Ferns. and Allied Plants

Transcription

1 Ferns and Allied Plants

2 Rolla M. Tryon Alice F. Tryon Harvard University Ferns and Allied Plants With Special Reference to Tropical America Habitat Photography Principally by Walter H. Hodge Springer-Verlag New York Heidelberg Berlin

3 Rolla M. Tryon Alice F. Tryon Harvard University Herbaria 22 Divinity Avenue Cambridge, MA 02138, U.S.A. With 2028 figures Library of Congress Cataloging in Publication Data Tryon, Rolla Milton, date Ferns and allied plants. Bibliography: p. Includes index. 1. Ferns-America. 2. Pteridophyta-America. 3. Ferns-Tropics. 4. Pteridophyta-Tropics. I. Tryon, Alice F. Goint author) II. Hodge, W. H. (Walter Henricks), III. Title. QK524.4.T78 587' by Springer-Verlag New York Inc. Softcover reprint of the hardcover 1 st edition 1982 All rights reserved. No part of this book may be translated or reproduced in any form without written permission from Springer-Verlag, 175 Fifth Avenue, New York, New York loolo, U.S.A. The use of general descriptive names, trade names, trademarks, etc. in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Typesetting: Progressive Typographers, Inc., Emigsville, P A ISBN-13: e-isbn-13: DOl: /

4 Preface This systematic treatment of the ferns and allied plants provides a modern classification of the Pteridophyta based on an assessment of the wealth of new data published during the last few decades as well as on our own research. The accounts of the genera include systematics, ecology, geography, spores and cytology and often other aspects of their biology. The scope of the work has involved all genera and the Old World species of those genera represented in America. For a few American genera, that are especially complex in the Old World, it has been necessary to limit their treatment to the American representatives and this is specifically indicated in those cases. The eight American genera that do not occur in the tropics are treated in less detail. They are Camptosorus, Gymnocarpium, Matteuccia and Onoclea of North America, Arthropteris and Thyrsopteris of the Juan Fernandez Islands, Pleurosorus of southern South America, and the amphitropical Cryptogramma. The complete generic nomenclature is included, except in a few specified cases where taxonomic complexities of the Old World elements have not been resolved. The synonyms and accepted names of subgeneric and sectional taxa are included only when pertinent to the generic nomenclature or to an infrageneric classification. Only the name of a publishing author is cited. There is no bibliographic purpose served by including the name of a person who was the source of, but did not validly publish, a name or epithet. Combinations published prior to January 1, 1953 (International Code of Botanical Nomenclature, Article 33) are accepted if there is an indication of the basionym even though it was not directly cited. Geographic records are based on the collections of the Gray Herbarium and Arnold Arboretum, Harvard University, the New York Botanical Garden, the United States National Herbarium, Smithsonian Institution, and on monographic and floristic literature. The maps present the general distribution of the genera, based on countries and their major political subdivisions. The scale employed is shown in common and widely distributed genera such as Elaphoglossum (Fig. 96.4), Asplenium (Fig. 97.4), Blechnum (Fig ) and Lycopodium (Fig ). New names of some countries and islands have not been adopted because the previous name is prevalent in the literature and herbarium: Galapagos Islands is used for ArchipieIago de Colon, British Honduras for Belize, British Guiana for Guyana, Staten Island for Isla del Estado, and Falkland Islands for Islas del Malvinas. All of the spore figures are documented by a specimen citation, and these are in the collections at Harvard University unless another herbarium is cited. Publications that have,come to our attention prior to December, 1981 are included in the cited literature. The text has been prepar;ed jointly, with each author amending initial drafts prepared by the other. The book thus developed as a synthesis of our studies, although the nomenclature is

5 VI Preface primarily the work of Rolla M. Tryon and the sections on spores and cytology of Alice F. Tryon. Most of the materials in this work were developed with the financial support of National Science Foundation Grants G1064, G15949, G18906, G81693, GB4184, GB31170, DEB , and DEB However, any opinions, findings, conclusions, recommendations expressed herein are those of the authors and do not necessarily reflect the views of the Foundation. In addition to this essential support, we are indebted to the collaboration and assistance of many persons who have aided our work. The treatment of the genus Thelypteris has been prepared with the collaboration of Alan R. Smith and Lycopodium and Isoetes with the collaboration of R. James Hickey. E. Hennipman has reviewed the treatment of Platycerium, K. U. Kramer the treatments of Odontosoria, Lindsaea and Ormoloma, B. <l>llgaard, Lycopodium, M. Price, Loxogramme, and K. Iwatsuki the Hymenophyllaceae. Stimulating discussions of fern classification with E. Hennipman, F. Jarrett, C. Jermy, and K. U. Kramer have aided in the resolution of many problems. We are indebted to the courtesies extended, during visits to major herbaria, by F. Badn~, Paris, F. Jarrett, Kew, C. Jermy, British Museum (Natural History), D. B. Lellinger, Smithsonian Institution, J. T. Mickel, New York Botanical Garden, and R. G. Stolze, Field Museum of Natural History. Walter H. Hodge has contributed the major portion of the field photographs, most of them obtained on several trips to the American tropics for this special purpose. His contribution is of major significance to our work. Other habitat photographs, acknowledged in the captions, have been contributed by D. S. Conant, L. D. GOmez, C. Haufler, K. U. Kramer, T. Lucansky, F. G. Meyer, B. <l>llgaard, T. Reeves, C. S. Sperling, J. A. Steyermark, W. A. Weber, L. Wells, and P. G. Windisch. We are indebted for this collaboration, which has made it possible to illustrate all but a few genera as they grow in nature. Other photography and art work has been by Lydia Vickers and principally by Emily Hoffman and Sarah Landry, whose talents have made it possible to provide extensive illustrations of the genera and many of the species. Edward Seling is responsible for the exceptionally fine quality of the SEM micrographs of the spores. We are also indebted to Carolyn Hesterberg, Helen Roca Garcia, and Brooke Thompson-Mills for providing secretarial and editorial assistance. Some of the figures of Adiantopsis, Adiantum, Cheilanthes, Cyatheaceae, Doryopteris, Eriosorus,jamesonia, Notholaena, Pellaea, Selaginella and Vittariaceae have been previously published by the authors in Annals of the Missouri Botanical Garden and ; in Contributions from the Gray Herbarium of Harvard University , , , and ; and in Rhodora Fig (right) is from G. J. Gastony, Contributions from the Gray Herbarium of Harvard University ; Figs

