American Genera

of Ferns and Lycophytes

a guide for students

Robbin C. Moran
The New York Botanical Garden

Prepared for
Tropical Ferns and Lycophytes
4–20 January 2019
Organization for Tropical Studies
versión 2.1
cover page, Diplazium ceratolepis

ii
 American Genera
of Ferns and Lycophytes
A guide for students

Robbin C. Moran
The New York Botanical Garden

version 2.1 (January 2019)

Not for sale or redistribution

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 Lycopodiaceae (16/388) Lycopodiales
Lycopodiopsida (lycophytes)
Isoëtaceae (1/250) Isoëtales

(tracheophytes) Selaginellaceae (1/700) Selaginellales

SEED PLANTS
Equisetidae (horsetails)
Equisetaceae (1/15) Equisetales
(euphyllophytes) Ophioglossidae Psilotaceae (2/17) Psilotales
Ophioglossaceae (10/125) Ophioglossales
Polypodiopsida Marattiidae
(ferns) Marattiaceae (6/111) Marattiales
Osmundaceae (6/18) Osmundales
Hymenophyllaceae (9/434) Hymenophyllales
Polypodiidae Matoniaceae (2/4)
(leptosporangiates) Dipteridaceae (2/11) Gleicheniales
Gleicheniaceae (6/157)
Lygodiaceae (1/40)
Schizaeaceae (2/35) Schizaeales
Anemiaceae (1/115)
Salviniaceae (2/21)
Salviniales
Marsileaceae (3/61)
Thyrsopteridaceae (1/1)
Loxsomataceae (2/2)
Culcitaceae (1/2)
Plagiogyriaceae (1/15)
Cibotiaceae (1/9)
Cyatheales
Metaxyaceae (1/6)
Dicksoniaceae (3/35)

Cyatheaceae (3/643)

Saccolomataceae (1/18) Saccolomatineae

Cystodiaceae (1/1)
Lonchitidaceae (1/2) Lindsaeineae
Lindsaeaceae (7/234)

Pteridaceae (53/1211) Pteridineae

Polypodiales
(cathetogyrates)
Dennstaedtiaceae (10/265) Dennstaedtiineae
Cystopteridaceae (3/37)
Rhachidosoraceae (1/8)
Diplaziopsidaceae (2/4)
Desmophlebiaceae (1/2)
Hemidictyaceae (1/1)

Aspleniaceae (2/730)

Woodsiaceae (1/39)
Onocleaceae (4/5) Aspleniineae (eupolypods II)
Blechnaceae (24/265)

Athyriaceae (3/650)
(eupolypods)

Thelypteridaceae (30/1034)

Didymochlaenaceae (1/1)
Hypodematiaceae (2/22)

Dryopteridaceae (26/2115)

Nephrolepidaceae (1/19)
Polypodiineae (eupolypods I)
Lomariopsidaceae (4/69)
Tectariaceae (7/250)
Oleandraceae (1/15)
Davalliaceae (1/65)

Polypodiaceae (65/1652)

A phylogeny of the families of ferns and lycophytes (PPG I, 2016)

Interactive key to the neotropical genera of ferns and lycophytes available at
www.plantsystematics.org

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Examples of petiole vasculature in ferns, as seen in cross section. A. Eupodium laeve (Marat-
tiaceae). B. Dicksonia sellowiana (Dicksoniaceae). C. Saccoloma chartaceum (Saccolomataceae).
D. Acrostichum danaeifolium (Pteridaceae). E. Diplazium hians (Eupolypods II, Athyriaceae). F.
Polystichum concinnum (Eupolypods I, Dryopteridaceae). From Vasco et al. (2013).

Petiole vasculature can be extremely useful for identification purposes when faced to an unknown fern.
Refer to Appendix 6 (page 616) for common vascular patterns in ferns.

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 About the author

Robbin C. Moran
Dr. Robbin Moran is the Nathaniel Lord Britton Curator of Botany at the New York Botanical Garden.
He studies ferns, horsetails, and lycophytes and has published four books and over 140 scientific
papers about these plants. For his book, A Natural History of Ferns, he received the Best Writing
award in 2004 from the Garden Writer’s Association. He was the main writer, editor, and organizer
for the fern and lycophyte volume of Flora Mesoamericana. Since 1998 he has taught Tropical Plant
Systematics, a five-week course in Costa Rica, for the Organization for Tropical Studies.

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 Contents
Index to genera......................................................................................................................................................................... xiii
Acknowledgements ................................................................................................................................................................ xvii
Introduction to Ferns and Lycophytes ................................................................................................................................... 1
The Pteridophyte Life Cycle...................................................................................................................................................... 4
The typical life cycle ..............................................................................................................................................................................................4
Observing gametangia (antheridia and archegonia)............................................................................................................................6
Antheridiogens..........................................................................................................................................................................................................7
Apogamy .................................................................................................................................................................................................................. 10
Hybridization .......................................................................................................................................................................................................... 13
Branching patterns in Ferns & Lycophytes ............................................................................................................................................ 16
Lycophylls versus Euphylls (or Microphylls versus Megaphylls).................................................................................................. 21
Eusporangia versus Leptosporangia........................................................................................................................................................... 23
Spores ........................................................................................................................................................................................................................ 24
Sterile-fertile leaf dimorphism in ferns .................................................................................................................................................... 29
Tips on Collecting Ferns................................................................................................................................................................................... 31
Overview of land plant phylogeny ......................................................................................................................................... 32
Lycophytes .................................................................................................................................................................................. 34
1. Lycopodiaceae Mirbel | Club Moss Family........................................................................................................................................ 35
2. Isoëtaceae Reichenbach | Quillwort Family...................................................................................................................................... 39
3. Selaginellaceae Willkomm | Spike-moss Family............................................................................................................................... 41
Ferns (Class Polypodiopsida)................................................................................................................................................... 47
Equisetales.........................................................................................................................................................................................................................49
4. Equisetaceae Michx. ex DC. | Horsetail Family............................................................................................................................... 49
Psilotales.............................................................................................................................................................................................................................52
5. Psilotaceae Kanitz | (Whisk Fern Family)............................................................................................................................................ 52
Ophioglossales................................................................................................................................................................................................................54
6. Ophioglossaceae Agardh | Adder’s-tongue Family........................................................................................................................ 54
Marattiales.........................................................................................................................................................................................................................58
7. Marattiaceae Bercht. & J. S. Presl | Marattia Family........................................................................................................................ 58
Leptosporangiate Ferns............................................................................................................................................................ 61
Osmundales......................................................................................................................................................................................................................61
8. Osmundaceae Bercht. & J. C. Presl | Royal Fern Family.............................................................................................................. 61
Hymenophyllales............................................................................................................................................................................................................65
9. Hymenophyllaceae Link | Filmy Fern Family..................................................................................................................................... 65
Gleicheniales....................................................................................................................................................................................................................70
10. Matoniaceae C. Presl | Matonia family.............................................................................................................................................. 71
11. Dipteridaceae Diels | Dipteris family................................................................................................................................................. 72
12. Gleicheniaceae (R. Brown) C. Pres l Forked Fern Family....................................................................................................... 73

ix
Schizaeales A. B. Frank in Leunis...........................................................................................................................................................................74
13. Lygodiaceae M. Roem. | Climbing Fern Family............................................................................................................................. 74
14. Schizaeaceae Kaulf. | Curly Grass Family.......................................................................................................................................... 75
15. Anemiaceae Link............................................................................................................................................................................................ 76
Salviniales | Water Ferns............................................................................................................................................................................................78
16. Salviniaceae Dumortier............................................................................................................................................................................. 79
17. Marsileaceae Mirbel | Clover Fern Family....................................................................................................................................... 81
Cyatheales A. B. Frank in Leunis | Tree Fern Order...................................................................................................................................83
18. Thyrsopteridaceae C. Presl...................................................................................................................................................................... 85
19. Loxomataceae C. Presl.............................................................................................................................................................................. 85
20. Culcitaceae Korall.......................................................................................................................................................................................... 86
21. Plagiogyriaceae Bower................................................................................................................................................................................ 86
22. Cibotiaceae (Nayar) Korall...................................................................................................................................................................... 88
23. Metaxyaceae Pic. Serm............................................................................................................................................................................... 88
24. Dicksoniaceae Bower.................................................................................................................................................................................. 89
25. Cyatheaceae Kaulf. | Scaly Tree Fern Family................................................................................................................................... 91
Polypodiales......................................................................................................................................................................................................................94
26. Saccolomataceae Doweld........................................................................................................................................................................ 94
27. Cystodiaceae J.R.Croft [to be done; Old World]........................................................................................................................ 95
28. Lonchitidaceae Doweld [to be done]............................................................................................................................................... 95
29. Lindsaeaceae Pic. Serm............................................................................................................................................................................... 95
30. Pteridaceae Reichenbach | Maidenhair Family.............................................................................................................................. 96
Eupolypods.....................................................................................................................................................................................................................103
Eupolypods II.................................................................................................................................................................................................................104
32. Cystopteridaceae (Payer) Schmakov...............................................................................................................................................104
33. Rhachidosoraceae X. C. Zhang...........................................................................................................................................................106
34. Diplaziopsidaceae X. C. Zhang & Christenh................................................................................................................................106
35. Desmophlebiaceae Mynssen, A. Vasco, R. C. Moran, Sylvestre & Rouhan...................................................................107
36. Hemidictyaceae Christenh. & Schneid.............................................................................................................................................107
37. Aspleniaceae Newman | Spleenwort Family...............................................................................................................................108
38. Woodsiaceae Herter.................................................................................................................................................................................109
40. Blechnaceae Newman | Chain Fern Family; Deer Ferns......................................................................................................112
41. Athyriaceae Alston | Lady ferns [to be done].............................................................................................................................114
42. Thelypteridaceae Ching ex Pichi Sermolli | Beech Ferns, Marsh Fern Family...........................................................114
Eupolypods I..................................................................................................................................................................................................................118
43. Didymochlaenaceae Ching ex Li Bing Zhang & Liang Zhang............................................................................................118
44. Hypodematiaceae Ching.........................................................................................................................................................................118
45. Dryopteridaceae Herter | Wood Fern Family...........................................................................................................................119
46. Nephrolepidaceae Pic.Serm..................................................................................................................................................................122
47. Lomariopsidaceae Alston.......................................................................................................................................................................123

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 48. Tectariaceae Lellinger.................................................................................................................................................................................123
49. Oleandraceae (J. Sm.) Ching ex Pichi Serm.................................................................................................................................125
50. Davalliaceae Mett. ex Frank | Rabbit’s foot Fern Family........................................................................................................127
51. Polypodiaceae Berchtold & J. Presl | Polypody Family............................................................................................................127
Keys to Families and Genera of Neotropical Ferns and Lycophytes............................................................................129
American genera of FERNS and LICOPHYTES................................................................................................................ 147
Literature Cited........................................................................................................................................................................578
Appendices................................................................................................................................................................................596
Appendix 1 | Squash Technique for Fern and Lycophyte Chromosomes..........................................................................596
Appendix 2 | Principal Pteridophyte Floras for the Neotropics..............................................................................................600
Appendix 3 | Families and Genera of American Pteridophytes..............................................................................................602
Appendix 4 | accepted genera of ferns and lycophytes world-wide....................................................................................604
Appendix 5 | A phylogeny for 400 species of leptosporangiate ferns.................................................................................608
Appendix 6 | Petioles in cross section....................................................................................................................................................613

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xii
 Index to genera
page numbers in bold face indicate genera main entries
A Astrolepis 3, 11, 97, 184, 194, 195, 222, Ctenitis 119, 236, 237, 320, 388, 398,
Abrodictyum 65, 66, 68, 132, 148, 149, 456 , 602 , 604 564, 603, 605
556, 604 Athyrium 114, 180, 196, 197, 338, 603, Culcita 83, 85, 86, 89, 226, 238, 239,
Acrostichum 26, 75, 96, 109, 123, 150, 604 , 606 264, 372, 552, 602, 605
151, 220, 602, 604 Austroblechnum 198, 199, 448, 540, 603, Cyathea 83, 90, 91, 92, 93, 134, 240,
Actinostachys 75, 133, 152, 153, 518, 604 241, 242, 243, 244, 264, 326, 372, 524,
602, 604 Austrolycopodium 130, 200, 201, 274, 525, 602, 605, 607

Adetogramma 154, 155, 603, 604 384 C. clade of Cnemidaria 242

Adiantopsis 97, 156, 157, 158, 222, 602, Azolla 2, 3, 25, 78, 79, 80, 82, 133, 202,
604
Adiantum 3, 14, 15, 18, 26, 95, 96, 97,
101, 156, 158, 159, 358, 506, 602, 604, Blechnum 14, 18, 86, 87, 93, 112, 198,
605
Alansmia 128, 145, 160, 161, 314, 356,
414, 548, 603, 604
Aleuritopteris 97, 162, 163, 222, 604,
607
Alsophila 91, 92, 134, 164, 165, 240,
264, 326, 372, 602, 604, 606
Amauropelta 115, 166, 167, 320, 394,
530, 536, 604
Amblovenatum 168, 169, 250, 603, 604
Ananthocorus 97, 170
Anchistea 172, 173, 374, 574, 603, 604
Anemia 8, 9, 26, 74, 76, 133, 174, 175,
176, 177, 436, 516, 602, 604, 606
Anetium 97, 492, 604
Anisocampium 114, 140, 180, 181, 603,
604
Anogramma 26, 96, 97, 178, 179, 234,
316, 470, 602, 604
Arachniodes 119, 182, 183, 246, 290,
352, 396, 488, 490, 494, 508, 564, 603, Cheiroglossa 54, 55, 130, 604
604, 605, 606
Argyrochosma 3, 96, 97, 184, 185, 222,
416, 424, 456, 470, 602, 604
Ascogrammitis 128, 145, 160, 186, 187,
414, 603, 604
Asplenium 2, 8, 9, 10, 11, 15, 24, 25, 27,
28, 108, 109, 188, 189, 190, 191, 192,
193, 196, 278, 330, 603, 604, 605, 606, 233, 482, 605
607
A. sect. Hymenasplenium 192, 193
Loxoscaphe 108, 190, 191, 605
Hymenophyllopsis 92, 244, 245, 605
203 , 392 , 602, 604 Cyclodium 119, 246, 247, 396, 480, 488,
B 494, 538, 603, 605
Cyclopeltis 123, 143, 236, 248, 420, 603,
204, 205, 362, 364, 366, 472, 474, 603, 605
604 Cyclosorus 115, 141, 166, 250, 251, 530,
Blotiella 102, 206, 207, 336, 370, 426, 536, 550, 603, 605, 606
603 , 604 Cystopteris 9, 11, 17, 104, 178, 234, 252,
Bolbitis 27, 119, 120, 208, 209, 344, 253, 324, 572, 603, 605
368 , 406 , 430, 544, 603 , 604 , 605 D
Bommeria 3, 8, 9, 11, 12, 28, 97, 210, Danaea 58, 59, 131, 254, 255, 310, 390,
211, 284, 334, 602, 604 404, 602, 605
Botrychium 28, 54, 55, 130, 174, 214, Dendrolycopodium 35, 200, 256, 257,
516, 602, 604 384, 602, 605
C Dennstaedtia 18, 102, 258, 259, 346,
Campyloneurum 128 , 144 , 216 , 217, 510, 603, 605
292, 294, 380, 408, 422, 570, 603, 604, Deparia 114, 140, 260, 261, 338, 603,
605
604, 605, 606, 607
Ceradenia 128, 145, 218, 219, 228, 306, Dicksonia 83, 89, 91, 102, 238, 240, 264,
322, 414, 576, 603, 604 265, 372, 552, 602, 605
Ceratopteris 8, 9, 96, 150, 220, 221, 602, Dicranoglossum 478, 479, 605
604, 606
Dicranopteris 73, 133, 266, 267, 282,
Cheilanthes 3, 10, 15, 96, 97, 156, 178, 318, 534, 602, 605
184, 222, 223, 224, 312, 316, 332, 416,
424, 456, 554, 572, 602, 604 Didymochlaena 118, 119, 268, 269, 603,
605
Didymoglossum 65, 66, 67, 68, 132, 232,
Cibotium 83, 88, 89, 226, 227, 238, 372, 270, 271, 272, 273, 482, 605, 606
602, 605
D. sect. Didymoglossum 270, 272
Cochlidium 118, 128, 145, 218, 228,
229, 322, 354, 380, 603, 605, 607 D. sect. Microgonium 270, 272
Diphasiastrum 17, 35, 130, 274, 275,
Cranfillia 139, 230, 231, 418, 603
277, 384, 594
Crepidomanes 65, 66, 67, 68, 132, 232,
Diphasium 35, 130, 200, 274, 276, 277,
384
C. subgen. Crepidomanes 232
Diplazium 108, 114, 188, 192, 196, 278,
Cryptogramma 96, 234, 235, 360, 602, 280, 281, 330, 338, 603, 604, 605, 606
605
D. Callipteris clade 280

xiii
Diplopterygium 70, 73, 133, 266, 282,
283, 602, 605
Doryopteris 96, 97, 210, 284, 285, 554,
602, 604, 605
Dracoglossum 286
Dryopteris 2, 7, 8, 10, 11, 15, 17, 103,
115, 119, 166, 182, 236, 246, 290, 291,
352, 450, 488, 490, 494, 564, 603, 605,
606
E
Elaphoglossum 19, 27, 31, 119, 120, 128,
208, 216, 292, 293, 294, 295, 296, 297,
298, 300, 301, 302, 303, 304, 305, 368,
406, 428, 570, 603, 605, 606
E. sect. Amygdalifolium 292
E. sect. Elaphoglossum 294
E. sect. Lepidoglossa 296, 298
E. sect. Polytricha 298
E. sect. Setosa 300, 301
E. sect. Squamipedia 302
E. sect. Wrightiana 304
Ephemeropteris 308
Enterosora 128, 145, 218, 306, 307, 576,
603, 605
Equisetum 2, 18, 24, 33, 47, 49, 50, 51,
308, 309, 602, 605
Eriosorus 26, 96, 97, 350, 426, 566, 605
Eupodium 59, 131, 310, 311, 390, 602,
605
G 532, 603, 605
Gaga 312, 313, 602, 605
Galactodenia 128, 145, 314, 315, 603,
605
Gleichenella 19, 73, 133, 282, 318, 319,
534, 602, 605
Goniopteris 115, 166, 320, 321, 530,
605, 606
Grammitis 128, 145, 218, 228, 322, 323,
354, 380, 532, 603, 605
Gymnocarpium 2, 104, 324, 325, 603,
605
Gymnosphaera 91, 92, 134, 164, 326,
327, 602, 605
H Lonchitis 95, 96, 206, 336, 370, 371,
Hecistopteris 96, 97, 136, 328, 329, 506,
602, 605
Hemidictyum 107, 108, 330, 331, 603,
605
Hemionanthes 332, 333, 605
Hemionitis 3, 15, 97, 210, 284, 332, 334,
335, 504, 602, 605
Histiopteris 17, 18, 102, 206, 258, 336,
337, 370, 426, 603, 605
Homalosorus 106, 338, 605
Huperzia 3, 17, 26, 35, 37, 340, 602
Hymenophyllopsis 92, 244, 245, 605
Hymenophyllum 65, 66, 132, 148, 244,
342, 343, 560, 568, 602, 604, 605, 606,
607
Hypoderris 123, 124, 143, 344, 345,
603, 605
Hypolepis 17, 18, 20, 96, 102, 137, 206,
258, 336, 344, 346, 347, 426, 603, 605
I Marsilea 25, 78, 79, 81, 82, 134, 392,
Isoëtes 1, 23, 24, 25, 34, 39, 40, 348,
349, 602
Stylites 40, 348
J Maxonia 119, 246, 396, 397, 480, 494,
Jamesonia 96, 350, 504, 566, 602, 605
L Megalastrum 119, 120, 236, 352, 388,
Lastreopsis 119, 120, 182, 236, 290,
352, 388, 396, 398, 450, 488, 494, 508,
603, 605
Lellingeria 128, 145, 186, 218, 228, 314,
322, 354, 355, 356, 380, 400, 412, 414,
Leucotrichum 128, 145, 160, 356, 357,
532, 603, 605
Lindsaea 95, 135, 158, 268, 358, 602,
605, 606
Llavea 96, 234, 360, 361, 602, 605
Lomaria 139, 362, 363, 364, 366, 368,
605
Lomaridium 86, 112, 139, 198, 230, 364,
365, 418, 448, 603, 605
Lomariocycas 87, 112, 139, 198, 366,
367, 448, 603, 605
Lomariopsis 27, 123, 143, 248, 254, 364,
368, 369, 406, 480, 603, 605
602, 605
Lophosoria 83, 88, 89, 91, 226, 372, 373,
602, 605
xiv
Lorinseria 139, 172, 374, 375, 574, 603,
605
Loxogramme 128, 144, 145, 376, 377,
603, 604, 605
Loxoscaphe 108, 190, 191, 605
Loxsomopsis 83, 85, 378, 602, 605
Luisma 218, 228, 322, 380, 381, 603,
605
Lycopodiella 35, 382, 602
Lycopodium 28, 35, 38, 41, 52, 384, 602
Lygodium 2, 14, 17, 26, 74, 75, 133, 386,
387, 512, 602, 605
M
Macrothelypteris 115, 140, 346, 388,
389, 398, 462, 603, 606
Marattia 58, 59, 131, 310, 390, 391,
602, 606
393, 602, 606
Matteuccia 12, 110, 111, 138, 394, 395,
432, 434, 603, 606
603, 606
398, 399, 450, 508, 564, 603, 606
Melpomene 128, 145, 186, 218, 314,
400, 401, 412, 414, 603, 606
Metaxya 83, 88, 89, 254, 404, 405, 602,
606
Mickelia 2, 27, 119, 120, 208, 304, 406,
407, 603, 606
Microgramma 2, 18, 20, 128, 144, 216,
292, 294, 376, 408, 409, 410, 422, 428,
476, 603, 606, 607
M. subgen. Solanopteris 2, 128, 410
Moranopteris 128, 145, 412, 413, 603,
606
Mycopteris 186, 414, 415, 603, 606
Myriopteris 96, 97, 125, 416, 417, 602,
606
N
Neoblechnum 112, 139, 230, 418, 419,
606
Nephopteris 606
Nephrolepis 122, 123, 248, 420, 421,
460, 603, 606
Neurodium 606
Niphidium 144, 216, 408, 422, 423, 603,
606
Notholaena 3, 96, 97, 184, 194, 222,
424, 602, 606
O Pleopeltis 128, 144, 216, 292, 294, 376,
Odontosoria 95, 135, 358, 426, 427,
602, 606
Oleandra 125, 126, 128, 428, 429, 603,
606
Olfersia 119, 246, 396, 430, 431, 480,
603, 606
Onoclea 73, 110, 111, 138, 394, 432,
433, 434, 603, 606
Onocleopsis 110, 138, 394, 432, 434,
435, 603, 606
Ophioglossum 28, 54, 55, 57, 74, 130,
436, 437, 602, 606
Cheiroglossa 54, 55, 130, 604
Oreopteris 438, 439, 603, 606
Osmunda 3, 61, 62, 63, 76, 131, 440,
441, 442, 602, 606
Osmundastrum 61, 63, 131, 440, 441,
442, 443, 445, 447, 602
P Polytaenium 96, 97, 376, 492, 493, 520,
Paesia 18, 102, 137, 258, 444, 445, 447,
603, 606
Palhinhaea 35, 129, 446, 447
Parablechnum 450
Parapolystichum 119, 120, 352, 450,
451, 603, 606
Parathelypteris 250, 452, 453, 606
Pecluma 11, 128, 144, 160, 186, 420,
454, 455, 464, 486, 522, 548, 603, 606
Pellaea 3, 10, 11, 12, 28, 96, 97, 101,
184, 284, 416, 456, 457, 459, 602, 606
Pentagramma 97, 458, 459, 470, 602,
606
Phanerophlebia 9, 119, 460, 461, 603,
606
Phegopteris 17, 115, 116, 140, 388, 462,
463, 603, 606
Phlebodium 25, 128, 144, 464, 465, 603,
606
Phlegmariurus 129, 340, 466, 467
Pilularia 25, 78, 81, 82, 134, 348, 468,
469, 602, 606
Pityrogramma 8, 26, 96, 97, 178, 316,
470, 471, 504, 602, 606, 607
Plagiogyria 83, 85, 86, 204, 238, 366,
472, 473, 474, 602, 606
Platycerium 474
408, 422, 476, 477, 478, 522, 542, 603,
605, 606
Dicranoglossum 478, 479, 605
Neurodium 606
Polybotrya 2, 27, 29, 30, 119, 246, 290,
368, 396, 430, 480, 481, 508, 564, 603,
606
Polyphlebium 65, 66, 67, 68, 132, 232,
270, 272, 482, 483, 556, 562, 568, 606
Polypodium 18, 19, 91, 104, 114, 118,
119, 127, 128, 144, 204, 228, 322, 364,
408, 410, 420, 454, 464, 476, 484, 485,
486, 487, 522, 542, 603, 606
P. dulce Group 144, 486
Polystichopsis 119, 246, 480, 488, 603,
606
Polystichum 4, 15, 119, 182, 290, 450,
460, 488, 490, 491, 508, 603, 604, 606
603, 606
Pradopteris 494, 495
Pseudolycopodiella 35, 129, 446, 496,
497
Psilotum 14, 17, 20, 49, 52, 53, 61, 498,
499, 602, 606
Pteridium 3, 7, 8, 18, 102, 137, 258, 500,
501, 603, 606
Pteris 10, 11, 12, 15, 17, 18, 96, 100,
150, 254, 500, 502, 503, 603, 604, 605,
606
Pterozonium 96, 504, 505, 566, 603, 606
R T. subgen. Amauropelta 536
Radiovittaria 96, 97, 103, 136, 170, 328,
506, 507, 520, 570, 603, 606
Rumohra 119, 182, 396, 398, 450, 488,
490, 494, 508, 509, 603, 606
S T. subgen. Steiropteris 536
Saccoloma 94, 182, 258, 290, 378, 508,
510, 511, 602, 606
Salpichlaena 2, 19, 29, 75, 386, 512, 513,
603, 606
Salvinia 2, 3, 25, 78, 79, 80, 82, 133, 468,
514, 515, 602, 606
xv
Sceptridium 54, 55, 57, 59, 130, 212,
214, 516, 517, 602, 606
Schizaea 27, 75, 133, 152, 154, 518,
519, 602, 606
Scoliosorus 96, 97, 136, 492, 520, 521,
603, 606
Serpocaulon 128, 144, 402, 464, 522,
523, 603, 606
Sphaeropteris 91, 92, 134, 164, 240,
241, 264, 326, 372, 524, 525, 526, 602,
605, 607
Sphenomeris 95, 135, 426, 526, 527,
602, 607
Spinulum 200, 256, 384, 528, 529, 602,
607
Stenogrammitis 128, 145, 314, 354, 356,
532, 533, 603, 607
Sticherus 73, 103, 133, 266, 282, 318,
534, 535, 602, 607
Stigmatopteris 115, 119, 246, 538, 539,
603, 607
Stylites 40, 348
Synammia 128, 542, 543, 607
T
Tectaria 14, 123, 124, 143, 208, 344,
406, 544, 545, 564, 603, 604, 605, 606,
607
Telmatoblechnum 546, 547, 603, 607
Terpsichore 128, 145, 160, 186, 218,
314, 412, 414, 454, 548, 549, 603, 607
Thelypteris 17, 103, 114, 115, 116, 140,
166, 167, 168, 250, 251, 320, 321, 388,
394, 402, 403, 438, 442, 452, 530, 536,
537, 550, 551, 603, 607
Cyclosorus 115, 141, 166, 250, 530,
536, 605, 606
T. subgen. Cyclosorus 250
T. subgen. Goniopteris 320
T. subgen. Meniscium 402, 403
T. subgen. Stegnogramma 530
Thyrsopteris 83, 89, 552, 553, 602, 607
Trachypteris 97, 284, 554, 555, 603, 607
Trichomanes 20, 65, 66, 67, 68, 93, 127,
132, 148, 232, 270, 272, 342, 378, 482,
556, 558, 559, 560, 562, 563, 568, 602,
607
T. subgen. Davalliopsis 556
Trichomanes (cont)
T. subgen. Feea 68, 132, 556, 558, 562
T. subgen. Lacostea 132, 556, 560, 562
T. subgen. Trichomanes 562
Triplophyllum 123, 124, 143, 236, 344, 488, 544,
564, 565, 603, 607
Tryonia 566
V
Vandenboschia 65, 66, 68, 482, 556, 560, 562,
568, 569, 607
Vittaria 8, 96, 97, 136, 170, 506, 520, 570, 571,
603, 607
W
Woodsia 2, 109, 252, 286, 572, 573, 603, 605,
607
Woodwardia 9, 19, 112, 432, 574, 575, 603, 604,
607
Z
Zygophlebia 128, 145, 218, 306, 414, 576, 603,
607

xvi
 Acknowledgements
This guide is not a finished product. I revise it every time I teach another course in Costa Rica for the Oragnization
of Tropical Studies. After each revision, it’s humbling to sit down and tally your debts to others. You suddenly realize
how much poorer the final product would have been if you had to go it alone. Above all, I thank the students who
participated in my short pteridology courses taught in Costa Rica, Venezuela, Ecuador, and Bolivia, and those in the
Tropical Plant Systematics course taught in Costa Rica under the auspices of the Organization for Tropical Studies.
These students offered many suggestions on previous versions of this work over the past 25 years. Seeing ferns anew
through their eyes was often an enlightening experience in learning what helps or confuses beginning students—the
kind of person this work is intended for. Also, I thank my colleagues who made these pteridology courses possible:,
Brad Boyle, Enrique Forero, Emilia García, Michael Grayum, Amanda Grusz, Barry Hammel, Daniel Janzen, Barbara
Lewis, David Neill, Benjamin Øllgaard, Michael Sundue, and Nelson Zamora. I have had three wonderful graduate stu-
detns: Michael Sundue, Alejandra Vasco, and Fenando Matos, and over the years they have provided many comments
for this guide.

Many Latin American botanists helped with the production of this book. Some revised the descriptions, and others
corrected my Spanish translations. For their interest, patience, and willingness to help, I thank Mercedes Asanza, Mau-
ricio Bonifacino, Jasivia González, Myra Guzmán-Teare, Francisco Lorea, Blanca León, Leticia Pacheco, Blanca Pérez-
García, Katya Romoleroux, Ricardo Rueda, Carmen Ulloa Ulloa, and Renato Valencia.

Illustrations are a vital part of this book. Most were done by Mr. Haruto Fukuda, a talented artist who lives in New
York City. Many of his illustrations were done specifically for this guide. He also did llustrations for the Pteridophytes
of Mexico (Mickel & Smith, 2004), and these were heavily used for this guide. For these, I thank John T. Mickel, my col-
league and predecessor at the New York Botanical Garden, and the New York Botanical Garden Press for generously
giving me permission to use illustrations from this book.To a lesser extent, illustrations were used from other sources.
William Burger, of the Field Museum of Natural History, Chicago, gave me permission to use illustrations from Rolla
M. Tryon and Robert G. Stolze’s Pteridophyta of Peru, published in five separate fascicles in Fieldiana, Botany. I thank the
Missouri Botanical Garden Press for permission to use illustrations from Novon of several genera of grammitid ferns.

Special thanks goes to my friend and colleague, Mauricio Bonifacino. He has contributed enormously to this guide by
migrating older versions of it from Microsoft Word to Adobe InDesign.The work is now much more manageable and
has internal links so that generic treatments can be readily accessed. Mauricio’s talent for graphic design, his extraor-
dinary computer skills, and his genuine enthusiasm for the green world have given new life to this project.

Robbin C. Moran
The New York Botanical Garden
16 December 2019

xvii
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xviii
 Introduction to Ferns and Lycophytes
There are several reasons to study ferns and lycophytes. First, they represent two of the six major clades of land
plants (the other clades being the liverworts, hornworts, mosses, and seed plants). They have a life cycle distinct from
these other clades, one that lends peculiar characteristics to their biology. It is therefore important to know something
about ferns and lycophytes to be a well informed botanist and to be able to teach introductory botany. Also, ferns
and lycophytes number about 12,000 species worldwide, with the most of these in the tropics. In wet tropical forests
they abound on the forest floor, lower portions of tree trunks, and as high canopy epiphytes. In Costa Rica, ferns and
lycophytes compose about 12% of the vascular plant flora—a large percentage compared to any flowering plant
family. A group this large and abundant cannot be ignored.

