Structural megadiversity in leaf litter predators - the head anatomy of Pselaphus heisei (Pselaphinae, Staphylinidae, Coleoptera)

Abstract

The head anatomy of Pselaphus heisei (Pselaphitae) is described and documented. The structural features are evaluated in comparison with findings presented in earlier studies on the subfamily, with a special focus on correlations with predacious habits and the groundplan of Pselaphinae. We found the tentorium, labrum, maxillary palps, shape of head, and a system of dorsal pits and sulci highly variable within the subfamily, reflecting multiple transformations, including many homoplasious changes. The following major characters are identified as groundplan features of Pselaphinae: falciform mandibles; small mola; semiglobular neck; ventrolateral antennal articulation; steep clypeal region; setiform labial palpomere 3; tentorium with nearly vertical main branches and lacking laminatentoria; separation of tentorial bridge from tentorial arms; fusion of dorsal tentorial arms with the head capsule; large brain placed in the posterior third of the head; and a triple cluster of well-developed cephalic glands. The last feature supports a hypothesis that multiple and independent cases of adaptations to myrmecophilous habits observed in various lineages of Pselaphinae were possible by re-programming already existing glands to produce appeasement secretions. The cephalic muscle apparatus of P. heisei is similar to what is found in other staphylinoid groups, with some exceptions, whereas it is strongly modified in the myrmecophile Claviger testaceus. We propose that the unparalleled structural megadiversity in Pselaphinae is primarily linked with life in the upper soil layers combined with specialized carnivorous habits, with small and agile or mechanically protected arthropods as prey. Within the group, various specialized life habits have evolved, including myrmecophily, termitophily, and also life in deep soil or caves, each with unique morphological adaptations.