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South American subtropical landscapes are dominated by open grasslands and mosaics of forest–grassland formations. Forests are often restricted to riverine margins with sharp forest–grassland ecotones. Understanding the mechanisms maintaining forest–grassland ecotones is important to anticipate the effects of changing climate and disturbance regimes on the extent of these biomes and the ecosystem services they provide.We used a combination of field surveys and long‐term field experiments to explore the mechanisms that explain tree cover expansion at the ecotone of riverine forests and grasslands in central Uruguay, within the South American Campos. We assessed the role of tree seed dispersal and seedling establishment limitations, and experimentally tested for the effects of cattle, nurse tree cover and grasses on the recruitment of forest and grassland tree species at the forest–grassland ecotone.We found that forest expansion depends on the interplay between cattle and nurse trees. Vachellia caven trees colonize the grassland successfully and facilitate the formation of forest patches by enhancing seed accumulation and seedling establishment of forest tree species. Surprisingly, grass cover had mostly positive effects on early seedling survival of forest tree seedlings. However, cattle limits tree seedling growth and survival, especially of forest tree species. This results in a nucleated vegetation pattern of tree patches that ultimately limits forest expansion.Synthesis and applications. Tree cover can potentially expand on the subtropical South American grasslands. Reductions in cattle densities and increases in rainfall levels associated with climate change could facilitate forest expansion in this region. [ABSTRACT FROM AUTHOR]