FeminizingWolbachiainZyginidia pullula(Insecta, Hemiptera), a leafhopper with an XX/X0 sex-determination system

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Abstract
Zyginidia pullulais a grass-dwelling leafhopper characterized by a bisexual reproduction mode. In this species, some females collected in Northern Italy, when mated with males, gave origin to an exclusively female brood. Here, we demonstrated that in these lineages an infection by a new strain ofWolbachia pipientis—designated aswZygpul—was detected by amplifying and sequencing thewspand 16S rRNA genes. About half of the female progeny were characterized by intersexual phenotypes, i.e. showing upper pygofer appendages, a typical male secondary sexual feature. The karyological analysis proved that while phenotypically normal females had a female genotype, those with upper pygofer appendages had a male genotype and were thus feminized males.The complete removal ofW. pipientisafter tetracycline treatment of morphologically normal females, and the consequent re-appearance of males in the brood, permitted us to connect the feminizing effect with the presence of the bacterium. This is the first case of feminization byW. pipientisin an XX/X0 sex-determination system, and is the second case reported in insects.