Imperfect eggs and oviform nymphs: a history of ideas about the origins of insect metamorphosis

The problem of insect metamorphosis has inspired naturalists for centuries. One question that often arises is why some insects, such as butterflies and bees, undergo a fairly radical metamorphosis while others, such as crickets and lice, do not. Even before the concept of homology emerged scientists speculated which stage found in more direct-developing insects would correspond with the pupal stage of metamorphosing insects. William Harvey (1651) considered the pupal stage to be a continuation of embryonic events, calling it a "second egg." Since then variations of this idea have emerged over the centuries of scientific research and have been supported by a wide variety of methods and rationales. This review will follow those ideas and the ideas that emerged in opposition to them to the present state of the field.