Interactions between all pairs of neighboring trees in 16 forests worldwide reveal details of unique ecological processes in each forest, and provide windows into their evolutionary histories

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands’ physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees’ distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions.
Author summary Worldwide, ecosystems are collapsing or in danger of collapse, but the precise causes of these collapses are often unknown. Observational and experimental evidence shows that all ecosystems are characterized by strong interactions between and among species, and that these webs of interactions can be important contributors to the preservation of ecosystem diversity. But many of the interactions–such as those involving pathogenic microorganisms and the chemical defenses that are mounted by their prey–are not easily identified and analyzed in ecosystems that may have hundreds or thousands of species. Here we use our equal-area-annulus analytical method to examine census data from over three million trees in forest plots from around the world. We show how the method can be used to flag pairs and groups of species that exhibit unusual levels of interaction and that are likely on further investigation to yield information about their causative mechanisms. We give a detailed example showing how some of these interactions can be traced to defense mechanisms that are possessed by one of the tree species. We explore how our method can be used to identify the between-species interactions that play the largest roles in the maintenance of ecosystems and their diversity.