Relationships among genera in Saniculoideae and selected Apioideae (Umbelliferae) inferred from nrITS sequences

The internal transcribed spacers (ITS1 and ITS2) of 18S­26S nuclear ribosomal DNA were newly sequenced for eight species of Umbelliferae (six species from subfamily Saniculoideae: Actinolema macrolema, Astrantia minor, Eryngium giganteum, E. coeruleum, Hacquetia epipactis , and Lagoecia cuminoides , two species from subfamily Hydrocotyloideae: Dickinsia hydrocotyloides and Azorella trifurcata ), as well as Hohenackeria exscapa , a species of uncertain position in the family. Phylogenetic analyses of new data, plus previously reported sequences of 52 other species using neighbor­joining, maximum parsimony, and maximum likelihood methods yielded similar results: (1) Actinolema is sister to Astrantia corresponding to Drude's treatments; (2) in Astrantia, molecular divergence is revealed between sects. Astrantiella ( A. minor ) and Astrantia (A. major, A. maxima) ; (3) Eryngium appears to be paraphyletic; (4) Hacquetia might be treated as a part of Sanicula ; and (5) Lagoecia is very distant from all other Saniculoideae and close to some genera of Apioideae. Our results correspond to matK data previously published: (1) Hohenackeria forms a clade with Bupleurum , in a position near the base of the Apioideae tree; (2) Azorella is sister to a large cluster uniting all Saniculoideae and Apioideae, being slightly closer to them than to the Hydrocotyle­Araliaceae clade; (3) Dickinsia is very distant from phenetically similar Hydrocotyle , falling within a large cluster of Apioideae, but also including Lagoecia and Naufraga .