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Generic and Subtribal Classification of American Vernonieae SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 89 Generic and Subtribal Classification of American Vernonieae Harold Robinson Smithsonian Institution Press Washington, D.C. 1999 ABSTRACT Robinson, Harold. Generic and Subtribal Classification of American Vernonieae. Smithso- niun Contributions to Botany, number 89, 116 pages, 1999.-The Vernonieae in America is herein defined to exclude the Liabeae and Pseudostifftiu (Moquinieae), and to include elements sometimes placed in the Heliantheae (Trichospiru) or Lactuceae (Stokesiu). Pollen, style bases, raphids, inflorescence form, involucre, anther appendage, and chemistry are some characters used in the reclassification. Tables 1-12 indicate the distribution of these characteristics in most American genera. Vernoniu s.s., with type A pollen, is typified by K noveborucensis (L.) and occurs in the Bahamas, eastern North America, south to central Mexico and has two spe- cies in temperate South America. All other species previously placed in Vernoniu need to be removed from the genus, a process that is nearly complete for neotropical species. Most Amer- ican Vernonieae seem to form a single related subgroup in the tribe. Subtribes included in the related subgroup are the Lychnophorinae (x = 15, 17, 18) and Centratherinae (x = 16) with type A pollen and hroheliangolides; Piptocarphinae (x= 17) with type A pollen, deciduous inner involucre, and sometimes opposite leaves; Vernoniinae (x = 17), many with glanduliferous anther appendages (including the Lepiduplou complex mostly with echinolophate pollen); and the newly proposed subtribes Sipolisiinae with type A pollen, armed receptacles, and carbon- ized achenes; Chrestinae with echinolophate pollen; and Leiboldiinae (x= 19) with type A pol- len, large heads, and a modified callus at the top of the achene. Of uncertain relationship to the preceding related subgroup are Stokesiu (x = 7), the Elephantopodinae (x = 1 1, 13), and the Rolandrinae (x = 8) with echinolophate pollen; three genera with type E pollen (Pucourinu, Acilepidopsis, and Mesunthophoru); and the Trichospirinae with type A pollen and flattened bicornute achenes. Ten subtribes are recognized, plus four unplaced generic groups. Three of the subtribal names, Leiboldiinae, Chrestinae, and Sipolisiinae are validated in the treatment. The genus Cuutingunthus of eastern Brazil, a possible relative of the Elephantopinae, is described as new. A list of the 76 accepted genera of Vernonieae, native or introduced in the New World, is provided, and a complete list of names of American Vernonieae is given with an indication of their present taxonomic dispositions and geographical distributions. Fifteen new combinations are provided. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Annuls ofrhe Smithsonian Znstifution. SERIESCOVER DESIGN: Leaf clearing from the katsura tree Cercidiphyllumjuponicum Siebold and Zuccarini. Library of Congress Cataloging-in-Publication Data Robinson, Harold Ernest, 1932- Generic and subtribal classification of American Vernonieae / Harold Robinson. p. cm. - (Smithsonian contributions to botany ; no. 89) Includes bibliographical references. 1. Vernonia-United States Classification. I. Title. 11. Series. QKIS2747 no. 89 [QK49S.C74] 580 sac21 [S83'.99] 99-3 1024 CIP @ The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials 239.48-1984. Contents Page Introduction ............................................................ 1 Acknowledgments ..................................................... 3 Relationships of the Vernonieae ............................................ 3 Distinguishing Features of the Vernonieae .................................... 3 Summary of the Subtribes and Genera of the Neotropical Vernonieae ...............7 Key to the Subtribes and Genera of the Neotropical Vernonieae ................. 8 LEIBOLDIINAE........................................................ 8 VERNONIINAE ......................................................... 9 PIPTOCARPHINAE..................................................... 11 CHRESTINAE ......................................................... 11 CENTRATHERINAE.................................................... 12 LYCHNOPHORINAE.................................................... 12 SIPOLISIINAE ........................................................ 13 ELEPHANTOPODINAE.................................................. 13 ROLANDRINAE....................................................... 14 TTUCHOSPIRINAE..................................................... 14 American Genera Not Placed in Subtribes ................................. 14 Caatinganthus ..................................................... 