Using network analysis to characterize forest structure

Network analysis quantifies different structural properties of systems of interrelated parts using a single analytical framework. Many ecological phenomena have network-like properties, such as the trophic relationships of food webs, geographic structure of metapopulations, and species interactions in communities. Therefore, our ability to understand and manage such systems may benefit from the use of network-analysis techniques. But network analysis has not been applied extensively to ecological problems, and its suitability for ecological studies is uncertain. Here, we investigate the ability of network analysis to detect spatial patterns of species association in a tropical forest. We use three common graph-theoretic measures of network structure to quantify the effect of understory tree size on the spatial association of understory species with trees in the canopy: the node degree distribution (NDD), characteristic path length (CPL), and clustering coefficient (CC). We compute the NDD, CPL, and CC for each of seven size classes of understory trees. For significance testing, we compare the observed values to frequency distributions of each statistic computed from randomized data. We find that the ability of network analysis to distinguish observed patterns from those representing randomized data strongly depends on which aspects of structure are investigated. Analysis of NDD finds no significant difference between random and observed networks. However, analysis of CPL and CC detected nonrandom patterns in three and one of the seven size classes, respectively. Network analysis is a very flexible approach that holds promise for ecological studies, but more research is needed to better understand its advantages and limitations.