A quantitative exploration of the meso-scale structure of ecological networks

Analysing ecological communities as complex networks of interactions has become an important tool for ecologists. Understanding how these networks change through time, over landscapes, or in response to disturbances is a primary goal of community ecology. The number of interactions and the way in which those interactions organise themselves as individuals, small groups, and the whole community can play an important role in predicting how ecological communities will respond to disturbances. In this thesis, we investigated variation in network structure at several scales both empirically and in a theoretical context. Our first hypothesis was that the structural role of species in a variable system would show little variation, despite high levels of species turnover and a fragmented landscape. In a collaboration with Riikkaa Kaartinen and Tomas Roslin, we studied the distribution of species’ roles at three scales in host-parasitoid networks collected from a fragmented forest in Finland. We found that species’ roles were remarkably consistent through time and in the presence of species turnover. These results suggest that species’ roles may be an intrinsic property of species and may be predictable over spatial and temporal scales. Our second study investigated the structural variation of simulated ecological networks and the relationship between structural variation and whole-network measures of network organization, such as connectance, nestedness, and modularity. We quantified structural variation of networks at three scales, macro-scale, motif-scale, and participation scale. These scales represent whole-network measures (macro-scale), sub-network measures (motifs – small groups of interacting species), and individual measures (motif participation). We compared the variation in these structures to connectance, nestedness, and modularity. We found that at fixed levels of connectance, nestedness, and modularity, the motif profiles of networks and the distribution of species across those profiles showed remarkable dissimilarity. This result suggests that networks displaying similar macro-scale structural measures can be composed of vastly different motif- and participation-scale structures. Together, the work that makes up this thesis suggests that we should give more attention to the meso-scale structures of ecological networks. As the more detailed perspective of motifs can capture additional detail about the structure of empirical networks, and as a result, provide a clearer picture of ecological communities. In addition, we found that the particular species themselves can have a significant impact on the meso-scale structure and, in some cases, may impose strict limitations on what interactions can occur within a community. This has important implications for our understanding of how ecological networks are built and maintained, and thereby for our understanding of the stability and resilience of ecological communities