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[Vol. 89 
much smaller, and no pronounced reservoir could be detected. Lep- 
togenvs workers have two well developed sternal glands between the 
5th and 6th, and the 6th and 7th sternites respectively (Holldobler 
and Engel 1978; Jessen et al 1979). Leptogenvs males lack these 
structures but do possess sternal glands in the 7th and 8th sternites. 
In males of the ponerine ant Palt hot hy reus tarsatus we found large 
paired clusters of glandular cells in the 8th sternite. The glandular 
ducts open through the intersegmental membrane between the 8th 
and 9th sternites. In addition Paltothvreus males possess unpaired 
intersegmental sternal glands, similar to those found in Paltothvreus 
workers (Holldobler and Engel 1978), but smaller, between the 5th 
and 6th, and 6th and 7th sternites. 
Other abdominal glands; 
As indicated in table 1 we found several other abdominal glands 
in males which cannot directly be assigned to the group of tergal or 
sternal glands. In a few species ( Novomessor , Leptogenvs) we 
detected glandular cell clusters in the petiole. In Ectatomma, Dia- 
camma, Paltothvreus we found small tergo-sternal glands. The 
ducts of the glandular cells composing them open laterally through 
the pleural membrane. We found similar small glandular cell 
bunches in males of Pachycondvla and Rhvtidoponera, but we 
could not clearly identify the glandular ducts. These tergo-sternal 
glands resemble closely similar structures described by Jessen et al 
(1979) in workers of several ponerine species. Finally we found 
small groups of glandular cells directly at the anus of males in 
Pachycondvla, Ectatomma, Neivamyrmex, Eciton, Mvrmecocystus 
and Liometopum (Fig. 7b). These anus glands vary considerably in 
size, and it is possible that they are present in more species than we 
were able to document. We first found them in workers of Dorv/us 
(Holldobler and Engel 1978). The anus glands should not be mis- 
taken for the rectal gland, an invagination of a glandular epithelium 
of the rectum, recently discovered in Oecophvlla workers by Holl- 
obler and Wilson (1978). It is interesting to note that males of 
Oecophvlla also possess a small rectal gland. 
Discussion: 
Except for the glands associated with the ovipositor and sting 
apparatus, which the males lack, ant males are as richly endowed 
with exocrine glands as the females. In many species of ants the 
males have well developed mandibular glands, pro- and post- 
