Quantitative measure of fitness in tri-trophic interactions and its influence on diet breadth of insect herbivores.

Herbivore-plant interactions should be studied using a tri-trophic approach, but we lack a quantitative measure of the combined effect of top-down and bottom-up forces on herbivore fitness. We propose the combination of the bi-trophic fitness slopes as a tri-trophic fitness measure. We use the relationship between fitness associated with top-down and bottom-up forces and the frequency of host plant use to calculate the top-down and bottom-up fitness slopes, which we then combine to obtain three possible directions of tri-trophic slopes. A positive tri-trophic slope indicates that herbivores have overall greater tri-trophic fitness on the more frequently used hosts. A null tri-trophic fitness slope indicates that herbivores have similar fitness on all host plants. A negative tri-trophic slope indicates that herbivores have generally lower fitness on the more frequently used hosts. We tested the explanation power of our method using data from the literature that tested herbivore host shifts and experimentally using a generalist herbivore with variable diet breadth across populations. We found that in host shifts, herbivores have higher tri-trophic fitness on the novel host, while in generalist populations, herbivores use most frequently the best host available. We present applications in other research areas and consider the limitations of our approach. Our approach is a first step towards a comprehensive model of multiple selective forces acting on the evolution of interactions.