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O .. X^ti>??X o 7 \.:vA5Viix rn XivdsK^ m CO — to £ to 1ITHS0NIAN INSTITUTION NoiiniiiSNi NviNosHiii^s S 3 iavaan ^ ^ - W '/0// X O V. JOAN W. NOWICKE and JOHN J. SKVARLA SMITHSONIAN CONTRIBUTIONS TO BOTANY v NUMBER 50 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge” was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 50 Pollen Morphology and Phylogenetic Relationships of the Berberidaceae Joan W. Nowicke andJohn J. Skvarla ISSUED OCT mnmu rubiicATiONS SMITHSONIAN INSTITUTION PRESS City of Washington 1981 ABSTRACT Nowicke, Joan W., and John J. Skvarla. Pollen Morphology and Phylogenetic Relationships of the Berberidaceae. Smithsonian Contributions to Botany, number 50, 83 pages, 215 figures, 3 tables, 1981. —Pollen from 68 collections repre- senting 14 genera and 40 species of the family Berberidaceae was examined by light microscopy, SEM, and TEM. In part, the pollen data reinforce the traditional view of closely related pairs or small groups of genera. In Berberis and Mahonia the pollen morphology would support separate family status as well as congeneric treatment. The unusual exine structure in Nandina would reinforce its treatment as a monotypic family, Nandinaceae. The distinction of Bongardia from Leontice and of Dysosma from Podophyllum is confirmed by pollen data. The presence of a fundamentally similar tectum in Achlys, Dysosma, Epimedium,Jeffersonia. Podophyllum peltatum, P. hispidum, and Vancouveria suggests closer relationship among these genera than has been previously thought. The close similarity of the pollen in Jeffersoma and Plagio- rhegma confirms their congeneric treatment. Palynologically, Bongardia, Caulo- phyllum, and Leontice are more closely related to each other than to any remaining genera. In three taxa, Diphylleia, Podophyllum hexandrum, and Ranza- nia, certain characteristic(s) of the pollen render it unique and for the most part nullify any systematic value within the family. The pollen morphology of the Berberidaceae s.l. is not similar to that of the Ranunculaceae, Hydrastis excepted, nor to Lardizabalaceae. There appear to be unusual examples of parallelism between the Berberidaceae and Gistaceae, and between Podophyllum and Croton. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. Series cover design; Leaf clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data Nowicke, Joan W Pollen morphology and phylogenetic relationships of the Berberidaceae. (Smithsonian contributions to botany no. ; 50) Bibliography: p. Title. III. 1. Berberidaceae. 2. Palynotaxonomy. I. Skvarla, John J., joint author. II. 50. Series: Smithsonian Institution. Smithsonian contributions to botany ; no. Q,K1.S2747 no. 50 [Q,K495.B45| 581s[583Ml7] 80-21960 Contents Page Introduction 1 Acknowledgments 2 Previous Treatments of the Family 2 Materials and Methods 6 Results and Discussion of Pollen Analyses 7 Ektexine Relationships 19 Genera! Discussion 21 Summary 26 Literature Cited 28 Tables and Figures 31 iii Pollen Morphology and Phylogenetic Relationships of the Berberidaceae Joan W. Nowicke andJohn J. Skvarla Introduction 600 species, with as many as 500 of these belong- ing to Berberis L. The geographical distribution of the flowering The genera most commonly regarded as be- plants presents some perplexing problems, but longing to the Berberidaceae are the following. none more so than that of disjunct or discontin- Achlys De Candolle, Berberis L., Bongardia C. A. uous genera that, by definition, occupy widely Meyer, Caulophyllum Michaux, Diphylleia Mi- separated regions. If these taxa are regarded as chaux, Dysosma R. E. Woodson, Epirnedium L., monophyletic, then it follows, largely by assump- Jeffersoma Barton (Plagiorhegma Maximowicz), tion, that either at one time their range must Leontice L., Mahonia Nuttall, Nandina Thunberg, have included intervening areas or the taxa have Podophyllum L., Ranzama Ito, and Vancouveria C. fruits and/or seeds with adaptations for long Morren and Decaisne. These genera are not con- distance dispersal. There are, in fact, floristic sidered to be a single closely related group, and relationships based upon two regions (rarely this is reflected in the fact that few systematists three) having a high number of disjunct genera have included all in one family. common to both, and it is more logical and Most modern generalists consider the genera as probable that the high number is an indication primitive or at least unspecialized and place them of a previously more continuous range. The floras with the Rannuculaceae, Menispermaceae, Lar- of eastern North America and eastern Asia are dizabalaceae, and several very small families as one of the classic examples, with as many as 80 the Order Ranunculales or Berberidales. genera having species in both regions. These gen- even era are not randomly spread among the dicots or This study of pollen morphology in the Ber- monocots, but tend to be concentrated in the beridaceae is part of an extensive and continuing research project the relationships more primitive families, one of which is the Ber- on phylogenetic beridaceae. Even in the widest sense this is a small of the Order Centrospermae. This unusual group family, consisting of 10 to 12 genera and about of families with the unique nitrogen-containing pigments, the betalains, and distinctive sieve tube Joan W.