What, then, are ferns and lycophytes? They are vascular plants that disperse by dust-like spores and have a life cycle
characterized by independent, free-living gametophyte and sporophyte generations. They might also be characterized
by what they lack: flowers, fruits, and seeds. The ferns are now known to include the horsetails (Equisetaceae) and
whisk ferns (Psilotaceae). In contrast, the lycophytes consist of club mosses (Lycopodiaceae), spike mosses (Selaginel-
la), and quillworts (Isoëtes).These two groups were previously classified as the Pteridophyta.The ferns, however, share
a more recent common ancestor with the seed plants than they do with the lycophytes. Therefore, “pteridophytes”
is a paraphyletic group and is no longer used (see phylogeny part of this Guide).

Figure 1. The number of fern and lycophyte species in various regions of the world.

The ferns and lycophytes boast an old and distinguished pedigree. The lycophytes appeared at the end of the Silurian
Period 400 million years ago, and ferns belonging to extant groups (Marattiaceae) first appeared at the beginning of
the Carboniferous, about 340 million years ago. In contrast, angiosperms first appeared about 145 million years ago,
during the Jurassic. Nevertheless, about 80% of the extant ferns species belong to families that first appear in the fossil
record during the early Cretaceous 140 million years ago—about the same time that angiosperms first appeared.

Today pteridophytes occur worldwide, from icy tundras above the Arctic Circle to sweltering tropical forests at the
equator (Figure 1).They illustrate well the Latitudinal Diversity Gradient, which is the major pattern in the distribution

1
of life’s diversity on earth. That is, as you go from the poles toward the equator, the number of species per unit area
increases. For instance, 30 species of ferns and lycophytes grow in Greenland, 100 in New England, 130 in Florida, 652
in Guatemala, 1160 in Costa Rica, and about 1,250 in Ecuador (Figure 2). Exceptions to this gradient can be found in
certain smaller groups, such as Dryopteris, Equisetum, Gymnocarpium, and Woodsia, which are temperate-zone genera.
But on the whole, the ferns and lycophytes are most diverse in the tropics.

The tropics are most diverse for ferns and lycophytes not only in numbers of species, but also in different growth
forms. Common in the tropics are arborescent ferns, epiphytes, secondary hemiepiphytes, scramblers, and climbers—
growth forms almost entirely absent from the temperate zones. Entire taxonomic groups of ferns are abundant in the
tropics but are absent or nearly so from the temperate zones, such as the tree ferns (Cyatheaceae and Dicksoniace-
ae), forked ferns (Gleicheniaceae), and dwarf polypodies (grammitids). Because the temperate zones harbor fewer life
forms and taxonomic groups, to get an overall view of fern and lycophyte diversity, you must visit the tropics

In the tropics, one aspect of this diversity that is readily seen is variation in size. Azolla, a floating aquatic fern, has leaves
about one millimeter long, and the whole plant is generally no bigger than a nickel. In contrast, there are tropical tree
ferns whose trunks soar 18 meters tall and bear leaves more than four meters long. But the prize for the longest
leaves go to the climbing ferns (Lygodium, Schizaeaceae) and Salpichlaena (Blechnaceae). Their leaf apices never stop
growing, and leaves can reach over 30 meters long.

Some tropical ferns have adaptations not found among the temperate species. The potato ferns (Microgramma sub-
gen. Solanopteris), a neotropical clade, bear modified rhizomes that resemble small potatoes. These are chambered
within and harbor ants that fiercely attack anything that disturbs the plant. Another fern, Salvinia, has rounded leaves

Figure 2. The latitudinal diversity gradient. Number of fern and lycophyte in selected areas of the world. From Moran (2004).

that float on water, and some species have on their upper surface tiny eggbeater-shaped hairs. If the plant is pushed
below the water, air is trapped in the cage-like part of the eggbeater hairs, and this buoys the plant back to the surface.
The bird’s nest fern (Asplenium serratum) is an epiphyte that forms its own soil in the tree tops. It has entire, elongate
leaves arranged like a funnel or basket. Falling organic debris is caught in the basket and decomposes, providing the
plant with nutrient-rich, water-holding organic humus into which to grow its roots. Some ferns climb tree trunks.
Mickelia and Polybotrya begin growth on the forest floor, but upon encountering a tree trunk, they turn upward and
climb. On the climbing part of their rhizome are produced fertile (spore-bearing) leaves that resemble skeletonized
versions of the photosynthetic sterile ones. After the spores are shed, the fertile leaves wilt, but the vegetative ones
persist. None of these adaptations occur in temperate ferns.

2
Although ferns and lycophytes reach their greatest frequency and abundance in wet forests, they also grow in dry
habitats. In fact, the dry regions of northern Mexico are a center for species richness and endemism of certain groups.
Especially prominent in dry habitats are genera of Pteridaceae such as Argyrochosma, Astrolepis, Bommeria, Hemionitis,
Myriopteris, Notholaena, and Pellaea. Most of their species reproduce asexually by diploid spores—a mode of repro-
duction called apogamy. The spores germinate, grow into a prothallus, and then vegetatively proliferate a new sporo-
phyte. This asexual reproduction obviates need for water in the life cycle—a distinct advantage in dry habitats. Other
adaptations to dry habitats are stiff, thick leaves with a layer of wax to prevent drying, or scales that absorb moisture.
In dry weather, some pteridophytes, such as Selaginella pallescens y Pleopeltis polypodioides, dry and curl, loosing over
95% of their normal moisture content. When it rains, these plants absorb water, uncurl, and begin to photosynthesize.
Ferns and lycophytes do occur in dry habitats, often at the bases of rocks or in protected crevices, but they are less
conspicuous in such habitats than in wet forests.

Compared to seed plants, pteridophytes are of little economic importance. One reason is that they lack alkaloids, a
group of chemicals physiologically active in mammals (e.g., caffeine, nicotine, quinine, strychnine). There is, however,
one exception: Huperzia produces huperzine, an alkaloid reported to slow the onset of Alzheimer’s disease.

But several pteridophytes are economically important, the most significant being Azolla. For centuries it has been
used as an organic fertilizer for rice in southeastern Asia (especially southern China and Vietnam). This small, floating
aquatic fern harbors a cyanobacterium in its leaves. The cyanobacterium fixes nitrogen from the air and converts it
into soluble nitrogenous compounds absorbed by the plant. Thus Azolla is a rich source of nitrogen. Other econom-
ically important ferns are the ornamentals, particulary Adiantum (maindenhair fern), Platycerium (staghorn fern), and
Davallia (rabbit’s-foot fern). Cultivated ferns have a host of enthusiastic growers in many parts of the world.The dense,
fibrous root mats of certain species of Osmunda and tree ferns are used by horticulturists to grow orchids. Lastly--and
something we take for granted--is that much of our electricity comes from burning the fossil ferns and lycophytes that
grew in the coal-forming swamps of the Carboniferous.

Other ferns are important economically, but in a harmful way. Bracken (Pteridium spp.) invades pastures and aban-
doned agricultural land in many parts of the world, sometimes excluding all other plants. It is poisonous to livestock
because it contains thiaminase, an enzyme that destroys thiamine (Vitamin B1). The plants are nearly impossible to
eradicate because of their deep, underground rhizomes. The Kariba weed (Salvinia molesta), a floating fern native to
southern Brazil, has escaped in the Old World where its natural insect enemies are absent. It has become terrible
weed, carpeting waterways and preventing navigation and fishing. It is also becoming a problem in the southeastern
United States from Florida to Texas.

3
 The Pteridophyte Life Cycle
The typical life cycle

Life cycle refers to the series of developmental and reproductive events that take place from one generation to
the next. All land plants (and many green algae) share a haplodiplontic life cycle, one that involves the alternation of
generations, in this case the alternation of a haploid gametophyte with a diploid sporophyte. This basic green plant
life cycle is modified in ferns and lycophytes—a modification usually denoted by the name “pteridophyte life cycle.”
To appreciate the biology of ferns and lycophytes, these modifications must be understood. It is one way that these
plants differ from the other major clades of land plants, and its properties generate and explain many peculiar aspects
of their biology.

To describe the typical pteridophyte life cycle, we’ll use a fern (Figure 3). We’ll start with a plant that has stems, roots,
and leaves. On the lower surface or margins of the leaves are black dots or lines called sori (singular sorus), which are
clusters of sporangia (singular sporangium) that contain the spores. A fern sporangium typically consists of a thin stalk
and a globose capsule that is encircled by a row of thickened, darkened cells called the annulus. Inside the capsule are
the spores, each one a single cell. The spores are barely visible to the naked eye. En masse they appear as fine dust.

Figure 3. The pteridophyte life cycle, as illustrated by the Christmas fern (Polystichum acrostichoides), a temperate North
American species. (From Carolina Biological Supply)

4
The annulus functions in hurling the spores out of the sporangia. As its cells dry, the annulus bends backward. This
splits the capsule transversely. The annulus continues bending until the elastic force of the cell walls overcomes the
drying force. At this point, the annulus, suddenly, snaps back to its original position. The movement is so fast that it
cannot be followed with the eye. This sudden, forceful movement hurls the spores out of the capsule, and they are
carried away by wind currents.

After landing, the spores germinate and develop into plantlets only 0.2–2 cm long. These plantlets, called prothalli
(singular prothallus), are typically bilobed or heart-shaped, thin, and flat. (Prothalli can usually be found in flowerpots in
greenhouses, and in wet forests on disturbed soils along trails, road cuts, or tip-up mounds of fallen trees. They often
have a slightly greasy luster, which helps distinguish them from mosses and liverworts.)

On the lower (ventral) surface of the prothalli are the sex organs, or gametangia.The male gametangium is the anther-
idium (plural, antheridia), which produces the sperm; the female gametangium is the archegonium (plural, archegonia),
which produces the egg. Fertilization takes place when there’s water in the environment. The antheridia absorb the
water, swell, and then burst to release the sperm. These swim to the egg and fertilize it. The sperm “know” where to
swim to because they are attracted by malic acid released from the decay of the archegonia neck cells. The single,
fertilized egg cell is called a zygote. This grows and develops into an embryonic plant, which further develops into a
“typical” plant with stems, roots, and leaves. The prothallus eventually withers and dies, leaving the young plant on its
own. This plant continues growing and producing new organs until it reaches full size and maturity. It then produces
leaves with sori and spores and repeats the process. The life cycle has now come full circle.

Note the two phases of the life cycle. One phase is a plant with roots, stems, and leaves. This phase, or generation,
produces the spores and is therefore called the sporophyte. The other phase produces the gametes (i.e., egg and
sperm) and is therefore called the gametophyte. Each phase starts from a single cell: the sporophyte from the zygote,
and the gametophyte from the spore., This alternation of sporophyte and gametophyte phases is called alternation
of generations. It characterizes all plants, not just pteridophytes. The way the main groups of plants alternate their
generations defines those groups and can have profound biological consequences (for example, the need for water in
fertilization in ferns is one reason why many ferns are restricted to moist habitats, and the fact that both gametangia
may occur on the same gametophyte can result in a self fertilization that is completely homozygous).

Besides functional and morphological differences, the two phases of the life cycle differ in the number of chromosome
sets (genomes). The sporophyte has two sets (diploid) and the gametophyte has one set (haploid). The two sets of
chromosomes in the sporophyte are separated to make one set. This happens during meiosis, the kind of cell division
that produces the spores, so that each spore contains only one set of chromosomes. Because the prothallus develops
from the spore, it too has only one set of chromosomes. When a haploid sperm and haploid egg fuse during fertiliza-
tion, the diploid condition is restored in the resulting zygote.Thus, alternation of generations is not only an alternation
in function, but also in chromosome number (most of the time).

The main groups of vascular plants differ in how they modify their life cycles, and these modifications characterize
those groups. For example, in comparison with other plants, pteridophytes differ by the combination of dispersal by
spores, a dominant sporophyte generation, and separate, free-living gametophyte and sporophyte generations. In con-
trast, mosses and liverworts, although they also disperse by spores, have the sporophyte attached to, and nutritionally
dependent upon, the gametophyte. Furthermore, the gametophyte is the dominant generation. Seed plants (angio-
sperms and gymnosperms) resemble pteridophytes by having a dominant sporophyte generation, but they differ by
separate male and female spores (heterospory), and only the male spores are dispersed, the female ones being held
within the tissues of the sporophyte and dependent upon it for its nutrition.

5
Observing gametangia (antheridia and archegonia)

Archegonia and antheridia can be difficult to recognize in living, unstained gametophytes. They can be most easily
observed on clean gametophytes grown on agar, but if those are unavailable, field collected material should we rinsed
and cleaned with forceps and dissecting needs to remove soil particles.

Antheridia may be formed anywhere on the gametophyte and are often frequent on small, irregularly shaped plants.
Antheridia of most common ferns (Polypodiales) are composed of three cells stacked on top of one another. From
the bottom up they are the basal cell, ring cell, and cap cell (Figure 4). These cells enclose a sphere of sperm cells.
(Eusporangiate and basal leptosporangiates differ by having more ring cells.)

Figure 4. Antheridia before and after sperm release. Note how view from the top changes because of the swelling
of the ring cell. (Courtesy of Donald Farrar)

When there is free water in the environment, the ring cells absorb water and swell, like a tire filling with air. The pres-
sure exerted by this swelling dislodges the cap cell and ejects the sperm. After ejection, the sperm remain motionless
for several seconds to a minute before rupturing their cell wall and swimming away.

The archegonia are usually found just behind the apical notch along the middle of the gametophyte. In contrast to
the antheridia of most ferns, archegonia are composed of many cells. Several cells, including the egg, are submerged in
the gametophyte thallus. The other cells project from the thallus and form the neck, which is in all ferns is composed
of four rows of cells, each row usually four cells high. The neck cells enclose one or more neck canal cells (Figure 5).

At maturity they behave like the antheridia. When there is free water in the environment, the upper neck cells swell
and break apart, the contents of the neck canal cells being exuded into the surrounding medium.This usually happens
within a minute or two after the gametophytes have been placed in water for microscopic examination. The material
exuded acts as a chemical attractant to the sperm, inducing the sperm nearby to swim down the archegonial neck

6
Figure 5. Archegonia before and after opening and fertilization. (Courtesy of Donald Farrar).

to the egg. Although only one sperm fertilizes the egg, many sperm may be attracted and the neck may eventually
become clogged with sperm.

To observe fertilization, select a number of fertile gametophytes. One or two should be large with many archegonia
in all stages of maturity. Quickly mount the gametophytes in water on a microscope slide and then focus the micro-
scope on a group of archegonia. Those farthest from the apex, but still green and unopened, are the ones most likely
to open. When an archegonium opens, focus on that archegonium using higher magnification and wait for the sperm
from a neighboring gametophyte to swim into the neck canal.

Antheridiogens

The gametophytes of homosporous ferns are generally capable of forming both antheridia and archegonia; that is,
they can become bisexual. No genetic regulation (such as sex chromosomes) has ever been demonstrated that de-
termines whether a gametophyte will become either male or female. It is known, however, that environmental factors
such as soil, light, and temperature influence sex expression. This shows that sex expression in ferns is labile. One of
the most powerful environmental factors determining the sex of an individual gametophyte is the presence of a class
of chemicals known as antheridiogens.

Antheridiogens are the hormone-like metabolic products of mature prothalli. When diffused into the surrounding
substrate, antheridiogens stimulate nearby younger prothalli to grow slowly and form antheridia precociously. Anthe-
ridiogens were first discovered by Döpp (1950), who demonstrated that substrate from mature prothallial cultures
of bracken (Pteridium aquilinum) induced antheridia formation in young prothalli of its own species and those of the
male fern (Dryopteris filix-mas). Döpp (1959) also observed that if a prothallus of P. aquilinum becomes older and
develops a central, multilayered cushion (i.e., becomes “meristic”), it produces antheridiogen but no longer reacts to
it. Only young prothalli react.

7
Since Döpp’s original discovery, four other kinds of antheridiogens have been demonstrated, these in Anemia (Näf,
1956), Ceratopteris (Schedlbauer & Klekowski, 1972), Vittaria (Emigh & Farrar, 1977), and Asplenium (Schneller & Hess,
1995). It has since been shown that the Pteridium antheridiogen is responded to in a wide range of families such as
Blechnaceae, Cyatheaceae, Davalliaceae, Dennstaedtiaceae, Dicksoniaceae, Dryopteridaceae, Polypodiaceae, Pterida-
ceae, and Thelypteridaceae. Antheridiogen from Anemia phyllitidis also affects several species of Schizaeaceae. Thus,
the term antheridiogen characterizes a function, not a chemical composition. The chemical structure of only one
group of antheridiogens is known—that of the Anemia—and it is a giberellin.

The Osmundaceae are unusual among ferns in that their gametophytes first become male when grown in isolation,
and they lack any known antheridiogen system (Greer et al., 2009).

By stimulating the production of antheridia, antheridiogens increase the probability of cross-fertilization. In homospo-
rous ferns, self-fertilization results in complete homozygosity (this cannot happen in heterosporous plants).This brings
up the problem of lethal recessive alleles being expressed in the homozygous condition. By promoting outcrossing,
the bad homozygotizing effects of selfing are avoided, and heterozygosity in populations is maintained. One might
argue that antheridiogen is unnecessary for species with bisexual prothalli where inbreeding predominates. Anther-
idiogens, however, have been detected in Asplenium ruta-muraria, which is predominantly an intragametophytic selfer
(Schneller & Hess, 1995).

Another function of antheridiogens is bypassing the need for light in spore germination. Most fern spores remain
dormant in the dark and need light to germinate; however, if kept in the dark and exposed to antheridiogen, spores
germinate. Antheridiogen-induced germination is important for spores buried in the soil (the spore bank). Antheridio-
gens can penetrate the soil to at least 1 cm and stimulate buried spores to germinate.The resulting prothalli are often
composed of several elongate cells that grow toward the soil surface. They also precociously produce antheridia. The
antheridia release sperm available for out-crossing with the older female prothalli on the surface. Epiphytic species
of Polypodiaceae have an antheridiogen system, which suggests that dark germination may play a role in competition
with bryophytes and seed plants (Chiou & Farrar, 1997). On the soil surface, the antheridiogens have been detected
up to 25 cm away from the source gametophyte (Voeller & Weinberg, 1969).

Apogamous ferns might be expected to lack an antheridiogen system because fertilization is not involved in their life
cycle. In the few species studied, some produce and respond to antheridiogens and others do not. The apogamous
diploid and triploid plants in the Dryopteris affinis group produce and react to their own antheridiogens (Schneller,
2008), as does the apogamous triploid Bommeria pedata (Haufler & Gastony, 1978). Cyrtomium, however, is variable.
The apogamous species C. falcatum neither produces its own antheridiogen nor responds to the Pteridium-type, yet
two other apogamous species, C. fortunei and C. macrophyllum, produce antheridiogen but do not react to it (Yatski-
evych, 1993).

Some evidence indicates that the antheridiogen response might actually be caused by two substances. The re-
sponse actually has two components: 1) precocious antheridia production, and 2) stunted growth (the prothalli
usually remaining as protonemata or becoming merely spatulate). Naf (1956) suggested that gametophytes sub-
jected to antheridiogen divert their potential vegetative growth into antheridia formation; however, some game-
tophytes respond to antheridiogens by stunting their growth and not producing antheridia (Chiou & Farrar, 1997).
This suggests the involvement of a growth-inhibiting substance and an antheridiogen. Antheridiogens might act by
slowing growth and reducing gametophyte size, which favors maleness (Quintanilla et al., 2007).

Antheridiogens | Selected References

Chiou, W.-L. & D. R. Farrar. 1997. Antheridiogen production and response in Polypodiaceae species. American
Journal of Botany 84: 633–640.
Döpp, W. 1959. Eine die antheridienbildung bei Farnen fördernde Substanz in den Prothallien von Pteridium aquilinum (L.)
Kuhn. Berichte der Deutschen Botanisschen Gesellschaft 63: 139–147.
Dubey, J. P. & S. K. Roy. 1985. A new antheridiogen from the fern Pityrogramma calomelanos. Proceedings of the Indian
Academy of Science (Plant Science) 95: 173–179.
Emigh, V. D. & D. R. Farrar. 1977. Gemmae: a role in sexual reproduction in the fern genus Vittaria. Science 198: 297–298.
Greer, G. K., M. A. Dietrich, S. Stewart, J. Devol & A. Rebert. 2009. Morphological functions of gibberellins in
8
 leptosporangiate fern gametophytes: insights into the evolution of form and gender expression. Botanical Journal of
the Linnean Society 159: 599–615.
Hamilton, R. G. & R. M. Lloyd. 1991. Antheridiogen in the wild: the development of fern gametophyte communities.
Functional Ecology. 5: 804–809.
Haufler, C. H. 1994. Antheridiogen, dark germination, and outcrossing mechanisms in Bommeria (Adiantaceae). American
Journal of Botany 81: 616–621.
_____ & G. J. Gastony. 1978. Antheridiogen and the breeding system in the fern genus Bommeria. Canadian Journal of Botany
56: 1594–1601.
_____ & T. A. Ranker. 1985. Differential antheridiogen response and evolutionary mechanisms in Cystopteris. American
Journal of Botany 72: 659–665.
Lloyd, R. M. 1974. Reproductive biology and evolution in the Pteridophyta. Annals of the Missouri Botanical Garden 61:
318–331.
Näf, U. 1956. The demonstration of a factor concerned with the initiation of antheridia in polypodiaceous ferns. Growth 20:
91–105.
_____. 1962. Developmental physiology of lower archegoniates. Annual Review of Plant Physiology 13: 507–532 [especially
pages 513–521].
_____. 1963. On dark germination and antheridium formation--a model study of the developmental change. Journal of the
Linnean Society (Botany) 58: 3321–331.
_____. 1979. Antheridiogens and antheridial development. Pages 435–470. In: The experimental biology of ferns [Experimental
Botany, An International Series of Monographs, Vol. 14: 1657 + xviii], A. F. Dyer, editor. Academic Press, London.
_____, K. Nakanishi, & M. Endo. 1975. On the physiology and chemistry of fern antheridiogens. Botanical Review (Lancaster)
41: 315359.
Nester, J. E. & M. D. Schedlbauer. 1982. Antheridogen activity of Anemia mexicana. Canadian Journal of Botany 60: 1606–
1610.
Quintanilla, L. G., L. de Soto, A. Jiménez & M. Méndez. 2007. Do antheridiogens act via gametophyte size? A study of
Woodwardia radicans (Blechnaceae). American Journal of Botany 94: 986–990.
Schedlbauer, M. D. & E. J. Klekowski. 1972. Antheridiogen activity in the fern Ceratopteris thalictroides (L.) Brogn. Botanical
Journal of the Linnean Society 65: 399–413.
Schraudolf, H. 1985. Action and phylogeny of antheridiogens. Proceedings of the Royal Society of Edinburgh, B, 86: 7580.
Schneller, J. J. 2008. Antheridiogens. Pages 136–160. In: T. A. Ranker & C. H. Haufler (eds.). The biology and evolution of ferns
and lycophytes. Cambridge Univ. Press.
_____, C. H. Haufler & T. A. Ranker. 1990. Antheridiogen and natural gametophyte populations. American Fern Journal 80:
143–152.
_____ & A. Hess. 1995. Antheridiogen system in the fern Asplenium ruta-muraria (Aspleniaceae: Pteridophyta). Fern Gazette
15: 64–70.
Tryon, R. M. & G. Vitale. 1977. Evidence for antheridogen production and its mediation of a mating system in natural popula-
tions of fern gametophytes. Bot. J. Linn. Soc. 74: 243–249.
Voeller, B. R. 1964. Antheridiogens in ferns. Pages 665–684. In: Régulateurs Naturels de la Croissance Végétale (Colloq. Intern.
Centr. Nat. Rech. Sci.), No. 23. [International Conference on Plant Growth Regulation, Proceedings]
_____ & E. S. Weinberg. 1967. Antheridium induction and the number of sperms per antheridium in Anemia phyllitidis. Amer-
ican Fern Journal 57: 107–112.
_____ & _____. 1969. Evolutionary and physiological aspects of antheridium induction in ferns. Pages 77–93. In: J. E. Gunckel,
editor. Current topics in plant science. Academic Press, London.
Yatskievych, G. 1993. Antheridiogen response in Phanerophlebia and related fern genera. American Fern Journal 83: 30–36.

9
Apogamy

The apogamous life cycle lacks fusion of gametes; it is asexual. Instead of producing 64 haploid spores per sporangium
(like most ferns), apogamous ferns produce, by a glitch in meiosis, 32 diploid spores (Figure 6). When these spores are
sown, they germinate and develop into smaller than normal prothallia. Instead of producing sex organs, the prothallia
soon proliferate a plantlet consisting of a root, stem, and leaf. As the plantlet grows and develops, the prothallus with-
ers. Eventually the new plant becomes independent and assumes a life of its own. When large enough, it sends forth
spore-bearing leaves, at which point the apogamous life cycle has repeated itself.

Figure 6. Normal meiosis (upper left) and different types of cell division that give rise to apogamous spores.

The apogamous life cycle is a fancy form of asexual reproduction, akin to dividing a plant’s stem and replanting both
halves, or detaching bulblets from a leaf and planting them to make new individuals. The only difference is that, in
apogamy, the structure propagating the plant is a single-celled spore.

Apogamy has been estimated to occur in about five to ten percent of ferns worldwide. In Japan, where the flora
is well studied, 13% of the fern species are apogamous. Apogamy is more frequent in some ferns than in others,
being especially common in spleenworts (Asplenium), lip ferns (Cheilanthes), wood ferns (Dryopteris), brakes (Pteris),
holly ferns (Cyrtomium), brakes (Pteris), and cliff brakes (Pellaea). On the other hand, it is noticeably absent from the
Blechnaceae, Cyatheales, and Thelypteridaceae.

10
Many fern species growing in dry or seasonally dry habitats, such as deserts, chaparral, and exposed cliff faces, are
apogamous. For these ferns, apogamy has two advantages. First, there is no need for water in reproduction (there is
no egg to be fertilized by swimming sperm). Second, the prothalli of apogamous ferns mature faster than those of
sexually reproducing ones. This means that the prothallus spends less time exposed to the hazard of desiccation. In
contrast, sexually reproducing fern species have the double disadvantage in dry habitats of needing water for fertiliza-
tion and a longer prothallial development during which time it’s susceptible to desiccation.

Nearly 75% of apogamous fern species are polyploidy, and many of these are triploid. In North America,
some familiar examples are the black-stemmed spleenwort (Asplenium resiliens), purple cliff brake (Pellaea atropur-
purea), slender lip-fern (Myriopteris feei), and star-scaled cloak fern (Astrolepis sinuata var. sinuata). Among cultivated
ferns, examples are the variegated brake (Pteris cretica var. albolineata), Fortune’s holly fern (Cyrtomium fortunei), and
shaggy wood-fern (Dryopteris atrata). These cultivated ferns tend to be weedy in greenhouses, often popping up be-
tween cracks in bricks or cement. Presumably, this happens because their apogamous mode of reproduction is quick
and efficient.

Triploid ferns must be apogamous to reproduce by spores. They cannot reproduce sexually because of
the way their three chromosomes sets behave during meiosis. Of the three sets, only two pair; leaving the third
set unpaired. The chromosomes of the paired sets are distributed evenly to the daughter cells of meiosis, but the
chromosomes in the unpaired set become distributed unequally. One daughter cell might receive, for example, 10
chromosomes, another with 16, and so on.The result of this imbalance is spore abortion—irregular, misshapen, usually
blackened spores that will not germinate (Figure 7). In apogamous ferns, however, and imbalance in the distribution
of chromosomes does not occur because there is no chromosome pairing. As a result, triploid ferns that reproduce
apogamously produce viable spores.

Figure 7. Aborted (left) and normal (right) spores of Cystopteris.

Viable spores can be picked up by air currents and dispersed long distances. This is true for the spores of apogamous
and sexually reproducing ferns. Nevertheless, compared to their sexually reproducing relatives, apogamous ferns tend
to have wider geographic distributions. For instance, the smooth cliff brake (Pellaea glabella) consists of a sexually re-
producing diploid race (with two sets of chromosomes) and an apogamously reproducing tetraploid race (with four
sets of chromosomes). The two races are indistinguishable to the naked eye (the diploid gave rise to the tetraploid
by the process of chromosome doubling, or “polyploidy”). The diploid is found only in southeastern Missouri where
it is relatively rare, but the tetraploid is common and widespread throughout much of eastern United States. Similar
range relationships are found in genera such as the copper ferns (Bommeria; Gastony & Haufler, 1976), brakes (Pteris),
star-scaled cloak ferns (Astrolepis), and comb ferns (Pecluma). It is a mystery why apogamous ferns tend to be more
widely distributed than their sexually reproducing relatives.

11
Apogamy | Selected References

Braithwaite, A. F. 1964. A new type of apogamy in ferns. New Phytologist 63: 293305.

Evans, A. M. 1964. Ameiotic alternation of generations: a new life cycle in the ferns. Science 143: 261–263.

_____. 1969. Problems of apomixis and the treatment of agamic complexes. Bioscience 19: 708–711.

Gastony, G. J. & C. H. Haufler. 1976. Chromosome numbers and apomixis in the fern genus Bommeria (Gymnogrammaceae).
Biotropica 8: 141.

_____ & M. D. Windham. 1989. Species concepts in pteridophytes: the treatment and definition of agamosporous species.
American Fern Journal 79: 6577.

Lloyd, R. M. 1973. Facultative apomixis and polyploidy in Matteuccia orientalis. American Fern Journal 63: 43–48.

Morzenti, V. 1962. A first report of pseudomeiotic sporogenesis, a new type of spore reproduction by which sterile ferns pro-
duce gametophytes. American Fern Journal 52: 6978.

Rigby, S. J. 1975. Meiosis and sporogenesis in a haploid plant of Pellaea glabella var. occidentalis. Canadian Journal of Botany
53: 894900.

Sheffield, E. & P. R. Bell. 1987. Current studies of the pteridophyte life cycle. Botanical Review 53: 442–490.

Tilquin, J. P. 1981. Note on apomixis in ferns. Acta Soc. Bot. Poloniae 50: 217222.

Tryon, A. F. 1968. Comparisons of sexual and apogamous races in the fern genus Pellaea. Rhodora 70: 124.

Walker, T. G. 1962. Cytology and evolution in the fern genus Pteris L. Evolution 16: 2743.

_____. 1966. Apomixis and vegetative reproduction in ferns. Pages 152161. In: J. G. Hawkes, editor. Reproductive biology and
taxonomy of vascular plants. Conference Report Botanical Society of Brittain. I, no. 9.

_____. 1985. Some aspects of agamospermy in ferns the Braithwaite system. Proceedings of the Royal Society of Edinburgh, B,
86: 5966.

12
Hybridization

Hybridization is the crossing of two species to form offspring that combine the characteristics of the two parents.This
occurs when the sperm from the prothallus of one species fertilizes the egg on the prothallus of another. Hybridiza-
tion has been well studied in the temperate zones, and from these studies it’s known that hybridization (followed by
polyploidy) has been an important evolutionary mechanism in ferns.

How can hybrids be detected? They are usually first noticed because they are morphologically intermediate between
their parents. This does not mean they look precisely fifty-fifty in the morphological middle between their parents,
but they exhibit a range of variation between parental extremes. For example, a hybrid between a simple-leaved and
a 1-pinnate-leaved species will exhibit leaves varying from lobed to deeply pinnate to fully 1-pinnate at the base and
pinnatifid above (Figure 9). The main exception to hybrid intermediacy is that hybrids tend to be larger than their
parents.

Another way used to detect hybrids is to look for aborted spores. Under a microscope, such spores appear mis-
shapen, irregular, and blackened (Figure 7). In some cases this irregularity is so extreme that the spores resemble dirt.
Aborted spores are caused by failure of the chromosomes to pair properly during meiosis. This happens because the
chromosomes from the two parental genomes differ either structurally or physiologically, enough so that they cannot
pair with their homeologue during meiosis. In either case, unequal distribution of the chromosomes to the daughter
cells (i.e., spores) causes abortion (Figure 8).

