Behavioral Changes in Japanese Beetle and Masked Chafer Grubs (Coleoptera: Scarabaeidae) After Parasitism by Tiphiid Wasps (Hymenoptera: Tiphiidae)

We studied effects of parasitism by two Tiphia species (Hymenoptera: Tiphiidae) on burrowing behavior of their respective scarabaeid hosts and tested the hypothesis that parasitoid alteration of host hormones accounts for the observed behavioral change. In turf Þeld-plots, thirdinstar masked chafer grubs, Cyclocephala sp., parasitized by native Tiphia pygidialis Allen burrowed to depths of 12Ð16 cm within 21 d, whereas nonparasitized grubs remained in the upper 4 cm of soil. Grubs bearing an egg or early-instar Tiphia larva were found relatively close to the surface, whereas those with a fourthor Þfth-instar parasitoid were deeper. Experiments in soil-Þlled, vertical observation chambers conÞrmed downward movement by parasitized Cyclocephala, and similar response in third-instar Japanese beetle, Popillia japonica Newman, parasitized by Tiphia vernalis Rohwer. Removal of the Tiphia egg frommasked chafers was followed by initial downwardmovement of grubs and then a gradual ascent, suggesting that host behavior is affected both by the venomous sting and feeding by thewasp larva. SigniÞcant but inconsistent differenceswere found in juvenile hormone-III (JH III) titers inhemolymph taken fromnonparasitizedgrubs or grubsparasitized for 1 or 14d.Topical application of JH III and methoprene acid resulted in grubs moving deeper into the soil, whereas injection of 20-hydroxyecdysone did not affect grub behavior. Failure to dig deeply enough in soil when sampling for parasitized scarab grubs or Tiphia cocoons likely will result in underestimation of parasitism rates.