Plant species, such as Steinchisma hians, that exhibit intermediacy between the C3 and C4 photosynthetic types offer the opportunity to study the evolution of adaptive complexes. We conducted a molecular phylogenetic study of this and related species with two markers: 1) the grass-specific insert sequence in the chloroplast RNA polymerase locus (rpoC2); and 2) the externally transcribed spacer region in the nuclear ribosomal repeat (ETS). To our knowledge this is the first report of the use of ETS in grass phylogenetics. Separate analyses indicated no substantial incongruence between these two loci so data combination was performed. Combined phylogenetic analysis resolved exclusive clades of two photosynthetic subtypes. A reversion of C4 to C3 photosynthesis during the evolution of the lineage that includes S. hians was identified. This and other recent results suggest that C4 to C3 reversions are more common than previously supposed. The C3 species, Panicum laxum, had a strongly supported sister group relationship to S. hians. The most parsimonious interpretation is that S. hians represents an incipient reversal from C3 back to C4 photosynthesis beginning with the capacity to compartmentalize photorespiratory metabolism in the bundle sheath tissue. Our results further confirmed the multiple origins of C4 photosynthesis within panic grasses.

Key words: C3 - C4 intermediacy, ETS, external transcribed spacer, Paniceae, photosynthesis, rpoC2, Steinchisma hians