Figure 8. Chromosome behavior during meiosis of species and hybrids. Letters represent genomes of the parent species. Left,
offspring of same-species crosses results in normal meiosis and viable spores. Right, hybrid offspring of different species have
abnormal meiosis and aborted spores (AB) because chromosomes do not pair properly. Chromosome doubling (polyploidy)
restores fertility because chromosomes now have homologues with which to pair during meiosis.

13
Circumstantial evidence is also important for determining a hybrid. In most cases, hybrids are found growing together
with their parents.

A powerful tool for detecting hybridization is isozyme analysis.Typically when the isozymes of the parental species are
separated electrophoretically, they have different banding patterns. Hybrids combine the different banding patterns of
their parents. In other words, the hybrid’s banding patterns are additive.

Asymmetrical hybridization has been documented for only a few fern hybrids but is probably of wide occurrence
(Testo et al. 2015). This phenomenon is where one species serves as the paternal (or maternal) parent more fre-
quently than would be expected from chance alone. For hybrids between diploids and polyploids, the polyploid is
usually the maternal parent (Testo et al., 2015). This is probably because diploids tend to outcross and have longer
sperm viability compared to polyploids. Also, most diploids have an antheridiogen system that would promote the
formation of male gametophytes, thus increasing the supply of spermatozoids potentially available for fertilization. In
contrast, polyploids are more often selfing and lack an antheridiogen system. Asymmetrical hybridization has been
well documented in Asplenium alternifolium, A. X protomajoricum, Dryopteris boottii, and D. X triploidea.

Hybrids may be designated by either a formula name or a binomial. A formula name consist of the binomials of the
two parents with an “X” placed between them; for instance, Adiantum latifolium X A. petiolatum, Psilotum complanatum
X P. nudum, and Tectaria incisa X T. panamensis. In contrast, a hybrid binomial resembles a scientific name except that
an “X” is placed between the genus name and the specific epithet; for instance, Adiantum X variopinnatum (=Adiantum
latifolium X A. petiolatum), Blechnum X caudatum (= B. gracile X B. occidentale), and Lygodium X lancetillanum (= L.
heterodoxum X L. venustum).

Figure 9. Parental species and their hybrids. The hybrids are variably intermediate between the two forms of the parental
species. Both examples are from Costa Rica.

14
Although formula names have the advantage of stating the hybrid’s parentage, such names are less stable nomencla-
turally than binomials. A formula name changes if one or both parents suffers a name change, as can happen when
an earlier name is found. This creates a situation where two (or more) formula names exist in the botanical literature
and refer to the same hybrid—a confusing situation to say the least! Also, formula names must change if the parent-
age of the hybrid is reinterpreted. The parentage of Asplenum X ebenoides, one of the most frequent hybrids in the
eastern United States, has undergone three different interpretations. Neither changes to the names of the parents nor
reinterpretations of the hybrid’s parentage affect binomials. They stay the same no matter what. Thus, giving binomial
names to hybrids promotes nomenclatural stability.

When a hybrid occurs between two genera, a hybrid genus name may be coined by combining the names of the two
parental genera and placing an “X” in front, such as XAsplenosorus (=Asplenium X Camptosorus) and XHemionanthes
(=Hemionitis X Cheilanthes). Thanks to recent DNA phylogenetic studies, the parents of most intergeneric hybrids
have been shown to belong to the same genus. XAsplenosorus is an example: Camptosorus nests within Asplenium and
therefore the hybrid genus is not necessary.

Intergeneric hybrids are rare in ferns (Rothfels et al., 2015). Only four that are currently accepted have been de-
scribed: XCyclobotrya Engels & Canestraro, XCystocarpium Fraser-Jenk, XDryostichum W. H. Wagner, and XLindsaeoso-
ria W. H. Wagner. Respectively, these represent: Cyclodium X Polybotrya (Dryopteridaceae), Cystopteris X Gymnocarpi-
um (Woodsiaceae), Dryopteris X Polystichum (Dryopteridaceae), and Lindsaea X Odontosoria (Dennstaedtiaceae)..
Another intergeneric hybrid occurs between Campyloneurum and Microgramma , but it has yet to be recognized as
such (pers. obs.). All other hybrid genera in ferns have now been shown by recent molecular phylogenetic studies to
occur between species of the same genus.

Selected references | Hybridization

Barrington, D. S. 1985. Hybridisation in Costa Rican Polystichum. Proceedings of the Royal Society of Edinburgh 86B: 335–340.
_____. 1986. Variability in intermediate character states in fern hybrids. American Journal of Botany 73: 733
_____, C. H. Haufler & C. R. Werth. 1989. Hybridization, reticulation, and species concepts in the ferns. American Fern Journal
79: 55–64.
Knobloch, I. W. 1976. Pteridophyte hybrids. Publications from the Museum of Michigan State University, Biol. Ser. 5: 273–352..
Moran, R. C. & J. E. Watkins, Jr. 2002.The occurrence and morphology of Adiantum X variopinnatum (Pteridaceae). Brittonia
54: 57–60.
Rothfels, C. J., A. K. Johnson, P. H. Hovenkamp, D. L. Swofford, H. C. Roskam, C. R. Fraser-Jenkins, M. D. Windham
& K. M. Pryer. 2015. Natural hybridization between genera that diverged from each other approximately 60 million years
ago. The American Naturalist 185: 433–442.
Testo, W. L., J. E. Watkins Jr. & D. S. Barrington. 2014. Dynamics of asymmetrical hybridization in North American wood
ferns: reconciling patterns of inheritance with gametophyte reproductive biology. New Phytologist doi: 10.1111/nph.13213
Wagner, W. H. Jr. 1962. Irregular morphological development in hybrid ferns. Phytomorphology 12: 87–100.
_____. 1965. Abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. American Fern Journal 55: 9–29.
_____. 1969. The role and taxonomic treatment of hybrids. Bioscience 19: 785–789.
_____. 1968. Hybridization, taxonomy and evolution. Pages 113–138. In: V. H. Heywood, editor. Modern Methods in Plant Taxon-
omy. Academic Press, London.
_____. 1983. Reticulistics: the recognition of hybrids and their role in cladistics and classification. Pages 63–79. In: N. I. Platnick &
V. A. Funk, editors. Advances in cladistics, vol. 2. Colombia University Press, New York.
_____. 1987. Some questions about natural hybrids in ferns. Botanica Helvetica 97: 195–205.
_____ & K. L. Chen. 1965. Abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. American Fern
Journal 55: 9–29.
_____ & F. S. Wagner. 1975. A hybrid polypody from the New World tropics. Fern Gazette 11: 125–135.
_____, _____ & L. D. Gómez. 1978. The singular origin of a Central American fern, Pleuroderris michleriana. Biotropica 10:
254–264.
_____, _____ & W. C. Taylor. 1986. Detecting abortive spores in herbarium specimens of sterile hybrids. American Fern Journal
76: 129–140.
Walker, T. 1958. Hybridization in some species of Pteris L. Evolution 12: 82–92.
Valentine, D. H. 1963. The treatment of hybrids in Flora Europaea. Webbia 18: 47–55.

15
Branching patterns in Ferns & Lycophytes

Ferns and lycophytes exhibit a variety of stem branching patterns. In fact, ferns exhibit more branching patterns than
any other group of vascular plants. Unlike seed plants (cycads excepted), which are monotonously uniform in exhib-
iting axillary branching, the lycophytes and nearly all ferns have extra-axillary branching.

Two types of branching are generally distinguished in vascular plants. The first is adventitious branching where new
shoots arise from fully expanded mature organs or organ parts, such as cut stems or the margins of leaves, often in
unexpected places. The second type is acrogenous, characterized by new shoots arising from the apical meristem.
Acrogenous branching is the type important for most ferns.

Acrogenous branching may be either dichotomous or lateral. If dichotomous, the apical meristem divides equally (or
nearly so) into two derivative branch meristems (Figure 10). During the branching process, the parent meristem gives
up its existence, leaving no remainder. The two resulting daughter shoots are mirror images of each other. The parent
shoot axis can now be thought of as a kind of pillar or podium upon which the two new branches rest. In contrast,
lateral branching initiates new branches from the periphery, or shoulders, of the apical meristem. The apical meristem
remains intact, keeping its identity throughout the branching process. It grows growing continuously and uninterrupt-
edly as the branch below it develops.

Figure 10. The two types of acrogenous branching, which takes place at the apical meristem. (From Kaplan, unpublished).

These types of acrogenous branching—dichotomous and lateral—can be characterized more precisely by how the
cellular zonation of apical meristems behave during the branching process. In ferns and lycophytes, the apical mer-
istem consists of two zones. The first is the initial zone dominated by a conspicuously enlarged tetrahedral cell that
produces vacuolated prismatic cells from its lateral cutting faces. Bordering this is the morphogenetic zone, whose cells
are small and densely cytoplasmic, where the leaf and branch primordia first appear. In dichotomous branching, the
initial zone widens and then divides in two. In lateral branching, however, the initial zone stays intact, and branches
arise below from primordia produced in the morphogenetic zone.

Dichotomous branching can be either isotomous or anisotomous. In the former, branches arise from the equal di-
vision of the initial zone of the meristem, and branches of equal length are produced. In the latter, the initial zone of
the meristem divides unequally and produces branches of different lengths. Conceptually, anisotomy is intermediate
between isotomy and lateral branching.

16
Sometimes it is useful to distinguish whether a dichotomy is cruciate or flabellate. If cruciate (Latin, cruciatus, cross
shaped), the successive dichotomies are at right angles to each another, producing a three-dimensional shoot system.
Examples are Huperzia (sens. str.) and Psilotum nudum. If flabellate (Latin, flabellatus, fan), the successive dichotomies
lie in the same plane, producing a flattened shoot system, such as in Psilotum complanatum or the ultimate branchlets
in the upright shoots of Diphasiastrum.

Depending on the location of the bud, lateral branching can be characterized as either axillary or extra-axillary. Nearly
all ferns have extra-axillary buds, with axillary ones known only in the trichomanoid genera (Hébant-Mauri, 1984)
and Meniscium reticulata (Hagemann, 1989). Plants with axillary buds may extend their shoots by either sympodial or
monopodial growth, but in ferns (even those with axillary buds) all shoot extension growth is monopodial.

A given species may exhibit more than one type of branching.The Gleicheniaceae exhibit isotomous branching at the
rhizome apex and lateral branching from buds on the dorsal surface (Hagemann & Schultz, 1978). Similarly, a given
species may have both kinds of dichotomy. Diphasiastrum has horizontal runners that branch anisotomously to give
rise to erect (orthotropic) shoots whose ultimate branchlets (usually flattened) branch isotomously. Thus, branching
patterns may vary within a given species.

Isotomy has been documented in Psilotum (Hagemann, 1980) and Lygodium japonicum (Mueller, 1982). It also appears
to have evolved independently in the Dennstaedtiaceae (e.g., Hypolepis, Dennsteaditia, Histiopteris), Thelypteridaceae
(Phegopteris, Thelypteris palustris), and Cystopteridaceae (Cystopteris, Gymnocarpoium). Isotomy in dennstaedtioid ferns
is atypical because one branch continues as the main stem and the other forms a leaf. Because two different organs—
rhizome and leaf—are produced, this is not isotomy in the usual sense. The stellar patterns of the rhizome below
the bifurcations are identical (Gruber, 1981; Imaichi & Nishida, 1973). Therefore, which branch of the bifurcation will
become the leaf, and which the rhizome, is not possible to predict anatomically.

Figure 11. Some rhizome branching types in ferns. A. Hypolepis, epipetiolar. B. Davallia, branch bud on a radius ca. 30
degrees below the leaves. C. Meniscium reticulata, apparently axillary D. Dryopteris, adventitious bud on adaxial surface. E.
Pteris. F. Gleichenia, at right is a branch bud, at left is a leaf primordium). (Modified from Hagemann 1989.)

17
Ferns with non-axillary branching (i.e., most ferns) have leaves and branch buds that occupy different radii on the
rhizome (unlike axillary branching where the bud and subtending leaf are on the same radius). In Davallia (Croxdale,
1976) and Microgramma (Hirsch & Kaplan, 1974), the leaf and branch bud are separated by a divergence angle of
about 30 degrees, with the bud being more ventral. In Equisetum, branch buds are produced between the leaves on
alternate radii. They break through the leaf sheath when they grow out to form conspicuous whorled branches.

Ferns with lateral branching display several kinds of spatial relationships between the leaves and branch buds. These
types have been named for genera that prominently display them (Hagemann, 1988).

The Hypolepis type has branch buds produced along the sides of the petiole bases (epipetiolar buds; Figure 11). This
is sometimes called phyllogenous branching because the branch buds are produced from the leaf. Besides Hypolepis,
epipetiolar buds are present in other dennstaedtioid ferns such as Dennstaedtia, Histiopteris, Paesia, and Pteridium.
The number of epipetiolar buds may vary per petiole (Troop & Mickel, 1968). Hypolepis repens may have up to four
(Gruber, 1981) and H. bogotensis up to six (pers. obs.). The first (proximal) bud on the shoot is always produced on
the side opposite to the continuing main rhizome, and the subsequent buds are produced on alternate sides. The
epipetiolar buds form after the leaves are initiated from the apical meristem (Imaichi & Nishida, 1973). Thus they are
adventitious, not acrogenous.

The Davallia branching type has a bud positioned near the leaf on the same side of the rhizome (Figure 12). The bud
occupies a radius about 30 degrees below (ventral) to the leaf (Croxdale, 1976).

In the Pteris type, the branch bud is produced below (proximal) the leaf base and along (or nearly so) the same radius.
This is characteristic of Pteris, Adiantum, and some species of Blechnum. Ontogenetically, the leaf and branch primordia
originate in pairs and branch (Hagemann, 1988).

Figure 12. Davallia and Polypodium rhizome branching.

18
The Polypodium type is characterized by branch buds alternating with the leaves and on the opposite side of the rhi-
zome. The buds occupy a different radius than the leaves, about 30 degrees lower on the ventral side (Hovenkamp,
1990). Elaphoglossum also has this type of branching, although the buds in most species are suppressed and thus the
rhizomes do not branch. In Elaphoglossum only two rows of buds are produced on the ventro-lateral side of the
rhizome regardless of how many rows of leaves (orthostichies) are present.

Another branching pattern is the Salpichlaena type. The bud and the leaf appear at the same node, but the bud occu-
pies a different radius, usually about 40 degrees to the left of the leaf (as viewed when the rhizome apex is oriented
upward). The vascular traces to the branch bud give rise to roots so that the roots appear to be localized, clumped,
in association with the branch bud. This pattern occurs in Salpichlaena volubilis and Woodwardia areolata (pers. obs.).

The Gleicheniaceae have a distinctive type of rhizome branching. The rhizomes are monostichous, with one rank
(orthostichy) of leaves on the dorsal surface. Branching is both dichotomous and lateral, with the lateral branches
occurring in the same dorsal row as the leaves (Figure 13). The leaves and branch buds might alternate in a regular
sequence, but this needs to be confirmed (Hageman & Shultz, 1978). The apical and dorsal branches have a different
vascular anatomy (Figure 13).

Probably more branching types occur in ferns than listed here. In the past, little attention has been paid to fern rhi-
zomes besides their use in keys (i.e., whether the rhizome is creeping, erect, or decumbent). Any such study should
first examine the external morphology of bud and leaf positional relationships on the rhizome. It should then be
followed with an anatomical study of the vasculature supply to the leaves and buds. The subject of fern branching
patterns is wide-open for original observation.

Figure 13. Rhizome of Gleichenella pectinata showing anatomy of the lateral (top) and apical branches (bottom). The apical
branching is isotomous.

19
Selected references | Branching patterns in Ferns & Lycophytes

Bell, A. D. & P. B. Tomlinson. 1980. Adaptive architecture in rhizomatous plants. Botanical Journal of the Linnean Society 80:
125–160.
Croxdale, J. G. 1976. Origin and early morphogenesis of lateral buds in the fern Davallia. American Journal of Botany 63:
226–238
Gruber, H. 1981. The branching patterns of Hypolepis repens. American Fern Journal 71: 41–47.
Hagemann, W. 1980. Ueber den Verzweigungsvorgang bei Psilotum und Selaginella mit Anmerkungen zum Begriff der Dichot-
omie. Plant Systematics and Evolution 133: 181–197.
_____. 1988. Acrogenous branching in pteridophytes. Pages 245–258. In: K. H. Shing & K. U. Kramer (eds.). Proceedings of the
international symposium on systematic pteridology. China Science and Technology Press, Beijing.
______ & U. Schulz. 1978. Wedelanlegung und Rhizomverzweigung bei einigen Gleicheniaceen. Botanscher Jahrbücher für
Systematik 99: 380–399.
Hebant-Marui, R. 1973. Branching patterns in Trichomanes and Cardiomanes (Hymenophyllaceous ferns). Canadian Journal
of Botany 62: 1336–1343.
Hirsch, A. M. & D. R. Kaplan. 1974. Organography, branching, and the problem of leaf versus bud differentiation in the vining
epiphytic fern genus Microgramma. American Journal of Botany 61: 217–229.
Hovenkamp, P. 1990. The significance of rhizome morphology in the systematics of the polypodiaceous ferns (sensu stricto).
American Fern Journal 80: 33–43.
Imaichi, R. 2008. Meristem organization and shoot diversity. Pages 75–106. In: T. A. Ranker & C. H. Haufler (eds.). Biology and
evolution of ferns and lycophytes. Cambridge University Press, London.
_____ & M. Nishida. 1983. Studies on the extra-axillary buds of Hypolepis punctata. Japanese Journal of Botany 48: 268–279.
Troop, J. E. & J. T. Mickel. 1968. Petiolar shoots in the dennstaedtioid and related ferns. American Fern Journal 58: 64—69.

20
Lycophylls versus Euphylls (or Microphylls versus Megaphylls)

The lycophytes and euphyllophytes (ferns and seed plants) have two fundamentally different types of leaves: lycophylls
and euphylls (sometimes called microphylls and megaphylls, respectively). Lycophylls are characterized by being sim-
ple, entire, and single-veined.They are sessile, not stalked like the leaves of most ferns and dicotyledonous seed plants.
(Although firmly entrenched in botanical literature, the term microphyll is slightly misleading. It means “small leaf,”
and although most are small---usually less than two centimeters long--- those of some quillworts reach up to one
meter long. Similar lengths were attained by the microphylls of tree lycophytes (lepidodendrids) that dominated the
coal-forming swamps of the Carboniferous. The term lycophyll has the advantage of not implying size and referring
to the group it characterizes.)

No one is sure how lycophylls evolved. One theory, the enation theory, claims that they are vascularized enations; that
is, an enation that became supplied with a vein (Figure 14). Another theory is that they represent the transformed
lateral sporangia of some zosterophyll-like ancestor. However they evolved, lycophylls are believed to have had a
completely different evolutionary origin from the type of leaf that characterizes ferns and seed plants, a type of leaf
called a euphyll.

Figure 14. Evolution of a lycophyll according to the enation theory. A. Naked axis of an early vascular plant. B. An enation forms.
C. A vein extends from the central vascular bundle to the base of the enation. D. The vein extends into the enation, forming a
lycophyll.

Euphylls evolved from the three-dimensional, photosynthetic branching systems of early vascular plants by a three-
step process (Figure 15). First, the branching system became flattened into one plane. Second, a green laminar tissue
developed between the flattened branching system, filling in the space between the branches and forming a web-
like lamina. Finally, some of the branches elongated more than other branches, overtopping them and producing a
prominent central branch with subordinate lateral ones.The result was a broad leaf with many veins---the veins being
derived from the vascular tissue of the branches. The sequence of flattening, webbing, and overtopping can be seen
in the fossil record in a series of intermediates. In fact, some fossils are so intermediate that it’s difficult to determine
whether they represent stem or leaf. Botanists are currently debating whether euphylls evolved only once in the
common ancestor of ferns and seed plants or evolved several times independently in those groups. Evidence from
development and genetic pathways evidence suggests more than one origin (Tomescu, 2008).

For the first 40 million years of their existence, land plants got along quite well without euphylls.They were leafless or
with only small spinelike appendages (enations). Euphylls first appeared during the Devonian and became widespread
when the concentration of carbon dioxide in the atmosphere dropped by about 90 percent. This drop might have
fostered the evolution of euphylls, which have the adaptive advantage of presenting a large surface area capable of
efficiently absorbing carbon dioxide (Kenrick, 2001).

21
Figure 15. Evolution of a megaphyll. From left to right: a three-dimensional branching system of an early vascular plant; flattening
of the branching system into one plane; webbing between the branches; overtopping to produce central vein and subordinate
lateral ones. For different plant groups, the actual sequence of steps may have varied from the one shown here. (From Moran,
2004)

Lycophytes and euphyllophytes also show anatomical differences in the stem stele, and these differences relate to
their leaves (Freeberg & Wetmore, 1967). In vascular plants, leaf traces (veins) are not something that are “given off ”
from the vascular bundle of the stem and extend into the leaf. Instead, the procambium of a leaf trace first differen-
tiates in the outer cortex of the stem and then extends acropetally into the leaf primordium and basipetally into the
stele. Lycophytes and euphyllophytes differ in the timing of this differentiation, and the effect on the stele is profound.
In lycophytes, the leaf trace differentiates late in development.There may be up to 10 leaf primordia around the shoot
apex before the lowest primordium is vascularized. When the lycophyll leaf trace finally joins the stele, the tissues of
the stele have completely differentiated so that the joining leaf trace has no effect on their patterning. Similarly, if a leaf
primordium of a lycophyte is experimentally punctured or removed, there is no effect of the shape and patterning of
tissues in the stele. In euphyllophytes, however, the leaf trace differentiates soon after the leaf primordium is formed.
Sometimes the youngest primordium at a shoot apex contains a differentiating procambial strand (Esau, 1965). As the
trace differentiates basipetally it merges with a young stele that consists mostly or entirely of undifferentiated tissue.
This merging trace influences the patterning of the tissues in the stele. It creates a “leaf gap,” or parenchyma-filled
region in the stele above the insertion of the leaf trace. If the leaf primordium of a euphyll is experimentally removed,
the procambial leaf trace does not develop, and a leaf gap does not form. Thus, in lycophytes, the leaf has no effect
on the patterning of the stele, whereas in euphyllophytes it causes a leaf gap. The differences result from the relative
timing in the development of the leaf trace: much latter in lycophytes when the pattern of stele tissues is determined,
and earlier on in euphyllophytes when the pattern of the stele tissues is undetermined.

Lastly, lycophylls and euphylls differ in the location of their sporangia. In lycophylls it is on the adaxial (upper) surface;
in euphylls, it is on the margin or abaxial (lower) surface.

22
Eusporangia versus Leptosporangia

There are two principal types of sporangia in land plants: eusporangia (“true sporangia”) and leptosporangia (“thin
sporangia”). Eusporangia are found in mosses, hornworts, liverworts, lycophytes, seed plants, and several basal families
of ferns (Ophioglossaceae, Psilotaceae, and Marattiaceae). Leptosporangia are found only within the ferns (Osmun-
daceae and above on the phylogeny). On the basis of outgroup comparison, leptosporangia are derived.

The two types of sporangia are distinguished by the number of cells involved in their initiation and the thickness of
the mature sporangium wall (Figure 16). Eusporangia originate by simultaneous divisions of several initial cells on the
surface of the developing leaf. Their divisions are parallel to the leaf surface (periclinal), resulting in an inner and outer
layer of cells. The inner layer forms the sporogenous cells that eventually give rise to the spore mother cells (sporo-
cytes) that divide meiotically to form spore tetrads. The outer layer forms a sporangium wall several cells thick. The
tapetum, or nutritive layer, may be derived from either the inner primary wall layers or the primary sporogenous cells.

In contrast, leptosporangia originate from a single cell on the surface of the developing leaf.This cell divides to form an
inner and outer cell, the latter protruding from the surface and appearing as a slightly raised bump.This outer cell gives

Figure 16. Development of eusporangia. A–E. and leptosporangia. A’–G’. (From Gifford & Foster, The Morphology of Vascular
Plants, 1989).
rise to the sporangium, whereas the inner cell plays no part. The outer cell acts as an apical initial and after dividing
several times forms a basal stalk and distal capsule. The mature wall of the capsule consists of a single layer of cells
within which there is a concentric tapetal layer and a central group of sporogenous cells.

Eusporangia are much larger and more massive than leptosporangia. Accordingly, they produce more spores. Depend-
ing on the genus or species, eusporangiate ferns can produce from 1000 to (in microsporangia of Isoëtes) 1,000,000
spores.The output in leptosporangiate ferns is variable but less than that for eusporangiates, generally being a multiple
of four: 16, 32, 64, 128, 256, and 512. The vast majority of leptosporangiates—the Polypodiales—have 64 spores per
sporangium. The lower spore output of leptosporangia is compensated for by their greater number per leaf.

23
The Osmundaceae and Equisetaceae are intermediate between the eusporangiate and leptosporangiate condition. In
Osmundaceae, the sporangia develop from several initial cells resulting in thicker stalks and greater spore output, but
their walls are only one cell thick. In Equisetaceae, the sporangium develops from a single cell and its wall is only one
cell thick, but thousands of spores are produced per sporangium. Despite these intermediate conditions, it is helpful
to recognize two types of sporangia into which all other ferns can be easily placed.

Spores

All fern spores are small. The largest ones are only about the size of a sand grain or the head of a pin—a distinction
that goes to the female spores of Selaginella and Isoëtes, which measure only 1 mm long. Most spores range from 30
to 50 microns long (a micron is one one-thousandth of a millimeter). Individual spores of this size are extremely hard
to see, but when viewed en masse they appear as a fine powder.

When examined under a microscope, spores can be seen to come in two basic shapes: globose-tetrahedral and
bean-shaped (Figure 17, Figure 20, Figure 21)These shapes result from the different ways the cell walls orient them-
selves during meiosis (Figure 17). After a spore mother cell divides by meiosis, the resulting four spores are stuck to-
gether in a tetrad. They soon separate, but each carries a scar indicating where it was attached to the other spores in
the tetrad (these scars are called “haptotypic markings”). In bean-shaped spores the scar is a short, straight line on the
concave side, and in globose-tetrahedral spores the scar is Y-shaped. Because bean-shaped spores have a single scar
they are called “monolete,” and tetrahedral spores with their three short lines are called “trilete.” The lines represent
a weakness in the cell wall through which the spore contents protrude upon germination.The spores of all ferns bear
haptotypic markings with the sole exception of Equisetum. Its spores are neither monolete nor trilete; they are “alete.”

Figure 17. The types of cell wall orientations during meiosis leading to (left) monolete and (right) trilete spores.

Fern spores have two protective layers around them that account for much of the spore’s external form and beauty.
The inner layer (exospore) is secreted by the living contents of the cell, whereas the outer layer (perispore) is de-
posited from the outside and represents the disintegrated remains of a nutritive layer of cells in the sporangium. In
some ferns the perispore is barely noticeable because it is extremely thin and tightly adheres to the exospore (Figure
20D, trilete). In other ferns the perispore is loosely attached, highly elaborated, and ornamented with folds, knobs, or
spines (Figure 20B, C). Sometimes the perispore is wing-like and perforated, appearing doily-like; in others it is spiny,
resembling a bur.

The spores of hybrids are almost always aborted and appear misshapen, irregular, and blackened (Figure 7 left).This is
extremely helpful in detecting most hybrids. Aborted spores result from the total or partial inability of the chromo-
somes from the two parental species to pair during meiosis (Figure 8).This failure to pair is caused by physiological or
structural differences in the chromosome sets.The inability to pair results in unequal distribution of the chromosomes
to the daughter cells of meiosis, and the spores end up with differing numbers of chromosomes (for example, a hy-
brid Asplenium (x=36) might produce a tetrad of spores have a with one spore containing 25 chromosomes, another
47, another 30, and another 42).

Another use of ferns spores is detecting polyploidy. In general, polyploid ferns have larger spores compared to their
diploid relatives. For instance, spores of the diploid (2x) race of Asplenium trichomanes are generally 27–31 microm-
eters long, whereas those of the tetraploid (4x) race are 38–44 micrometers long (Figure 18).

24
Figure 18. Differences in spore size of diploid and tetraploid Asplenium trichomanes. From Moran (1982).

Most pteridophytes produce only one kind of spore; that is, they are homosporous. Others produce two kinds of
spores, male and female, and are termed heterosporous. Of the approximately 300 genera of ferns and lycophytes,
only seven are heterosporous: Azolla, Isoëtes, Marsilea, Pilularia, Regnellidium, Salvinia, and Selaginella. In these genera
the female spores are usually 10 to 30 times larger than the males. Why the difference?

Female spores are larger because they need to store food for a future embryo. Unlike spores of homosporous ferns,
female spores germinate and develop into gametophytes within the spore wall, not outside of it—a condition called
endosporic (Figure 19). This exposes little surface area so that photosynthesis (if it were present, which it is not)
would be insufficient to manufacture enough food for the gametophyte and, after fertilization, the growing embryo.
Thus, female spores must store food, which necessitates a larger size. The male spores are small because they are
ephemeral.They produce the sperm, liberate it, then die. Unlike the female spores, they have no need for stored food.

Besides size and shape, fern spores differ in color. Most are brown or black, but some are yellow or green. The latter
colors characterize certain groups. For instance, the polypody family (Polypodiaceae) and forked ferns (Gleichenia-
ceae) have yellow spores. These can be seen on the commonly cultivated golden polypody (Phlebodium aureum). Its
spores are bright yellow and impart their color to the entire sorus (the “golden” in its name, however, refers to the
rhizome scales, not the spores).

Green spores contain the photosynthetic pigment chlorophyll. They occur in about 7% of ferns worldwide and are
characteristic of certain groups, such as the Equisetaceae, Grammitidaceae, Hymenophyllaceae, and Osmundaceae.

Figure 19. Endosporic female gametophytes of Selaginella.The filamentous structures are rhizoids. At left, note how the spore
has germinated through the trilete markings and the archegonia are located in the center. At right is a longitudinal section
showing megagametophyte development within the spore wall, and a developing embryo.

25
Figure 20. Trilete spores. A. Pityrogramma trifoliata. B. Lygodium microstachyum. C. Huperzia lucidula. D. Anogramma
leptophylla, left spore shows proximal face, right, distal. E. Anemia intermedia. F. Acrostichum aureum. G. Adiantum pedatum.
H. Eriosorus novogranatensis, left spore shows proximal face, right, distal. I. Selaginella exaltata. Scale bars = 10 micrometers.
Photos by Judith Garrison Hanks and Robbin Moran.

Green spores differ from non-green spores in two ways: they remain viable for shorter periods and they germinate
faster. Typically, green spores remain viable for a few days to a few months, but non-green ones remain so for three
to many years. Furthermore, green spores germinate in one to three days after sowing, but non-green spores take
their time, germinating at a more leisurely pace, usually 10 to 14 days after sowing.These differences result from green
spores remaining metabolically active, unlike brown or black spores that exhibit dormancy. Because they remain ac-
tive, green spores are constantly metabolizing stored food, and when this is gone, they lack energy to germinate. This
accounts for their short viability. Their continuous metabolic activity also means that they can germinate as soon as
conditions are favorable. They don’t waste time breaking dormancy as non-green spores do.

Dormancy, however, has its advantages. It allows spores that have been washed into the darkness of the soil to remain
there, viable, for many years. These spores form a “spore bank,” a storage reserve for a population’s spores. These
spores might one day be brought to the surface by the activities of burrowing animals, or falling trees up-rooted at

26
Figure 21. Monolete spores. A. Actinostachys laevigata. B. Schizaea pusilla. C. Asplenium truncorum. D. Polybotrya osmundacea. E.
Lomariopsis guineensis. F. Unknown dryopteroid; perispore broken and exposing the smooth endospore beneath. Laesura clearly
visible in depression on right side of spore. G. Elaphoglossum setigerum. H. Mickelia lindigii. I. Bolbitis appendiculata. Scale bars =
10 micrometers. Photos by Judith Garrison Hanks and Robbin Moran.

their base, or erosion. Once on the surface and exposed to light, the spores germinate. Light is the necessary trigger.
Thus, spores deposited in the soil bank have the potential to form new plants and contribute to future populations.

