Glands, muscles and genitalia. Morphological and phylogenetic implications of histological characters in the male genitalia of Fritillary butterflies (Lepidoptera: Nymphalidae: Argynnini)

The histology and internal morphology of the male valve in a broad sample of Fritillary butterflies are examined. A large internal ridge to which the extensor of the valve muscle fastens is probably an autapomorphy for the tribe Argynnini. A large glandular pocket is found in the valves of all examined Heliconiinae, but is missing in other examined Nymphalidae. The pocket shows considerable variation between the higher taxa. In the higher Fritillaries (subtribe Argynnina) the pocket probably has an effective closing mechanism allowing the male to store a considerable amount of glandular secretion. It has earlier been suggested that the secreta from the pocket glands are transferred to the female during copulation, where they probably have an antiaphrodisiac effect, preventing other males from mating a once mated female. All examined male Fritillaries also have glands in the anellus, the terminal membrane between the valves. In Boloria these glands are extremely well developed and abundant, a condition that is probably an autapomorphy for that genus. The obtained characters are tested against a known phylogeny of the tribe Argynnini.