1982] Holldobler & Engel- Siegel — Glands in Male Ants 1 17 
abdominal tergites. In recent investigations this gland was found in 
workers of representative species belonging to all subfamilies, 
except in the Formicinae. Although the structure and size of the 
gland varies greatly, its wide distribution led us to conjecture that 
this gland might be a primitive monophyletic trait in ants generally, 
perhaps reaching back to the typhioid (or mutilloid) wasp ancestors 
of ants. In fact, we have recently found first indications that this 
gland is also present in some living typhiid wasps. 
Since this gland is anatomically closely associated with the last 
exposed tergite in female ants (7th abdominal tergite = pygidium) 
Kugler (1978) suggested that it be called the pvgidial gland. Of the 
several tergal glands recently discovered, the pygidial gland appears 
to be the most frequent in occurrence. Moreover, in several species 
its secretions have been found to serve as pheromones (Holldobler 
et al 1976; Holldobler and Haskins 1977; Maschwitz and Schonegge 
1977; Kugler 1979; Holldobler and Traniello 1980 a,b; Traniello and 
Jayasuriya 1981). The pygidial gland seems to be homologous with 
the “anal glands” of the dolichoderine ants described by Pavan and 
Ronchetti (1955). As we pointed out previously (Holldobler and 
Engel 1978) the term “anal gland” is misleading, because the gland 
does not exit from the anal or cloacal opening of the gaster, but 
between the 6th and 7th abdominal tergites. We therefore suggested 
to refer the dolichoderine structure to the pygidial gland. Recently 
Jessen and Maschwitz (in press) proposed to name the pygidial 
gland in honor of its discoverer Charles Janet. Thus we have now 
three names for this tergal gland: anal gland, pygidial gland and 
Janet’s gland. 
Because the anatomical designation of the organ in ant workers (a 
criterion we prefer) has been used in several recent publications, we 
will continue to call the tergal gland opening between the 6th and 
7th abdominal tergites pygidial gland. 
Table 1. (Following pages) List of species that were investigated 
histologically, and of the types of tergal and sternal glands found. When 
the histological series was incomplete and we could not make a definite 
statement, or when we could not clearly identify glandular ducts, the 
column is marked with a “?”. r=with reservoir; c=with cuticular 
structure. 
