. In the latter 
case the female participated in excavating behavior. In the first 
case she followed him in loose formation on the search for digging 
terrain. However, no rigid pattern such as that referred to as “tan- 
dem run” was ever observed. In suitable ground the digging pair 
disappeared from the surface within minutes. If the substrate was 
too hard or if the pair was attacked by ants, the male went on in 
search of another place, followed by the female. In the postflight 
behavior the male was always the attracting and leading partner 
and the female actively hunted for him. If the female was experi- 
mentally removed from the pair, the male resumed calling behavior. 
However, reproductives collected in petridishes did not resume sex 
attraction behavior when separated and tested the day after flight 
in the laboratory nor did a crushed male sternal gland under those 
conditions attract any female. 
