title: Evolution of breeding systems and fruits in New World Galium and relatives (Rubiaceae).
abstract: OBJECTIVE
  Dioecy occurs in only about 6% of angiosperms, yet it has evolved many times from hermaphroditism. Polygamy is an even more uncommon condition within angiosperms, in which both unisexual and bisexual flowers occur within a species. Polygamy, dioecy, and hermaphroditism all occur within a New World clade of Galium (Rubiaceae), in which dioecy is hypothesized to have evolved from hermaphroditism via polygamy. At least five sections of Galium as traditionally defined by fruit morphology occur within this group. We tested the monophyly of sections defined by fruit morphology and sought to determine origins and pathways of breeding systems within this group. •
  
  
  METHODS
  We obtained chloroplast (rpoB-trnC, trnC-psbM, trnL-ndhJ) and nuclear ribosomal (external transcribed spacer) DNA sequences for 89 taxa from the Cruciata-Galium-Valantia (CGV) clade to estimate the phylogeny. Ancestral states for breeding systems, fruit types, and fruit hairs were reconstructed using parsimony and likelihood analyses. •
  
  
  RESULTS
  We identified nine well-supported lineages of New World Galium taxa. However, none of the sections traditionally defined by fruit morphology are monophyletic. Dioecy is inferred to have arisen at least three times from hermaphroditism; polygamy is inferred to have arisen at least twice from dioecy and at least six times from hermaphroditism. •
  
  
  CONCLUSIONS
  Polygamy appears to be a terminal condition in the CGV clade and not a pathway to dioecy. Fruit characters traditionally used in the taxonomy of this group have arisen multiple times within this clade of Galium and are not reliable indicators of shared evolutionary history.
authors: Department of Biology, University of Washington, Box 355325, Seattle, Washington 98195-5325 USA.
authors: Valerie L
authors: VL
authors: Soza
authors: Richard G
authors: RG
authors: Olmstead