Buried spores can also contribute to future populations when induced to germinate in the dark by hormones known
as “antheridiogens.” These are secreted by mature, usually female, prothalli on the soil surface. They diffuse or are
washed into the soil where they stimulate buried spores to germinate and develop precociously into dwarf male
prothalli studded with antheridia (normally, antheridia form on older, well developed prothalli). When there’s enough
water in the soil, the antheridia burst and release their sperm that swim to the archegonia of the female prothalli
lying at the soil surface—the prothalli that released the antheridiogens—and fertilize its egg. Thus cross-fertilization is
achieved, and in this manner buried spores can contribute to a population’s gene pool.

In contrast to the light requirement needed by most fern spores to germinate, the spores of some ferns germinate

27
only in the dark. Examples are the grape ferns (Botrychium), adder’s-tongues (Ophioglossum), and clubmosses (Lycopo-
dium). These spores germinate in the darkness of the soil and develop into prothalli underground. The prothalli are
generally whitish or tan, having no need for chlorophyll, which would be useless without light. Subterranean prothalli
are also fleshy and carrot- or potato-shaped, usually up to half an inch (1.2 cm) long, and embedded in their tissues
is a symbiotic fungus that absorbs nutrients from the soil and translocates these to the plant. Little is known about
subterranean prothalli, which are rarely seen.

All spores—especially those that germinate in darkness underground—contain stored food in the form of high-en-
ergy oils. In fact, the spores of some species contain so much oil that they exhibit unusual properties. The spores of
Lycopodium are so oily that they will ignite. This made them ideal for use in flash powder during the early days of
photography. Their oiliness also repels water, and for this reason early apothecaries kept of jar of Lycopodium spores
to coat pills that they had made by hand to prevent them from sticking. Nowadays Lycopodium spores are used com-
mercially to dust latex surgical gloves and condoms to prevent them from sticking together.

The spores of pteridophytes are not the villains in hay fever. That malady is caused primarily by pollen from wind-pol-
linated flowering plants. Allergies to pollen result from a reaction of your immune system to proteins on the surface
of the pollen grain. These proteins signal the pollen to germinate if they are compatible with proteins on the stigma,
the pollen-receiving tip of the seed-producing organ. If proteins of the pollen and stigma are incompatible, then the
pollen does not germinate or subsequent growth of the pollen tube (which contains the sperm) is irregular or incom-
plete and fertilization does not occur. This protein recognition system prevents cross-fertilization between different
species of flowering plants. In ferns, however, there are no flowers, no stigmas, no surface proteins, and, consequently,
no hay-fever-causing spores.

Selected references | Fern and Lycophyte Spores

Balick, M. J. & J. M. Beitel. 1989. Lycopodium spores used in condom manufacture: associated healthhazards. Economic Botany
43: 373–377.
_____ & _____. 1988. Lycopodium spores found in condom dusting agent. Nature 332: 591.
Barrington, D. S., C. A. Paris, & T. A. Ranker. 1986. Systematic inferences from spore and stomate size in the ferns. Amer-
ican Fern Journal 76: 149–159.
Dyer, A. F. & S. Lindsay. 1992. Soil spore banks of temperate ferns. American Fern Journal 82: 89–12.
Haufler C. H. & C. B. Welling. 1994. Antheridiogen, dark germination, and outcrossing mechanisms in Bommeria (Adiantace-
ae). American Journal of Botany 81: 616–621.
Lloyd R. M. & E. J. Klekowski, Jr. 1970. Spore germination and viability in Pteridophyta: evolutionary significance of chloro-
phyllous spores. Biotropica 2: 129–137.
Moran, R. C. 1982. The Asplenium trichomanes complex in the United States and adjacent Canada. American Fern Journal 72:
5–11.
Raghavan, V. 1992. Germination of fern spores. American Scientist 80: 176485.
Sundue, M., A. Vasco & R. C. Moran. 2011. Cryptochlorophyllous spores in ferns: nongreen spores that contain chlorophyll.
International Journal of Plant Science 172: 1110–1119.
Tryon, A. F. 1985. Spores of myrmecophytic ferns. Proceedings of the Royal Society of Edinburgh, B 86: 105–110.
_____ & B. Lugardon. 1991. Spores of the Pteridophyta. Springer-Verlag, New York. [contains nearly 2800 photomicrographs
of spores]
_____ & R. C. Moran. 1997. The Ferns and Allied Plants of New England. Massachusetts Audubon Society, Lincoln, Massachu-
setts. [contains SEM photomicrographs of New England species]
Tryon, R. M. & A. F. Tryon. 1982. Ferns and Allied Plants, with Special Reference to Tropical America. Springer-Verlag, New York.
Wagner, W. H., Jr. 1974. Structure of spores in relation of fern phylogeny. Annals of the Missouri Botanical Garden 61: 332–353.
_____, F. S. Wagner & W. C. Taylor. 1986. Detecting abortive spores in herbarium specimens of sterile hybrids. American
Fern Journal 76: 129440.
Walker, T. G. 1985. Spore filaments in the antfern Lecanopteris mirabilis—an alternative viewpoint. Proceedings of the Royal
Society of Edinburgh, B 86: 111414.
Windham, M. D., P. G. Wolf & T. A. Ranker. 1986. Factors affecting prolonged spore viability in herbarium collections of three
species of Pellaea. American Fern Journal 76: 141–148.

28
Sterile-fertile leaf dimorphism in ferns

Sterile-fertile leaf dimorphism is generally thought of as narrower and taller fertile leaves compared to the sterile, but
it is much more than that. It is a syndrome of different characters (Table 1).

These character state differences maximize spore dispersal and energy used in construction of the fertile leaf. For
example, the reduced lamina tissue—in extreme forms resulting in a skeletonized lamina—decreases the thickness
of the boundary layer of still air over the lamina, thus promoting faster drying of the sporangia. The longer and more
erect petioles elevate sporangia higher above the ground where air currents are likely to be stronger. Because their

character fertile leaf sterile leaf

Seasonality seasonal and year-round year-round, long-lived
Duration ephemeral, ca. 1-3 months 1 year or more
Orientation erect spreading
Placement on climbing portion of the stem only on both climbing and terrestrial portions
Lamina size smaller larger
Petiole length shorter in climbing species, longer in terrestrial species longer in climbing species, shorter in terrestrial
species
Petiolo color pale reddish darkened or stramineous
Petiole texture fleshy, relatively soft stiff, hard
Lamina skeletonized, reduced to a thin green wing above the expanded
axes
Diplodesmic veins present absent
Mesophyll parenchyma compact, with very few air spaces spongy, with numerous, large air spaces
Segment margins inrolled with exposed sporangia plane

Table 1. Summary of character states differences associated with sterile and fertile leaves of Polybotrya (from Moran 1987).

function is finished once they have shed their spores, fertile leaves tend to be of “cheaper” construction, built with
energetically less expensive tissues such as parenchyma and collenchyma, not sclerenchyma. They need not last as
long as sterile leaves.

Little is known about the seasonality of fertile leaf production in tropical ferns. Of course, it may differ for different
species, with some species showing marked seasonality and others none at all. But for almost all ferns nothing is
known. It takes observation of many plants over a long time to gather enough data to make definitive statements
about seasonality. Given this, little research has been done on the topic. It is a wide-open field for research.

Three terms are sometimes used to distinguish different types of sterile-fertile dimorphy (Figure 22). Where none
occurs—that is, where the sterile and fertile leaves are the same—the condition is said to be monomorphic. Some-
times only a part of the leaf is fertile—the hemidimorphic condition.The extreme is holodimorphic, where the lamina
of the fertile leaf is soriferous throughout, not just in part. Such leaves differ completely from the sterile ones.

A final note: be careful not to use “dimorphy” by itself for what should properly be called “sterile-fertile leaf di-
morphy.” Because sterile-fertile leaf dimorphy is common we tend to think it is the only kind of leaf dimorphy. But
dimorphy may also occur (albeit rarely) among the sterile leaves of a given species. For instance, the non-climbing vs.
climbing leaves of Salpichlaena look completely different (the non-climbing are 1-pinnate, determinate, up to about
half a meter tall, the rachis straight (not twining), and the pinnae 3–5 cm distant, whereas the climbing are two pinnate,
indeterminate, up to 13 m long, the rachis twining, and the pinnae 20–30 cm distant along the rachis).

29
Figure 22. Different types of sterile-fertile leaf dimorphy. The thick dark lines represent fertile pinnae. (Modified from Wagner
& Wagner, 1977).

Selected References | Sterile-fertile Leaf Dimorphy

Britton, M. R. & J. E. Watkins. 2016. The economy of reproduction in dimorphic ferns. Annals of Botany doi: 10.1093/aob/
mcw177
Moran, R. C. 1987. Sterile-fertile leaf dimorphy and evolution of soral types in Polybotrya (Dryopteridaceae). Systematic Botany
12: 617–628.
Nozu, Y. 1968. Studies on the leaves of ferns. I. The dissimilarities of the fertile and sterile fronds in some ferns. Journal of the
Faculty of Science, University of Tokyo, sec. 3, 10: 13–27, pl. 2.
Wagner, W. H., Jr. & F. S. Wagner. 1977. Fertile-sterile leaf dimorphy in ferns. Gardens’ Bulletin Straits Settlements 30: 251–267.

30
Tips on Collecting Ferns

Techniques for collecting ferns differ little from those for seed plants, but a few points should be kept in mind. The
most important is to collect the rhizome. This structure bears two characters helpful in distinguishing taxa. The first
is its habit, whether erect, decumbent, or creeping, and if the latter, whether short- or long-creeping. Rhizome habit
should always be recorded in field notes and stated on the herbarium label. The second is the kind of rhizome indu-
ment, whether of hairs or scales, and the particular characteristics of these, such as color, habit, margin, and kind of
attachment. It is often impossible to identify a fern to species using a key if rhizome characters are absent.

If a rhizome is too large or bulky to be conveniently mounted on a herbarium sheet, it should be cut longitudinally
through the middle. Besides reducing bulk, this exposes the scales at the apex, making them easier to observe. If still
too thick, the rhizome can be cut longitudinally again. A quartered rhizome is better than none at all.

In general, collect specimens with fertile leaves. Several characteristics of the sori are helpful in identification such as
their shape and position and the presence or absence of an indusium. Also, the fertile leaves of some ferns are dimor-
phic; that is, they have a different form and appearance than the sterile ones. In such cases, it is important to collect
both fertile and sterile leaves. When the fertile leaves are mounted on a herbarium sheet, be sure that the side bearing
the sori is displayed up. Actually, both sides of the leaf can be important in identification. Sometimes it’s necessary
to see the upper (adaxial) surface to check for the presence or absence of hairs or the structure of the rachis-costa
juncture. If possible, collect a fragment that can be mounted to show the upper surface.

Although fertile leaves enhance the specimen’s value, some ferns can be identified to species without them. This is
the case with Elaphoglossum, one of the largest fern genera. Its taxonomy is based largely on rhizome habit, leaf shape,
and scale characteristics. Therefore sterile collections make acceptable specimens. Of course, the problem is that you
have to know before-hand which fern genera can be collected in sterile condition. It is probably best to collect a fern
you are uncertain about rather than pass it by because sori are absent.

Collecting tree ferns and other big ferns (i.e., ones with leaves over two meters tall) presents a problem. Pressing the
complete leaf is impractical and time-consuming, and the redundancy of parts (pinnae, pinnules) provides little extra
information for botanists studying the specimen. How much of the leaf should be collected? For large ferns, collect the
following three parts: 1) the petiole base and basal pinnae, 2) the medial pinnae, and 3) the apex. Each part should
go into a separate newspaper, and the newspaper should be labeled that they are from the same leaf. These three
parts provide different information about the plant. The petiole base often bears scales or hairs or spines for identi-
fying species, and the basal pinnae, by their size, give an idea about how much reduced the lamina is toward the base.
The medial pinnae give an idea of the degree of cutting and how wide the blade is. The apical portion is important
because it can either be gradually tapered to the tip, abruptly tapered, or “conform,” that is, having more or less the
same shape as the lateral pinnae. Be sure to get all three parts. Also, take notes on the length and width of the leaf,
diameter of the trunk, and any other pertinent observations that cannot be seen on the collected material.The notes
should be added to the herbarium label.

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 Overview of land plant phylogeny
Where do lycophytes and ferns “fit” on the tree of life? Among the Eukaryotes, they belong to a branch that includes
all the green algae and land plants—a branch often recognized as Kingdom Plantae. This branch is characterized by
cellulose as the principal constituent of the cell walls, chlorophyll a and b as the main photosynthetic pigments, and
starch as the main food-storage product.

Within Kingdom Plantae there is a branch that includes only the land plants (i.e., green algae are excluded), and this
branch is called the embryophytes (Figure 23). Their distinctive characteristics involve adaptations to the dry environ-
ment of the land (that is, drier compared to the aquatic habitat of their green algal ancestors), such as diplobiontic
alternation of generations, desiccation-resistant haploid spores, cuticles, sporangia, and distinctive male and female
gametangia (antheridia and archegonia, respectively), certain ultrastructural details of male gametogenesis, and unique
type of cell division. The groups that compose the embryophytes are the bryophytes (mosses, hornworts, and liver-
worts), lycophytes, ferns, and seed plants (gymnosperms and angiosperms).

Figure 23. Phylogenetic relationships of the major groups of land plants. Modified from Schneider et al. (2002).

32
Within the embryophytes, the first three basal branches are the bryophytes (Figure 23).They are a paraphyletic group
because they exclude some descendents of their common ancestor (i.e., the rest of the land plants). As a group the
bryophytes can are defined only by what they lack (i.e., lignified vascular tissue) and by certain ancestral characteristics
that do not indicate relationships, such as the dominant gametophyte generation and unbranched sporophytes with
a single terminal sporangium.

Sister to bryophytes are the tracheophytes (Figure 23). As their name suggests, they have well developed conduct-
ing tissue with lignified tracheids in their stems. Unlike the bryophytes, their sporophyte generation is dominant,
and it is also branched and thus capable of producing several to many sporangia per sporophyte (not one as in the
bryophytes). For this reason the group is sometimes called the polypsporangiophytes—a term nearly synonymous
with tracheophytes (some Silurian and early Devonian polysporangiophytes lacked lignified conducting tissue, and
therefore cannot be called Tracheophytes. But if only the present-day plants are referred to, the two terms are syn-
onymous). The three major clades of tracheophytes are the lycophytes, ferns, and seed plants.

The basal group of tracheophytes is the lycophytes, which includes three extant families: Lycopodiaceae, Selaginellace-
ae, and Isoëtaceae (there are also fossil families).These possess the morphological synapomorphies of single reniform
adaxial sporangia, microphyllous leaves, and exarch protoxylem (Figure 24). Their monophyly is also supported by
molecular studies.

Sister to the lycophytes are the ferns and seed plants, collectively called the euphyllophytes (Figure 23). This relation-
ship is strongly supported by DNA sequences and genomic structural data. Ferns and seed plants share a 30,000
base-pair inversion in the chloroplast genome (Raubeson & Jansen, 1992; Pryer et al. 2001a). Morphologically, they
have multiflagellate sperm, although the latter is not applicable to the angiosperms with their highly reduced male
gametophytes (all lycophytes have biflagellate sperm except one species: Phylloglossum drummondii which as 16
flagellae). The ferns are now known to include the horsetails (Equisetum) and whisk ferns (Psilotaceae), which were
previously considered “fern allies” (Pryer et al., 2001b; Wikström & Pryer 2005).

The ferns are characterized by leaves of megaphyllous origin (fronds or “pteridophylls”) with circinate vernation.
They are also characterized by septate rhizoids on the gametophytes and spores with a three-layered exospore.
Seed plants also have megaphyllous leaves, but these probably evolved independently from those of ferns (Kenrick
& Crane, 1997). Of course, seed plants are also characterized by seeds (this involves several characters) and bifacial
cambia that produce wood and bark. Traditionally they are divided into angiosperms and gymnosperms. The latter
group, defined by “naked seeds,” might be paraphyletic. The angiosperms are characterized by flowers and by seeds
enclosed in a carpel, and all analyses show that they are monophyletic. The monocots form a monophyletic group
nested within the dicots.

Roots apparently evolved separately in not only evolved separately in lycophytes and euphyllophyts., but also more
than once in the lycophytes (Fujinami et al., 20017).

33
 Lycophytes
The lycophytes are the sister group to all other vascular plants (Figure 23). They are considered monophyletic based
on their microphyllous leaves, single adaxial reniform sporangia, and exarch protoxylem. Their monophyly is also sup-
ported by sequence data from rbcL (Korall et al. 1999; Manhart, 1994; Wikström & Kenrick, 2001), atpB (Wolf, 1997),
and mitochondrial nad5 (Vangerow et al. 1999).

Microphyllous leaves, which characterize the lycophytes, are simple, entire (rarely denticulate), and with one exception
single-veined (Wagner et al., 1982). Such leaves might have evolved by an enation becoming supplied by a vein (Figure
14), but there are two other hypotheses (see Kenrick & Crane, 1997). The fossil record shows intermediate forms
where a vein runs toward, but not into, the enation (e.g., Asteroxylon, from the Devonian). The sporangia are borne in
the axils of the leaves or on their adaxial surface at the base—in contrast to ferns and basal gymnosperms where the
sporangia are on the margin or lower surface of the leaves. It is uncertain how the sporangia of zosterophylls (fossil
precursors of modern lycophyte lineages) became developmentally associated with the leaf base (Kenrick & Crane,
1997).

Figure 24. Phylogeny of the three extant families of Lycophytes.

The lycophytes arose during the early Devonian (Figure 23) and have been distinct from all other vascular plants ever
since. They were of much greater historical importance than is suggested by their present-day diversity. Nowadays ly-
cophytes harbor less than 0.4 % of the world’s vascular plant species, but during the Carboniferous they accounted for
about 50% and were conspicuous part of the land flora. Extinction has removed several major groups and reduced
the rhizomorphic clade (the group containing the arborescent taxa that dominated the coal swamps of the Carbon-
iferous) to its sole, small, representative of today: Isoëtes. Selaginella boasts the longest fossil history of any extant
genus of plants, a distinction it receives from Selaginella fraipontii, an extinct species known from the Carboniferous.

The lycophytes contains three present-day families: the Lycopodiaceae, Selaginellaceae, and Isoëtaceae (Figure 24).
The Lycopodiaceae is sister to the other two families that are defined by ligulate leaves, heterospory, and roots con-
taining a central air canal and eccentric stele (Stevenson & Loconte, 1996). DNA sequence data also support this
relationship (Wikström et al., 1999). The ligule, which comes from the Latin word ligula, meaning “little tongue,” is of
unknown function. It is initiated precociously on the adaxial surface of the leaf primordia and matures long before its
associated leaf. Its stains darkly with proteophilic stains and secrete mucilage, but the function of the mucilage, if any,
is unknown (Horner et al., 1975). Korall et al. (1999) estimated that the divergence between the Selaginellaceae and
the Isoëtaceae may have occurred by the upper Devonian (370 Ma). A symposium volume on the Lycopodiopsida
contains much information on the evolution, taxonomy, and morphology of the living and fossil members (DiMichele
& Skog, 1992).

34
1. Lycopodiaceae Mirbel | Club Moss Family

Plants terrestrial, epiphytic, or rarely rupestral; habit erect or pendulous. Stems dichotomously branched, occasionally
with lateral branching, plectostelic. Leaves simple, entire or rarely denticulate, one-veined, homophyllous (leaves all
alike) or anisophyllous (with reduced leaves, usually spore-bearing, in the terminal divisions); ligules absent. Sporophylls
dimorphic or not, usually aggregated in terminal clusters (strobili). Sporangia solitary in the leaf axils or on the upper
side of the leaf, reniform or nearly globose, 1-locular, short-stalked, dehiscing over the top and dividing the sporangium
into two clam-like valves. Spores trilete, tetrahedral-globose, of one kind (homosporous). Gametophytes epigeal and
green (Lycopodielloideae) or subterranean, mycotrophic, and non-green (Huperziodeae and Lycopodioideae). Chromo-
some numbers varies, but is constant with a given genus. The ancestral base number was probably x=11 (Wagner,
1992).

Type species: Lycopodium clavatum L., one of the most widespread pteridophytes in the world.

Distribution and ecology: Cosmopolitan, from Arctic islands to the Antarctic, from sea level to snow line. Habitats
extremely diverse. About half (185) of the species occur in the American tropics (Øllgaard, 1979a, 1992, 1995). Slight-
ly more than half of the Huperzioideae, or about half of Phlegmariurus species, are epiphytes (Beitel, 1979), whereas
Lycopodioideae and Lycopodiellioideae (except for a single epiphytic species of Palhinhaea – P. steyermarkii) are terres-
trial. The gametophytes of Huperzia and Lycopodium are subterranean, acholorophylous, and mycotrophic, but those
of Lycopodiella and Phylloglossum are green and epigeal.

Pseudolycopodiella caroliniana and Diphasiastrum complantatum are extremely widespread species. Huperzia is primar-
ily north-temperate and circumboreal. Phylloglossum is endemic to Australia, Tasmania, and New Zealand. (Wikström
& Kenrick, 2001,Testo et al., 2018) Phlegmariurus, excluding the H. selago group (see below), contains two large clades,
one neotropical, the other paleotropical. These clades are thought to reflect a vicariant pattern from the final break-
up of South American and Africa about 95 My. .

Genera/species: 16/388. Subfam. Lycopodielloideae: Lateristachys (4), Lycopodiella (15), Palhinhaea (25), Pseudo-
lycopodiella (10); Subfam. Lycopodioideae: Austrolycopodium (8), Dendrolycopodium (4), Diphasiastrum (20); Dipha-
sium (5), Lycopodiastrum (1), Lycopodium (15), Pseudodiphasium (1), Spinulum (3); Subfam. Huperzioideae: Huperzia
(25), Phlegmariurus (250), Phylloglossum (1)

Economic plants and products: The spores of Lycopodium contain volatile oils and are highly flammable.They were
used in flash powder during the early days of photography. Nowadays Lycopodium spores are used to dust latex
surgical gloves).

Discussion: Recondite but helpful taxonomic characters for the subfamilies are shape of the sporangial epidermis
cells (Øllgaard, 1975), and gametophyte morphology (Bruce 1976b).

Unlike ferns, the Lycopodiaceae exhitits complete absence of apogamy and a paucity of alloploidy. Allohomoploidy
is known in Diphasiastrum and Lycopodiella (Wagner, 1992). Allopolyploidy has been demonstrated or the origin of
Huperzia porophila (Waterway, 1986) and Diphasiastrum wightianum and D. zanclophyllum (Aagaard et al., 2009). It also
occurs in Lycopodiella subappressa and L. margueritae, although the parentage of these taxa have not been confirmed.

Huperzia is the only genus in the family in which abortive spores are regularly produced. Hybrids in Lycopodiella and
Diphasiastrum have normal chromosome pairing and produce apparently normal (not misshapen), viable spores. In
one case (D. X habereri), the spores have been germinated and grown into gametophytes that produce normal ar-
chegonia and antheridia (Whittier & Britton, 1995). One hybrid, D. X sabinifolium, hybridizes with D. digitatum, showing
that the hybrids produces viable sperm (Gilman, 1994).

35
The Lycopodioideae consist of about 40 species worldwide. The subfamily is diverse vegetatively, with some species
short and tree-like (Dendrolycopodium), some vine-like (Pseudodiphasium), and others typically with flattened branches
(Diphasium, Diphasiastrum). The latter group was monographed by Wilce (1965) as Lycopodium sect. Complanata. She
recognized only one species in this group in the Neotropics: D. thyoides.

Lycopodielloideae has about 54 species. All have green surficial gametophytes, rugulose spores, erect stems produced
dorsal (not dorso-laterally) from the main horizontal rhizome, sporangial epidermal cell walls straight (not sinuate)
and non-lignified, sporangia borne on the sporophyll stalk (not in the axil), and mucilage canals along the veins. Four
genera are recognized in the subfamily: Lateristachys, Lycopodiella, Palhinhaea, and Pseudolycopodiella.

Figure 25. Relationships of the three subfamilies of Lycopodiaceae. .

Figure 26. Growth patterns in four genera of Lycopodiaceae subfam. Lycopodioideae. All have an indeterminate main horizonal
shoots that branch anisodichotomously and alternately to produce short, determinate, erect shoots.This pattern differs from than
that found in subfamilies Huperzioideae (isodichotomously branched shoots) and Lycopodielloideae (anisotomously branched
shoots with the erect shoot arising from the dorsal side of the horizontal shoot (from Zier et al., 2015).

36
Figure 27. Lycopodiaceae, Huperzioid clade A. Huperzia selago, 1) Habit and bulbil. 2) Subterranean gametophyte. B.
Phlegmariurus pithyoides, stem cross-section, with stele in the center surrounded by many roots growing down through
the cortex (cortical roots). C. Phlegmariurus linifolius. D. Phlegmariurus phyllicifolius, modified sporophylls and sporangia.
E. Phlegmariurus varius, stem cross-section. F. Phlegmariurus squarrosus, stem cross-section. G. Phlegmariurus eversus,
unmodified sporophylls and sporangia..

37
Figure 28. Lycopodiaceae. A. Lycopodium clavatum, habit. 1) Sporophyll with sporangium. 2) Spores. 3) Gametophyte. 4)
Longitudinal section of gametophyte showing (k) young sporophyte. 5) Antheridia. 6) Archegonia. 7) Biflagellate sperm. 8) Embryo
with suspensor. B. Diphasiastrum complanatum, gametophyte. C. Pseudodiphasiastrum volubile, habit. 1) Anisophyllous
branchlet. 2) Strobilus. D. Diphasiastrum tristachyum, cross-section of stem showing plectostele.

38
2. Isoëtaceae Reichenbach | Quillwort Family

Plants generally less than 30 cm tall, tufted, evergreen aquatics or ephemeral terrestrials. Stems usually globose,
fleshy, 2- or 3-lobed, apparently protostelic. Roots thick, fleshy, with a single central air chamber and eccentric ste-
le, emerging from grooves between the lobes of the stem. Leaves linear, subulate, one-veined, terete or slightly
flattened adaxially, often winged toward the base, monomorphic, containing four air chambers, the air chambers
septate; ligules adaxial, immediately distal to the sporangia, ephemeral; sporangia single, usually ovoid, embedded in
the leaf base, lacking an annulus (indehiscent), usually covered to various degrees by a hyaline membrane
(the velum), with internal incomplete walls (trabeculae), of two kinds: megasporangia and microsporangia;
megaspores tetrahedral-globose, trilete, often with a prominent equatorial ridge and three radial ridges converging at
the pole on the proximal face, smooth or highly ornamented, several hundred per sporangium; microspores ellipsoid,
monolete, the surface diversely ornamented, several thousand per sporangium. Gametophytes endosporic. x=11.

Type species: Isoëtes lacustris L., a circumboreal species.

Distribution and ecology: Nearly worldwide. Typically submerged or emergent aquatics. Others are temporary
aquatics that go into dormancy as their habitats dry. A few are seasonal terrestrials, actively growing in the spring.

Genera/species: 1/150. Genera: Isoëtes (150 spp.).

Economic plants and products: none.

Discussion: The monophyly of the family is well supported by DNA sequence data and the following morphological
synapomorphies: reduced axial growth, lacunate leaves, labia, vela, sporangia sunken and indehiscent, trabeculae, and
multiflagellate sperm. Its sister group relationship to the Selaginellaceae is supported by ligules, heterospory, and DNA
sequence data. Rydin and Wikström (2002) used rbcL to study the phylogeny of Isoëtes, but resolution was in general
poor. No satisfactory infrageneric taxonomy has been proposed for the genus. Some infrageneric classifications em-
phasize spore sculpturing, number of lobes on the stem, and habitat, but these characteristics have apparently been
subject to frequent parallelisms and reversals (Taylor & Hickey, 1992).

Isoëtes can be distinguished from all other vascular plants by the four air chambers in its leaves.

Isoëtes is characterized by a unique “corm” (stem and rooting structure) that consists of two halves (Figure 31A1).
The upper half bears a tuft of leaves, and the lower half (called the “rhizomorph”) bears roots. Each half is produced
by growth from an apical meristem, and because two meristems are involved and leading to growth in opposite
directions, the stem has bipolar growth. In contrast, the stems of all other extant ferns and lycophytes that have uni-
polar growth; that is, they grow in one direction from the action of a single apical meristem. Also, all other ferns and
lycophytes produce adventitious roots, ones not produced from an apical meristem as in Isoëtes. The stem of Isoëtes
has secondary growth and a highly unusual anatomy. The roots are unique among extant plants by having a definite
arrangement on the stem, or rhizotaxy (in all other plants the roots are produced irregularly on the stem, not in a
definite sequence). The rhizomorph is believed to be homologous with the dichotomously forked, underground axes
of the Carboniferous tree lycopsids (Stigmarian axes). Many fossil groups of the late Paleozoic and Mesozoic show a
rhizomorph-like structure, and these plus Isoëtes are referred to as the “rhizomorphic clade.”

Covering the adaxial wall of the sporangium of Isoëtes is a thin flap of tissue called the “velum” (Figure 31 A2). The
extent of its coverage has diagnostic value. It is usually durable and can be examined by lifting it with forceps off the
sporangium wall.

All leaves of Isoëtes bear sporangia, and the micro- and megasporophylls are alike. This contrasts with seed plants
where the two kinds of leaves are distinct and the difference can be detected early in their development. The me-
gasporophylls occur at the periphery of the rosette, and the microphylls on the inside. The sporangia of Isoëtes are
the largest of any extant plant, with some reaching up to one centimeter long.They lack a dehiscence mechanism and
apparently release the spores as they rot. Up to 300 spores have been counted in megasporangia, whereas 150,000

39
to 300,000 are reported for some microsporangia. Within the sporangium, these spores are separated by incomplete
partitions called “trabeculae” which are a structure unique to the family.

Another synapomorphy for Isoëtes is multiflagellate sperm, a character that evolved independently in euphyllophytes.
The outgroups of Isoëtes (hornworts, liverworts, mosses, club mosses, and spike mosses) all have biflagellate sperm.

Hybridization and polyploidy are extensive in the north-temperate species of Isoëtes, especially in those of aquatic
habitats (Taylor & Hickey, 1992). These phenomena have not been investigated in the tropical species of Isoëtes. Hy-
brids can be distinguished by malformed megaspores that are usually flattened, irregular, or dumbell-shaped.

Many species of Isoëtes grow in oligotrophic lakes and display two unusual physiological adaptations for life in these
environments (Moran, 2004). One is Crassulacean Acid Metabolism (CAM), a method of carbon assimilation usually
found in desert plants. CAM allows carbon dioxide to be taken up during the night and stored as malic acid in the
vacuoles of the cells. During the day, the malic acid is decarboxylated and the carbon dioxide is shunted into the nor-
mal photosynthetic pathway. This allows the plant to take up carbon all day long instead of only during the daytime.
The other adaptation is that the roots are capable of absorbing carbon dioxide the lake sediments where this nutrient
is in greatest supply because of microbial activity (most other aquatic plants absorb carbon dioxide from the water).
Both CAM and the uptake of carbon by the roots are distinct advantages in oligotrophic lakes where carbon is the
main limiting nutrient.

Stylites has sometimes segregated from Isoëtes; however, it is now known to be nested within Isoëtes and therefore
no longer recognized (Larsén & Rydin, 2015).The two species previously recognized as “Stylites” grow in the Andes of
Peru and Bolivia around the margins of alpine pools on decaying mounds of Distichia. Their stems are elongate, erect,
and often dichotomously branched. Their roots arise from a single lateral furrow. The leaves are flat, rigid, thick-cuti-
cled, and lack stomata.The sporangia are nearly superficial and often borne several centimeters above the base of the
leaf—not deeply sunken and at the base as in other species of Isoëtes. The plants photosynthesize via CAM.

Figure 29. . Isoëtaceae, Isoëtes. A. I. lacustris, habit. 1) Longitudinal section of corm. 2) Sporophyll, showing ligule (L), vellum
(V), and sporangium (S). B. I. japonica, cross section of root, showing central air canal and excentric stele.

