NEPOKROEFF, MOLLY1, WARREN L. WAGNER2*, STEVEN G. WELLER3, PAMELA S. SOLTIS4, ELIZABETH A. ZIMMER2, ANN K. SAKAI3, and DOUGLAS E. SOLTIS4. 1Department of Biology, 177 Churchill-Haines, University of South Dakota, 414 East Clark St., Vermillion, SD 57069-2390; 2Department of Systematic Biology-Botany, MRC-166, Smithsonian Institution, P. O. Box 37012, Washington, DC 20013-7012; 3Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697; 4Florida Museum of Natural History, Dickinson Hall, P. O. Box 117800, University of Florida, Gainesville, Florida 32611-7800. - Origin and diversification of the endemic Hawaiian genus Schiedea (Caryophyllaceae subfamily Alsinoideae) inferred from combined molecular and morphological data.
Schiedea exhibits an extraordinary range of morphological
diversity, breeding system variation, and habitat preference, making
it among the most spectacular adaptive radiations in the Hawaiian
Islands. We explored phylogenetic relationships of the endemic
Hawaiian genus Schiedea (33 species) using combined molecular
(nr ITS and ETS sequences) and morphological data. Schiedea is
monophyletic and several natural groups can be recognized
corresponding to six sections. Schiedea membranacea and S.
helleri comprise a clade sister to the remaining species of
Schiedea. Species of the formerly separate genus
Alsinidendron are monophyletic and are treated as second and
third well-marked sections. These taxa are weakly associated with
Schiedea verticillata from Nihoa, treated as a fourth section.
The fifth section includes 12 species characterized by large,
one-nerved leaves. The sixth clade contains all species of two
primarily dimorphic clades identified in previous morphological
analyses (adamantis and globosa clades), suggesting a
single origin of dimorphic breeding systems in Schiedea and
several reversals to hermaphroditism. There have been at least two
apparent shifts from mesic to dry habitats, one in the basal,
morphologically divergent S. verticillata, and at least one
other in the large clade containing primarily dimorphic species. The
correlation of breeding system and wind pollination with dry habitats
indicates that breeding systems may have evolved in response to shifts
to dry habitats during the course of adaptive radiation. In contrast,
shifts to wet habitats are associated with the evolution of autogamy,
which occurs in eight species. The diversity of breeding systems is
greater in Schiedea than any other monophyletic lineage in the
Hawaiian Islands.
Key words: Caryophyllaceae, ETS, Hawaiian Islands, ITS, phylogeny, Schiedea