Abstract
C4 is one of three known photosynthetic processes of
carbon fixation in flowering plants. It evolved independently more than
61 times in multiple angiosperm lineages and consists of a series of
anatomical and biochemical modifications to the ancestral
C3 pathway increasing plant productivity under warm and
light-rich conditions. The C4 lineages of eudicots
belong to seven orders and 15 families, are phylogenetically less
clustered than those of monocots, and entail an enormous structural and
ecological diversity. Eudicot C4 lineages likely evolved
the C4 syndrome along different evolutionary paths.
Therefore, a better understanding of this diversity is key to
understanding the evolution of this complex trait as a whole. Compiling
1,207 recognized C4 eudicots species described in the
literature and presenting trait data among these species, we identify
global centres of species richness and of high phylogenetic diversity.
Furthermore, we discuss climatic preferences in the context of plant
functional traits. We identify two hotspots of C4
eudicot diversity: arid regions of Mexico/Southern United States and
Australia, where several C4 eudicot lineages diversified
independently. Further eudicot C4 hotspots with many
different families and genera represented are in South Africa, West
Africa, Patagonia, Central Asia and the Mediterranean. In general,
C4 eudicots were abundant in deserts and xeric
shrublands, tropical and subtropical grasslands, savannas and
shrublands. We found C4 eudicots to occur in areas with
less annual precipitation than C4 grasses which can be
explained by frequently associated adaptations to drought stress such as
among others succulence and salt tolerance. We conclude that in most
eudicot lineages C4 evolved in ancestrally drought
adapted clades and enabled these to further spread in these habitats and
colonise even drier areas.