40
3. Selaginellaceae Willkomm | Spike-moss Family

Plants terrestrial, on rocks, or rarely epiphytic. Stems creeping or erect, dichotomous, branched regularly or irregularly,
protostelic, siphonostelic or actino-plectostelic, the stele suspended in a central cavity. Rhizophores (modified leaf-
less shoots producing roots) present or rarely absent, produced at the branch fork, usually whitish and bearing branched
roots at the base. Leaves usually less than 10 mm long, one-veined, on a single plant either monomorphic and spirally
arranged or (more commonly) 4-ranked and anisophyllous; ligules present (but evanescent and hard to see) on the
adaxial surface of the leaf base. Stomata anomocytic. Strobili (clusters of overlapping sporophylls) compact, cylindri-
cal or quadrangular or flattened, usually borne at the branch tips. Sporophylls monomorphic or dimorphic. Sporangia
short-stalked, solitary in the axil of sporophylls, opening by a slit across the top; microsporangia globose. Spores
of two types (heterosporous), trilete, megaspores 4 per sporangium (rarely 1 or more than 4), mostly 200—100
micrometers; microspores numerous per sporangium, mostly 2—30 micrometers. Gametophytes endosporic.

Type species: Selaginella selaginoides (L.) Link (basionym: Lycopodium selaginoides L.), of northern North America,
Eurasia, nw. Africa, and Canary Islands.

Distribution and ecology: Cosmopolitan, primarily tropical. In many habitats.

Genera/species: 1/800. Genus: Selaginella.

Economic plants and products: Several species are cultivated as ornamentals. Selaginella kraussiana is perhaps the
most widely cultivated and has several attractively colored cultivars, including variegated forms. Selaginella willdenovii
and S. uncinata are prized for their blue-iridescent leaves, a characteristic best developed when the plants are grown
in shade. Various species known as “resurrection plants,” are sold as novelty items. They curl into a dull-brown ball
when dry and expand into a bright green rosette when wet. Selaginella erythropus is often cultivated because of the
attractive red lower surfaces of its branchlets.

Discussion: The Selaginellaceae are a well-supported monophyletic group based on morphology (Kenrick & Crane,
1997; Stevenson & Loconte, 1996) and both nuclear and chloroplast sequence data (Korall & Kenrick, 2002; Korall et
al., 1999; Zhou & Zhang, 2015). Morphological synapomorphies are stem steles suspended by endoderm cells (tra-
beculae) in a cavity (Figure 31E), epidermal cells with a reduced number of chloroplasts (often only one), the presence
of strobili, megaspores reduced to 4 per sporangium (sometimes less), and microsporangia globose.

Many species of Selaginella have a single chloroplast at the bottom of their dorsal-facing epidermal cells (Jagels
1970a,b). These cells have convex outer walls that refract light toward the bottom of the cell, thus increasing the
likelihood of the chloroplast capturing light. In one such species, S. erythropus, the chloroplast of the dorsal-facing
epidermal cells are differentiated into two zones that differ in the density and placement of the granae and thyka-
loids. This type of chloroplast is termed a bizonoplast (Sheue et al., 2007). The mesophyll and ventral-facing cells of S.
erythropus, however, contain normal discoid chloroplasts. The systematic occurrence of bizonoplasts in Selaginella has
yet to be determined.

The following discussion of Selaginella phylogeny is based on Zhou & Zhang (2015). They found that subgen. Selag-
inella is sister to the rest of the genus (Figure 32). The subgenus consists of only two species: S. selaginoides (Figure
31B), cicrumboreal and type of the genus, and S. deflexa, endemic to Hawaii.These two species have helically arranged
monomorphic trophophylls and sporophylls, unlike most other species the genus which have four-ranked and dimor-
phic leaves. Rhizophoes are absent, unlike the rest of the genus where they are present. The roots, which number less
than 8, are produced from a thickened "hypocotyl" the region below the first two sporeling leaves; the roots are not
produced elsewhere along the stems (Karrfalt, 1981).The dichotomies of the stem are unequal, with the larger branch
erect, thicker, and determinate. The axis of the strobilus elongates before spore dispersal—a unique character in the
genus. The type of megaspore dispersal mechanism is also unique and has been termed “compression and slingshot
megaspore ejection” (Page, 1989). The Carboniferous fossil S. fraipontii resembles this subgenus.

41
Figure 30. Typical strobilus of Selaginella sect. Heterostachys (S. chrysocaulos) The strobilus is dorsiventrally flattened, and
the size of the leaves is reversed from that of the vegetative shoot, with the larger leaves on the upper surface (A, C) and
the smaller ones on the ventral (B, C). The larger upper sporophylls (D) have winglike extension (w) that resembles basal half
of the leaf, but the true basal leaf half is represented by a small, ciliate flap (bh). A. Strobilus, dorsal view. B. Strobilus, ventral
view. C. Strobilus cross section. D. Sporophyll. In B, D is the dorsal vegetative leaf, V is the ventral vegetative leaf, Sp is the
sporophyll on the ventral side of the strobilus (Modified from Goebel, 1905).

Sister to subgen. Selaginella is the “rhizophore clade” consisting of all other species in the genus. The rhizophore is
a unique structure produced at the branch forks by a meristem in the bifurcation, called an “angle meristem.” The
rhizophore grows downward and branches dichotomously when it touches ground, producing all the roots. Having
characteristics of both root and stem, the rhizophore and its homology with those organs has been much debated
(see Bierhorst (1971) for a review).

The next diverging clade is subgen. Boreoselaginella, consisting of about five species in the Old World. The sterile
leaves in this group are monomorphic and four-ranked. The megaspore have densely contiguous tubercles, and mi-
crospore are rugulate and tuberculate. The rhizophore is dorsal. Examples of species belonging to this clade are S.
nummularifolia, S. rossii, S. sanguinolenta, and S. yemensis.

The next clade is Selaginella subg. Ericetorum. It consists of about 100 species and is pantropical, being most diverse
in the Neotropics. It is characterized by dorsal rhizophores restricted to the base of main stem, sterile leaves mono-
morphic and decussately arranged (at least in lower portion of plants), and reticulate megaspores with wing-like and
highly convoluted laesurae.

Subgenus Tetragonostachys consists of 40 species that typically grow in dry habitats or deserts. Nearly all are New
World. Their sterile leaves are helically arranged, same-shaped (isophyllous), thickly cutinized, and filiform-tipped. A
groove runs longitudinally on the abaxial surface of the leaves, and within it are localized the stomata.The rhizophores
occur between branches along the stem, in contrast to the other subgenera where they occur at the branch forks.

42
Figure 31. Selaginellaceae. A1-5. Selaginella lepidophylla. A. Habit when wet. A1. Habit when dry. A2. Heterophylly. A3.
Microsporangium. A4. Megasporangium. A5. Strobilus. B. S. sellaginoides, habit. B1. Microsporophyll. B2. Megasporophyll. C. S.
umbrosa, habit. C1. Strobilus and branch tip, seen from above. C2. Branch tip seen from below. D. S. lyallii, longitudinal section of
leaf base with ligule (li) (st is stem epidermis). E. S. kraussiana. E1. Stem with steles. E2. Stem cross-section showing suspended
vascular bundles in central cavity..

43
The plants are low-growing and often mat-forming. In general appearance this is the easiest subgenus to recognize.
Because of its distinctness, it is sometimes elevated to generic rank as Bryodesma. The group is most diverse in arid
regions of Mexico and the southwestern United States. It was monographed by Tryon (1955).

Subgenus Ericetorum consists of three species: two in the proteaceous heathlands of Australia on one in southern
Africa. Their monophyly is supported by the synapomorphy of decussate phyllotaxy.

Subgenus Articulatae consists of about 40 species, nearly all of which are neotropical. The group takes its name from
the articulations immediately below the branch forks.The articulations take on two forms, one is a swelling (collapsing
and darkening upon drying) and other is as swollen joints often with a dark line running through the middle.The artic-
ulations are nonfunctional; the stems do not break cleanly at these points. Other synapomorphies that unite the group
are: 1) each strobilus has a single (rarely 2) megasporangium at its base (this occurs in other species of Selaginella and
might be a synapomorphy at a more inclusive level or have evolved more than once; Korall & Kenrick, 2002); 2) each
megasporophyll is enlarged relative to the microsporophylls and has one or two enlarged sterile leaves subtending
it; 3) the megaspores are exceptionally large and have high reticulate crests; 4) the microspores are echinate and
dark yellowish or colored; 5) the microsporangia open by a distinctive type of annulus and are ejected from the leaf
axil during dehiscence (Koller & Scheckler, 1986; Somers, 1982). These characteristics make this subgenus one of the
easiest of the anisophyllous groups to recognize.
Subgenus Heterostachys is pantropical and contains about 150 species (Valdespino, 1995). Its synapomorphies are
dorsiventrally flattened strobili and anisophyllous sporophylls (Figure 30A). The anisophylly of the strobili is the re-
verse of the vegetative branches; that is, the dorsal sporophylls are larger and laterally spreading, whereas the ventral
ones are smaller and ascending (only two cases are known where the anisophylly follows that of the vegetative
branches; Goebel, 1905). This reversal in dorsi-ventrality has lead to the strobili being described as resupinate, but
there is no twisting of the stem, and thus no resupination. The sporophylls on the upper side of the strobili bear an-
other synapomorphy: a prominent wing-like extension from the upper surface.The wing partially enfolds and protects
the sporangium and lower sporophylls (Figure 30A, D).This wing is sometimes termed a sporophyll-pteryx (Quansah
& Thomas, 1985). It looks like half of the sporophyll, but the true basiscopic half of the sporophyll is represented by
a narrow, less prominent wing. Marginal cilia are borne along this less prominent “wing”, thus indicating its homology
with the basiscopic half of the leaf (Figure 30A, d). The large wing extension does not bear cilia, indicating it is a new
structure not homologous with the basiscopic half of the leaf. Many neotropical species in this subgenus have minutely
pubescent leaves (Valdespino, 1995), but it is unknown whether this is a snynapomorphy. Molecular results suggest
the subgenus is polyphyletic (Figure 32).The species in Central and South America were monographed by Valdespino
(1995).
Subgenus Stachygynandrum is by far the largest subgroup of Selaginella, containing about 300 species. Its shoot system
is usually branched in one plane to form a pinnate-like arrangement, and the whole shoot system resembles a frond.
The sporophylls are uniform and 4-ranked, often forming quadrangular strobili. Molecular results suggest the subgenus
is polyphyletic (Figure 32).
Selaginella has the oldest fossil history of any extant genus, extening back to the Late Paleozoic (Thomas, 1992, 1997).
The earliest fossil is Selaginellites resimus, a herbaceous, isophyllous plant from the early Carboniferous (Mississippian,
ca. 345 Ma). Anisophyllous species are also known from the Late Carboniferous (310 Ma).
The rhizophore, a special root-bearing organ, is unique to the Selaginellaceae. It usually arises at the branch forks,
although in subgenus Tetragonostachys it is scattered between the branches. It grows toward the soil, occasionally
branching dichotomously along the way, and its aerial parts are long, thin, whitish, and cutinized. Upon reaching the
soil, it branches profusely (again by dichotomies) and gives rise to roots of endogenous origin. Its stelar anatomy
resembles that of the roots of the Isoëtaceae, Lepidodendraceae (an extinct fossil group), and most Lycopodiaceae
(Bierhorst, 1971).
The arrangement of mega- and microsporangia in the strobilus has been studied (Horner & Arnott, 1963), but there
seems to be little correlation with the current taxonomic subgroups.
In Selaginella martensii, a apecies with anisophyllous creeping shoots, the leaves are initiated around the apical meri-
stem in a diagonal decussate pattern: A medial leaf is opposite a ventral leaf on the other side of the stem (Dengler,
1999). This is not always obvious because the two leaves may become displaced and appear non-opposite after stem

44
elongation. The apical meristems of this species are dorsiventrally flattened, a prerequisite condition in plants for
anisophylly and asymmetrical leaves. In erect stems of Selaginella—such as those found in “arborescent” species or
the erect strobili of many species—the stems are terete and therefore the leaves are isophyllous and symmetrical. An
index to all Selaginella names was prepared by Reed (1966).

The dry-area isophyllous species of Selaginella (i.e., the S. rupicola group) are remarkably drought-tolerant. Webster
and Steeves (1964) reported taking a dried plant collected two years and nine months earlier, reviving it for two
hours in tap water, and placing it on sand in a clay pot. Three months later it was thriving and producing strobili.
Tryon (1955) reported "A specimen of S. watsonii was planted three months after it had been pressed and dried
and it put forth new growth. Part of a mat of S. densa var. densa was planted six months after it had been collected
and stored as an herbarium specimen and it also grew."

45
Figure 32. Phylogeny of the Selaginellaceae based on rbcL ( Korall & Kenrick, 2002). Strict concensus tree. Branches with
bootstrap support above 80% are thicker.

46
 Ferns (Class Polypodiopsida)
The ferns are considered monophyletic based on sequence studies of several genes: rbcL (Hasebe et al., 1993), atpB
(Wolf, 1997), and the mitochondrial small sub-unit rDNA (Duff & Nickrent, 1999). Morphological synapomorphies
that might support their monophyly are the possession of 3-layered exospores and septate rhizoids on the gameto-
phyte.

The leaves of ferns and seed plants are not homologous with the leaves of lycophytes. Although they share certain
ontogenetic similarities such as primordia differentiation into an upper and lower leaf zone (Kaplan, pers. com.), ev-
idence from the fossil record suggests that the leaves of both groups have had separate evolutionary origins from
early Devonian ancestors that lacked leaves. The microphylls of lycophytes probably evolved by an enation on the
surface of an ancient stem becoming supplied by a vein (Figure 14). In contrast, the megaphylls (or euphylls) of ferns
and seed plants are thought to be highly modified branch systems. The distal branch systems of early vascular plants
are believed to have become overtopped, planated, and webbed (Figure 15). Megaphylls might have been inde-
pendently derived in ferns and seed plants (Kendrick & Crane, 1997). The megaphylls of ferns (pteridophylls) differ
those of nearly all seed plants by circinate vernation and growth predominated by a marginal meristem (vs. diffuse,
intercalary growth). Except for the Ophioglossaceae, Psilotaceae, and Equisetaceae, ferns leaves have special aerating
tissue, called aerophores or pneumatophores, and in most ferns these are apparent as two light-colored lines on the
dorso-lateral sides of the petiole (Davies, 1991).

Also distinctive are septate rhizoids on the gametophyte, which are present in the Psilotaceae and in some Ophio-
glossaceae, Osmundaceae, Schizaeaceae, Stromatopteridaceae (Bierhorst, 1971), and Marattiales (Camus, 1990; Hill
& Camus, 1986). The lycophytes apparently lack septate rhizoids.

The number of spore layers also appears to be a synapomorphy for ferns. TEM work on extant Psilotaceae, Marat-
tiaceae, and Ophioglossaceae shows that their exospore is three-layered, with cavities in the middle layer (Tryon &
Lugardon, 1991). Preliminary studies of Trimerophytes (Gensel, 1980) shows that the exospore is two-layered, the
outer sculptured layer readily separating from the inner one. In cycads the exposore is five-layered.

Based on outgroup comparison to lycophytes and seed plants, sunken gametangia are plesiomorphic, whereas super-
ficial ones (borne on or projecting from the surface) are derived. The antheridia of Ophioglossales, Equisetales, and
Marattiales are large and sunken, with many sperms. Those of other ferns are small and projecting, with few sperms
(Eames, 1936, p. 301). Apparently superficial gametangia evolved twice within the ferns: in Psilotaceae and again in the
leptosporangiates.

Ferns are sometimes referred to as the “Moniliformopses,” a term that originated with the cladistic analyses of Kenrick
and Crane (1997). In their analyses, they used extinct Devonian species as representatives of ferns and Equisetum.
Specifically, they used Rhacophyton and Pseudosporochnus to represent ferns, and Ibyka to represent Equisetum. Un-
fortunately, there are problems using these genera as respresentatives. Subsequent cladistic studies have found that
Pseudosporochnos and Cladoxylon are not ferns; they form a part of a paraphyletic grade near the base of the vascular
plant tree (Rothwell, 1999). Ibyka has not been reconstructed as a whole plant and is not well enough understood
to be used in broad-scale phylogenetic analysis. Thus, “Moniliformopses” should not be used for ferns. An alternative,
“Polypodiophyta,” can be used and has the advantage of being based on a well known extant genus.

47
 Lycopodiaceae (16/388) Lycopodiales
Lycopodiopsida (lycophytes)
Isoëtaceae (1/250) Isoëtales

(tracheophytes) Selaginellaceae (1/700) Selaginellales

SEED PLANTS
Equisetidae (horsetails)
Equisetaceae (1/15) Equisetales
(euphyllophytes) Ophioglossidae Psilotaceae (2/17) Psilotales
Ophioglossaceae (10/125) Ophioglossales
Polypodiopsida Marattiidae
(ferns) Marattiaceae (6/111) Marattiales
Osmundaceae (6/18) Osmundales
Hymenophyllaceae (9/434) Hymenophyllales
Polypodiidae Matoniaceae (2/4)
(leptosporangiates) Dipteridaceae (2/11) Gleicheniales
Gleicheniaceae (6/157)
Lygodiaceae (1/40)
Schizaeaceae (2/35) Schizaeales
Anemiaceae (1/115)
Salviniaceae (2/21)
Salviniales
Marsileaceae (3/61)
Thyrsopteridaceae (1/1)
Loxsomataceae (2/2)
Culcitaceae (1/2)
Plagiogyriaceae (1/15)
Cibotiaceae (1/9)
Cyatheales
Metaxyaceae (1/6)
Dicksoniaceae (3/35)

Cyatheaceae (3/643)

Saccolomataceae (1/18) Saccolomatineae

Cystodiaceae (1/1)
Lonchitidaceae (1/2) Lindsaeineae
Lindsaeaceae (7/234)

Pteridaceae (53/1211) Pteridineae

Polypodiales
(cathetogyrates)
Dennstaedtiaceae (10/265) Dennstaedtiineae
Cystopteridaceae (3/37)
Rhachidosoraceae (1/8)
Diplaziopsidaceae (2/4)
Desmophlebiaceae (1/2)
Hemidictyaceae (1/1)

Aspleniaceae (2/730)

Woodsiaceae (1/39)
Onocleaceae (4/5) Aspleniineae (eupolypods II)
Blechnaceae (24/265)

Athyriaceae (3/650)
(eupolypods)

Thelypteridaceae (30/1034)

Didymochlaenaceae (1/1)
Hypodematiaceae (2/22)

Dryopteridaceae (26/2115)

Nephrolepidaceae (1/19)
Polypodiineae (eupolypods I)
Lomariopsidaceae (4/69)
Tectariaceae (7/250)
Oleandraceae (1/15)
Davalliaceae (1/65)

Polypodiaceae (65/1652)

Figure 32. A phylogeny of fern and lycophyte families (PPG1, 2016)

48
 Equisetales
4. Equisetaceae Michx. ex DC. | Horsetail Family
Plants terrestrial, occasionally emergent aquatics, rhizomatous. Stems green, jointed, longitudinally grooved,
usually hollow in the center and with a series of smaller canals under the stem ridges (carinal canals) and
larger ones under the valleys (vallecular canals). Branches (if present) whorled at the nodes, erupting through
the base of the leaf sheath. Leaves whorled, fused into a sheath, the tips free, tooth-like, often deciduous.
Sporophylls aggregated in terminal strobili, peltate, tan or yellowish, bearing 5–10 sporangia on the inner surface;
sporangia oblong, dehiscing longitudinally. Spores without haptotypic markings (i.e., neither trilete no monolete), glo-
bose, green, with 4 arm-like appendages (elaters). Gametophytes epigeal, green, with erect lamellae, unisexual but
becoming bisexual with age. Sperm multiflagellate. x=108.

Type species: Equisetum fluviatile L., a circumboreal species.

Distribution and ecology: Cosmopolitan, except New Zealand and Australia. Unlike most groups, Equisetaceae has
more species in the temperate zones than in the tropics.

Genera/species: 1/15. Genera: Equisetum (15 spp.).

Economic plants and products: The silica-roughened stems of most species have been used for scouring pots and
pans, thus giving rise to the common name “scouring rush.” The stems are also used to polish the reeds of woodwind
instruments. Throughout Latin America, bundles of stems are sold in open markets and used to make tea for kidney
and other health problems.

Discussion: One the biggest surprises that DNA evidence has served up is that Equisetum nests squarely among the
ferns. It has been resolved near the Marattiaceae and Osmundaceae (Des Marais, 2003; Pryer et al. 2001) or, based on
78 plastid genes, is sister to Psilotum (Ruhfel et al., 2014). One study based on structural changes of the entire plastid
genome resolved Equisetum sister to the Ophioglossaceae + Psilotaceae (Grewe et al., 2013).

It’s hard to imagine Equisetum as a fern because its anatomy and morphology is unlike that of any extant fern—or
for that matter any other vascular plant. It has green, jointed, longitudinally ribbed stems, whorled branches, highly
reduced leaves fused into sheaths, and terminal strobili. Its stomatal complex is unique among plants, with two sub-
sidiary cells that overlie the guard cells completely, whereas in other plants these cells are superficial, and the inner
tangential wall of each subsidiary cell develops 7–24 ridges-like thickenings, a character not found in any other genus.
The terminal strobili and peltate sporophylls are unlike anything elsewhere in ferns. Their spores have strap-shaped,
hygroscopic elaters—a character unique among present-day ferns. When exposed to humid air (the humidity in your
breath being sufficient), the elaters coil tightly around the spore; when dry, they uncoil and spread, catching the wind
and creating drag that helps keep the spores afloat in the air currents. The spores have no haptotypic markings; they
are neither trilete nor monolete.This is unique among ferns.The sperm cells of Equisetum share several characteristics
with other ferns that support the inclusion of the genus in the ferns (Renzaglia et al., 2000).

Silica, not lignin, plays the major mechanical role in maintaining the erectness of Equisetum stems. The silica is localized
in the epidermis (Parsons & Cuthbertson, 1992; Sapei et al., 2007). In E. palustre it accounts for 25% of the dry weight
(Timell, 1964). Presumably, the silica also defends against herbivorous insects and fungal attacks.

The horsetail clade (i.e., the Equisetaceae and extinct Archaeocalamitaceae and Calamitaceae) has a long fossil history
extending back to the Devonian.The fossil genus Ibyka from the middle Devonian was shown by Kendrick and Crane
(1997) to be sister to this clade based on whorled branching and protoxylem disintegration to form lacunae. During
the Carboniferous, the tree-like Archaeocalamitaceae and Calamitaceae abounded in swamps. These differed from
modern-day Equisetum primarily by the presence of secondary growth and whorled bracts subtending or between
the whorls of sporangiophores in the cones. Mesozoic fossils are placed in Equisetites if they cannot be assigned to

49
any modern-day species of Equisetum. It is possible that some Equisetites actually represent Equisetum. All Cenozoic
fossils, however, are placed in Equisetum. The 15 modern-day species appear to have radiated from a common ances-
tor present during the Eocene (Figure 333; Des Marais, 2003; Soltis et al. 2002). A comparative study (Renzaglia et
al. 2002) of sperm in E. arvense (subgen. Equisetum) and E. hyemale (subgen. Hippochaete) demonstrated remarkable
similarities in ultrastructure, a result consistent with recent radiation in the Eocene.

Two subgenera are usually recognized: subgen. Equisetum (superficial stomates and branched stems, non-apiculate
strobili; 7 species) and subgen. Hippochaete (sunken stomates and usually unbranched stems, apiculate strobili; 8 spe-
cies). Both subgenera are strongly supported by rbcL and trnL–F sequences, although the placement of E. bogotense,
a tropical American species, is ambiguous (Des Marais et al., 2003; Guillon, 2004). Also supporting their distinction is
the lack of hybrids between the two subgenera (although hybrids are common between species within each subge-
nus). The two subgenera differ in nuclear DNA C-values as determined by flow cytometry (Obermayer et al., 2002).
Eight of the 15 species have been analyzed for such values. Subgenus Equisetum has ranges from 1C = 12.5–14.2 pg,
and subgen. Hippochaete has 21.3–31.6 pg. Manton (1950) claimed that there was a difference of chromosome size
between the two subgenera, with subgen. Hippochaete having larger chromosomes.

Equisetum hybrids produce aborted spores that are non-green and irregularly shaped. They also produce a small
amount of large, globose, green spores.These represent diplospores—unreduced spores, or 2n—and they have been
detected in all hybrids examined for them within subgen. Hippochaete. Diplospores are generally 80–140 microme-
ters long, whereas normal meiospores are generally 40–60 micrometers (Duckett, 1970; Hauke, 1978). In some cases
diplospores have germinated and produced gametophytes (Dubois-Tylski & Girerd, 1986; Krahulec et al., 1996). Dip-
lospores are probably involved in the production of triploid taxa known in subgen. Hippochaete (Bennert et al., 2005).

Although they produce aborted spores, hybrids can become common locally by means of vegetative reproduction
and can even extend beyond the ranges of one or both parents. Detached stem segments can root at the nodes, thus
vegetatively propagating both hybrids and parents (Praeger, 1934; Schaffner, 1931; Wagner & Hammit, 1970).

The reason Equisetum is capable of rooting at the nodes is because, at each node, there is an intercalary meristem—a
ring of cells that remains capable of division. Intercalary meristems are ones located (“intercalated”) between adjacent
regions of non-meristematic tissue.The meristems give rise to lateral branches and new stem tissue distally. When the
aerial stems of Equisetum are pulled apart segment by segment, the base of each segment can be seen to consist of
lighter green, younger, softer tissue that is easily broken.This younger tissue was produced by the intercalary meristem.
The meristem functions only in one direction; that is, it produces new tissue distally, or upward, and tissue maturation
is therefore basipetal. This softer, younger tissue needs support, and such support is provided by the leaves. Each leaf
is connate laterally with its neighbors into a sheath surrounding the stem. The leaf sheath supports the weak young
tissue produced by the intercalary meristem. This also happens in many monocots such as grasses, sedges, and Com-
melinaceae (Fisher & French, 1976). Intercalary meristems are present in the aerial shoots of all species of Equisetum.
In the underground rhizomes, intercalary meristems are present in subgen. Hippochaetae and absent in subgen. Equi-
setum (Golub & Wetmore, 1948; French, 1984).

In the temperate species of Equisetum, the apical meristem ceases to exist during the late summer of the year before
they emerge from the ground. Thus, most of the extension growth of the stem in the following spring comes from
the action of intercalary meristems. Similarly, the strobili are formed underground during the late summer the year
before they emerge.Their spores even become green in the season before emergence. If stobili-bearing buds are dug
and cut longitudinally, green spores can be seen within the sporangia. At this time, the apical meristem of the shoot
has ceased to exist, differentiating as collenchymas (Hauke, 1985).

Equisetum is characterized by air canals that run longitudinally throughout the stems. There are three types of canals:
central, vallecular (located on the same radius as the valleys between the ridges), and carinal (located on the same
radius as the ridges, or “keels”). The size and arrangement of these canals are distinctive enough to distinguish the
species. The carinal canals conduct water (Bierhorst, 1971), but the vallecular canals conduct gases.

Of nine species of horsetails studied, four exhibited internal pressurized convection (Armstrong & Armstrong, 2010).
The four represented both subgenera. Gas movement by pressurized convection is especially important for species
that typically grow in waterlogged, anoxic soil. Internal mass flow of “fresh” air through the vallecular canals has been

50
demonstrated for E. telmateia (Armstrong & Armstrong, 2009). Mass flow (convection) occurs when a humidity
gradient has been established within the plant. Conduction rates can be as high as 120 cm per minute. The internal
air is vented through the older, broken aerial stems. Mass flow does not occur in the central canals because of high
pressure-flow resistance from the nodal diaphragms.

The stomata of Equisetum are apparently unique among land plants (Dayanandan & Kaufman, 1973). Incredibly, the
mature stomata appear to be permanently closed (Cullen & Rudall, 2016). Morphologically, the stomatal complex
consists of guard cells lying below the subsidiary cells, the two forming a pore or pit. Also, the inner surfaces of the
subsidiary cells bear silicified ridges that radiate outward from the pore. The function of these ridges is unknown.
These characteristics of the stomatal complex were present in fossil relatives such as the archeocalamites, calamites,
and sphenophylls (Thomassen, 1980; Channing et al., 2011). In the modern-day E. myriochaetum, the subsidiary cells
have an interlocking outer ledge, derived from the cuticle, that delimits the pore. Also, the guard cells produce an inner
ledge. The subsidiary cells appear to lock shut in mature stomata (Cullen & Rudall, 2016).

Equisetum has a high and presumably polyploidy base number of x=108. Genetically, however, it behaves like a diploid
(Soltis, 1986). The 12 species that have been counted represent diploids with 2n=216.

An excellent overview of the biology of Equisetum was given by Husby (2013).

Figure 33. Phylogeny of the Equisetaceae plotted against geological time. From Des Marias et al. (2003).

51
 Psilotales
5. Psilotaceae Kanitz | (Whisk Fern Family)

Plants terrestrial or epiphytic. Roots absent. Stems protoselic. Subterranean stems creeping, nongreen, mycorrhizal,
branched dichotomously, sometimes gearing gemmae; aerial stems erect, green, non-mycorrhizal, glabrous, dichoto-
mously branched; leaves (in Psilotum) small (1–2 mm), scale-like, without veins, or (in Tmesipteris) toungue-shaped
to lanceolate, one-veined; sporangia 2- or 3-locular, sessile in the fork of bifid leaves; spores monolete, reniform,
hyaline. Gametophytes subterranean, cylindrical, elongate, often branched, nongreen, mycotrophic, bearing gametangia
on all sides; archegonia sunken; antheridia superficial. x=52.

Type species: Psilotum nudum (L.) P. Beauv. (basionym: Lycopodium nudum L.)

Distribution and ecology: Tropics and subtropics worldwide, generally at low to middle elevations but absent
from dry areas. Tmesipteris is restricted to Australasia and islands of the Pacific. Both genera often grow as hanging
or arching epiphytes on the fibrous root mantles tree fern trunks or in humus-filled axils of old palm leaves. They are
found less frequently on soil, and Psilotum occasionally grows on rocks. The plants growing on soil or rock are erect.

Genera/species: 2/12. Sole genera: Psilotum (2 spp.), Tmesipteris (10).

Economic plants and products: none.

Discussion: The phylogenetic relationships of this family have long been controversial, and in 1976 a symposium was
held on the subject (White, 1977). Nowadays the Psilotaceae is believed to be sister to the Ophioglossaceae based
DNA studies (Hauke et al., 2003; Schuettpelz & Pryer, 2007).

Psilotum contains two species: P. nudum (L.) Beauv. and P. complanatum Sw. The two hybridize to produce P. X interme-
dium W. H. Wagner. Several ploidy levels are known within P. nudum, but it is unknown whether these levels correlate
with ecology, geography, or spore size.

The leaves of Tmesipteris are unusual. Their blades are oriented vertically so that their edge, not their flat adaxial
surface, is oriented toward the stem. They grow from a basal meristem, not an apical or marginal one. Consequently,
instead of maturing toward their apex like a fern fiddlehead, they differentiate first at the apex, and then mature to-
ward the base. A complete serial transition can be seen between short leaves at the base of a shoot and the longest
leaves at the apex.There is no worldwide taxonomic treatment of Tmesipteris, but four species occur in New Zealand,
and these were studied by Chinnock (1975). Most species commonly grow on the root mantles of tree ferns.

Sister to the Ophioglossales are the rest of the ferns. This clade supported by molecular studies and apparently in-
volved the evolution of green, surficial, thalloid gametophytes, archegonia borne ventrally, and root hairs (the Ophio-
glossaceae lacks root hairs, and the Psilotaceae lacks roots).

Key to the genera of Psilotaceae

1. Leaves scale-like, without a mid-vein; sporangia 3-locular, sessile in the axil of a bifid sporophyll ................... Psilotum
1. Leaves tongue-shaped to lanceolate, with a mid-vein; sporangia 2-locular, borne away from the
axil on the base of a bifid sporophyll............................................................................................................................................... Tmesipteris

52
Figure 34. Psilotaceae. Psilotum nudum. A. Habit. B. detail fo the stem. C. Sporangium. D. Section of sproangium. E Spores.
Taken from Flora Vascular de Andalucía Occidental (http://www.rjb.csic.es/floraiberica/).

