Spiracles of Adult Scarabaeoidea (Coleoptera) and Their Phylogenetic Significance. I. the Abdominal Spiracles1

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Abstract
A comparative study was made of the number, location, and morphology of abdominal spiracles of adult Passalidae, Lucanidae, and Scarabaeidae, using material of more than 150 genera. Major evolutionary trends are a reduction from 8 to 7 or fewer pairs of functional spiracles, a reduction in size of the posterior spiracles or of their openings, migration of 1 or more posterior pairs of abdominal spiracles into the tergites or sternites, and changes in the filter apparatus from simple atrial spines to a solid wall with imbedded trabeculae surrounding a slitlike subatrial opening. These characters are useful for assessing relationships between genera, tribes, and subfamilies, but division of the Scarabaeoidea into 2 great series, the Laparosticti and Pleurosticti, should be abandoned, since repeated evolutionary changes in location and size of the spiracles and in their number and morphology as well have occurred in different phyletic lines.

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