Why are there no C₄ forests?

C₄ photosynthesis is absent from the arborescent life form, with the exception of seven Hawaiian Euphorbia species and a few desert shrubs that become arborescent with age. As a consequence, wherever C₃ trees can establish, their height advantage enables them to outcompete low stature C₄ vegetation. Had C₄ photosynthesis been able to evolve in an arborescent life form, forest cover (by C₄ trees) could have been much more extensive than today, with significant consequences for the biosphere. Here, we address why there are so few C₄ trees. Physiological explanations associated with low light performance of C₄ photosynthesis are not supported, because C₄ shade-tolerant species exhibit similar performance as shade-tolerant C₃ species in terms of quantum yield, steady-state photosynthetic and use of sunflecks. Hence, hypothetical C₄ trees could occur in the regeneration niche of forests. Constraints associated with the evolutionary history of the C₄ lineages are more plausible. Most C₄ species are grasses and sedges, which lack meristems needed for arborescence, while most C₄ eudicots are highly specialized for harsh (arid, saline, hot) or disturbed habitats where arborescence may be maladapted. Most C₄ eudicot clades are also young, and have not had sufficient time to radiate beyond the extreme environments where C₄ evolution is favored. In the case of the Hawaiian Euphorbia species, they belong to one of the oldest and most diverse C₄ lineages, which primed this group to evolve arborescence in a low-competition environment that appeared on the remote Hawaiian Islands.