The missing stase in spider mites (Acari : Tetranychidae) : when the adult is not the imago

Apart from a calyptostatic prelarva, the ontogeny of spider mites is limited to one six-legged immature followed by three eight-legged instars. The usage consists in naming them: larva, protonymph, deutonymph and adult. In this study, the "missing stase" and different associated hypotheses are explored using diverse approaches: light microscopy, electron microscopy, comparative chaetotaxy, ontogenetic trajectories, and interspecific comparisons. It turns out that the imago (new name for the last stase) is missing in Tetranychus urticae and that its post-embryonic develop- ment may be summarized as follows: prelarva (calyptostase), larva (six-legged stase), protonymph (eight-legged stase),deutonymph (eight-legged stase), tritonymph (with paedogenesis, i.e. precocious development of sexual maturity). The respiratory role of the prelarva is emphasized. The current terminology of post-embryonic development is confusing as it pertains to three approaches: the state or form description, the instar standard and the stase paradigm.