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[Vol. 89 
pharyngeal glands, maxillary glands, salivary glands, and meta- 
pleural glands, although the size of these various structures can vary 
considerably between the female castes and males. In this paper we 
surveyed specifically the abdominal sternal and tergal glands in ant 
males. 
In almost all species studied we encountered two major glandular 
structures that Janet (1907) had already described in Myrmica 
rubra, the penis glands and the subgenital plate glands. Also quite 
generally present in males (except in the Formicinae) are the pygi- 
dial glands. The males share these organs with the females, although 
less well developed in some species. An interesting case is Notho- 
myrmecia: here the males have a rudimentary pygidial gland but a 
weil developed postpygidial gland (between the 7th and 8th tergites). 
The males of the doryline ants are unusually well endowed with 
abdominal glands, in which they differ markedly from the workers. 
Although doryline workers have well developed pygidial- and post- 
pygidial glands (Holldobler and Engel 1978), the males have mas- 
sive glandular structures in each segment. In this context the 
findings by Whelden (1963) are of considerable interest. Whelden 
described a series of exocrine glands in the gaster of Eciton queens 
as follows: “Each of the segments of the gaster, including those 
telescoped together in the posterior part, contains a pair of these 
glands which are smaller in the anterior segment than those in the 
following segments”. We were not yet able to section a doryline 
queen and therefore cannot compare the queen organs with those 
we found in males. It appears, however, that the males possess a 
glandular equipment very similar to that of the queens. Presumably 
in doryline queens these massively developed exocrine glands play 
an important role in the queen’s chemical control of the worker ants 
and in her high attractiveness to workers, (Watkins and Cole 1966). 
We hypothesize that the males imitate queen pheromones, which 
might enable them to penetrate a foreign colony in order to get 
access to the wingless virgin female reproductives (Franks and Holl- 
dobler unpublished). In fact, this might also be the function of the 
massively developed sternal gland in Leptogenvs males. In this genus, 
as in the dorylines ergatoid reproductive females presumably mate 
in the nest, so that males flying in from other nests have to penetrate 
a foreign colony. 