6 Preface vii 35.6 are from C. Haufler, Journal of the Arnold Arboretum ; Fig is from E. Scamman, Contributions from the Gray Herbarium of Harvard University ; Figs. Cyath. 8,10-13 are from A. F. Tryon and L. Feldman, Canadian Journal of Botany ; Figs are from E. Hennipman, Leiden Bot. Ser. 2. Leiden University Press. 1977; and Fig is from L. D. Gomez, Brenesia The base maps used are from the Goode Series, Copyright by the University of Chicago Geography Department, with permission of the copyright holder. Cambridge, Massachusetts December, 1981

7 Contents Preface V Introduction Key to familes of Pteridophyta in America 22 Family l. 1. Botrychium 26 Ophioglossaceae Ophioglossum 33 Family 2. Marattiaceae Marattia Danaea 45 Family 3. Osmundaceae smunda 52 Family 4. Schizaeaceae Anemia Lygodium Schizaea 76 Family 5. Gleicheniaceae Gleichenia Dicranopteris 92 Family 6. Hymenophyllaceae Hymenophyllum Trichomanes 114 Family 7. Loxomataceae Loxsomopsis 125 Family 8. Hymenophyllopsidaceae Hymenophyllopsis 130 Family 9. Plagiogyriaceae Plagiogyria 134 Family 10. Dicksoniaceae Culcita Dicksonia Cibotium Thyrsopteris 153 Family 11. Lophosoriaceae Lophosoria 157 Family 12. Metaxyaceae Metaxya 162

8 X Contents Family 13. Cyatheaceae Sphaeropteris Alsophila Nephelea Trichipteris Cyathea Cnemidaria 208 Family 14. Pteridaceae Pityrogramma Anogramma Eriosorus ] amesonia Pterozonium Nephopteris Cheilanthes Bommeria Adiantopsis Notholaena Hemionitis Pellaea Doryopteris Trachypteris Cryptogramma Llavea Ceratopteris Adiantum Pteris Anopteris Neurocallis Acrostichum 348 Family 15. Vittariaceae Hecistopteris Antrophyum Vittaria Anetium 368 Family 16. Dennstaedtiaceae M icrolepia Dennstaedtia Saccoloma Pteridium Paesia Hypolepis Blotiella Lonchitis Histiopteris Odontosoria Lindsaea Ormoloma 429 Family 17. Thelypteridaceae Thelypteris 433

9 Contents Xl Family 18. Dryopteridaceae Ctenitis Atalopteris Tectaria Hypoderris Cyclopeltis Rumohra Lastreopsis Dryopteris Cyrtomium Didymochlaena Stigmatopteris Polystichum Maxonia Polybotrya Diplazium Athyrium Hemidictyum Adenoderris Cystopteris Gymnocarpium Woodsia Onocleopsis Matteuccia Onoclea A rth ropteris Oleandra Bolbitis Lomariopsis Lomagramma Elaphoglossum 617 Family 19. Aspleniaceae Asplenium Pleurosorus Camptosorus Holodictyum Schaffneria 653 Family 20. Davalliaceae Nephrolepis 656 Family 21. Blechnaceae Woodwardia Blechnum Salpichlaena 681 Family 22. Polypodiaceae Polypodium Pleopeltis Dicranoglossum Marginariopsis Microgramma 715

10 .. Xli Contents Ill. Campyloneurum Niphidium Neurodium Solanopteris Platycerium Loxogramme Grammitis 747 Family 23. Marsileaceae Marsilea Regnellidium Pilularia 767 Family 24. Salviniaceae l. S alvinia Azolla 776 Family 25. Psilotaceae Psilotum 783 Family 26. Equisetaceae Equisetum 789 Family 27. Lycopodiaceae Lycopodium 797 Family 28. Selaginellaceae Selaginella 813 Family 29. Isoetaceae Isoetes 826 Index to Names 835

11 Ferns and Allied Plants