Ferns and Allied Plants
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
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), 1912- III. Title. QK524.4.T78 587'097 82-3248 1982 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 9 8 7 6 543 2 1 ISBN-13: 978-1-4613-8164-8 e-isbn-13: 978-1-4613-8162-4 DOl: 10.1007/978-1-4613-8162-4
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. 104.4) and Lycopodium (Fig. 125.6). 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
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, DEB74-12319, and DEB78-03148. 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 42.1955 and 44. 1957; in Contributions from the Gray Herbarium of Harvard University 143.1942,179.1956,191. 1962, 194. 1964 and 200. 1970; and in Rhodora 73. 1971. Fig. 24.3 (right) is from G. J. Gastony, Contributions from the Gray Herbarium of Harvard University 203. 1973; Figs. 35.3-
Preface vii 35.6 are from C. Haufler, Journal of the Arnold Arboretum 60. 1979; Fig. 45.6 is from E. Scamman, Contributions from the Gray Herbarium of Harvard University 187. 1960; Figs. Cyath. 8,10-13 are from A. F. Tryon and L. Feldman, Canadian Journal of Botany 53.1975; Figs. 93.11-93.17 are from E. Hennipman, Leiden Bot. Ser. 2. Leiden University Press. 1977; and Fig. 114.4 is from L. D. Gomez, Brenesia 4. 1974. 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
Contents Preface V Introduction Key to familes of Pteridophyta in America 22 Family l. 1. Botrychium 26 Ophioglossaceae 25 2. Ophioglossum 33 Family 2. Marattiaceae 39 3. Marattia 41 4. Danaea 45 Family 3. Osmundaceae 50 5.0smunda 52 Family 4. Schizaeaceae 58 6. Anemia 60 7. Lygodium 69 8. Schizaea 76 Family 5. Gleicheniaceae 83 9. Gleichenia 85 10. Dicranopteris 92 Family 6. Hymenophyllaceae 97 11. Hymenophyllum 103 12. Trichomanes 114 Family 7. Loxomataceae 125 13. Loxsomopsis 125 Family 8. Hymenophyllopsidaceae 129 14. Hymenophyllopsis 130 Family 9. Plagiogyriaceae 134 15. Plagiogyria 134 Family 10. Dicksoniaceae 138 16. Culcita 140 17. Dicksonia 144 18. Cibotium 149 19. Thyrsopteris 153 Family 11. Lophosoriaceae 156 20. Lophosoria 157 Family 12. Metaxyaceae 162 21. Metaxya 162
X Contents Family 13. Cyatheaceae 166 22. Sphaeropteris 175 23. Alsophila 182 24. Nephelea 188 25. Trichipteris 195 26. Cyathea 201 27. Cnemidaria 208 Family 14. Pteridaceae 213 28. Pityrogramma 216 29. Anogramma 224 30. Eriosorus 228 31. ] amesonia 235 32. Pterozonium 241 33. Nephopteris 246 34. Cheilanthes 249 35. Bommeria 262 36. Adiantopsis 266 37. Notholaena 270 38. Hemionitis 278 39. Pellaea 284 40. Doryopteris 293 41. Trachypteris 302 42. Cryptogramma 306 43. Llavea 309 44. Ceratopteris 312 45. Adiantum 319 46. Pteris 332 47. Anopteris 341 48. Neurocallis 345 49. Acrostichum 348 Family 15. Vittariaceae 354 50. Hecistopteris 356 51. Antrophyum 357 52. Vittaria 362 53. Anetium 368 Family 16. Dennstaedtiaceae 373 54. M icrolepia 373 55. Dennstaedtia 377 56. Saccoloma 383 57. Pteridium 387 58. Paesia 395 59. Hypolepis 398 60. Blotiella 403 61. Lonchitis 406 62. Histiopteris 410 63. Odontosoria 414 64. Lindsaea 421 65. Ormoloma 429 Family 17. Thelypteridaceae 432 66. Thelypteris 433
Contents Xl Family 18. Dryopteridaceae 454 67. Ctenitis 459 68. Atalopteris 468 69. Tectaria 470 70. Hypoderris 481 71. Cyclopeltis 484 72. Rumohra 487 73. Lastreopsis 491 74. Dryopteris 496 75. Cyrtomium 510 76. Didymochlaena 515 77. Stigmatopteris 519 78. Polystichum 524 79. Maxonia 533 80. Polybotrya 536 81. Diplazium 543 82. Athyrium 555 83. Hemidictyum 561 84. Adenoderris 565 85. Cystopteris 568 86. Gymnocarpium 574 87. Woodsia 577 88. Onocleopsis 582 89. Matteuccia 585 90. Onoclea 587 91. A rth ropteris 590 92. Oleandra 593 93. Bolbitis 600 94. Lomariopsis 607 95. Lomagramma 613 96. Elaphoglossum 617 Family 19. Aspleniaceae 627 97. Asplenium 629 98. Pleurosorus 646 99. Camptosorus 648 100. Holodictyum 650 101. Schaffneria 653 Family 20. Davalliaceae 654 102. Nephrolepis 656 Family 21. Blechnaceae 662 103. Woodwardia 663 104. Blechnum 669 105. Salpichlaena 681 Family 22. Polypodiaceae 684 106. Polypodium 688 107. Pleopeltis 702 108. Dicranoglossum 708 109. Marginariopsis 712 110. Microgramma 715
.. Xli Contents Ill. Campyloneurum 722 112. Niphidium 727 113. Neurodium 732 114. Solanopteris 735 115. Platycerium 739 116. Loxogramme 743 11 7. Grammitis 747 Family 23. Marsileaceae 759 118. Marsilea 760 119. Regnellidium 765 120. Pilularia 767 Family 24. Salviniaceae 770 12l. S alvinia 771 122. Azolla 776 Family 25. Psilotaceae 782 123. Psilotum 783 Family 26. Equisetaceae 788 124. Equisetum 789 Family 27. Lycopodiaceae 796 125. Lycopodium 797 Family 28. Selaginellaceae 812 126. Selaginella 813 Family 29. Isoetaceae 826 127. Isoetes 826 Index to Names 835
Ferns and Allied Plants