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(American Journal of Botany. 2009;96:1462-1486.)
doi: 10.3732/ajb.0800396
© 2009 Botanical Society of America, Inc.
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Evolution and Phylogeny

Evolution of lamina anatomy in the palm family (Arecaceae)1

James W. Horn2,6, Jack B. Fisher2, P. Barry Tomlinson2–4, Carl E. Lewis2 and Karen Laubengayer5

2 Fairchild Tropical Botanic Garden 11935 Old Cutler Rd., Coral Gables, Florida 33156 USA 3 The Kampong of the National Tropical Botanical Gardens, 4013 Douglas Road, Miami, Florida 33133 USA 4 Harvard Forest, Harvard University, Petersham, Massachusetts 01366 USA 5 Department of Biological Sciences, Florida International University, Miami, Florida 33199 USA

ABSTRACT

The unique properties of tree building in Arecaceae strongly constrain their architectural lability. Potentially compensating for this limitation, the extensive diversification of leaf anatomical structure within palms involves many characters whose alternate states may confer disparate mechanical or physiological capabilities. In the context of a recent global palm phylogeny, we analyzed the evolution of 10 such lamina anatomical characters and leaf morphology of 161 genera, conducting parsimony and maximum likelihood ancestral state reconstructions, as well as tests of correlated evolution. Lamina morphology evolves independently from anatomy. Although many characters do optimize as synapomorphic for major clades, anatomical evolution is highly homoplasious. Nevertheless, it is not random: analyses indicate the recurrent evolution of different cohorts of correlated character states. Notable are two surface layer (epidermis and hypodermis) types: (1) a parallel-laminated type of rectangular epidermal cells with sinuous anticlinal walls, with fibers present in the hypodermis and (2) a cross-laminated type of hexagonal cells in both layers. Correlated with the cross-laminated type is a remarkable decrease in the volume fraction of fibers, accompanied by changes in the architecture and sheath cell type of the transverse veins. We discuss these and other major patterns of anatomical evolution in relation to their biomechanical and ecophysiological significance.

Key Words: Arecaceae • correlated evolution • epidermis • homoplasy • hypodermis • lamina anatomy • leaf biomechanics • macroevolution • palm phylogeny • venation architecture

Received for publication 26 November 2008. Accepted for publication 1 April 2009.

FOOTNOTES

1 The authors thank B. Whitney for technical assistance, C. Asmussen for supplying the data matrix used in Asmussen et al. (2006), W. J. Baker for dried leaf samples from Kew (K), R. Dirig for dried leaf samples from Cornell (BH), M. Bourell for leaf samples from the San Francisco Botanical Garden, and H. Forbes for leaf samples from University of California Botanical Garden at Berkeley. P. Griffith generously allowed full access to the collections of the Montgomery Botanical Center throughout the course of this study. C. Jones Leiva, curator of the living palm and cycad collections at FTBG, facilitated our collection efforts with unstinting patience. L. Woodbury and FTBG volunteers provided herbarium support. Research was supported in part by NSF award 0515683 and the Crum Professorship in Tropical Botany of the National Tropical Botanical Garden.

6 Author for correspondence (e-mail: hornj{at}si.edu); present address: Department of Botany and Laboratories of Analytical Biology, Smithsonian Institution, P.O. Box 37012, NMNH MRC-0166, Washington, D.C. 20013-7012 USA


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