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The tribe Neillieae, a small group of about 18 species in the Rosaceae, comprises three taxonomically difficult genera, Neillia, Physocarpus, and Stephanandra. The tribe, characterized by lobed leaves with persistent or deciduous stipules and ovoid, shiny seeds with copious endosperm, is strongly supported as a monophyletic group by a variety of lines of molecular evidence. Due to the high amount of morphological variation across the three genera and the species in tribe Neillieae, conflicting classification schemes and numerous species have been proposed over the past three centuries. However, no comprehensive systematic study of the group, including all species across their geographic ranges, has ever been undertaken. As part of a taxonomic revision of tribe Neillieae, a revision of Phys- ocarpus based on the morphological examination of herbarium specimens, including types, and field observation is presented. Artificial keys, comprehensive nomenclatural treatments, descriptions, distribution maps, and lists of spec- imens examined are provided. Six species in Physocarpus are recognized. A lectotype is here designated for the fol- lowing species: Opulaster pubescens, Opulaster ramaleyi, Spiraea opulifolia var. parvifolia, Spiraea opulifolia var. tomentella, Physocarpus michiganensis, and Physocarpus missouriensis.

Recent phylogenetic analyses of tribe Neillieae in Rosaceae, which comprises Neillia, Physocarpus, and Stephanandra, based on both nuclear and chloroplast DNA sequences revealed that Neillia and Stephanandra together form a strongly supported monophyletic group and that species of Stephanandra are nested within Neillia. The close relationship between Neillia and Stephanandra is also supported by leaf morphology, inflorescence type, and carpel number. In order to better reflect the evolutionary relationships among species of Neillia and Stephanandra, based on molecular phylogenetic and morphological evidence, Stephanandra is united with Neillia, and three traditionally recognized species and a hybrid of Stephanandra are transferred to Neillia. This treatment results in new combinations as follows: N. hanceana (Kuntze) S. Oh, N. incisa (Thunberg) S. Oh, and N.×nakatsu-riparia (H. Takahashi) S. Oh. Stephanandra incisa (Thunberg) Zabel var. macrophylla Hideo Takahashi is treated as a new synonym of N...

A phylogeny of the tribe Neillieae (Rosaceae), which comprises Neillia, Stephanandra, and Physocarpus, was reconstructed based on nucleotide sequences of several regions of cpDNA, the ITS and ETS regions of rDNA, and the second intron of LEAFY, to elucidate relationships among genera and species in Neillieae and to assess the historical biogeography of the tribe. Phylogenetic analyses indicated that Physocarpus and Neillia-Stephanandra were strongly supported as monophyletic and suggested that Stephanandra may have originated by hybridization between two lineages of Neillia. Dispersal-vicariance analyses suggested that the most recent common ancestor of Neillieae may have occupied eastern Asia and western North America and that Physocarpus and Neillia-Stephanandra may have been split by an intercontinental vicariance event in the early Miocene. The biogeographic analyses also suggested that species of Neillia and Stephanandra diversified in eastern Asia, whereas in Physocarpus one dispersal event from western North America to eastern Asia occurred. Two divergent types of LEAFY sequences were found in the eastern North American species, P. opulifolius, but only one type was present in each plant. The two types of sequences may represent homeologous genes that originated by hybridization between P. capitatus and P. monogynus, both western North American species.

A homeotic gene, LEAFY, has been suggested to be a single-copy gene in diploid angiosperms. Nucleotide sequences of the second intron of this gene, along with those of several regions of the chloroplast genome (trnL-trnF, trnD-trnY-trnE-trnT, and matK-trnK) and nuclear ribosomal ITS, were obtained from the species of Neillia and Stephanandra to examine the phylogenetic utility of the intron and to elucidate the phylogenetic relationships among species of the two genera. PCR amplification of the second intron of LEAFY using universal degenerate primers produced PCR products in sufficient quantity for successful direct sequencing. The length of the intron ranged from 591 to 622 base pairs (bp) in Neillia and Stephanandra, except in N. thibetica (ca. 1370 bp), and sequence analysis of this region from multiple accessions revealed low levels of infraspecific variation. Comparison of the LEAFY data with ITS and cpDNA data demonstrated that the LEAFY intron was the most variable and useful for phylogenetic analysis at the species level, providing many more phylogenetically informative characters per 100 bp (7.4) than either ITS (3.2) or cpDNA (0.7). Phylogenetic analyses of LEAFY data using both maximum parsimony and likelihood methods generated well supported and highly resolved gene trees with few homoplasies (CI=0.97). Stephanandra is monophyletic and is nested within Neillia in both LEAFY and cpDNA trees, while the relationship is poorly resolved by ITS data. LEAFY and cpDNA data, however, strongly conflicted with each other with respect to the position of Stephanandra: LEAFY trees placed Stephanandra as sister to the ((N. affinis, N. gracilis), N. thyrsiflora) clade whereas cpDNA data suggested Stephanandra is sister to N. uekii. Both gene trees, however, are nearly identical to each other when Stephanandra is excluded. A hybrid origin of Stephanandra is suggested as a plausible hypothesis to explain the incongruence between LEAFY and cpDNA data sets, though gene duplication/loss and lineage sorting events cannot be ruled out as possibilities.