AbstractIn the stomatal lineage, cells make fate transitions from asymmetrically dividing and self-renewing meristemoids, to commitment to the guard mother cell identity, and finally though a single division to create mature, post-mitotic stomatal guard cells. Flexibility in the stomatal lineage allows plants to alter leaf size and stomatal density in response to environmental conditions; however, transitions must be clean and unidirectional in order to produce functional and correctly patterned stomata. Among direct transcriptional targets of the stomatal initiating factor, SPEECHLESS, we found a pair of genes, SOL1 and SOL2, required for effective transitions in the lineage. Here we show that these two genes, which are homologues of the LIN54 DNA-binding components of the mammalian DREAM complex, are expressed in a cell cycle dependent manner and regulate cell fate and division properties in the self-renewing early lineage. In the terminal division of the stomatal lineage, however, these two proteins appear to act in opposition to their closest paralogue, TSO1, revealing complexity in the gene family may enable customization of cell divisions in coordination with development.