Abstract
Many unusual features of scale insects (Hemiptera: Coccoidea) can be explained as historical legacy. Developmental specializations in ancestral coccoids resulted in a neotenous adult female and a drastic metamorphosis of the male. Subsequent evolution led to numerous, often convergently derived, adaptations to parasitic life on higher plants. The sedentary lifestyle of female scale insects has favored the evolution of appendage reduction or loss, gross changes in body shape, and protective wax secretions, tests, and other scale covers. Morphological peculiarities of adult males relate to flight or to mating with concealed females. Scale insects have diverse egg-protecting methods, a range of chromosome behaviors (including several methods of sex determination), marked sexual dimorphism [even sometimes in first-instar nymphs (crawlers)], and more rarely sexual dichronism. Crawlers have evolved as the main agents of dispersal. The biotic interactions of scale insects include diverse endosymbioses with microorganisms, sometimes morphological and behavioral adaptations for obligate association with ants, and often highly specific host-plant associations that may lead to demic adaptation or the evolution of complex galls.
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