53
 Ophioglossales
The monophyly of the Ophioglossales (Figure 32) is supported by five genes: rbcL (Hasebe et al., 1995; Hauk et
al. 2003; Manhart, 1994; Pryer et al., 1995), atpB (Wolf, 1997), chloroplast SSU rDNA (16S) (Nickrent et al., 2000),
nuclear SSU of rDNA (18S) (Hedderson et al. 1998), mitochonrial SSU r DNA (19S) (Duff & Nickrent, 1999), and
mitochondrial nad5 (Vangerow et al., 1999). Further support for the clade was found by Wolf (1997), who did a
combined analysis of atpB and rbcL and found there was stronger support than for either molecule alone.

Morphological synapomorphies for the order are subterranean, fleshy, cylindrical, mycotrophic, achlorophyllous ga-
metophytes (Figure 36A, A11 & A12) and adaxial position of the sporophore on the leaf. The synangia are supplied
individually with a vein running to their base. In no other fern group are the sporangia similarly supplied.The order ex-
hibits a tendency toward reduction of the root system, with the Ophioglossaceae having simple or sparsely branched
roots, and the Psilotaceae lacking roots completely (Schneider et al., 2002). Despite the sister relationship of the order
to other ferns (Figure 32), its fossil record dates only to the Tertiary (Carpenter, 1988). Molecular age estimates put
the divergence between the Ophioglossaceae and Psilotaceae at about 305 MA, in the late Carboniferous (Pryer et
al., 2004).

6. Ophioglossaceae Agardh | Adder’s-tongue Family

Plants terrestrial or (a few species) epiphytic, fleshy, without sclerenchyma, generally small. Roots thick, simple or
rarely branched, glabrous, mycorrhizal, lacking root hairs. Stems subterranean, short, erect, typically unbranched, lacking
hairs or scales, containing an ectophloic siphonostele. Fronds circinate or folded (bent) or conduplicate in bud, usually
less than 30 cm long. Petiole bases dilated and sheathing up to 5 nested primordia. Fertile leaves divided into a
blade-like photosynthetic portion (trophophore) and a spike-like or paniculiform sporangium-bearing
portion (sporophore). Trophophore entire to decompound. Sporophore borne adaxially and medially, diver-
gent from the common petiole base with the trophophore, erect. Veins free or netted. Sporangia globose, sessile or
subsessile or (in Ophioglossum) sunken in two rows on either side of the sporophore, containing hundreds of spores;
annulus absent. Spores trilete, globose-tetrahedral, usually verrucose. Gametophytes subterranean, nongreen, mycor-
rhizal, fleshy, irregularly elongate, cylindrical. Gametangia sunken, distributed on all sides, the antheridium of many cells.
x=30 in Ophioglossum; x=45 in Botrychium.

Type species: Ophioglossum vulgatum L.

Distribution and ecology: Cosmopolitan; generally absent from dry regions. Most species are terrestrial in wet
forests or slightly disturbed, moist places such as thickets, meadows, pastures, or lawns. Because of their small size
they are easily overlooked. They tend to form genus communities, with more than one species occurring together
(Wagner & Wagner, 1983). Ophioglossum tends to form colonies by long, proliferous roots. Cheiroglossa palmata and
C. pendula are the only epiphytes in the family.

Genera/species: 8/90. Genera: Botrychium (30 spp.), Botrypus (2), Cheiroglossa (2), Ophioglossum (30), Helmintho-
stachys (1), Japanobotrychium (1), Mankyua (1), Sceptridium (25).

Economic plants and products: none.

Discussion: The Ophioglossaceae are strongly supported as monophyletic by molecular studies (Hauke et al., 2003;
Schuettpelz & Pryer, 2007). The one (or, rarely) few leaves per year or growing season is a synapomorphy for the
group, as is the fertile leaf divided into a sterile blade (trophophore) and fertile spike (sporophore). Other distinctive
characters are non-circinate leaves (except Botrychium sect. Sceptridium), sheathing leaf bases that surround several
successive leaf primordia and the stem apical meristem, eusteles, circular-bordered pits, and absence of root hairs
and sclerenchyma. Some pteridologists has suspected a relationship of the family to progymnosperms on the basis of

54
circular-bordered pits in the tracheids, axillary buds (in Botrychium, s.l.), limited secondary grown, and eusteles (Bier-
horst, 1971; Kato, 1987, 1988); however, molecular studies (cited above) support its sister-group relationship to the
Psilotaceae. The circular-bordered pits were studied by Morrow and Dute (1998, 1999). The presence of these pits in
the tracheids of the Ophioglossaceae is unique among ferns.

As currently circumscribed, the Ophioglossaceae constitute eight genera:, one of which, Mankyua, was recently de-
scribed (Sun et al., 2001). Ophioglossum is nearly cosmopolitan, Botrychium is mainly temperate and boreal, and the
monotypic Helminthostachys is limited to lowland Indo-Malayan and Austalasian regions. Mankyua is reported only
from Cheju Island off the coast of Korea. Sometimes these genera are subdivided. Kato (1987) divided Botrychium
into four separate genera: Botrychium (s.s.), Sceptridium, Japanobotrychium, and Botrypus (=subgen. Osmundopteris), a
classification followed herein. Hauk et al. (2003) found that these subgroups are monophyletic except for Botrypus,
which is paraphyletic to a clade consisting of Botrychium + Sceptridium (Figure 35). A phylogeny of Botrychium was
presented by Hauk et al. (2012). Presl (1845) treated Ophioglossum as four separate genera: Ophioglossum (s.s.), Rhi-
zoglossum, Ophioderma, and Cheiroglossa.

At the species level, Clausen (1938) published a worldwide monograph that is now out-of-date. In the past two
decades many new species of Botrychium (s.s.) have been discovered, especially in western North America, and re-
ticulate relationships and ranges have become much better known. Nowadays, about 30 species are recognized in
Botrychium, of which 16 are of putative allopolyploids. For an up-to-date account, consult Farrar (2006).

Ophioglossum reticulatum, a pantropical species, has the highest chromosome number of living organism: 2n = 1,262
(Abraham & Ninan, 1954). Higher counts have sometimes been reported, but these are not photographically docu-
mented and therefore should not be accepted.The trophophore of Ophioglossum lacks a costa, which is a helpful field
characteristic to distinguish the plant from look-alikes. The trophophore of Ophioglossum is thought to be an expand-
ed rachis, or phyllode, developed by intercalary growth. Some species, such as O. lusitanicum, have the trophophore
unexpanded, presumably representing the ancestral character state.

The family is known in the fossil record from a single species, Botrychium (Botrypus) wightonii, from the Paleocene of
western Canada (Rothwell & Stockey, 1989). It resembles Botrypus virginianus, an extant species of temperate and
tropical Americ

55
Figure 35. A phylogeny of the genera of Ophioglossaceae.The relationships of the Asian Mankyua are unknown. Ophioderma
and Rhizoglossa, both Old World genera, are not shown.. (Based on Hauk et al., 2003.)

56
Figure 36. Ophioglossaceae. A1–3. Sceptridium multifidum. A1. Habit. A2, 3. Sporangia. A4–12. Botrypus virginianus. A4–10.
Cross section through various levels of the leaf showing departing vascular bundles to the fertile spike. A11. Gametophyte
(a=antheridia; b=archegonia; c=cortex with mycorrhizal fungus). A12. Gametophyte, top view. B1. Cheiroglossa palmatu. B2.
Ophioglossum vulgatum. C1, 2. Helminthostachys zeylanica. C1. Habit. C2. Sporangia.

57
 Marattiales
7. Marattiaceae Bercht. & J. S. Presl | Marattia Family
Plants terrestrial, with mucilage canals in the root, stem, and leaf; roots polyarch, succulent; rhizomes creeping or
erect, in some species massive and subarborescent, fleshy, lacking sclerenchyma, bearing two large, fleshy, ear-like
stipules on either side of the fronds; rhizome scales generally peltate and inconspicuous; sterile and fertile leaves
monomorphic or (in Danaea) dimorphic; petioles swollen at the very base where it joins the rhizome, containing
two or more concentric rings of separate vascular bundles; laminae simple to 5-pinnate; rachises or costae
swollen at their junctions with pulvinus-like thickening; veins free or (in Christensenia) netted; stomata more
or less strongly cyclocytic; sori abaxial, non-indusiate, elongate to linear or (in Christensenia) round, fused laterally
into synangia or (in Angiopteris) free but clustered closely together, opening by a terminal pore or slit; spores >
1000 per sporangium, trilete and globose or monolete and ellipsoid; gametophytes surficial, green, thalloid, commonly
mycorrhizal, with endoscopic embryo; antheridia borne on both surfaces; archegonia borne ventrally. x=40, 39.

Figure 37. Marattiaceae. A. Synangium of Marattia. B. Ptisana fraxinea, pinna. Note swollen node at base. C. Stipules at base
of petiole. D. Stele of Angiopteris evecta.

58
Type species: Marattia alata Sw., of the West Indies.

Distribution and ecology: Pantropical. Three genera occur in the New World: Marattia, Eupodium, and Danaea (the
latter two are entirely neotropical). All members of the family grow on wet, shaded, forest floors.

Genera/species: 6/120. Genera: Danaea (50 spp.), Angiopteris (ca. 30?), Ptisana (27), Marattia (7), Eupodium (2),
Christensenia (2).

Economic plants and products: none.

Discussion: The Marattiaceae can be easily identified by pulvinus-like swellings on the leaf axes, ear-like stipules at the
stem-leaf junction, and sporangia opening by a terminal pore or slit (Figure 377). The pulvinus-like swellings are more
properly termed "pulvinuloids" becuase they lack the reversible movement of true pulvini, as in legumes. The stipules
are fleshy and if cut and placed in soil will often form plantlets along the margin. Lehn & Lopes (2017) reported
stipular buds in Danaea nodosa from Brazil, but from their photos, these appear to be branch buds, not stipular.

Two synapomorphies of the family are internal: the mucilage canals and the petiole vasculature. The mucilage canals
can be detected after the stem or petiole is cut: thick mucilage oozes out of points (the canals) on the cut surface.
The petiole vasculature, when seen in cross-section, consists of separate vascular bundles are arranged in several
concentric circles. In three dimensions the vasculature would resemble a series of nested and linked cones. No other
extant ferns even remotely have this kind of petiole vasculature.

The Marattiaceae has the oldest fossil record of any fern family. They appears in the late Carboniferous represented
by Psaronius, an arborescent genus with a well-developed root mantle. Psaronius was prominent in swamps of the
Carboniferous and went extinct in the early Permian.

The Marattiaceae have an endoscopic embryo, a character found elsewhere among fern only in Sceptridium. The first
embryonic leaf penetrates and emerges through the tissue of the gametophyte; it does not emerge around the side
of the thallus or between the apical notch. The first leaf, therefore, appears to spring directly from the upper surface
, which is a helpful character for identifying the gametophyte of this family in the field.

Six genera (see above) were recognized in the most recent phylogenetic study of the family (Murdock, 2008). The
study described one new genus, Ptisana (paleotropical), and resurrected one old name, Eupodium (neotropical), that
had not been previously widely used. It was found that Danaea, which is entirely neotropical, is sister the rest of the
genus. The remaining genera form two clades: Eupodium + Ptisana, and Christensenia + Marattia + Angiopteris (Figure
38).These results differ markedly from previous a phylogenetic study of the family based on morphology and anatomy,
and one that also included fossils (Hill & Camus, 1986). This study showed Christensenia sister to an unresolved clade
consisting of the remaining extant genera.

A phylogenetic study of 31 species of Danaea revealed three well supported clades: the leprieurii clade, nodosa clade,
and alata clade (Christenhusz et al., 2008). These clades overlapped ecologically and geographically.

An assessment of the geography and morphological characters of the Old World genera is given by Camus (1988).
Christensenia, a genus of Asia and Malesia, has stomata that measure about 80 micrometers—the largest of any plant.
They are visible to the naked eye and always remain open.

Hovenkamp et al. (2009) reported an unusual spore ejection mechanism from the sporangia of Angiopteris cf. evecta.
The spores suddenly “jump” several millimeters caused by the cavitation of water held between the exospore and
perispore.

59
Figure 38. Phylogeny of the Marattiaceae based on maximum likelihood analysis of rps4–trns and trnSGG. Support values:
Bayesian posterior probabilities/maximum parsimony/decay index “+” = 100, “-“ = < 50 (0 for decay index values). (from
Murdock, 2008)

60
 Leptosporangiate Ferns
The leptosporangiate ferns are well-supported by molecular evidence from several genes or gene regions (Pryer et
al., 1995, 2004; Schneider et al. 2004; Wolf, 1997). They are also well supported by a unique type of sporangial devel-
opment (Figure 16) that gives the group its name.The type of sporangial development found in hornworts, liverworts,
mosses, lycophytes, Ophioglossales, and Marattiales (and possibly the Equisetaceae) results in a capsule wall two to
several cells thick and generally has high spore output (500+).This type of sporangium is called a “eusporangium.”The
leptosporangium is clearly the derived condition, having a wall only one cell thick and fewer spores per capsule. Other
synapomorphies for leptosporangiate ferns are a differentiated annulus, sporangial stalks 4–6 cells wide, superficial
(not sunken) gametangia, and exarch protoxylem. Even the roots of leptosporangiate ferns exhibit a synapomorphy: a
single apical initial, instead of up to four as in the Ophioglossaceae (and Psilotum?) and Marattiaceae (Schneider, 1996).

Osmundales
8. Osmundaceae Bercht. & J. C. Presl | Royal Fern Family
Plants terrestrial. Roots abundant, fibrous, often thickly investing the rhizome, diarch. Rhizomes decumbent or erect,
hairy, surrounded by overlapping persistent petiole bases and roots; the stele a ectophloic dictyoxylic
siphonostele. Fronds to 1.5 m long, monomorphic or partly to completely dimorphic, hairy but becoming glabrous
with age. Petioles gradually expanded toward the base into sheathing wings, containing a single C-shaped
vascular bundle with the open end oriented adaxially. Laminae 1- to 2-pinnate, cespitose. Veins free. Stomata
anomocytic. Sori abaxial or on naked axes. Sporangia globose, short-stalked, with a single lateral annular patch not
greatly differentiated from the other cells of the wall, dehiscing vertically. Spores trilete, globose-tetrahedral, green,
125–512 per sporangium. Gametophytes green, epigeal, thalloid, with thickened central midrib. Archegonia borne
on the ventral surface in rows along either side of the thickened midrib. Antheridia borne mostly on the ventral sur-
face of the margins or at the edge. x=22.

Type species: Osmunda regalis L., a nearly cosmopolitan species.

Distribution and ecology: Cosmopolitan. Osmunda and Osmundastrum are the only representatives of the family
in the New World and Northern Hemisphere. Leptopteris and Todea are Australasian. Wet habitats, either in forests
or open areas.

Genera/species: 4/18. Genera: Osmunda (9 spp.), Leptopteris (6), Todea (2), Osmundastrum (1).

Economic plants and products: The fibrous roots of Osmunda are occasionally used as a substrate for growing
orchids. Osmundastrum cinnamomea (cinnamon fern) and O. regalis (royal fern) are grown as ornamentals.

Discussion: The Osmundaceae is well-supported based on seven plastid loci and morphological characteristics
such as its rudimentary annulus (Figure 399,A4; Metzgar et al., 2008; Bobrov, 1967). Molecular phylogenetic analyses
(Metzgar et al., 2008) reveal four clades within the family, and pteridologists are now recognize these clades as gen-
era. The main change to pre-existing classifications is that Osmunda cinnamomea is now placed in its own monotypic
genus, Osmundastrum, to preserve the monophyly of Osmunda s.s. (Figure 4040).

The family has a long fossil history. About 150 extinct species have been described from all continents, with the fossils
dating from at least the Permian onward (Miller, 1967, 1971). Triassic fossils of Osmunda claytoniana are known from
Antarctica and have shown little change over the past 220 million years from the present-day plants (Phipps et al.,
1998); however, Yatabe et al. (1999) thought the fossils represented O. cinnamomea, not O. claytoniana. Undisputed
fossils of O. cinnamomea are known from the Late Cretaceous of western North America (Serbet & Rothwell, 1999).

61
Many fossil species were arborescent and are known from beautifully preserved trunks. These trunks can be confi-
dently assigned to the family because of their distinctive cross-sectional anatomy of C-shaped vascular petiolar traces
and many overlapping, winged petiole bases that encase the rhizome like armor. The whole structure was permeated
and invested by wiry roots. This method of effectively increasing the width of the rhizome to become a tree can be
seen in the extant genera Leptopteris and Todea, which have trunks up to 1–2 meters tall.

The rhizome of the Osmundaceae (Figure 399B) is an “ectophloic dictyoxylic siphonostle.” It is basically an ectophloic
siphonostele; that is, one with the endodermis and phloem on the outside of the xylem cylinder only, not on the inside
(in which case it is “amphiphloic”). It is unique, however, because the leaf gaps interrupt only the xylem cylinder, not
the phloem or endodermis. Thus the term dictyoxylic, which means “net xylem.”

Hewitson (1962) studied the anatomy of extant species and found that the arrangement of sclerenchyma strands
in the petioles is diagnostic of certain groups—a finding that helps classify some fossil species where this feature is
preserved. The gametophytes of the family were described by Stokey & Atkinson (1956a).

Osmunda (in the traditional sense) is the most species-rich and widespread genus of the family. It exhibits dimorphy
of the sterile and fertile parts, and this dimorphy may be either complete or partial. In the latter, only a few pinnae are
fertile on an otherwise sterile frond, a condition sometimes called “hemidimorphic.” In either case, the fertile parts are
usually highly contracted with little or no green tissue. In Todea, an exapanded lamina is present on the fertile leaves.

Leptopteris is unusual because its lamina is only a few cell layers thick and without stomata.This imparts a filmy appear-
ance to the fronds. Sequence data from rbcL indicates that Leptopteris is sister to Todea and that these two genera
are nested within Osmunda (Yatebe et al., 1999).

Hybrids are extremely rare in the family. Brownsey (1981) reported one in Leptopteris. In the temperate zones,
species of Osmunda grow together frequently, but only one hybrid combination is known: Osmunda X ruggii (=O. clay-
toniana X O. cinnamomea) (Tryon, 1940; Wagner et al., 1978). It occurs in the eastern United States, and is currently
known from only one locality. Hybrids in Osmunda and the geographic distribution of the genus were reviewed by
Kato (2007, 2009).

In the family phylogeny of ferns (Figure 32), one striking morphological change occurred along the internode from the
Osmundaceae to the Gleicheniaceae: the elaboration of the annulus into a ring-like structure completely encircling
the sporangium. The annulus by-passes the stalk and is therefore said to be “oblique.” In the Polypodiales, the annulus
is vertical and interrupted at the stalk.

Between the Osmundaceae and Schizaeaceae (Figure 32) another synapomorphy appears. It involves two structural
changes to the chloroplast genome: a duplication of the psbA gene and an inversion rRNA order in the inverted
repeat region (Stein et al., 1992; Raubeson & Stein, 1995).

The Osmundaceae are unusual among ferns in that their gametophytes first become male when grown in isolation,
and they lack any known antheridiogen system (Greer et al., 2009).

Osmunda can form extremely old clones. A study of O. claytoniana in West Virginia (USA) found growth rates of
0.1–0.6 cm per year (Faust & Petersen, 2015). This is perhahps the slowest of any fern (pers. obs.). Because the
sheathing leaf bases are persistent throughout the rhizome and the average number of leaves is known per year, the
age of clones can be calculated. It was estimated that some clones more than 2 m wide were at least 414 years old.

62
Figure 39. Osmundaceae. A1. Osmunda regalis, note winged petiole base. A2–A3. Developing sporangia. A4. Open
sporangium with lateral annulus patch. B1. Osmundastrum cinnamomeum, habit. B2. Cross-section of rhizome showing
dictyoxylic stele and overlapping leaf bases. C1–2. Osmunda japonica, cross-section of petiole showing c-shaped vascular
bundle with enrolled arms.

63
Figure 40. Phylogeny of the Osmundaceae. Fifty percent majority rule consensus tree from a Bayesian analysis of seven plastid
loci (Metzgar et al., 2008).

64
 Hymenophyllales
9. Hymenophyllaceae Link | Filmy Fern Family
Plants epiphytic or terrestrial. Rhizomes protostelic, pubescent (not scaly), usually creeping and bearing distichous
leaves, less commonly erect and radially symmetrical. Leaves generally small, 1–40 cm long, monomorphic or (less
commonly) dimorphic. Petioles with a single vascular bundle. Laminae almost always one cell thick between the
veins, without stomata. Veins free. Sori marginal, borne at the vein tips. Indusia formed by the green laminar tissue,
either urceolate, tubular or bivalvate. Sporangia borne on all sides of the receptacle, short-stalked (the stalk about
6 rows), with annulus oblique, not interrupted at the stalk. Spores green, trilete, tetrahedral-globose. Gametophytes
filamentous or thallose or both, often gemmiferous; antheridia with 5 to many cells. x=36 (with 32 and 34 derived).

Type species: Trichomanes crispum L., nom. cons.

Distribution and ecology: Tropics and (less commonly) subtropics, worldwide. Most species are either terrestrial
or epiphytic, but a few are hemiepiphytes or lianas.

Unlike other ferns, the leaves of filmy ferns are one-cell thick between the veins (with few exceptions) and lack dif-
ferentiated epidermises and stomata. Their cuticle is highly reduced or absent (Haertel, 1940). This means that their
leaves are prone to dry quickly and completely.Yet upon dehydration, they start metabolizing almost immediately (i.e.,
they are poikilohydric). They are thus not delicate plants that need constant humidity and shade. In fact, Hymenophyl-
lum tunbridgense and H. wilsonii grow poorly in constant high humidity; they need periodic drying to maintain optimal
photosynthesis (Proctor, 2003). Physiological tolerance to drying may allow certain filmy ferns to grow in the xeric
environment of the canopy (Krömer & Kessler, 2006).

Genera/species: 9/600. Major genera: Hymenophyllum (250 spp.), Trichomanes (60), Crepidomanes (30), Didymo-
glossum (30), Abrodictyum (25), Polyphlebium (15), Vandenboschia (15).

Key to the traditional Genera of Hymenophyllaceae

1. Indusium two-valved; sporangia borne within the indusium valves (i.e., not exert) on a short receptacle;
rhizomes long-creeping................................................................................................................................................................... Hymenophyllum
1. Indusium cylindrical or funnel-shaped; sporangia exerted from indusia on a long receptacle;
hizomes long- to short-creeping or erect......................................................................................................................................Trichomanes

65
Economic plants and products: none.

Discussion: The monophyly of this family has never been questioned and is well supported by molecular studies
(Ebihara et al., 2007; Pryer et al. 2001b; Schuettpelz & Pryer, 2006, 2007) and the morphological synapomorphies of
laminae one-cell-layer thick, marginal sori, and green spores. Two main clades roughly equal in size occur in the family:
Hymenophyllum and the trichomanoid clade (eight genera). With rare exceptions, Hymenophyllum has two-valved
indusia whereas the trichomanoid clade has tubular indusia,. The base chromosome number for the family is x=36,
from which were derived 32 and (twice, separately) 34 (Ebihara et. al., 2007).

Hymenophyllum (Figure 42) diversified more recently than Trichomanes and exhibits slower evolutionary rates
(Schuettpelz & Pryer, 2006). It is far less diverse ecologically and morphologically than Trichomanes, consisting almost
entirely of epiphytes with long-creeping rhizomes. It has been the focus of several phylogenetic studies by Hennequin
et al. (2003, 2006a, 2006b, 2010).

The trichomanoid clade is remarkably varied in growth habit (Figure 41). It can be terrestrial, lianescent, hemiepiphytic,
or epiphytic, and the rhizomes are correspondingly varied (Dubuisson et al., 2003). (In lianas, the sporophyte starts
growth terrestrially on the forest floor and then climbs; in hemiepiphytes, the sporophyte starts growth as an epi-
phyte (usually on the base of a trunk) and becomes secondarily terrestrial by growing long roots into the soil.) This
variation has lead pteridologists to propose as many as 42 genera (Copeland, 1933, 1938, 1947; Morton, 1968; Pichi
Sermolli, 1977; Iwastsuki, 1990). The most recent classification recognized Hymenophyllum and split Trichomanes into
eight genera (with subgenera): Didymoglossum, Crepidomanes, Polyphlebium, Vandenboschia, Abrodictyum, Trichomanes,
Cephalomanes, and Callistopteris (Ebihara et al., 2006).

Didymoglossum (including Microgonium) contains about 40 species, most of which are neotropical. Its monophyly is
strongly supported by analyses of rbcL, morphology, and anatomy (Dubuisson (1997a, b, 1998; Dubuisson et al. 2003).
It distinguishing characteristics are creeping rhizomes, leaves less than 5 cm long, laminae simple to lobed (rarely pin-
natifid), highly reduced vascular anatomy, and lack of roots (Wessels Boer, 1962). Many species have false veins, which
appear as faint streaks unconnected to the main veins and not composed of vascular tissue. Cytologically, the group
is uniformly x=34 (Tryon & Tryon, 1982), a number derived within the family (Ebihara et al., 2007).

Polyphlebium is pantropical and consists of about 15 species, either epiphytic or epilithic. Many have long-creeping fili-
form rhizomes densely covered by dark adhesive hairs. Also present are root-like rhizomes (Schneider, 2000). Unlike
most neotropical groups, the venation is anadromous and the pinnae inequilateral. An unusual species is P. capillaceum
(L.) Ebihara & Dubuisson, which is restricted to tree fern root mantles (Moran et al., 2001) and has a three-dimen-
sional leaf consisting of capillary segments.

Vandenboschia is a pantropical genus of about 15 species. All appear to be primary hemiepiphytes; that is, the sporo-
phytes start growth on the bases of trunks or rocks and become secondarily terrestrial by growing thick roots into
the soil, where they branch profusely. This has been well documented for V. collariata (Nitta & Epps, 2009). The climb-
ing rhizome have long internodes and are covered, at least ventrally, by adhesive roots (Schneider, 2000). The leaves
are arranged distichously and have relatively short petioles.

Abrodictyum is primarily an Old World genus, but its subgen. Pachychaetum has about half of its 10 species in the New
World.They are terrestrial and have erect rhizomes supported by thick roots. Multicellular, reddish hairs occur on the
petioles, and the lamina segments are narrow with only a few rows of cells on either side of the midribs. The lamina
cells are large and easily visible with a hand lens, except in the most widespread neotropical species, A. rigidum (Sw.)
Ebihara & Dubuisson, which has occluded cells. A synapomorphy for the genus is x=33 (Ebihara et al., 2007).

Trichomanes has four main clades, these most recently recognized as subgenera (Ebihara et al., 2007). The first, subgen.
Trichomanes, contains approximately 40 species and is entirely neotropical except for one species in western Africa
(T. crenatum Bosch). Typically the plants are terrestrial with 1-pinnate leaves (Dubuisson et al. 2003a,b; Ebihara et al.,
2007). Most species bear sori at the apices of the pinnae, not the sides. The second clade, subgen. Feea, consists of
five species, all neotropical. All (except T. mougeotii Bosch) have dimorphic sterile and fertile leaves, and their laminae
are pinnatifid or 1-pinnate with proliferous, flagellate apices.They typically grow on steep clay banks or on rocks; none
are epiphytic. The third clade, subgen. Davalliopsis, consists of two neotropical species, T. elegans Rich. and T. resinosum

66
R. C. Moran. These have erect rhizomes supported by thick roots, and finely divided blades three-cells thick between
the veins. When growing in shade, the laminae are strongly greenish blue iridescent on the upper surface.The physical
basis of the iridescence is thin-film interference, and the thin layers that produce it lie within the chloroplasts of the
upper epidermal cells. The chloroplasts contain grana stacks, which have five thylakoids each and are connected by
extremely short stromal lamellae, for a repeating series of filters of just the right thickness to produce the remarkable
interference color (Graham et al., 1993; Lee, 1997). (See Moran (2004) for a popular account of iridescence in ferns.)
Other apparent synapomorphies of the subgenus include sori bent downward out of the plane of the lamina and
x=32. The last clade, subgen. Lacostea, consists of four species, all neotropical. They are said to be lianas (Dubuisson
et al., 2003b). All are unusual among ferns by having their leaves pressed flat against their support trunks and the sori
protruding perpendicularly from the substrate.

Didymoglossum and most species of Crepidomanes (paleotropical) lack roots (Schneider, 2000), and this loss evolved
independently in both genera (Dubuisson et al., 2003a,b; Ebihara, 2007). The function of the roots has been trans-
ferred to short root-like rhizomes and adhere to the substrate. That these are short modified rhizomes, not true
roots, is evinced by intermediate forms (with the typical elongated rhizomes) that can be found on the same plant.
Also, these root-like rhizomes occasionally bear reduced leaves or leaf buds, they lack a root-cap, they possess a
cuticle, and their origin is exogenous, not endogenous as in roots. They are borne in association with leaf nodes, not
scattered along the rhizome as roots would be. Several other criteria are listed by Schneider (2000).

Another unusual adaptation is adhesive hairs found in Didymoglossum, Crepidomanes, Polyphlebium, and some groups
of Trichomanes (Schneider, 2000). Adhesive hairs resemble root hairs but differ in ontogeny and function. They are
cut off from the subtending epidermal cell by a septum, and they themselves can be 0–3-septate (root hairs are
non-septate extensions of the epidermal cell). Adhesive hairs anchor the plant and may enlarge or branch at the tip
upon contact with the substrate. They have been reported as being covered by a cuticle and therefore incapable of
absorbing water or nutrients; however, experiments with water-soluble dyes have shown that some species of Tricho-
manes readily absorb water through their adhesive hairs (M. Sundue, pers. comm.). Moran and Vidal (2004) postulated
adhesive hairs were one reason Didymoglossum godmanii is one of the few vascular plants able to colonize the smooth
trunks of the palm Welfia georgii in Costa Rica.

A survey of hair types in the family was given by Duckett et al. (1996). Besides adhesive hairs, they found two other
types. One was a glandular hair consisting of two cells: a dark and persistent basal cell and an ephemeral apical cell.
Glandular hairs are often found along the veins, and ones with elongate basal cells were found overarching the rhi-
zome apex. These might protect the apical meristem from desiccation. The second type of hair was single-celled, dark
(usually black), and acicular. It occurs only in Didymoglossum and is frequently found in pairs or clusters, imparting the
appearance a stellate hair.

Didymoglossum is highly unusual among ferns because it young leaves are planar, not circinnate (pers. obs.). The apex
and margins of the leaves show no tendency to enroll. In sect. Didymoglossum, the margin is protected by dark stellate
hairs that develop precociously from some of the cells along the marginal meristem. The fringe of hairs on the lamina
margin is noticable even at very young stages of leaf develpment. The hairs are absent in sect. Microgonium.

This family has a sparse fossil record. The earliest fossil, Hopetedia praetermissa, is from the upper Triassic of North
Carolina (Axsmith et al., 2001).

67
Figure 41. Phylogeny of the Trichomanes (sens. lat.) clade. · = Asia. ¨ = Africa. o = Neotropics. Ab = Abrodictyum. Ac =
Trichomanes subgen. Trichomanes; Cr = Crepidomanes. Da = Trichomanes subgen. Davalliopsis; Di = Didymoglossum; Fe =
Trichomanes subgen. Feea. Le = Lecanium. Mg = Microgonium. NT = Trichomanes, all neotropical; Pa = Abrodictyum clade; PT
= paleotropical clade. Se = Abrodictyum subgen. Pachychaetum. Tp = Polyphlebium; Va = Vandenboschia. AS = Asian clade..
From Dubuisson et al. (2003a).

68
Figure 42. A phylogeny of Hymenophyllum (sens. lat.). (From Schuettpelz & Pryer, 2006.)

69
 Gleicheniales
This order consists of the Dipteridaceae, Gleicheniaceae, and Matoniaceae. The families were abundant and cosmo-
politan during much of the Mesozoic. Although previous molecular phylogentic studies found only weak support for
the group (Hasebe et al., 1994, 1995; Kato & Setoguchi, 1999; Pryer et al., 1995), a recent study has found it strongly
supported (Pryer et al. 2004). This study also suggested that the Hymenophyllaceae were sister to the Gleicheniales
(Figure 43). The chloroplast genome of one species of Gleicheniaceae, Diplopterygium bancroftii (reported as Gle-
ichenia), contains a structural rearrangement unique in ferns (Raubeson & Stein, 1995).

