Ecology and Evolution of mutualisms
I am broadly interested in the ecology and evolution of species interactions. Much of my research focuses on understanding how cooperation between species can persist. Mutualisms are present nearly everywhere. For example, many plants need pollinators to reproduce, and many animals (including humans) need gut bacteria to take up nutrients. But why would individuals from different species provide costly benefits to each other? What prevents partners from turning into cheaters that reap the benefits of the interaction without paying the costs? My research explores these, and other questions related to the ecology and evolution of mutualisms. I am also interested in behavioural ecology. Some of the projects I work on are described in more detail below.
Sanctions and cheating in the fig tree – fig wasp mutualism
My main study system is the mutualism between fig trees and their pollinating fig wasps. This is a very attractive system because cooperative behaviour can be manipulated, and lifetime reproductive success can be quantified in the field. Further, each wasp species is typically specific to a particular host species, and there are over 700 species of figs across the tropics, enabling comparative studies.
Fig wasps develop inside fig fruits, and transfer pollen from the fruit in which they are born to a different, flowering, fig tree. Most fig wasp species actively collect pollen with their front legs, carry it in special pollen pockets on their thorax, and then again use their front legs to actively deposit the pollen onto fig flowers. Some wasp individuals fail to carry pollen, and can therefore not perform the pollination service for the tree. The vast majority of wasps, however, do pick up and deposit pollen. Why do they do this?
Through field experiments we have demonstrated that fig trees (probably by allocating more resources to better-pollinated figs) reduce the fitness of wasps that do not pollinate: unpollinated figs are more likely to abort, and in unpollinated figs that mature, wasp larvae have a higher mortality. These host sanctions vary in strength across fig species. The proportion of fig wasps that are uncooperative also vary across species: pollen-free wasps are least common in the species with strong sanctions, where costs of not cooperating are high. In fig species with weaker sanctions, pollen-free wasps are more common. Because sanctions act on the level of the entire fig rather than on individual flowers within the fig, uncooperative wasps that do not pollinate can free-ride on the efforts of pollinating wasps.
Actors external to the mutualism
Ants frequently make their nests in fig trees. Although ants sometimes prey upon the pollinator wasps, they have easier access to the herbivores and parasitic wasps that lay eggs from the outside of the fig. In fact, having ants present on the tree increases the fitness of the tree and the pollinators dramatically. Ants therefore function as indirect mutualists to the fig tree – fig wasp mutualism.
Conflict within species: Fatal fighting among female fig wasps
Female fig wasps lay their eggs in the flowers that line the inside of the fig. Each wasp larva consumes one flower. This can create a competitive situation among the egg-laying females over limited egg-laying sites. Female wasps aggressively fight, sometimes maiming or killing each other.