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(American Journal of Botany. 2008;95:608-625.) doi: 10.3732/ajb.2007346 © 2008 Botanical Society of America, Inc.

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From annuals to perennials: phylogeny of subtribe Castillejinae (Orobanchaceae)¹

² Department of Biology, University of Washington, Box 355325, Seattle, Washington 98195-5325 USA ³ Herbarium, Burke Museum of Natural History, University of Washington, Box 355325, Seattle, Washington 98195-5325

ABSTRACT

Variation in life history strategies is a fundamental question in evolutionary biology, and the cooccurrence of annual and perennial habits in Castilleja and Castillejinae provides the opportunity to study the evolution of plant life history in a phylogenetic context. Molecular phylogenetic analysis of two chloroplast (rps16 and trnL/F) and two nuclear ribosomal (internal and external transcribed spacers) DNA regions support the monophyly of subtribe Castillejinae (Orobanchaceae). A well-supported phylogeny of the six genera (Castilleja [180 spp.], Clevelandia [1 sp.], Cordylanthus [18 spp.], Ophiocephalus [1 sp.], Orthocarpus [9 spp.], and Triphysaria [5 spp.]) comprising the subtribe is presented, and morphological synapomorphies are identified for the major lineages recovered. Orthocarpus and Triphysaria are both monophyletic; Cordylanthus is biphyletic. Clevelandia and Ophiocephalus are derived from within Castilleja. The perennial Castilleja clade (160 spp.) is derived from a grade of annual taxa including Castilleja sect. Oncorhynchus (16 spp.), Cordylanthus, Orthocarpus, and Triphysaria. This suggests that the perennial habit evolved a single time from an annual ancestral lineage that persisted throughout the diversification of Castillejinae, contrary to classical interpretations of life history evolution in plants. Given the prevalence of polyploidy among perennial Castilleja species, perenniality may have played an important role in the origin and establishment of polyploidy in Castilleja.

Key Words: annual • Castilleja • Castillejinae • Cordylanthus • Orobanchaceae • Orthocarpus • perennial • polyploidy

Received for publication 24 October 2007. Accepted for publication 28 February 2008.

FOOTNOTES

¹ The authors thank J. Ammirati, T. Bradshaw, M. Egger, and three anonymous reviewers for critical comments on earlier drafts of the manuscript; M. Donoghue, K. Karol, B. Moore, T. Near, and S. Smith for helpful discussions; and S. Collier, P. Lu-Irving, and P. Reeves for laboratory assistance. This research was supported by a Graduate Fellowship in Molecular Systematics from the University of Washington Department of Botany, the Karling Graduate Student Research Award from the Botanical Society of America, the Research Award for Graduate Students from the American Society of Plant Taxonomists, the Award for Graduate Student Research from the Society of Systematic Biologists, a Sigma-Xi Grants in Aid of Research from the University of Washington Chapter, and the Giles Award for Graduate Student Field Research from the University of Washington Department of Botany to D.C.T., and the NSF Doctoral Dissertation Improvement Grant DEB-0412653 to R.G.O. for D.C.T.

⁴ Author for correspondence (e-mail: david.tank{at}yale.edu); present address: Division of Botany, Peabody Museum of Natural History, Yale University, P.O. Box 208118, New Haven, CT 06520-8118 USA

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