The role of pre- and post-alighting detection mechanisms in the responses to patch size by specialist herbivores

Experimental data on the relationship between plant patch size and population density of herbivores within fields often deviates from predictions of the theory of island biogeography and the resource concentration hypothesis. Here we argue that basic features of foraging behaviour can explain different responses of specialist herbivores to habitat heterogeneity. In a combination of field and simulation studies, we applied basic knowledge on the foraging strategies of three specialist herbivores: the cabbage aphid (Brevicoryne brassicae), the cabbage butterfly (Pieris rapae L.) and the diamondback moth (Plutella xylostella L.), to explain differences in their responses to small scale fragmentation of their habitat. In our field study, populations of the three species responded to different sizes of host plant patches (9 plants and 100 plants) in different ways. Densities of winged cabbage aphids were independent of patch size. Egg-densities of the cabbage butterfly were higher in small than in large patches. Densities of diamondback moth adults were higher in large patches than in small patches. When patches in a background of barley were compared with those in grass, densities of the cabbage aphid and the diamondback moth were reduced, but not cabbage butterfly densities. To explore the role of foraging behaviour of herbivores on their response to patch size, a spatially explicit individual-based simulation framework was used. The sensory abilities of the insects to detect and respond to contact, olfactory or visual cues were varied. Species with a post-alighting host recognition behaviour (cabbage aphid) could only use contact cues from host plants encountered after landing. In contrast, species capable with a pre-alighting recognition behaviour, based on visual (cabbage butterfly) or olfactory (diamondback moth) cues, were able to recognise a preferred host plant whilst in flight. These three searching modalities were studied by varying the in flight detection abilities, the displacement speed and the arrestment response to host plants by individuals. Simulated patch size - density relationships were similar to those observed in the field. The importance of pre- and post- alighting detection in the responses of herbivores to spatial heterogeneity of the habitat is discussed.