Direct and Indirect Adverse Effects of Tomato on the Predatory Mite Neoseiulus californicus Feeding on the Spider Mite Tetranychus evansi

Plants may defend themselves against herbivores via morphological traits, chemical traits, or a combination of both. Herbivores that overcome the defensive echanisms of a plant tend to specialize on this plant due to enhanced protection from natural enemies. Well-known examples of plants possessing a suite of defensive mechanisms are found in nightshades (Solanaceae), especially in the tomato genus Lycopersicon . The spider mite Tetranychus evansi Baker and Pritchard (Acari: Tetranychidae) is specialized on solanaceous plants and is an invasive pest of tomato in Europe and Africa. Biological control of T. evansi with currently available natural enemies, such as the predatory mites Phytoseiulus persimilis Athias-Henriot and Neoseiulus californicus McGregor (both Acari: Phytoseiidae), is unsuccessful, with the underlying mechanisms only vaguely known. We hypothesized that T. evansi is a key pest of tomato because this host plant provides a two-pronged protection from natural enemies. Direct adverse effects of tomato on predators may arise from morphological traits and/or trichome exudates, whereas indirect effects are prey-mediated through the accumulation of toxic plant compounds. Using a 2 × 3 factorial design, we assessed and separated direct and indirect effects of tomato on the life history of N. californicus feeding on two strains of T. evansi (reared on bean or tomato) on three substrates (tomato leaf, bean leaf, and an artificial cage). Developmental time and oviposition rate of N. californicus were both directly and indirectly negatively affected by tomato whereas offspring sex ratio and survival of juveniles and adult females were unaffected. The direct and indirect, preymediated adverse effects of tomato on N. californicus with T. evansi prey had similar magnitudes and were additive. We conclude that T. evansi per se is a suitable prey species for N. californicus and discuss the results with respect to the potential use of N. californicus as biological control agent of T. evansi on tomato and other host plants.