EVOLUTIONARY REDUCTION OF COMPLEX LIFE CYCLES: LOSS OF HOST-ALTERNATION IN PEMPHIGUS (HOMOPTERA: APHIDIDAE).

In a Utah canyon, the aphid, Pemphigus betae, exhibits two life cycles: a cycle involving host-alternation between cottonwood trees and roots of herbaceous plants and a secondarily reduced cycle, in which the cottonwood generations are eliminated so that wingless forms live year round on roots. Relative frequencies of the two life-cycle types vary along a 30-km stretch of the canyon, with the reduced cycle predominating at upper sites. Factors underlying this life-cycle variation were examined with common-garden and transfer experiments. Results showed 1) a facultative increase in production of alternating forms in response to crowding in root colonies, 2) a genetic component to both within- and between-site variation in tendency to produce alternating morphs, and 3) site-specific environmental effects on level of investment in the reduced versus alternating life cycles. Thus, the variation in frequency of life-cycle reduction in this aphid is dependent on a complex of interdependent factors. These include adaptive phenotypic plasticity, microgeographically variable cues affecting mechanisms of morph determination, and genetically based variation in tendency to show reduction versus alternation. Genetic variation between sites corresponds to microgeographic variation in success of life-cycle phases. Where cottonwood hosts are absent (lower elevations) or where the cottonwood phase has low survival (upper elevations), clones tend to produce fewer migrating morphs, as compared to clones from middle elevations, where the cottonwood phase is relatively favorable. Such independence between conditions in alternate phases is a general feature of complex life cycles and can generate strong site-specific selection for permanent life-cycle reduction. Such life-cycle shifts have sometimes been followed by extensive radiations in aphids and other groups.