Morphology and closing mechanism of the mandibular gland orifice in ants (Hymenoptera: Formicidae).

The mandibular gland of ants releases chemical compounds with functions ranging from nestmate alarm and recognition to antimicrobial defense. While the morphology of this ethologically important gland is well investigated in several species, the mechanism of secretion release in ants was not explicitly addressed so far. To clarify this question, we examined the anatomy of the gland orifice in ant species from 14 different subfamilies employing different techniques. The orifice close to the mandibular base is located on an area called mandalus. Our investigations revealed variation in mandalar shape, with clear trends in different subfamilies. By contrast, the internal organization is remarkably congruent across all investigated species. The thin external mandalar cuticle is always connected to the mandibular gland duct by a cuticular lamella, visible as a characteristic anchor-shaped structure in cross section. The slit-like gland orifice at the distal end of the mandalus is usually crescent-shaped. In some ant species with specialized mandibles such as trap-jaws, the organization of the orifice area is adapted to the mandibular shape, but always retains the general internal organization. No muscles were found in association with the orifice, nor with any other part of the mandibular gland. However, the base of the mandalus is connected to the prepharyngeal sucking pump by a cuticular ligament. Additionally, it is continuous with the conjunctiva connecting the mandible to the head capsule. We propose that retraction of the sucking pump by the muscle M. tentoriobuccalis, potentially in concert with opening of the mandible, stretches out the ligament and thus pulls on the mandalus and mandalar lamella to open the gland orifice and allow for secretion release. This hypothesis is congruent with findings on other aculeate Hymenoptera and expands our knowledge on the function of an important gland of ants.