Monophyly of the order is strongly supported by morphology (Stevenson & Loconte, 1996).The synapomor-
phies (pers. obs.) that support the group are: 1) two-layered exospore on the distal face of the spore (three-layered
proximally), 2) a flange on the proximal face of the spore, 3) 3—5 protoxylem poles in the root, 4) involute ends of
the C-shaped petiolar vascular bundle, 5) sporangia arranged circularly around the receptacle, 6) leaves borne in two
ranks on the upper surface of the rhizome, 7) archegonial necks curved toward the apical notch of the prothallus, 8)
uplifted wings on the prothallus, and 9) ring-shaped annulus. The families also share a number of characteristics con-
sidered ancestral such as lack of an antheridiogen system, thick long-lived prothalli with (often) sex organs on both
surfaces. Unlike leptosporangiate ferns in general, the members of this order lack pinnate frond architecture.

Figure 43. A phylogeny of the genera of Gleicheniales. (After Pryer et al., 2004.)

70
10. Matoniaceae C. Presl | Matonia family

Plants terrestrial or epipetric. Rhizomes long-creeping, pubescent or bristly, containing 2 or 3 concentric solenoste-
les. Petioles containing a C-shaped vascular bundle with enrolled arms. Blades divided into two more or less equal
scorpioid halves (Matonia), or sympodial-elongate (Phanerosorus), glabrous. Veins free in the sterile segments, retic-
ulate around the sori. Sori abaxial, round, indusiate, the indusium umbrella-like, peltate, falling easily. Sporangia
few and large, sessile or nearly so, arranged in a circle, maturing simultaneously; annulus oblique, incomplete; stomium
none. Spores trilete, tetrahedral, 48-64 per sporangium. Gametophytes epigeal, thalloid, cordate, with ruffled wings;
gametangia borne on both surfaces. x=26 in Matonia, 25 in Phanerosorus.

Type species: Matonia pectinata R. Br.

Distribution and ecology: Malesia. Matonia grows in open or semi-open habitats, usually on mineral-poor soils.
Phanerosorus grows on exposed limestone (Walker & Jermy, 1982).

Genera/species: 2/4. Genera: Matonia (2 spp.), Phanerosorus (2).

Economic plants and products: none.

Discussion: The monophyly of the Matoniaceae is strongly supported by rbcL (100% bootstrap support; Kato &
Setoguchi, 1999) and morphology (Stevenson & Leconte, 1995). The family is sister to the Dipteridaceae (Kato &
Setoguchi, 1998; Pryer et al., 2004b). Kato and Setoguchi (1999) analyzed all four species in the family and found that
Matonia and Phanerosorus were sister to each other.

The family has an extensive fossil record; in fact, it was described from fossils before any extant species were discov-
ered. During the Mesozoic the family was cosmopolitan and abundant, and from that Era 14 fossil genera and many
species have been described. The family also has an extensive fossil spore record (van Konijnenburg-van Cittert,
1993). Compared to its worldwide distribution in the Mesozoic, the present-day distribution is greatly reduced and
fragmented. Like the Dipteridaceae, part of the reason for the restricted present-day distribution might be the rise to
dominance of deep-shade angiosperm forests toward the end of the Mesozoic (late Cretaceous).

The fronds of Matonia are quite distinct from those of Phanerosorus. In Matonia the petioles are erect and the blades
horizontal (or nearly so) and fan-shaped, the latter because of the repeatedly pedate leaf architecture at the base.
In contrast, the fronds of Phanerosorus are pendulous, elongate, and pinnate. Despite these differences, the leaves
on juvenile and mature plants of Phanerosorus (Kato & Iwatsuki, 1985) resemble fronds of juveniles of Matonia. Both
have segments that are linear-oblong, entire, and simple or forked. These similarities led Kato and Setoguchi (1999)
to suggest that the pinnate leaves of Phanerosorus evolved by paedomorphosis (the retention of the heteroblastically
young leaf form) from pedate ones like those of Matonia. This agrees with the fossil record because there are no
known fossils of Phanerosorus-like foliage.

Matonia contains two species: M. pectinata and M. foxworthyi (Kato, 1993). The two species of Phanerosorus are P.
sarmentosus and P. major.

The gametophytes of Matonia were described by Stokey and Atkinson (1952) and those of Phanerosorus by Yoroi
and Kato (1987). Both gametophytes provide evidence of a relationship to the other Gleicheniales (see discussion of
that Order).

Key to the Genera of Matoniaceae

1. Blades fan-shaped; segments pinnatisect; plants terrestrial....................................................................................................... Matonia

1. Blades long-trailing; segments entire or once-forked; plants epipetric................................................................. Phanerosorus

71
11. Dipteridaceae Diels | Dipteris family

Plants terrestrial or epipetric. Rhizomes creeping, solenostelic, two-ranked, bristly, the bristles dark, several cells
wide at the base. Fronds monomorphic. Petioles containing a C-shaped vascular bundle, the arms of the bundle
enrolled. Lamina divided to the top of the petiole into two equal halves (but entire or bifid in Cheiropleuria),
the halves flabellate, more or less deeply incised, the primary divisions anadromously divided on the acroscopic side
only, in living plants the base twisted through 90º (in pressed specimens this rotated 180º so that the abaxial surface
faces upward), chartaceous to subcoriaceous, glabrous. Veinlets branchig at right angles and joining to form a
fine mesh, included veinlets present. Sori soliatary in the meshes, sometimes appearing acrostichoid, nonindusiate.
Sporangia mixed with capitate paraphyses, the stalk short, of four rows of cells, annulus usually vertical, incomplete,
of about 12 cells, stomium ill developed. Spores monolete, nongreen, smooth, 64 per sporangium (ca. 128 in Cheirop-
leuria). Gametophytes epigeal, green, thalloid with thickened midrib, gametangia borne ventrally and sometimes dor-
sally; x=33.

Type species: Dipteris conjugata Reinw., of Malesia.

Distribution and ecology: Southeastern Asia, Malesia to Fiji. This present-day distribution is relictual; during the Me-
sozoic the family was cosmopolitan (Waterlot & Waterlot, 1979), more species-rich, and abundant in the vegetation.
Open areas, semi-open woods.

Genera/species: 2/11. Genera: Cheiropleuria (3 spp.), Dipteris (8 spp.).

Economic plants and products: none.

Discussion: The Dipteridaceae was once thought related to the Polypodiaceae on the basis of round non-indusiate
sori.This idea was refuted by Jarrett (1980) who demonstrated its distinctness from that family and the more derived
leptosporangiate ferns in general. Subsequent molecular studies have supported this conclusion (Hasebe et al. 1995;
Kato and Setoguchi, 1999; Pryer et al. 1995, 2004b).

The Dipteridaceae was cosmopolitan and abundant during the Mesozoic, even at high northern and southern lati-
tudes (Corsin & Waterlot, 1979; Ôishi & Yamasita, 1936; Waterlot & Waterlot, 1979). Its present-day range is relictual.
Part of the reason for the range restriction is probably the rise to dominance of deep-shade angiosperm forests
toward the end of the Mesozoic (late Cretaceous).The Dipteridaceae do not tolerate deep shade; they grow in open
or semi-open areas.

Cheiropleuria and Dipteris share the apparent morphological synapomorphies of capitate paraphyses, finely reticulate
venation with included veins, four-rowed sporangia stalks, and paracytic stomata. The two genera are distinguished
by few characters: protostelic rhizomes, frond dimorphy, soral shape, diplodesmic veins, and spore type. When more
species of Dipteris are analyzed cladistically, Cheiropleuria might be shown to be nested within Dipteris. Although the
fronds of Cheiropleuria look quite different from those of Dipteris, they are fundamentally similar. Both have the main
veins catadromously branched and similar finely reticulate venation. If a Dipteris were to evolve simple blades, they
would resemble those of Cheiropleuraia. The blades of Cheiropleuria vary from simple to bicuspid to quadricuspid
(the two pointed apices themselves are forked). Plant from Taiwan rarely have bi-cuspidate blades. Cheiropleuria was
thought to be monotypic, but Kato et al. (2001) recognized three species.

The gametophytes of Dipteris were described by Stokey (1945) and Nayar and Kaur (1971). The gametophyte of
Cheiropleuria was described by Stokey and Atkinson (1954).

72
12. Gleicheniaceae (R. Brown) C. Pres l Forked Fern Family

Plants terrestrial. Rhizomes long-creeping, frequently dichotomously branched, bristly or scaly, protostelic or (in one
species) solenostelic. Fronds monomorphic, indeterminate, discontinuous growth, clambering over surrounding vegetation,
borne in a single row on the dorsal surface of the rhizome. Petiole containing a C-shaped vascular bundle with enrolled
arms. Pinnae opposite, typically forked repeatedly with a resting bud between the forks, the ultimate divisions
with pectinate or pinnatifid. Rachises rounded or flattened adaxially (not grooved). Veins free. Stomata anomocytic
or diacytic. Sori round, abaxial, non-indusiate, with relatively few (2-10) sporangia. Sporangia opening across the top by
a vertical slit, the stalk very short and several cells wide; annulus oblique, not interrupted by the stalk. Spores mono-
lete or trilete, nongreen, smooth or sparsely ornamented, 100—800 per sporangium. Gametophytes epigeal, green,
thalloid, with thickened midrib and clavate hairs. Antheridia 6-12 celled. Chromosome number differing among the
genera: x=39, 43 in Dicranopteris; x=56 in Diplopterygium; x=20, 22 Gleichenia; x=34 in Sticherus.

Type species: Gleichenia polypodioides (L.) Sm., of South Africa (basionym: Onoclea polypodioides L.)

Distribution and ecology: Pantropical. Mostly open, sunny, disturbed habitats, especially compacted soils. Common
in roadsides, landslides, and pastures, especially at middle elevations, often forming dense colonies. Although weedy in
the wild, these ferns are extremely difficult to cultivate.

Genera/species: 6/130. Genera: Gleichenia (10 spp.), Dicranopteris (10), Diplopterygium (10), Sticherus (100), Gle-
ichenella (1), Stromatopteris (1).

Figure 44. Rhythmic leaf extension growth in the Gleicheniaceae. (From Moran, 2004)

Economic plants and products: In Asia the rachises of several species are tough and wiry and used for basket
weaving.

Discussion: The leaves of these ferns mostly climb through and on the surrounding vegetation, using it for support.
Their manner of frond elongation is ideally adapted for this.The Gleicheniaceae have an unusual growth habit adapted
for life in dense thicket vegetation (Moran, 2004). When a new leaf emerges from the rhizome, it grows straight up
through the surrounding vegetation and then stops to form a resting bud (Figure 444). Just below this bud develops

73
a pair of opposite pinnae. These unfurl and expand, eventually coming to rest on the surrounding vegetation. After
they have fully developed, the resting bud resumes growth, and the weight of the leaf is now supported by the pinnae
resting on the surrounding vegetation. This mode of growth provides not only support, but also easy passage for the
leaf apex to poke through dense vegetation because it is unencumbered by bulky lateral pinnae that would otherwise
snag surrounding twigs and vines. This growth habit is the secret to the group’s success in dense, scrubby vegetation
along roadsides. Because the rest-growth cycle repeats itself many times, some species of forked ferns have leaves
over 20 meters long—some of the longest leaves of any fern. Andersen and Øllgaard (1996) suggested a standard-

ized terminology that should be applied to the leaves of this family.

The family has received a fair amount of study. Chrysler (1943, 1944) examined the anatomy of the petioles and
stems. Holttum (1957a, 1957b, 1959) reviewed the morphology and growth habit and, along with Nakai (1950) pro-
posed a subfamilial classification. The gametophytes were described by Stokey (1950). The Gleicheniaceae has a fossil
recorded that extends back to the Permian (Collinson, 1996). It was common and widespread worldwide throughout
the Mesozoic.

Many species of Gleicheniaceae form dense impenetrable thickets. These can be controlled by cutting the leaves, al-
lowing them to dry, and then burning. Because the rhizomes are superficial (not subterranean) they are killed by fire.

Schizaeales A. B. Frank in Leunis

The most characteristic synapomorphy for this order is the complete subapical annulus. It is also characterized by dif-
ferentiation of the sterile and fertile parts of the leaf, and the absence of well defined sori.The order has been strongly
supported as monophyletic (Hasebe & al., 1995; Pryer & al., 2001a, 2004b; Skog & al., 2002; Wikström & al., 2002;
Labiak et al., 2015).Vegetatively the order is extremely diverse as is reflected by its three main clades, here recognized
as families.Various morphological and anatomical aspects of the order were reviewed by de la Sota & Morbelli (1987).

Within the order, the Lygodiaceae is sister to the Anemiaceae and Schizaeaceae (Figure 455).The estimated minimum
age for the split between Lygodiaceae and the wto other families is mid-Jurassic (Wikström & al., 2002).The family has
a fossil record that dates back to the early Jurassic (Collinson, 1996). The other genera have non-climbing fronds. In
Anemia the blades are horizontal or inclined to the soil, and the two basal pinnae are erect, lacking green laminar tissue
(or nearly so), and bear sporangia on their branched tips. The genus typically grows in dry or seasonally dry, open or
semi-open habitats. Most the genus has received monographic study by Mickel (1962, 1967, 2016).

13. Lygodiaceae M. Roem. | Climbing Fern Family

Terrestrial; rhizomes creeping, bearing fronds in a single orthostichy on the dorsal surface, branching dichoto-
mously at the apex, pubescent, protostelic; leaves indeterminate, the rachises twining around a support; pinnae
alternate, short-stalked, the stalk forked and often with a bud in the bifurcation; veins free or anastomosing; sori borne
marginally on the lobes, often in small finger-like projections (sorophores), but in some species the entire fertile seg-
ment is narrowed; sporangia solitary, one per vein tip, each sporangia covered by a green flap of tissue; spores
128-256 per sporangium, tetrahedral, trilete; gametophytes cordate, surficial, green; x=29, 30.

Type species: Lygodium scandens (L.) Sw. (basionym: Ophioglossum scandens L.)

Distribution and ecology: Tropical and warm-temperate regions worldwide.

74
Genera/species: 1/25. Sole genus: Lygodium (25 spp.)

Economic plants and products: In southern Thailand, Malaya, and Java, the rachises of Lygodium are used to weave
baskets, purses, and other small items. They are stripped of their pinnae, split lengthwise, and flattened, and then wo-
ven into a frame of other material.

Discussion: Lygodium has twinning rachises that use surrounding vegetation for support. Only Salpichlaena in the
Blechnaceae has a similar leaf habit (the twining rachis habit is not found in angiosperms). The pinnae of Lygodium
are born on short-stalks that soon fork and bear two opposite pinnules. A resting bud occurs in the fork, and it
normally does not develop unless the leaf apex is injured or dies. The sori are borne on short finger-like projections
(sorophores) along the margins of the pinnules, although the pinnules can be complete dimorphic in some species,
such as the eastern North American L. palmatum. Along the sorophore are two rows of sporangia, side by side, each
sporangium terminating a single vein and individually covered by a flap of green tissue. Actinostachys pennula has the
smallest plastome of all leptosporangiate ferns that have been fully sequenced thus far. (Labiak & Karol, 2017).

14. Schizaeaceae Kaulf. | Curly Grass Family

Plants terrestrial or epipetric. Rhizomes protostelic, bristly, the bristles multicellular. Petioles containing a single vas-
cular bundle. Blades simple, linear or fan-shaped, variously cleft; veins free, dichotomous; stomata in longitudinally
arranged rows; indusia absent; sporangia born on marginal, narrowed projections at the blade tips; sporangial capsules
pear-shaped to obovoid, sessile or short-stalked, the stalk with many rows of cells; annulus apical, complete. Spores
bilateral, monolete, nongreen, 128-256 per sporangium. Gameophytes epigeal, green, filamentous (Schizaea), or
subterranean and non-green (Actinostachys). x=77, 94, 103.

Type species: Schizaea dichotoma (L.) Sm. (basionym: Acrostichum dichotomum L.)

Distribution and ecology: Tropical and warm-temperate regions worldwide. Only one mainly temperate species:
Schizaea pusilla, primarily of northeastern North America. Many species are characteristic of sandy soils.

Genera/species: 2/30. Genera: Actinostachys (12 spp.), Schizaea (20).

Economic plants and products: None.

Discussion: The monophyly of the family is well supported by molecular phylogenetic studies (Hasebe et al., 1995;
Pryer et al., 1995; Wikstöm et al., 2002, Skog et. al., 2002; Labiak et al., 2015). Monolete spores are a synapomorphy
for the family because the closest outgroup families have trilete spores.The Schizaeaceae has a fossil record extending
back to the Jurassic (Wikström et al., 2002).

Actinostachys and Schizaea are both are monophyletic (Wikstöm et al., 2002; Labiak et al., 2015). Their leaves are
typically grass-like and bear sorophores apically. The sorophores in both genera are digitate. In Schizaea they are dis-
tinctly digitate, but in Actinostachys the fertile rachis is very short, imparting a digitate appearance. The fertile pinnae
in Schizaea are strongly ascending, and in Actinostachys they are vertical. The number of rows between the costa and
margin differ: in Schizaea has only one, whereas Actinostachys has two (i.e., two and four total, respectively).The spores
of both genera differ. In Schizaea they are foveolate or granulate with spherical deposits, whereas in Actinostachys they
are longitudinally parallel-striate (Fig. 21A,B).

The gametophytes of Actinostachys and Schizaea are mycotrophic but otherwise quite distinct. Those of Schizaea are
filamentous and branched--a character found elsewhere among ferns only in Trichomanes (Hymenophyllaceae). The
filaments bear rhizoides only on specialized globose cells, called rhizoidophores, that occur along the length of the fil-
aments (Britton, 1902; Kiss & Swatzell, 1996).The rhizoides are penetrated by fungi that grow through the rhizoid and
into the rhizoidophore. Once in the rhizoidophore, the fungus swells to form bladders that nearly fill their host cells.
The fungus rarely invades the filamentous cells. The development and morphology of Schizaea pusilla gametophytes

75
were reviewed by Davidson et al. (2008) and Swatzell et al (1996).The gametophytes of Actinostachys are, in contrast,
elongate-tuberous, subterranean, and achlorophyllous (Bierhorst, 1971).

Davidson et al. (2008)

Actinostachys oligostachya Bierh. is atypical by producing only one leaf at a time.

15. Anemiaceae Link

Terrestrial; rhizomes creeping to suberect, solenostelic (subgen. Anemiorrhiza) or dictyostelic (subgen. Anemia +
the Mohria clade) pubescent; leaves determinate, typically hemidimorphic, rarely holodimorphic; veins free or rarely
anastomosing; sporangia usually borne on a basal pair of erect pinnae; spores 128–256 per sporangium, tetra-
hedral, striate with prominent parallel ridges; gametophytes cordate, surficial, green; x=38.

Type species: Anemia phyllitidis (L.) Sw. (basionym: Osmunda phyllitidis L.)

Distribution and ecology: Primarily neotropical, but 16 in Africa-Madagascar, and one in southrn India and islands
in the Indian Ocean.

Genera/species: 1/115. Sole genus: Anemia.

Economic plants and products: None.

Discussion: The family has been strongly supported as monophyletic (Labiak et al., 2016).The rhizome anatomy--spe-
cifically, the presence of leaf gaps--is synapomorphic for the Anemiacae. It is either solenostelic (subgen. Anemiorrhiza)
or dictyostelic (subgen. Anemia + the Mohria clade); in contrast, the Lygodiaceae and Schizaeaceae are protostelic.

In most instances, the field identification of the family is easy because nearly all the species have two erect basal
pinnae that bear sporangia. Anemia colimensis Mickel, from Mexico, is unusual by having basal pinnae that are fertile
and spreading (not erect), with the green lamina tissue only slightly contracted. Also atypical are several species with
holodimorphic sterile and fertile leaves, such as A. millefolia (Gardner) C. Presl and A. wrightii Baker.

Anemia is divided into two subgenera (Figure 45). The first, subgen. Anemia (including Mohria Sw.), is characterized by
dictyostelic rhizome anatomy, red, brown, or orange hairs on the rhizomes, and polystichous leaves. It contains species
ranging from diploids to dodecaploids. The second, subgenus Anemiorrhiza Prantl, has solenstelic rhizomes, blackish
hairs on the rhizomes, and distichous leaves. Most of its species have small ranges and are restricted to the Caribbean
basin. The genus is well known taxonomically thanks to the work of Mickel (1962, 1967, 1982, 2016).

76
Figure 45. Phylogeny of the Anemiaceae. (Labiak et al., 2015)

77
 Salviniales | Water Ferns
The Salviniales, a group of about 70 species, are the only heterosporous ferns. In fact, they are the only plant group
to have evolved heterospory since the Paleozoic (Nangalingum et al., 2006). Their megasporangiate sori bear a single
megaspore (a condition sometimes referred to as monomegaspory), a character otherwise restricted, among extant
plants, to the seed plants (Bateman & DiMichele, 1994). As suggested by their common name, these plants grow ei-
ther floating or rooted in wet soils that may be dry part of the year.The clade (Figure 46) consists of two families: the
Salviniaceae (Azolla and Salvinia) and Marsileaceae (Marsilea, Regnellidium, Pilularia). The monophyly of the Salviniales
is strongly supported by DNA sequence data and morphology (Hasebe et al., 1995; Nangalingum et al., 2008; Pryer
et al., 1995; Stevenson & Loconte, 1996).

Figure 46. Phylogram of the Salviniales. The average branch lengths obtained by Bayesian inference using atpB, rbcL, and rps4.
Numbers above the branches are MP bootstrap/ML bootstrap/BI posterior probability. MPBS and MLBS with “*” = 100%; PP
with “*” = 1.0. (From Nagalingum et al., 2008.)

78
16. Salviniaceae Dumortier

Plants aquatic, free-floating, small. Rhizomes thin, long-creeping, frequently branched, protosetlic or solenostelic, gla-
brous. Fronds 1-30 mm long, not circinate. Sori on the submerged (ventral) side of the plant, of two types, either
megasporangiate or microsporangiate, borne at the vein tips; indusia thin, globose, whitish. Sporangia staked; annulus
absent. Spores trilete, nongreen. Megasporangia one-spored at maturity. Gametophytes endosporic, nongreen.
x=9 (Salvinia; the lowest known in any fern), 22, 26 (Azolla).

Type species: Salvinia natans (L.) All. (basionym: Marsilea natans L.), of Europe and Asia.

Distribution and ecology: Cosmopolitan, mainly tropical. Aquatic habitats, especially stagnant or slow-moving water,
muddy shores, vernal ponds.

Genera/species: 2/18. Genera: Azolla (7 spp.), Salvinia (11).

Economic plants and products: Azolla is a rich source of nitrogen because of a cyanobacterium (Anabaena azollae)
in its leaves. It has been used for centuries as an organic fertilizer in the rice paddies of southern China and Vietnam,
and sometimes it is fed as a dietary supplement to pigs and chickens. Salvinia molesta, a native of southern Brazil, is one
of the world’s worst aquatic weeds. It carpets the surface of waterways, forming dense mats that impede navigation,
clog intake valves, smother aquatic life, and prevent fishing. Successful control of S. molesta has been achieved in many
parts of the world by introducing a curculionid weevil that feeds only on Salvinia (Moran, 2004).

Discussion: Azolla and Salvinia are quite different structurally. Azolla has bilobed leaves about 1 mm long, with the
lower lobe thin, nongreen and resting on the water. The dorsal lobe is thick, green, arched upward, and contains a
cavity for the Anabaena symbiont. Sori are rarely produced, but when present are produced on the lower lobe, which
then becomes reduced to veins and bear sori at their tips. In contrast, Salvinia has at each node three leaves, two of
which are green, rounded, and floating, and the third submerged, nongreen, finely dissected and root-like. Sori are
produced only on the submerged leaf. Salvinia reussii, a fossil species, shows an intermediate condition with the sub-
merged leaf finely dissected and root-like, but with several expanded laminar segments (Figure 47).

Figure 47. Salvinia reussii, a Miocene fossil species with expanded laminae on submerged leaf. (from Collinson, 1991.)

79
The sori in this family have sometimes been referred to as “sporocarps,” but they are not homologous with true spo-
rocarps in the Marsileaceae and therefore should not be called as such.The sporocarp of the Marsileaceae consists of
many sori within a highly modified pinna, one that has become over the course of evolution folded lengthwise, sealed
along its margins, and sclerified. In contrast, the “sporocarp” of the Salviniaceae consists of a receptacle, sporangia, and
indusium. In other words, it is a sorus. There is no highly modified, enclosing pinna as in the Marsileaceae.

For a floating aquatic plant, wetting the upper surface of the leaves is a serious problem that could lead to water log-
ging and sinking, or to eventual damage by fungal growth. The floating leaves of Salvinia exhibit exquisite architecture
preventing this. Their upper surfaces are covered by hydrophobic (water-repelling) wax. When water splashes onto
the surface, it beads-up into nearly perfect spheres and readily rolls off. Thus, it does not wet the surface. This wa-
ter-shedding effect is enhanced by closely spaced, parallel rows of wax-covered hairs or papillae on the upper surfaces
of the leaves. Water droplets tend to sit on top of these structures; thus, contact of the droplet with the leaf surface
is further reduced (Barthlott et al., 2009).

In Salvinia, the hairs or papillae on the adaxial side of the floating leaves differ in height, shape, and composition. Four
types have been discerned, each named for a representative species (Barthlott et al., 2009). The first is the “cucullata
type.” These are solitary, multicellular slightly bent hairs of 6–10 cells. It is found in S. cucullata and S. hastata.

The second is the “oblongifolia type.” It consists of groups of two multicellular, uniseriate hairs bent in the same direc-
tion. The two hairs are joined by their penultimate cell. This type occurs only in S. oblongifolia.

The third is the “natans type.” It consists of four multicellular, uniserite, 6–8-celled hairs.These are perched on a papilla
(technically called an “emergence”) and spread away from each other. This is characteristic of S. natans and S. minima.

The fourth is the “molesta type.” It resembles and egg-beater and consists of a group of four hairs, each 6–8-celled,
fused near their tips by penultimate apical cells. Each group is elevated on a large papilla (emergence). These struc-
tures are the largest in the genus, generally 1–2 mm long. Unlike the other three types, the free apical cells are not
covered by epicuticular waxes. For that reason they are hydrophilic (water-attracting).This type is found in S. auriculata,
S. biloba, S. herzogii, and S. molesta.

The ontogeny of the “molesta type” structures (= hairs + emergence) was studied by Barthlott et al. (2009). They
found that the hairs develop first, fully within the folded conduplicate leaf before it opens. Each hair divides until it
is 6–8 celled. At this stage, the apical cells are larger than the basal ones. The emergence (papilla) then arises from
subepidermal tissue beneath the group of four hairs. As this happens, the subapical (penultimate) cells of the hairs
fuse, and the apical cells elongate horizontally, whereas all the basal cells elongate vertically. The apical cells soon die
and wither. The emergences continue to lengthen after the leaf opens.

Collinson (1991, 1992) reviewed the extensive fossil record of the Salviniaceae. Azolla is known from whole fertile
fossil plants from the Paleocene onwards, and from dispersed megaspores in the late Cretaceous. Over 30 fossil
species of Azolla have been recognized, in contrast to the six present-day ones. In the fossil record, Salvinia is mostly
known from dispersed spores, with whole-plant fossils being relatively rare.

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17. Marsileaceae Mirbel | Clover Fern Family

Plants aquatic or amphibious. Rhizomes creeping, slender, solenostelic, often with several internal air canals in the cor-
tex, hairy or scaly, often glabrescent. Fronds monomorphic, circinate in bud. Blades of 1 or 2 pairs of opposite pinnae
or (in Pilularia) absent. Pinnae cuneate, the apex rounded or truncate. Veins free or netted. Stomata diacytic or ano-
mocytic. Sori borne within hardened bean-like structures called “sporocarps”, these indehiscent, usually hairy, borne
singly to many at the base of the petioles, short-stalked. Indusium present. Sporangia of two types, megasporangia and
microsporangia, each born in the same sorus and sporocarp. Spores trilete, globose, nongreen; microspores 16—64
per sporangium, megaspores 1 per sporangium. Gametophytes endosporic, nongreen. x=20 and (in Regnellidium)
x=19.

Type species: Marsilea quadrifolia L., of Europe and Asia.

Distribution and ecology: Cosmopolitan. Although Marsilea and Pilularia are widespread, Regnellidium is known
from only three sites in southern Brazil and northern Argentina. Shores, seasonally ponds, shallow stagnant water. The
plants produce sporocarps only when the habitat dries; they usually remain sterile when growing in water.

Genera/species: 3/55. Genera: Marsilea (45 spp.), Pilularia (7), Regnellidium (1).

Economic plants and products: Several species of Marsilea are occasionally cultivated as ornamentals, especially M.
mutica, which has shiny variegated pinnae.

Discussion: Based on analyses of rbcL and morphology, Pryer (1999) found that Pilularia and Regnellidium are more
closely related to one another than either is to Marsilea (Figure 46).

The sporocarps represent fertile pinnae that have been folded longitudinally, fused along their margins, and sclerified
(Puri & Garg, 1953). They generally bear within 10 to 20 sori. The sporocarps germinate when their outer wall is
weakened and the gelatinous material within imbibes water, thus expanding and cracking the wall further. In Marsilea
the sori are carried out of the germinating sporocarp on a tail-like mucilaginous structure called the sorophore.
Pilularia and Regnellidium also have gelatinous sorophores, although these are not tail-like (Nagalingum et al., 2006).

Sporocarps up to 130 years old have been germinated and found to produce viable spores, and sporocarps contain-
ing spores capable of producing viable sporophytes have been documented at up to 100 years old (Johnson, 1985).
They might be an adaptation for life in arid regions where rainfall in infrequent or seasonal.

A thick, translucent gelatinous sheath (the perine) around the megaspore acts as a floating organ and is a synapomor-
phy for the family. In each of the family’s three genera, the sheath is slightly modified in various ways. Schneider and
Pryer (2002) found that floating megaspores of Marsilea form a meniscus at the water’s surface. The meniscus helps
funnel sperm into a hollow channel (sometimes called the “sperm lake”) at the apex of the gelatinous sheath, thus
propelling the sperm toward the archegonium.

A remarkable Late Cretaceous fossil fern, Hydropteris pinnata, combines characteristics of the Marsileaceae and Sal-
viniaceae (Rothwell & Stokey, 1994). It resembles Marsilea in by long-creeping rhizomes, leaves with several cuneate
pinnae, and a stalked sporocarp arising from the petiole base. It resembles the Salviniaceae, however, by bearing me-
gaspore complexes (megaspores + aborted megaspores or “floats,” and any attached indusial parts), glochidia on the
megaspore surfaces, and microsporangiate massulae. A cladistic study by Pryer (1999) found that the placement of
Hydropteris was unresolved with respect to the other genera of Pilulariales. This is a remarkable conclusion because
the sporocarps of Hydropteris and Marsileaceae both represent basal fertile pinnae that have become folded, sealed
along the margins, and sclerified (although apparently the sporocarps of Hydropteris were not sclerified). These steps
in the evolution of the sporocarp (i.e., the folding and sealing) were not, for some unknown reason, included in the
cladistic analyses of water ferns by Rothwell and Stockey (1994), Bateman (1996), or Pryer (1999).

Water clover, the common name of Marsilea, refers to the resemblance between the leaves of Marsilea and those of

81
clovers (Trifolium, Fabaceae).The leaves can be notoriously variable depending on whether they are submerged, float-
ing, or aerial. The submerged or floating leaves are typically entire, whereas the aerial ones are often lobed (Launert,
1968). The submerged and aerial leaves also show differences in venation and the development of air canals in the
rhizomes and petioles (Gaudet, 1964). Marsilea is the only fern genus with true pulvini (not pulvinuloids as in the
Marattaiceae) and the only nyctinastic fern. As documented by Darwin (1869), the action of the pulvini at the base
of the four pinnae result in a folding upward, so that the four pinnae forma “packet,” as darkness approaches in the
late afternoon.

An important phenomenon in the evolution of leaves in the Marsileaceae has been “juvenilization”, or accelerated
growth rate and early termination at a simplified leaf form (Pryer & Hearn, 2008).

