The evolutionary history of maternal plant-manipulation and larval feeding behaviours in attelabid weevils (Coleoptera; Curculionoidea)

Arms race between leaf rollers and parasitoids: diversification of plant‐manipulation behavior and its consequences

“…This hardness might enable trichogrammatid wasps to evolve various morphological and behavioral adaptations for parasitizing eggs in leaf rolls: in Poropoea, behaviors such as piercing leaf rolls with a long ovipositor or crawling into leaf rolls have been reported (Hirose 1963, Kobayashi andKato 2004b), and Ophioneurus sp. Considering that all attelabine species construct severalfold wrapped leaf rolls and no species lost the rolling behavior (Kobayashi et al 2012), there may be the continued existence of coevolution between attelabines and Poropoea wasps. Ophionerus wasps have been reported as specific egg parasitoids of Rhynchinae (Viggiani 1974) and Poropoea wasps mainly of Attelabinae and partly of Rhynchitinae (Hirose 1963, Subba Rao 1969, Kobayashi and Kato 2004b, both causing heavy mortality on attelabid hosts.…”

“…Plant-cutting behavior is characterized as general maternal manipulation in Attelabidae and is thought to originate in the common ancestor of this group (Kobayashi et al 2012). The most ancestral mode of larval feeding type in Attelabidae is the stem or petiole borer, which was almost parasitoid-free, and then leafblade feeders like leaf miners, which suffered high parasitism rates, and leaf-roll feeders, which tended to show relatively low parasitism rates, evolved from stem/ petiole borers (Morimoto 1964, Kobayashi et al 2012).…”

“…This contrasts with other concealed herbivorous insect groups in which larval or pupal parasitoids dominate, and fewer egg parasitoids are observed (Kato 1994, Mills 1994). In Attelabidae, female adults cut plants before and/or after oviposition in order to prevent plant chemical defenses from inflowing into larval feeding area, likely because of the nutritive or detoxification demand of larvae (Kobayashi et al 2012). However, eggs are a desirable host for parasitoids because they lack immune defenses and have no aggressive behavior against parasitoid attack (Waage and Greathead 1986).…”

Sexual shape dimorphism accelerated by male–male competition, but not prevented by sex‐indiscriminate parental care in dung beetles (Scarabaeidae)

“…To demonstrate this conclusion more clearly, leaf‐rolling beetles in Attelabidae and related Rhynchitidae may be another testable taxon, because various patterns of parental provisioning and male–male competition have been known (Kobayashi et al. ).…”

“…It indicates that even combination of crucial and sex-indiscriminate parental provisioning does not prevent the evolution of sexual shape dimorphism, but it limits SSD in dung beetles. To demonstrate this conclusion more clearly, leaf-rolling beetles in Attelabidae and related Rhynchitidae may be another testable taxon, because various patterns of parental provisioning and male-male competition have been known (Kobayashi et al 2012).…”