Nutlet micromorphology and character evolution of Lappula species (Boraginaceae) and its systematic implications

Background The macro/micro-morphology of nutlets in 11 species (and 22 accessions) of the Boraginaceae family was investigated using stereomicroscope and scanning electron microscopy to evaluate the taxonomic relevance of the traits. To evaluate the phylogenetic significance of the character evolution, phylogenetic analysis was carried out by comparing available DNA sequence data from GenBank with selected original nutlet data. Results The Rochelieae nutlets' shape varied from ovoid (ovoid, ovoid-triangular, and ovoid-rectangular) to pyramid. Six major patterns were recognized based on the nutlet ultrastructure characters. Rocheliae is characterized by a transition from "without appendage" to "with tubercles and prickles" on the nutlet disk, and also via a shift from "lack of prickles" to "glossy prickles". Conclusions The results show that the nutlet ultrastructure pattern of Rochelieae is systematically informative at the genus level, but not at the species level. Findings demonstrated that glochid is not an ancestral trait but is a synapomorphy and the transition to this trait occurred in the genus Lappula. The close boundary of nutlet microstructures between L. barbata and L. microcarpa has been discussed.