Geographic variation and evolution in the life cycle of the witch-hazel leaf gall aphid, Hormaphis hamamelidis

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Abstract
Two divergent life cycles associated with different elevations and latitudes have been documented for the witch-hazel leaf gall aphid, Hormaphis hamamelidis. At low elevation in northern Virginia, the aphid had seven distinct generations alternating between the primary host, witchhazel (Hamamelis virginiana), and a secondary host, river birch (Betula nigra). These findings confirm the original published life cycle description for the same locality. A second, abbreviated life cycle consisting of only three generations restricted to witch-hazel was discovered at high elevation (1000 m) in north central and northwestern Virginia. Aphids of both life cycles were sympatric at a middle elevation site. The life cycles and morphology suggest that the two forms are separate species. Although monoecious life cycles on primary hosts in aphids generally are thought to be ancestral to complex host-alternating ones, it is certainly possible that monoecious cycles are sometimes secondarily derived from complex cycles. By constructing a preliminary phylogeny of the described species in the tribe Hormaphidini, we propose that the abbreviated life cycle is derived from the complex one in the case of these witchhazel gall aphids. Our findings are discussed in the context of current theory regarding the evolutionary stability of complex life cycles.

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