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.