A fungal colony is a syncytium composed of a branched and interconnected network of cells. Chimerism endows colonies with increased virulence and ability to exploit nutritionally complex substrates. Moreover, chimera formation may be a driver for diversification at the species level by allowing lateral gene transfer between strains that are too distantly related to hybridize sexually. However, the processes by which genomic diversity develops and is maintained within a single colony are little understood. In particular, both theory and experiments show that genetically diverse colonies may be unstable and spontaneously segregate into genetically homogenous sectors. By directly measuring patterns of nuclear movement in the model ascomycete fungus Neurospora crassa, we show that genetic diversity is maintained by complex mixing flows of nuclei at all length scales within the hyphal network. Mathematical modeling and experiments in a morphological mutant reveal some of the exquisite hydraulic engineering necessary to create the mixing flows. In addition to illuminating multinucleate and multigenomic lifestyles, the adaptation of a hyphal network for mixing nuclear material provides a previously unexamined organizing principle for understanding morphological diversity in the more-thana-million species of filamentous fungi.heterokaryon | hydrodynamics | biological networks G enetic diversity between individuals is important to the resilience of species (1) and ecosystems (2). However, physical and genetic barriers constrain internal genetic diversity within single organisms: Cell walls limit nuclear movement between cells, whereas separation of germ and somatic cell lines means that diversity created by somatic mutations is not transmitted intergenerationally. However, in syncytial organisms, including filamentous fungi and plasmodial slime molds (3), populations of genetically different and mobile nuclei may share a common cytoplasm ( Fig. 1A and Movie S1). Internal diversity may be acquired by accumulation of mutations as the organism grows or by somatic fusion followed by genetic transfer between individuals. For filamentous fungi, intraorganismic diversity is ubiquitous (4, 5). Shifting nuclear ratios to suit changing or heterogeneous environments enhances growth on complex substrates such as plant cell walls (6) and increases fungal virulence (7). Fusion between different fungal individuals is limited by somatic (heterokaryon) compatibility barriers (8), and most internal genetic diversity results from mutations within a single, initially homokaryotic individual (4). However, somatic compatibility barriers are not absolute (9), and exchange of nuclei between heterospecific individuals is now believed to be a motor for fungal diversification (10-12).A fungal chimera must maintain its genetic richness during growth. Maintenance of richness is challenging because fungal mycelia, which are made up of a network of filamentous cells (hyphae), grow by extension of hyphal tips. A continual tipward flow of vesicles and n...