15 Key to the Species of Caatinganthus ................................. 16 Caatinganthus harleyi ............................................. 16 Caatinganthus rubropappus ........................................ 16 Figures ............................................................... 17 Tables ................................................................ 37 Appendix I: Accepted Generic Names of the Vernonieae Native or Introduced in the NewWorld .......................................................... 49 GenusExcluded ...................................................... 52 Appendix 11: List of Species Names of Western Hemisphere Vernonieae .......... 53 Literaturecited ....................................................... 111 ... 111 Generic and Subtribal Classification of American Vernonieae Harold Robinson Introduction ment had a brief mention of potentially useful pollen structure that was first used in the taxonomy of the tribe by Steetz The present paper is derived from text and illustrations origi- (1864). The Bentham classification was accepted almost with- nally prepared for presentation at the International Compositae out change by Hoffmann (1890-1894), differing only in the Conference, Royal Botanic Gardens, Kew, in July-August addition of the genus Gorceixia Baker (1882) and the transfer 1994 and for subsequent publication in the Symposium vol- of Gongrothamnus Steetz ex Peters from the Senecioneae to ume. The paper is expanded in the present version to include the synonymy of Vernonia. This classification, based on validation of new subtribes, illustrations of selected genera, Bentham in Bentham and Hooker f. and adopted by Hoffmann, scanning electron photomicrographs of pollen (Figures 1-8), il- was accepted as traditional by most workers in the tribe for the lustrations of various neotropical Vernonieae (Figures 9-1 8), next ninety years. tables of the subtribes and genera of Vernonieae (Tables 1-12), The review of the Vernonieae by Jones (1977) provided the a description of a new species of Caatinganthus H. Rob. (Fig- most recent basis for revisional studies in the tribe using pol- ures 19, 20), a list of the Western Hemisphere genera of the len, chromosome numbers, and chemistry. This work was part Vernonieae with their synonyms, authors, and places of publi- of a series of studies by Jones and various coauthors (1973- cation (Appendix I), and a complete as possible list of the spe- 1986), and various students (Chapman and Jones, 1978; Jones cies of American Vernonieae with accepted taxonomic disposi- and Faust, 1978; Coile and Jones, 1981, 1983; Keeley and var- tions and geographic distribution (Appendix 11). ious coauthors, 1978-1980; B.L. King, 1986; Kirkman, 1981; The treatment of the Vernonieae by Bentham in Bentham and MacLeish, 1984-1987; MacLeish and Schumacher, 1984; G.L. Hooker f. (1873) presented a pattern common in traditional Smith, 1981, 1982; and Stutts, 1980, 1981, 1983, 1988). Some concepts of Asteraceous tribes, namely a large core genus, Er- of these studies, and work of other recent authors, emphasized nonia Schreb., and various satellite genera based mostly on trichomes (Faust and Jones, 1973; Pope, 1983), underground variations in pappus and obvious features of the inflorescence. systems (Menezes et al., 1979); anatomy (Magalhiies and Alen- Some of the primary characters used by Bentham in the treat- castro, 1973; Alencastro, 1978; Carlquist, 1964; Petriella, ment of the tribe involved congestion of the heads, number of 1966), pollen (Stix, 1960; C.E. Smith, 1969; Kingham, 1976; florets in the heads, surface of the receptacle, and presence or Jones, 1979b, 1981a; Keeley and Jones, 1977b, 1979; Bolick, absence and texture of tails on the bases of the anther thecae. 1983; Isawumi et al., 1996), presence of latex (Lewinsohn, The 1873 treatment suffered from the comparatively few char- 199l), chromosome numbers (Jones, 1974, 1979a; Keeley, acters used at the time, the excessively paraphyletic nature of 1978; Dematteis, 1996), and nomenclatural details (Nicolson, the core genus, Vernonia, the artificial delimitation of many of 1991; Jeffrey and Hind, 1994; Grant, 1994). Jones (1977), Har- the segregate genera, and the failure to intercalate the segregate borne and Williams (1977), and Robinson, Bohlmann, and genera into the related parts of the core genus. The 1873 treat- King (1980) presented some early apparent correlations of chemical data with classification or geography, and the exten- sive subsequent studies of secondary metabolites were summa- Harold Robinson, Department of Botany, National Museum Of Natu- ral History, Smithsonian Institution, Washington,D. C. 20560-0166. rized by Bohlmann
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