Ultrastructural and phylogenetic assessment of wax glands in pit scales (Homoptera : Coccoidea)

TEM/SEM and computerized images of 5 wax glands for 3 type species of Coccoidea (Homoptera): Asterodiaspis variolosa (Asterolecaniidae), Cerococcus quercus (Cerococcidae) and Lecanodiaspis sardoa (Lecanodiaspididae) were studied. Their cuticular structures were compared with 142, 56, and 61 species in their respective families to determine relationships among pit scale taxa. Significant differences include: the morphology of the outer and inner ductule of the tubular duct gland, structure of the pores (8-shaped, multilocular and quinquelocular), and the absence or presence of cribriform plates and their structural variations. Three distinctive tubular duct shapes (asterform, ceroform and lecanoform) are common in pit scale species. Apomorphic characteristics of the asterform tubular ducts include an absence of the inner ductule and the progressive reduction of the outer ductule's diameter from the pore to its inner end. These characters easily separate asterolecaniids from the cerococcid-lecanodiaspidid lineage. The constricted lecanoform tubular ducts and the curved teeth on the rim at the inner end of the outer ductule in the ceroform tubular ducts are regarded as autapomorphic. The presence of 8-shaped pores is considered a plesiomorphic condition. Specific cuticular variations of the 8-shaped pores, characterizing familial taxa, include pores even with the surface in asterolecaniids, pores with raised walls in cerococcids, and bent pores in lecanodiaspidids. The dominant 8-shaped pore patterns in pit scales are those arranged in a marginal band in lecanodiaspidids, in a swirl-like pore pattern in the cerococcids, and in a marginal row in asterolecaniids. A divergent evolutionary trend is noted for the structure of the cribriform plate; they are with micro-orifices in cerococcids, but without micro-orifices in lecanodiaspidids. The former state is considered apomorphic. Cribriform plates arranged in clusters characterize the cerococcids, while plates in longitudinal rows characterize the lecanodiaspidids. These data confirm the concept that the pit scales constitute a paraphyletic group and the Asterolecaniidae, Cerococcidae and Lecanodiaspididae are monophyletic.