Natural Enemies in a Variable World

Abstract

Natural enemies are ubiquitous in nature. In many communities, natural enemies have a major effect on the diversity of their prey. Their effects are very diverse: they can promote or undermine the ability of their prey to coexist through a variety of mechanisms. As such, an important step in understanding how diversity is maintained will be to understand how different forms of predator behavior affect prey coexistence. In this dissertation, I study how two major types of predators affect plant coexistence in two different communities. First, I study natural enemies in tropical forests, using both theory and empirical work. In tropical forests, most natural enemies are thought have a narrow host range, and be distance-responsive (i.e., mainly harm seeds and seedlings that are near adults of their main host). Previous theoretical work has shown that specialized natural enemies can maintain diversity of their prey, whether or not they are distance-responsive. However, it is unknown whether specialist natural enemies are more or less able to promote prey coexistence if they are distance-responsive. Using theoretical models, I show that distance-responsive predators are less able to maintain diversity. Additionally, I show that habitat partitioning does not interfere with the ability of distance-responsive predators to maintain diversity, even if it causes seedling survival to be highest near conspecific adults. From an empirical aspect, I studied the host range of seed-associated fungi. Soilborne microbes, such as fungi, are thought to play an important role in maintaining diversity in tropical forests. However, the microbial community itself is often treated as a black box, and little is known about which microbes are causing major effects, or how 8 specialized seed-microbe associations are. Here I use experimental inoculations to examine the host range and effect of a guild of seed-associated fungi that are thought to be mainly pathogens. I show that fungal species are differentially able to colonize different seed species, and have species-specific effects on seed germination. I show that in many cases, plant phylogeny, and to a lesser extent fungus phylogeny, are good predictors of colonization. Finally, I study how an optimally foraging granivore can promote (or undermine) coexistence amongst annual plants, using theory. Optimal foraging theory is one of the major theories for how predators behave; despite this, little is known about whether an optimally foraging predator could promote coexistence amongst a diverse community of prey. Previous models have shown than two species can coexist due to optimal foraging, but did not test whether multiple prey can coexist, nor if the effect is altered by environmental variation. Here, I show that if the predators specialize on different prey at different times, the predators can allow multiple prey species to coexist. In this case, environmental variation has little effect on the ability of predators to maintain diversity. If the predators are generalists, they cannot maintain diversity. Additionally, I show that generalist predators will create a negative storage effect, undermining coexistence.