Structure and function of the eversible glands of the aquatic firefly Luciola leii (Coleoptera: Lampyridae)

Summary.  When attacked or otherwise disturbed, larvae of the aquatic firefly Luciola leii display fork-shaped glands laterally from the meso- and metathorax as well as each abdominal segment. Glandular eversion is accompanied by a strong pine oil-like odour, thanatosis and glowing from paired larval light organs. Observations by SEM and TEM revealed that there are numerous, almost spherical protuberances, measuring 9 μm in diameter on the surface of each gland. Each protuberance is connected to a well developed secretory cell via a 0.1 μm thick and 0.2 μm long stalk and bears three to six 7 μm long spines on its apex. The convoluted glandular walls measure 0.2 μm. The cytoplasm of the secretory cells is characterized by the presence of numerous mitochondria and an extensive system of cisternae and tubular endoplasmic reticulum. Preliminary GC-MS analysis of the glandular secretion revealed two volatile terpenoids: terpinolene and γ-terpinene. Choice and no-choice bioassays involving fish and ants as well as other predators demonstrate that the secretions serve as an effective deterrent against a range of ecologically relevant enemies. The larval postural adjustments, light emission, everted glands, and glandular release of distasteful or repellent compounds, appear to function as a multi-modal, aposematic larval defence. The tandem evolution of glandular secretions and conspicuousness in larval fireflies could partially explain their successful radiation into both aquatic and terrestrial habitats.