Theoretical models predict that brood guarding may evolve in situations where eggs are costly to produce or when handling times are long. This study reveals that females of the secondary hyperparasitoid Trichomalopsis apanteloctena guarded cocoon broods of Cotesia kariyai, a gregarious endoparasitoid. Hyperparasitoid females also monopolized host resources and protected their offspring by driving away other conspecific hyperparasitoid females. The females exhibited antagonistic behavior towards competitors through threatening body postures, biting and chasing. Using a video camera to determine how long a hyperparasitoid female attended and parasitized cocoons within a single host brood, it was found that after about 4 days, cocoon guarding behavior became much less apparent. Moreover, more than 90% of hosts were typically parasitized by a hyperparasitoid female over the course of 4 days after she commenced brood guarding. Observations of egg production during a female's lifetime revealed a physiological interval rhythm that typically lasted 3-4 days, which correlates almost exactly with the period during which the cocoons were guarded. To confirm the giving-up time for a host cocoon brood, hyperparasitoid females were given access to 24 h-old cocoon clusters, each containing 60-100 individual cocoons. Ninety percent of the females remained on cocoons for approximately 72 h. Furthermore, twenty-five percent of wasps continued attending and presumably guarding host cocoon broods for more than 138 h after the female first attended the brood. C. kariyai larvae pupate within a few hours of egression from their host and emerge as adults about 5 days (120 h) later. Therefore, many hyperparasitoid females continued to guard older host cocoons of greatly reduced quality as a resource for their progeny and some even after eclosion of the primary parasitoid. Late-brood guarding enabled a hyperparasitoid female to protect her own progeny from other hyperparasitoid females that readily attacked and killed them when she was removed. Our study thus reveals that extended guarding behavior is an adaptive mechanism that probably plays an important role in the survival of the original brood.
At maturity, the endoparasitoid larvae of several subfamilies of the Braconidae have to emerge from inside of the host to pupate. Although the hosts hormonal milieu and the timing of larval parasitoid emergence have been studied, no report has yet focused on the physiological state of the host in connection with the emergence behavior of endoparasitoids. We investigated the mechanism of larval emergence behavior in a gregarious endoparasitoid, Cotesia kariyai. The parasitoid larvae inserted their mandibles into the host cuticle and perforated the integument by moving their head-capsule backwards and forwards. The emerging parasitoid larva must have a physical support (an "anchor") with the terminal appendages in order to exert the necessary pressure to cut the host integument. Morphological observations revealed that each parasitoid larva was enveloped in a capsule just before emerging from their host. Eight and nine day-old parasitoid larvae secreted material around their bodies to form these capsules. This material consisted of acid-glycoproteins which coated the exuvium of the 2nd instar larvae. The haemolymph volume of the parasitised host also decreased in later stages and was dramatically reduced immediatly prior to parasitoid emergence. This final reduction of the host haemolymph volume is the result of absorption by parasitoid larvae. This mechanism allows the parasitoid larvae to create an anchor more easily. The parasitoid larvae could also adhere to each other with the glycoprotein. In addition, these capsules prevent the leaking of host haemolymph through the emergence hole; these holes on the host integument were plugged by the capsules after parasitoid emergence. Although the pressure acquired by the anchor was lost once the head of the parasitoid larvae emerges from the host integument, the parasitoid larvae crawls out of the host cavity using backward pointing spines which enable the parasitoid to grip the capsule and move forward via peristaltic contractions.