  Networks have become a key approach to understanding systems of interacting
objects, unifying the study of diverse phenomena including biological organisms
and human society. One crucial step when studying the structure and dynamics of
networks is to identify communities: groups of related nodes that correspond to
functional subunits such as protein complexes or social spheres. Communities in
networks often overlap such that nodes simultaneously belong to several groups.
Meanwhile, many networks are known to possess hierarchical organization, where
communities are recursively grouped into a hierarchical structure. However, the
fact that many real networks have communities with pervasive overlap, where
each and every node belongs to more than one group, has the consequence that a
global hierarchy of nodes cannot capture the relationships between overlapping
groups. Here we reinvent communities as groups of links rather than nodes and
show that this unorthodox approach successfully reconciles the antagonistic
organizing principles of overlapping communities and hierarchy. In contrast to
the existing literature, which has entirely focused on grouping nodes, link
communities naturally incorporate overlap while revealing hierarchical
organization. We find relevant link communities in many networks, including
major biological networks such as protein-protein interaction and metabolic
networks, and show that a large social network contains hierarchically
organized community structures spanning inner-city to regional scales while
maintaining pervasive overlap. Our results imply that link communities are
fundamental building blocks that reveal overlap and hierarchical organization
in networks to be two aspects of the same phenomenon.
