Head capsule stacking by caterpillars: morphology complements behaviour to provide a novel defence.

Herbivores employ a variety of chemical, behavioural and morphological defences to reduce mortality from natural enemies. In some caterpillars the head capsules of successive instars are retained and stacked on top of each other and it has been suggested that this could serve as a defence against natural enemies. We tested this hypothesis by comparing the survival of groups of the gumleaf skeletoniser Uraba lugens Walker caterpillars, allocated to one of three treatments: "-HC," where stacked head capsules were removed from all individuals, "+HC," where the caterpillars retained their stacked head capsules, and "mixed," where only half of the caterpillars in a group had their stacked head capsules removed. We found no difference in predation rate between the three treatments, but within the mixed treatment, caterpillars with head capsules were more than twice as likely to survive. During predator choice trials, conducted to observe how head capsule stacking acts as a defence, the predatory pentatomid bug attacked the -HC caterpillar in four out of six trials. The two attacks on +HC caterpillars took over 10 times longer because the bug would poke its rostrum through the head capsule stack, while the caterpillar used its head capsule stack to deflect the bug's rostrum. Our results support the hypothesis that the retention of moulted head capsules by U. lugens provides some protection against their natural enemies and suggest that this is because stacked head capsules can function as a false target for natural enemies as well as a weapon to fend off attackers. This represents the first demonstration of a defensive function.