A bodyguard or a tastier meal?: dying caterpillar indirectly protects parasitoid cocoons by offering alternate prey to a generalist predator

In some parasitic Hymenoptera the dying caterpillars remain attached or close to the parasitoid cocoons. It has been suggested that the caterpillars act as bodyguards' for the vulnerable cocoons and therefore protect them against predators and/or hyperparasitoids (the usurpation hypothesis'). This hypothesis has been demonstrated in associations where the caterpillars remain active and/or aggressive after parasitism. However, in other associations the caterpillars are so physiologically depleted after parasitism that they are unable to physically defend the cocoons and instead sit atop them in a moribund state. In this study a generalist predator, the spined soldier bug, Podisus maculiventris Say (Hemiptera: Pentatomidae), was provided with cocoons of the gregarious endoparasitoid Cotesia glomerata L. and the solitary endoparasitoid Microplitis mediator Haliday (both Hymenoptera: Braconidae), in turn attended by their hosts, Pieris brassicae L. (Lepidoptera: Pieridae) and Mamestra brassicae L. (Lepidoptera: Noctuidae), respectively. Cotesia glomerata produces broods of up to 40 cocoons and the dying caterpillars sit atop the cocoons where they exhibit little response to physical stimuli. Previous studies reported that dying P.brassicae caterpillars were ineffective bodyguards against two species of hyperparasitoids. In both associations, the dying host caterpillars were significantly preferred as food by P.maculiventris over the parasitoid cocoons. However, in absence of caterpillars, the bugs readily attacked the C.glomerata cocoons. Alternatively, the survival of M.mediator was very low, irrespective of whether a caterpillar was present or not. Caterpillars attacked by M.mediator are several times smaller than those attacked by C.glomerata. Consequently, the predators ran out of food much more quickly in the former and switched from one prey to the other. We show that in some host-parasitoid associations the dying caterpillars provide more visually apparent or nutritionally superior prey, rather than acting as bodyguards