Non‐trophic effects of litter reduce ant predation and determine caterpillar survival and distribution

The tritrophic model featuring plants consumed by herbivores consumed by parasitoids or predators has become the primary paradigm used to describe herbivore dynamics. However, interactions involving herbivores can be habitat‐ specific and plants often provide habitat, as well as food. Structural complexity of the habitat may favor predators or may allow herbivore prey to escape detection and capture. This study considered the spatial and temporal dynamics of an arctiid caterpillar, Platyprepia virginalis. The tritrophic model that includes only a tachinid parasitoid that attacks P. virginalis and the caterpillars’ primary host‐plant, Lupinus arboreus, has failed to provide much insight into this system. Instead, we found that ants killed and removed many small caterpillars. Protecting caterpillars from ants increased their survival three‐fold and five‐fold in assays conducted during two years. Caterpillars were more likely to survive in short‐term assays at sites that naturally had a deeper cover of dead and living plant material. Experiments with baits showed that ant recruitment declined as litter depth increased on average. These survey results indicated that ant predation was an important source of mortality for young caterpillars and that the presence of thick litter reduced this mortality. These results were corroborated in an experiment that manipulated litter depth and ant access to caterpillars. Previous findings that other defoliating caterpillars increased litter depth and benefitted P. virginalis are also consistent with this hypothesis. Litter acts as an important non‐trophic resource, allowing caterpillars to avoid predation by ants such that wet sites with deep litter act as source populations for caterpillars. Our results show strong effects of both trophic and non‐trophic interactions since plants indirectly provided limiting habitat and this heterogeneous habitat strongly affected risk of predation and ultimately caterpillar abundance and distribution.