Abstract Parasitic organisms are of interest in evolutionary biology, often displaying drastic modifications in morphology, physiology, genomes, and ecology. These properties, however, make them challenging from a systematics perspective. Mycoheterotrophy, in which plants become non-photosynthetic parasites on fungi, is an excellent example, and this unique life history has evolved numerous times in the orchid family. Here we focused on Stereosandra, a genus of mycoheterotrophic orchid comprising a single species, S. javanica, about which little is known. Stereosandra has been placed in the orchid tribe Nervilieae, along with the leafy, autotrophic Nervilia, and the leafless, mycoheterotrophic Epipogium. We characterized the first complete plastid genome for Stereosandra and used nuclear sequence capture to determine its relationships within Nervilieae. This study presents the first genetic data ever produced for Stereosandra. The plastid genome exhibits rampant gene losses, pseudogenes, and reduced size relative to Nervilia, but not to the extent seen in Epipogium. There is evidence of relaxed negative selection in six genes in Stereosandra, including matK, which functions in Group IIA intron removal of seven plastid genes, four of which have been lost or pseudogenized in this species. Applying mixture models, plastid genomes provided weak support for a sister position of Stereosandra to a clade of Epipogium + Nervilia. Nuclear phylogenomic analyses provided strong support for the same relationships. Ancestral state reconstruction revealed clear evidence that mycoheterotrophy evolved multiple times in the tribe from leafy ancestors. This study provides a previously unidentified, convergent instance of the evolution of full mycoheterotrophy in plants. We discuss the results in the context of proposed models of reductive plastid genome evolution and the genomic and evolutionary consequences of radical life history shifts in heterotrophic plants.
Barrett et al. (Wed,) studied this question.