RAI, HARDEEP S1*, PATRICK A REEVES2, RICHARD G OLMSTEAD2, and SEAN W GRAHAM1. 1Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada, T6G 2E9; 2Department of Botany, University of Washington, Seattle, Washington, USA 98195. - Conifer monophyly and higher-order relationships based on a large, multigene plastid data set.
Recent work addressing the phylogeny of the extant conifers has
yielded sharply conflicting views. For example, several studies have
provided robust support for a sister-group relationship between
Gnetales and Pinaceae, while others indicate that the extant conifers
are monophyletic. To address these conflicts and clarify conifer
relationships we sequenced a large portion of the plastid genome
(spanning 15-17 conservative genes) from numerous seed plants,
including 18 exemplar conifer taxa. The genes examined represent ca.
one-ninth of the conifer plastid genome. Most analyses using different
optimality criteria and data partitions produced similar results.
These were are all largely congruent with the relationships proposed
in a conifer-wide study by Stefanovic et al (1998) using 26S rDNA
data, and the study of Cupressaceae s.l. by Gadek et al. (2000) using
combined matK + rbcL sequences. We observed strong bootstrap support
across almost all of our plastid-based phylogeny. These results
suggest that the current level of character sampling is sufficient to
reliably recover most aspects of higher-order relationship among the
extant conifers. An unusual expansion in the open reading frame of one
of the genes provides a structural synapomorphy linking Araucariaceae
and Podocarpaceae. The conifers are consistently shown to be
monophyletic among living seed plants, with Pinaceae indicated as the
sister group of the other conifers, and with
Araucariaceae-Podocarpaceae as the sister group of (Sciadopityaceae +
Taxaceae + Cephalotaxaceae + Cupressaceae s.l.). The family
Sciadopityaceae is the sister-group of the remaining taxa in the
latter clade. Our results also suggest that the "Gnepine
hypothesis" found in several recent molecular studies is a severe
long-branch artifact that may be partly ameliorated by a sufficiently
dense taxon sampling within the conifers. The likely parallel
dissolution of several chloroplast NADH dehydrogenase genes in
Pinaceae and Gnetales is also discussed.
Key words: Araucariaceae, Chloroplast, deep phylogeny, Gnepine, ndh genes, Podocarpaceae