1977] Leuthold & Bruinsma — Behavior in Hodotermes 
115 
Nasutitermes lujae 1 mm 
pseudergate 
Kalotermes flavicollis 
<o 
Figure 3. Comparative sizes of termite sternal glands: the contours of the glands 
in top view of H. mossambicus, reproductive male and female (after flight) and 
major worker; those of T. bettonianus, characterized by Leuthold and Liischer 
(1974) as unusually hypertrophic in the reproductive caste; those of other species 
[N. lujae after Pasteels (1965), K. flavicollis from own drawings]. The glands were 
drawn from fresh whole-mount preparations in Ringer solution. 
postflight behavior had been assumed from calling behavior by 
several authors (reviewed by Stuart, 1969). A few species have 
been more closely analyzed: In Kalotermes flavicollis (Kalotermiti- 
dae) calling females have been only occasionally observed with 
exposed sternal glands (Wall, 1969). A more intensive relationship 
of the sexes was found in the tandem behavior in which either the 
male or the female could be leader. Both sternal and tergal glands 
were experimentally verified as sources of sex attraction from lab- 
oratory bioassays. The sternal gland of the female was exclusively 
male attractive. The male tergal gland was dominantly attractive 
to females and to a lesser extent to males, and the female tergal 
gland was slightly active towards both sexes (Wall, 1971). Pasteels 
(1972) reported calling courtship but no distinct tandem pattern in 
Zootermopsis nevadensis (Hodotermitidae). In most cases the fe- 
male was the calling partner but apparently for the first time also 
