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First published online December 24, 2008; doi:10.3732/ajb.0800320 American Journal of Botany 96: 216-227 (2009) © 2009 Botanical Society of America, Inc.

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Different gymnosperm outgroups have (mostly) congruent signal regarding the root of flowering plant phylogeny¹

UBC Botanical Garden & Centre for Plant Research (Faculty of Land and Food Systems), 2357 Main Mall, and Department of Botany, 6270 University Boulevard, University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada

ABSTRACT

We examined multiple plastid genes from a diversity of gymnosperm lineages to explore the consistency of signal among different outgroups for rooting flowering plant phylogeny. For maximum parsimony (MP), most outgroups attach on a branch of the underlying ingroup tree that leads to Amborella. Maximum likelihood (ML) analyses either root angiosperms on a nearby branch or find split support for these neighboring root placements, depending on the outgroup. The inclusion of two species of Hydatellaceae, recently recognized as an ancient line of angiosperms, does not aid in inference of the root. Cost profiles for placing the root in suboptimal locations are highly correlated across most outgroup comparisons, even comparing MP and ML profiles. Those for Gnetales are the most deviant of all those considered. This divergent outgroup either attaches on a long eudicot branch with moderate bootstrap support in MP analyses or supports no particular root location in ML analysis. Removing the most rapidly evolving sites in rate classifications based on two divergent angiosperm root placements with Gnetales yields strongly conflicting root placements in MP analysis, despite substantial overlap in the estimated sets of conservative sites. However, the generally high consistency in rooting signal among distantly related gymnosperm clades suggests that the long branch connecting angiosperms to their extant relatives may not interfere substantially with inference of the angiosperm root.

Key Words: Amborella • angiosperm phylogeny • gymnosperm outgroups • Nymphaeales • rate classes • root cost profiles • taxonomic sampling • tree rooting • Trithuria • water lilies

Received for publication 23 September 2008. Accepted for publication 30 November 2008.

FOOTNOTES

¹ The authors are grateful to T. S. Feild for providing material of Trithuria filamentosa and to S. Magallón and S. Stefanovi for helpful comments. This work was supported by a University of British Columbia Graduate Fellowship to W.J.D.I. and by an NSERC (Natural Sciences and Engineering Research Council of Canada) Discovery Grant to S.W.G.

² Author for correspondence (e-mail: swgraham{at}interchange.ubc.ca)

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