Abstract
Identifying and explaining the structure of complex networks at different scales has become an important problem across disciplines. At the mesoscale, modular architecture has attracted most of the attention. At the macroscale, other arrangements-e.g. nestedness or core-periphery-have been studied in parallel, but to a much lesser extent. However, empirical evidence increasingly suggests that characterizing a network with a unique pattern typology may be too simplistic, since a system can integrate properties from distinct organizations at different scales. Here, we explore the relationship between some of these different organizational patterns: two at the mesoscale (modularity and in-block nestedness); and one at the macroscale (nestedness). We show experimentally and analytically that nestedness imposes bounds to modularity, with exact analytical results in idealized scenarios. Specifically, we show that nestedness and modularity are interdependent. Furthermore, we analytically evidence that in-block nestedness provides a natural combination between nested and modular networks, taking structural properties of both. Far from a mere theoretical exercise, understanding the boundaries that discriminate each architecture is fundamental, to the extent that modularity and nestedness are known to place heavy dynamical effects on processes, such as species abundances and stability in ecology.