Herbivory in Myrtillocactus geometrizans (Cactaceae): Do Parasitoids Provide Indirect Defense or a Direct Advantage?

Plants respond to herbivory in diverse, complex ways, ranging from avoidance or tolerance to indirect defense mechanisms such as attracting natural enemies of herbivores, i.e., parasitoids or predators, to strengthen their defense. Defense provided by parasitoids to cultivated plants is well documented and is used in biological control programs. However, its effectiveness on wild plants under natural conditions has been little studied. Such is the case of the cactus Myrtilllocactus geometrizans (known in Mexico as garambullo), which is consumed by the soft-scale insect Toumeyella martinezae (herbivore) which, in turn, is host to the parasitoid wasp Mexidalgus toumeyellus, and mutualist with the ant Liometopum apiculatum, that tenders and protects it. This study explores the role of the parasitoid as an indirect defense, by examining its effect on both the herbivore and the plant, and how this interaction is affected by the presence of the mutualistic ant. We found that scales adversely affect the cactus' growth, flower, and fruit production, as well as its progeny's performance, as seedlings from scale-infested garambullo plants were shorter, and it also favors the presence of fungus (sooty mold). The parasitoid responded positively to herbivore abundance, but the presence of ants reduced the intensity of parasitism. Our results show that parasitoids can function as an indirect defense, but their effectiveness is reduced by the presence of the herbivore's mutualistic ant.