I. Foldi et P. Lambdin, ULTRASTRUCTURAL AND PHYLOGENETIC ASSESSMENT OF WAX GLANDS IN PIT SCALES (HOMOPTERA, COCCOIDEA), International journal of insect morphology & embryology, 24(1), 1995, pp. 35-49
TEM/SEM and computerized images of 5 wax glands for 3 type species of
Coccoidea (Homoptera): Asterodiaspis variolosa (Asterolecaniidae), Cer
ococcus quercus (Cerococcidae) and Lecanodiaspis sardoa (Lecanodiaspid
idae) were studied. Their cuticular structures were compared with 142,
56, and 61 species in their respective families to determine relation
ships among pit scale taxa. Significant differences include: the morph
ology of the outer and inner ductule of the tubular duct gland, struct
ure of the pores (8-shaped, multilocular and quinquelocular), and the
absence or presence of cribriform plates and their structural variatio
ns. Three distinctive tubular duct shapes (asterform, ceroform and lec
anoform) are common in pit scale species. Apomorphic characteristics o
f 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 asterolecanii
ds from the cerococcid-lecanodiaspidid lineage. The constricted lecano
form tubular ducts and the curved teeth on the rim at the inner end of
the outer ductile in the ceroform tubular ducts are regarded as autap
omorphic. The presence of 8-shaped pores is considered a plesiomorphic
condition. Specific cuticular variations of the 8-shaped pores, chara
cterizing familial taxa, include pores even with the surface in astero
lecaniids, pores with raised walls in cerococcids, and bent pores in l
ecanodiaspidids. The dominant 8-shaped pore patterns in pit scales are
those arranged in a marginal band in lecanodiaspidids, in a swirl-lik
e pore pattern in the cerococcids, and in a marginal row in asteroleca
niids. A divergent evolutionary trend is noted for the structure of th
e cribriform plate; they are with micro-orifices in cerococcids, but w
ithout micro-orifices in lecanodiaspidids. The former state is conside
red apomorphic. Cribriform plates arranged in clusters characterize th
e cerococcids, while plates in longitudinal rows characterize the leca
nodiaspidids. These data confirm the concept that the pit scales const
itute a paraphyletic group and the Asterolecaniidae, Cerococcidae and
Lecanodiaspididae are monophyletic.