BALDWIN, BRUCE G.* AND STACI MARKOS. Jepson Herbarium and Department of Integrative Biology, University of California, Berkeley, CA, 94720. - Phylogenetic utility of the external transcribed spacer (ETS) of 18S-26S rDNA: Congruence of ETS and ITS trees of Calycadenia (Compositae).
Universal primers based on plant, fungal, and animal sequences were
designed and used to amplify the intergenic spacer (IGS) of 18S-26S
rDNA and to sequence the 3' end of the external transcribed spacer
(ETS) in Compositae. Based on these sequences, an internal ETS primer
useful across Heliantheae was designed and used to amplify and
sequence directly a 3' region of the ETS in 21 representatives of
Calycadenia, Osmadenia, and two outgroups that were
previously studied phylogenetically using rDNA ITS sequences. Size
variation in the amplified ETS region varied across taxa of
Heliantheae from approximately 350 bp to 700 bp, in part attributable
to an approximately 200 bp tandom duplication in a common ancestor of
Calycadenia and Osmadenia. Phylogenetic analysis of the
repeat region alone and examination of apomorphic changes in the
repeat region demonstrate that the 200 bp repeats in
Calycadenia/Osmadenia have evolved divergently.
Phylogenetic analyses of the entire amplified ETS region yielded a
highly resolved strict consensus tree that is nearly identical in
topology to the ITS tree, with strong bootstrap and decay support on
most branches of the ETS tree. Combined parsimony analyses of ETS and
ITS data yielded a strict consensus tree that is better resolved and
generally better supported than trees based on either data set
analyzed separately. We calculated a 1.3 to 1.4 fold higher rate of
sequence evolution by nucleotide substitution in the ETS region
studied than in ITS-1 + ITS-2. A similar disparity in the proportion
of variable (1.3 ETS: 1 ITS) and potentially informative (1.5 ETS: 1
ITS) sites was observed for the ingroup. Levels of homoplasy are
similar in the ETS and ITS data. We conclude that the ETS holds great
promise for augmenting ITS data for phylogenetic studies of young
lineages.
Key words: Calycadenia, Compositae, ETS, external transcribed spacer, phylogeny, rDNA