  Large-scale structural patterns commonly occur in network models of complex
systems including a skewed node degree distribution and small-world topology.
These patterns suggest common organizational constraints and similar functional
consequences. Here, we investigate a structural pattern termed pathway
proliferation. Previous research enumerating pathways that link species
determined that as pathway length increases, the number of pathways tends to
increase without bound. We hypothesize that this pathway proliferation
influences the flow of energy, matter, and information in ecosystems. In this
paper, we clarify the pathway proliferation concept, introduce a measure of the
node--node proliferation rate, describe factors influencing the rate, and
characterize it in 17 large empirical food-webs. During this investigation, we
uncovered a modular organization within these systems. Over half of the
food-webs were composed of one or more subgroups that were strongly connected
internally, but weakly connected to the rest of the system. Further, these
modules had distinct proliferation rates. We conclude that pathway
proliferation in ecological networks reveals subgroups of species that will be
functionally integrated through cyclic indirect effects.
