- Ecology, evolution, and biogeography of plant-animal interactions, especially mutualism.
- Associations between mites and leaf domatia.
- Seed dispersal mutualisms in the genus Acacia.
- Tri-level trophic interactions, especially the effects of plant morphology on the third trophic level.
- The antiquity of protective mutualism between plants and arthropods.
- The ecology and control of environmental weed.
- Biodiversity education.
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