Figure 48. Spore-bearing structures of Marsiliaceae (A, B) and Salviniaceae (C. D). A, Longitudinal section of a Marsilea
sporocarp. B. Transverse section of Pilularia sporocarp. C. Fertile leaf (submerged) of Salvinia. D. Azolla nilotica, reproductive
structure (only this species bears 4 sori per structure). In = indusium. Me = megasporangia. Mes = megasporangiate sori. Mi =
microsporangia. Mis = microsporangiate sori. Pl (se) = parenchymatous layer or sporophore envelope. S= sorophore. V=vein.
Scale bars = 1 mm. (From Nagalingum et al., 2006.)

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 Cyatheales A. B. Frank in Leunis | Tree Fern Order
The compositionof and relationships within this order are an excellent example of the many surprises served up
by phylogenetic analyses of DNA data. Traditionally, the “tree ferns” consisted of the Cyatheaceae, Dicksoniaceae,
Lophosoriaceae, and Metaxyaceae. It has now been shown that these families form a clade with the Loxomataceae
and Plagiogyriaceae—a relationship previously completely unexpected (Hasebe et al., 1995; Pryer et al., 2004; Wolf
et al., 1999; Korall et al., 2006, 2007). Furthermore, it has been shown that the Lophosoriaceae should be classified
in the Dicksoniaceae, and that the Hymenophyllopsidaceae (endemic to the tepui region of southern Venezuela) is
nested in Cyathea.

Apparent morphological synapomorphies for the order are erect, radially symmetrical stems, and transverse cells
in the phloem (that is, a narrow band of phloem sieve-cells with their long axes oriented transversely to the other
phloem cells that are elongate longitudinally; Holttum & Sen, 1961). Spore germination (polar vs. equatorial) and ga-
metophytes might provide additional synapomorphies, but these need to be documented for more species.

Key to Genera of non-scaly Cyatheales

1. Sori abaxial
2. Leaves pinnatisect or 1-pinnate
3. Leaves 1-pinnate, apex conform; sterile and fertile leaves monomorphic; plants below 800 m...........Metaxya
3. Leaves pinnatisect to 1-pinnate basally, apex tapered; sterile and fertile leaves
dimorphic plants above 2000 m................................................................................................................................................Plagiogyria
2. Leaves 3- to 4-pinnate, sterile and fertile leaves dimorphic.......................................................................................Lophosoria
1. Sori marginal
4. Stems creeping, slender...............................................................................................................................................................Loxsomopsis
4. Stems erect, massive
5. Indusia cup-shaped; proximal fertile pinnae skeletonized. Juan Fernández Islands.............................Thyrsopteris
5. Indusia clam-shaped; proximal fertile pinnae with expanded green lamina.
6. Rachises and costae grooved adaxially.
7. Laminae 3- to 4-pinnate. Neotropics, Azores, Madeira, w. Spain................................................................. Culcita
7. Laminae 2-pinnate. Malesia.......................................................................................................................................Cystodium
6. Rachises and costae flat or slightly rounded adaxially.
8. Laminae anadromously dissected. Java and Philippines to Samoa...................................................Calochlaena
8. Laminae catadromously dissected.
9. Laminae narrowed toward the base; stems erect, arborescent......................................................Dicksonia
9. Laminae not or only slightly narrowed toward the base; stems not arborescent................. Cibotium

83
Figure 49. Phylogeny of the Cyatheales. Fifty percent majority rule consensus tree from Bayesian analysis of a combined atpA,
atpB, rbcL, and rsp4 data set. Numbers above branches represent support values from Bayesian, maximum likelihood, and
maximum parsimony analyses, respectively. A dash (-) represents bootstrap percentage <50%. A plus (+) denotes 100% support.
An X indicates reversal of arborescent habit. From Korall et al. (2006).

84
18. Thyrsopteridaceae C. Presl

Plants terrestrial; stems erect to decumbent, to ca. 2 m tall, massive, solenostelic, sometimes producing stolons, pubes-
cent, the hairs stiff, multicellular; leaves 2-3.5 m long, partially dimorphic, the fertile ones with strongly modified lower
pinnae; petioles with long, matted hairs; laminae 3-5-pinnate, broadly ovate, tapered to the tip; pinnae and pinnules
stalked, ascending; rachises and costae grooved adaxially; veins free; sori terminal on narrow, non-laminate
segments, often restricted to the proximal pinnae; receptacle columnar, clavate; paraphyses absent; indusia
cup-shaped, entire; spores tetrahedral-globose, trilete, with prominent angles. x= ca. 76–78.

Type and sole species: Thyrsopteirs elegans Kunze.

Distribution and ecology: Endemic to the Juan Fernández Islands off the coast of Chile, where it grows in moist
woods and thickets between 500–1000 m.

Genera/species: 1/1.

Economic plants and products: none.

Discussion: The phylogenetic placement of the Thyrsopteridaceae with respect to other families of Cyatheales is
uncertain (Korall et al., 2006). Fossil representatives of this family were common and widespread during the Mesozoic
(Harris, 1961).

19. Loxomataceae C. Presl

Plants terrestrial. Rhizomes long-creeping, siphonostelic, densely covered with dark, rigid bristles with mul-
ticellular base. Petioles containing a U-shaped vascular bundle. Fronds monomorphic. Blades 2- to 3-pinnate, de-
tlate. Veins free. Stomata cryptopore, paracytic. Sori marginal, terminal on the vein tips, with an enlarged cylindrical
receptacle. Indusia urceolate. Sporangia pyriform or nearly so, on short, thick, pluricellular stalks, dehiscing vertically
or transversely; annulus slightly oblique, continuous but only partly indurated. Spores trilete, tetrahedral-globose, non-
green. Gametophytes cordate to elongate, with multicellular and often pluriseriate hairs on the cushion; antheridia
pluricellular, often with a divided cap cell, sometimes present on the dorsal surface. x=46 (Loxsomopsis), 50 (Loxoma).

Type species: Loxoma cunninghamii R. Brown.

Distribution and ecology: Neotropics (Costa Rica, Colombia to Bolivia) and New Zealand (North Island). Loxso-
mopsis occurs in moist, open places (often disturbed) in cloud forests. Loxoma occurs in lowland forests, and its sole
species is endemic to North Island of New Zealand

Genera/species: 2/2. Genera: Loxoma (1 sp.), Loxsomopsis (1).

Economic plants and products: none.

Discussion: This family is sister to a clade containing Plagiogyria and Culcita (Korall et al., 2006). No known morpho-
logical synapomorphies have been noted for this clade. Fossils of the family are known from the Jurassic (Stachypteris)
and Cretaceous. The gametophytes were described by Stokey & Atkinson (1956b)..

85
20. Culcitaceae Korall

Plants terrestrial; rhizomes creeping or ascending, not arborescent, solenostelic, densely pubescent, the hairs 2-5 cm
long, golden-brown, multicellular; leaves monomorphic; petioles about half the length of the leaf, with an ome-
ga-shaped vascular bundle (the open end oriented adaxially); laminae deltate, 4-5-pinnate-pinnatifid; rachises
and costae grooved, the grooves deep and decurrent into those of the lesser order; veins free; sori ter-
minal on the veins, to 3 mm wide, paraphysate; indusia clam-shaped, outer indusia scarcely differentiated from
the laminar tissue, but the inner noticeably modified in color or texture, fitting inside the upper indusia lobe; spores
tetrahedral-globose, trilete; x=66.

Type species: Culcita macrocarpa C. Presl.

Distribution and ecology: Neotropics (C. coniifolia (Hook.) Maxon), Azores, Madeira, Tenerife, southwestern Eu-
rope (C. coniifolia). The neotropical species occurs mostly in cloud forests above 2000 m.

Genera/species: 1/2. Sole genus: Culcita.

Economic plants and products: none.

Discussion: Culcita was previously classified with the Dicksoniaceae, but it has recently been shown to be distinct
and sister to the Plagiogyriaceae (Korall, 2006). Similarly, Calochlaena (seven species of Dicksoniaceae from Malesia,
Australia, and Polynesia) was long associated with Culcita, but it was shown to be distinct on the basis of anatomy
(Schneider, 1996a; White & Turner, 1988).

21. Plagiogyriaceae Bower

Plants terrestrial. Stems short, erect or suberect, radially symmetrical, surrounded by persistant petiole bases, dic-
tyostelic, lacking hairs or scales, occasionally producing stolons. Fronds dimorphic, the fiddleheads mucilaginous, with
lateral peg-like aerophores (at least when young). Petioles with enlarged bases, more or less triangular in cross sec-
tion, flat adaxially, carinate abaxially, containing a single U- or V-shaped vascular bundle with laterally extended arms,
or with three separate vascular bundles that unite into one distally. Blades pinnatisect or 1-pinnate, glabrous. Pinnae
entire or serrate to biserrate. Rachis grooved adaxially, carinate abaxially. Veins free. Stomata anomocytic. Sporangia
along the veins, protected by the revolute margin, mixed among slightly capitate paraphyses; stalk of 4-6 rows of cells,
elongate; annulus oblique, nearly complete, not interrupted at the stalk, the stomium well-defined. Spores trilete,
tetrahedral-globose, nongreen, 48 per sporangium. Gametophytes epigeal, green, cordate, with strongly developed
wings; gametangia borne on both surfaces; antheridia with many cells. Chromosome number variable, with n=66, 75,
100, ca. 132 having been reported.

Type species: Plagiogyria semicordata (C. Presl) H. Christ (basionym: Lomaridium semicordatum C. Presl)

Distribution and ecology: Tropical and warm-temperate parts of the Neotropics and Asia (Figure 50), with most
species in southern China. Middle to high elevations in moist, open habitats or shaded forests.

Genera/species: 1/11. Sole genus: Plagiogyria.

Economic plants and products: none.

Discussion: Plagiogyria is sister to Culcita, with which it has little in common morphologically (Korall et al., 2006).
Plagiogyria has no fossil record. Superficially, it resembles a Blechnum with dimorphic leaves, but it differs by lack of

86
scales on the rhizomes and petiole bases, rachises carinate abaxially, three or fewer vascular bundles in the petioles
(Blechnum has more than three), and oblique annuli.

The mucilage covering the fiddleheads is secreted by tiny hairs with swollen terminal cells (Hennipman, 1969). The
aerophores protrude through this mucilage. Stokey and Atkinson (1956) described the gametophytes of two species.

The genus was monographed by Zhang and Nooteboom (1998), but they did not fully assess the neotropical species.
They recognized 10 species in the Old World and one in the Neotropics. Lellinger (1971) recognized six species
in the Neotropics. One species from Peru and Bolivia (perhaps undescribed?) has an arborescent trunk up to 1 m,
resembling an arborescent Blechnum (subg. Lomariocycas).

Figure 50. Distribution of the Plagiogyriaceae. From Zhang and Nooteboom (1998).

87
22. Cibotiaceae (Nayar) Korall

Plants terrestrial; rhizomes massive, either creeping or erect (to 6 m), solenostelic or dictyostelic, pubescent, the hairs
golden, long, soft; Fronds monomorphic, 2-4 m long; petiole bases densely pubescent, with three corrugated vascular
bundles arranged (more or less) in an omega shape; laminae 2-pinnate-pinnatifid to 3-pinnate; pinna rachises and
costules ridged adaxially; veins free; sori marginal, terminal on the vein tips; indusia bivalvate, each lobe with a strongly
differentiated, non-chlorophyllous outer indusium and a similarly modified tongue-like inner one; paraphyses filiform;
spores tetrahedral-globose, trilete, with prominent angles and a well-developed equatorial flange; anteridial walls
5-celled; x=68.

Type species: Cibotium chamissoi Kaulf.

Distribution and ecology: Central America (2 spp.), eastern Asia (3 spp.), Hawaii (6 spp.). In Central America it
occurs in middle elevation rain forests.

Genera/species: 1/11.

Economic plants and products: The stem apices and petiole bases of Cibotium barometz, which are thickly covered
by golden hairs, are fashioned into small lamb- or dog-like objects and sold to tourists. In parts of China, these crea-
tures are kept in households and the hairs are pinched off and used to staunch bleeding.

Discussion: Cibotum was formerly classified in the Dicksoniaceae, but phylogenetic studies show it to be distinct
(Korall et al., 2006). Its chromosome number of x=68 is unique in the Cyatheales. Also, its spores are distinctive in
the Cyatheales, having a prominent equatorial flange and thick, nearly parallel ridges on the distal face (although these
ridges are reduced or absent in the two neotropical species). Lophosoria also has a prominent equatorial flange, but
it lacks the distal ridges. The Hawaiian species were studied by Palmer (1994).

23. Metaxyaceae Pic. Serm.

Plants terrestrial or epipetric. Rhizomes short-creeping, dorsiventral with leaves borne on the upper surface, usually
4-ranked, solenostelic, hairy, the hairs several-celled, tawny to golden. Fronds monomorphic, fiddleheads circinate.
Petioles rarely with adventitious buds, containing a single corrugated horseshoe-shaped vascular bundle. Laminae
1-pinnate, with a conform terminal pinna, glabrous to sparsely hairs beneath. Pinnae entire, caudate-serrate api-
cally, the margins cartilaginous. Rachises and costae grooved adaxially.Veins free, closely parallel from costa to margin.
Sori round, borne on the abaxial surface (not marginal), 1-3 per vein, irregularly dispersed between the costa and
margin but mostly toward the costa; indusium absent. Stomata paracytic-laterocytic. Sporangia numerous, matur-
ing simultaneously, mixed with conspicuous hairs (i.e., paraphyses); stalks short, with 4 rows of cells; annulus slightly
oblique, bypassing the stalk, with a distinct several-celled stomium. Spores trilete, tetrahedral-globose, nongreen, 64
per sporangium. Gametophytes epigeal, green, cordate to ribbonlike, with a thickened central cushion, lacking thick
multicellular hairs; gametangia borne ventrally; antheridia usually with 5 or a few more cells. x= 95, 96.

Type species: Metaxya rostrata (H.B.K.) C. Presl (basionym: Aspidium rostratum H.B.K.)

Distribution and ecology: Neotropics. Moist, shaded, forest floors, usually under 800 m.

Genera/species: 1/2.

Economic plants and products: none.

88
Discussion: Although exact relationships are unclear, Metaxya is likely the sister group of a clade comprising Dickso-
niaceae, Cibotiaceae, and Cyatheaceae (see cladogram to fern families in front of this book; Korall et al., 2006).

Although a member of the tree fern clade (Cyatheales), Metaxya does not have a trunk; instead, it has a creeping
dorsiventral rhizome with usually four-ranked leaves (Qiu et al., 1995). It is unique among ferns by sometimes bearing
two (rarely three) sori per vein (one sorus per veins is most common, though). The two sori along a single vein arise
by subdivision of a single large patch of sporangia initial cells (Qiu et al., 1995). Another unusual aspect is that the small
leaves (< 15 cm long) are quite unlike longer ones, having deeply incised pinnae (Tuomisto & Groot, 1995).The family
has received considerable study of its morphology, anatomy, and spores (Gastony & Tryon 1976; Holttum & Sen, 1961;
Lucansky, 1974, 1982; Lucansky & White, 1974; Qiu et al., 1995; Roy & Holttum, 1965). For a long time it was thought
to consist of a single species, but Smith et al. (2001) have found a second species, M. lanosa, from Amazonian Venezu-
ela and Peru. Unlike some other Cyatheales (i.e., Cyatheaceae and Dicksoniaceae), the gametophytes of Metaxya lack
multiseptate hairs on the dorsal surface (Qiu et al., 1995).

24. Dicksoniaceae Bower

Plants terrestrial; stems erect, ascending or rarely creeping, often massive and arborescent, covered with roots and per-
sistent petiole bases, solenostelic (if creeping) or dictyostelic, hairy, the hairs generally 2-5 cm long, pluricellular,
golden to dark brown. Fronds monomorphic or commonly 1-3 m long; petioles wooly at the base, scales absent,
containing 3 corrugated vascular bundles; laminae 2- to 5-pinnate; rachises and costae not grooved adaxially; veins
free; stomata diacytic, paracytic; sori marginal terminal on the veins, or (in Lophosoria) superficial and sometimes more
than one along a vein (a character unique in ferns); indusia clam-shaped or (in Lophosoria) absent, when present the in-
ner (abaxial) valve membranous to chartaceous, the outer (adaxial) valve commonly thicker and green to light brown
modified lamina margin; sporangia numerous, maturing simultaneously, short- or long-stalked, the stalk 4—6 rows of
cells, mixed with hairs (i.e., paraphyses); annuli oblique, not interrupted at the stalk; spores trilete, tetrahedral-globose,
64 per sporangium; gametophytes epigeal, green, thick, obcordate to cordate-elongate; antheridia with 5 or more cells.
x=56 (Calochlaena), 65 (Dicksonia, Lophosoria).

Type species: Dicksonia arborescens L’Hértier, of St. Helena.

Distribution and ecology: Pantropical and warm-temperate areas, but absent from Africa. Most species occur in
forest, especially semi-open ones, usually above 1000 m.

Genera/species: 3/28. Genera: Dicksonia (20 spp.), Calochlaena (5), Lophosoria (3).

Economic plants and products: The root mantles of Dicksonia are used in horticulture as a substrate for growing
epiphytes, or sculpted into various forms and sold as handicrafts.

Discussion: The Dicksoniaceae has been drastically re-defined to conform to the phylogeny reported by Korall et
al. (2006) and Wolf et al. (1999). It now includes Lophosoria, formerly placed in its own monotypic family (Kramer,
1990; Tryon & Tryon, 1982), and excludes Cibotium, Culcita, and Thyrsopteris. The latter three families do not form a
monophyletic group and are now placed in their own families. Defined as such, the Dicksoniaceae has no apparent
morphological or anatomical synapomorphies.

Lophosoria is a large terrestrial fern with massive rhizomes usually less than one meter tall. The fronds are large and
dissected, glaucous benath, and bear abaxial, nonindusiate sori with hairs mixed among the sprangia. Its spores are
distinctive by having a prominent subequatorial flange, with the proximal face tuberculate and the distal face perforate.
The gametophytes were described by Pérez-García et al. (1995). The buds on the petiole bases have been exam-
ined anatomically (Lucansky, 1974, 1982) but not morphologically. The buds form stolons, which explains the plant’s
clumped distribution in the field. The buds and stolons need to be studied by excavation in the field to determine
their frequency, branching patterns, and habit.

89
Figure 51. Cyatheaceae. A1. Cyathea sp., habit. A2. Trunk and development of the root mantle toward the base. A3. Leaf
scars on trunk. A4. Cross-section of trunk showing meristeles surrounded by a dark sclerenchyma sheath. A5. Sporangium,
oblique annulus. A6. Sorus.

90
Formerly, Lophosoria was considered monotypic, but a second species (Lophosoria quesadae A. Rojas) occurs in Costa
Rica (Rojas-Alvarado, 1996) and a third (L. contracta (Hieron.) A. R. Sm.) in southern Ecuador and Peru.

Lophosoria was more widespread in the Southern Hemisphere during the Cretaceous. Fossil spores of Cyatheacidites,
which closely resemble those of Lophosoria quadripinnata, are known from the Cretaceous of Australia and Antarctica
(Dettmann, 1986). Foliage fossils and spores of L. cupulata D. J. Cantrill from Antarctica are considered closely related
to extant L. quadripinnata.

25. Cyatheaceae Kaulf. | Scaly Tree Fern Family

Plants terrestrial or rarely epiphytic or hemiepiphytic. Stems arborescent rarely short-erect, scaly, dictyostelic, the
meristeles surrounded by sclerenchyma. Leaves monomorphic, generally 1-4 m long. Blades 1- to 4-pinnate, but most
commonly 2-pinnate-pinnatifid, without buds, divided catadromically. Petioles scaly, containing three vascular bundles,
two adaxial and one abaxial, these corrugated or not. Rachises often with a dark gland abaxially at the point of
pinna attachment, grooved to slightly rounded adaxially, often pubescent adaxially, the hairs antrorse, several-celled.
Veins free or (less commonly) the basal pair uniting. Stomata anomocytic, polocytic or acyclicparacytic. Sori abaxial,
round, usually borne at the fork of a vein, the receptacle elevated or globose or subcylindrical. Sporangia many
per sorus, short-stalked (appearing sessile), the stalk with 4 rows of cells, the capsules often angular and obconic; in-
dusia present or absent, when present inserted at the base of the receptacle, and if not complete surrounding the
receptacle inserted on the costular side; annuli oblique, not interrupted by the stalk, the stomium well developed,
several-celled. Spores trilete, tetrahedral-globose, nongreen, usually 64 per sporangium, less commonly 16. Gameto-
phytes epigeal, green, cordate or elongate, midrib thickened; antheridia 5-celled. x=69.

Type species: Cyathea arborea (L.) Sm. (basionym: Polypodium arboreum L.), of the Antilles and northern South
America.

Distribution and ecology: Pantropical, with extensions into extending into warm-temperate regions in the South-
ern Hemisphere. Primarily in wet forests at middle elevations in the mountains.

Genera/species: 3/500. Major genera: Alsophila (235 spp.), Cyathea (160), Sphaeropteris (110).

Economic plants and products: In Australia, “ponga ware” is fashioned from the stems of several species that have
black sclerenchyma and cream-colored ground tissue. The contrast between these two tissues is strikingly beautiful.
Like Dicksonia, the root mantles of several species have been cut and used in horticulture for growing epiphytes, es-
pecially orchids, although it is now illegal to export tree-fern trunks for this purpose. The rigid trunks are sometimes
used as supports for houses or patios in the backwoods in the tropics.The Australian tree fern (Sphaeropteris cooperi)
is widely cultivated as an ornamental. In Malesia native people sometimes eat the fiddleheads of certain species and
the pith of young parts of the trunk. The latter was formerly eaten by the Maoris in New Zealand. Alsophila tricolor,
the silver fern, is the national plant of New Zealand and features prominently in design motifs, money, passports, etc.

Discussion: The Cyatheaceae can be distinguished from the other Cyatheales by scales instead of hairs on the trunks
and petioles. Given its large number of species, the family is notable for being uniformly based on x=69.

Three main clades have been supported on the basis of DNA evidence and morphology: Alsophila, Cyathea, and
Sphaeropteris (Conant et al., 1994, 1995, 1996; Stein et al., 1996; Korall et al., 2006, 2007; Korall & Pryer, 2014).
Sphaeropteris is sister to a trichotomy containing Cyathea, Alsophila, and Gymnosphaera (Figure 4952). The petiole
scales distinguish the genera. Sphaeropteris has conform scales; that is, the cells of the scale have the same shape and
orientation. In contrast, Alsophila, Cyathea, and Gymnosphaera have marginate scales; that is, with the marginal cells
differentiated, usually being shorter, divergent, and irregular.

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As currently construed, Cyathea contains four groups previously recognized as distinct (Korall et al., 2006, 2007, Korall
& Pryer 2014). The first is Hymenophyllopsis, a small-statured group with 8 species endemic to the tepui region of
southern Venezuela (Lellinger, 1984). (Goebel (1929) noted its close relationship to Cyathea after analyzing H. dejecta
and comparing its sporangium with a broad, oblique annulus to that of the Cyatheaceae. The scales of both families
are similar by marginal hairs or processes formed of a single cell, not from two adjacent cells as in, for example, the
eupolypod ferns.) Second, Trichipteris, distinguished from Cyathea solely by the absence of an indusium (Barrington,
1978), has been shown to be polyphyletic, the indusium having been lost several times independently. Third, although
monophyletic, Cnemidaria nests in Cyathea (Conant et al., 1994, 1995, 1996; Stein et al., 1996; Korall et al., 2007).
Cnemidaria is defined by trunks short or absent, laminae 1-pinnate to 1-pinnate-pinnatisect, and spores with three
equatorial pores (Stolze, 1974). Lastly, Sphaeropteris subgen. Sclephropteris (Windisch, 1977) is nested in Cyathea. It
has the characteristic scale type of Cyathea, not Sphaeropteris. This placement is also supported by DNA evidence
(Korall et al., 2007).

Similarly, as currently construed, Alsophila contains Nephelea (Gastony, 1973). This is supported by morphological
(Conant, 1983) and DNA evidence (Korall et al., 2007). It is here kept tentatively distinct from Gymnosphaera. Sphaer-
opteris is now recognized in the strict sense. It includes only subgen. Sphaeropteris; that is, the “S. horrida group,” whose
neotropical species were monographed by Tryon (1971). It excludes subgen. Sclephropteris (=Cyathea), which was
monographed by Windisch (1977).

92
Figure 52. Phylogeny of the Cyatheaceae. A 50% majority-rule consensus tree resulting from Bayesian analyses of five genes.
From Korall et al. (2007).

The trunks of many tree ferns are supported toward the base by well-developed root mantles (Figure 511, A2).
These can be up to half a meter wide and consist of darkly sclerotized, wiry, roots. Root mantles are an important
habitat for epiphytes. Certain species fern epiphytes (e.g., Blechnum ensiforme, B. fragile, and Trichomanes capillaceum)
are nearly restricted to root mantles or occur more frequently and abundantly on them compared to angiosperm
trunks (Moran et al., 2003). Also, root mantles harbor more species compared to angiosperm trunks (Moran et al.,
2003). In some instances where a tree fern grows in open grasslands (e.g., Cyathea gleichenioides and C. macregorii in
the high mountains of New Guinea), the root mantle insulates the stem within from the heat of periodic ground fires.

Few life history studies have been made on tree ferns. Little is known about how fast the trunks grow or the timing of
leaf production. Several that have looked at fertile leaf production and leaf turnover are Seiler (1981), Tanner (1983),
Holttum and Edwards (1983), Ortega (1984), and Bittner (1995).

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 Polypodiales
The monophyly of this group is strongly supported by molecular studies (Hasebe et al., 1995; Pryer et al., 1995,
2004b; Schneider et al., 2004c; Schuettpelz & Pryer, 2007; Wolf, 1997; Vangerow et al., 1998) and by two morpholog-
ical synapomorphies of the sporangium: the stalk is three-rowed, and the annulus is vertical and interrupted at the
stalk. In contrast to the Polypodiales, the basal leptosporangiate ferns typically have a greater number of stalk cells
and complete, ring-like annuli that bypass the sporangial stalk. Other apparent synapomorphies for the Polypodiales
include mixed sporangial maturation (the sporangia mature at different times, not simultaneously) and 64 spores per
sporangium. The gametophytes also show apparent synapomorphies: they are typically cordate (elongate or strap-
shaped in some epiphytic groups) with three-celled antheridia composed of a basal stalk cell, a ring cell, and an cap
cell (operculum). At the molecular level, phytochrome 3 (PHY3) has been found so far only in the Polypodiales, not
in the eusporangiates, basal leptosporangiates, or seed plants (Kawai et al., 2003).

It is often said that ferns are an old group, but this is an oversimplification. The Polypodiales comprises about 80%
of the extant species of ferns, and with the possible exception of the Lindsaeaceae, all the families of Polypodiaceae
have a fossil history dating back to the Early Cretaceous (140 Ma). Thus they are of relatively recent origin, on par
the oldest angiosperms (Archeofructus, the oldest angiosperm, is about 142 Ma). Why did polypodialean ferns evolve
recently, geologically speaking? Many of these ferns are epiphytes, and it seems likely that their radiation was fostered
by the rise of angiosperm-dominated forests during the late Cretaceous. Before then, forests were dominated by
gymnosperms, which generally support fewer epiphytes (this needs field verification). Also, earlier forests apparently
did not cast the dense shade required by many polypodialean ferns. In wet tropical forests today, generally 30 to 50%
of the fern species are epiphytic.

26. Saccolomataceae Doweld

Terrestrial or saxicolous; rhizomes decumbent to erect, scaly, surrounded by persistant petiole bases; steles dicyclic,
concentric, the outer solenostele giving rise to the leaf traces; sterile and fertile leaves monomorphous; petioles
without buds at the base, with an omega-shaped (Ω) vascular bundle (the open end oriented adaxially);
laminae 1-pinnate to 5-pinnate; rachises and costae grooved on the upper surface, the grooves glabrous within;
veins free; sori marginal or submarginal, each at the apex of a vein, often slightly sunken in the laminar tissue and
prominulous adaxially; indusia hyaline or scarious, usually obconic with a truncate apex, opening toward the lamina
margin; spores trilete, tetrahedral-globose, with distinctive pattern of parallel, anastomosed ridges; x = ca. 63.

Type species: Saccoloma elegans Kaulf, of the American tropics.

Distribution and ecology: Pantropical, in wet shaded forests.

Genera/species: 2/12. Sole genus: Saccoloma (12 spp.), Orthiopteris (8)

Economic plants and products: None.

Discussion: Saccoloma was previously treated in the Dennstaedtiaceae because of its omega-shaped vascular bun-
dles in the petiole and submarginal sori, but it was atypical there because of its short rhizome surrounded by per-
sistent petiole bases and lack of epipetiolar buds (dennstaedtioids typically have long-creeping, scaly rhizomes and
epipetiolar buds). The concentric, dicyclic metisteles are a synapomorphy for the family (pers. obs.)

The Asian Orthiopteris was monographed by Luong et al. (2015), who recognized eight species and five varieties. It is
unknown whether Orthiopteris is monophyletic or paraphyletic with respect to Saccoloma.

94
27. Cystodiaceae J. R. Croft

Terrestrial ferns of moderate size. Rhizome a stout caudex, radially symmetrical, prostrate to erect but generally not
arborescent, dictyostelic, young parts generally covered with long, fine golden hairs. Fronds long-stalked, the stipe
with 2 arcuate vascular bundles concave laterally, each with a short cross branch on the abaxial end and 2
separate smaller adaxial bundles uniting upwards to form 2 lateral strands with a simple branch at each end, below
the basal pinnae the two arcs uniting to form a single X-shaped strand, then separating to form a dorsiventral pair of
strands, still with the simple branches at each end, rachises and stipe adaxially grooved and ridged, not contin-
uous or open to admit the grooves of the higher order axes, lamina bipinnate, basally truncate, catadromous, slightly
dimorphic, the fertile pinnules narrower with larger teeth protecting the sori; veins in pinnules pinnately arranged with
a prominent costule, free, simple or forked. Sori submarginal terminal on the veins, receptacle short, slightly raised,
protected by a reflexed lobe of the lamina (outer or false indusium) and a thinner, smaller, cupped inner indusium
attached at the base of the receptacle; sporangia maturing basipetally long-stalked, annulus slightly oblique to almost
longitudinal and interrupted by the pedicel; paraphyses numerous, simple, multiseptate, with no apical glandular cell.
Spores 32 per sporangium, trilete, globose, granular; x=56. (Description from Croft, 1986).

Type species: Cystodium sorbifolium (Sm.) J. Sm.

Distribution and ecology: Borneo and Sulawesi (Celebes) to the Solomon Islands; wet forests.

Genus/species: 1/1

Economic plants and products: none.

Discussion: A monotypic family and genus with two geographically separated but contiguous subspecies. For a long
time thought related to the Cyatheales, particularly the Dicksoniaceae, on the basis of long yellowish hairs on rhi-
zomes and petioles, marginal sori, and bivalved indusia. Molecular phylogenetic studies, however, place it among the
Lindsaeoid ferns (Lindsaeineae), sister to Lonchitidaceae and Lindsaeaceae (Korall et al., 2006). Anatomically, the family
is unique among ferns in that the rachis is traversed by two elongated vascular bundles, one ab- and the other adaxial
(Croft, 1986). It is also unusual by having 32 spores per sporangium.

28. Lonchitidaceae Doweld [to be added]

29. Lindsaeaceae Pic. Serm.

Plants terrestrial or climbing from the soil by roots. Stems short- or long-creeping, hairy or scaly, protoselic or sole-
nostelic, if protostelic then with internal phloem. Petioles with a single vascular bundle, adaxially grooved. Lamina
architecture extremely variable. Veins free or anastomosing. Sori supramedial, close to the margin. Receptacular
hairs (paraphyses) present. Spores trilete or (rarely) monolete. Gametophytes cordate, epigeal, green, dorsiventrally
flattened. Base chromosome number variable: x=34, 38, 39, 44, 47, 48, 49, 50, 51.

Type species: Lindsaea lancea (L.) Bedd. (basionym: Adiantum lanceum L.)

Distribution and ecology: Pantropical. Mostly terrestrial in wet forests.

Genera/species: 7/234. Major Neotropical Genera: Lindsaea (180 spp.), Odontosoria (23), Tapeinidium (18),Osmo-
lindsaea (7),Sphenomeris (1).

95
Economic plants and products: none.

Discussion: The monophyly of the Lindsaeaceae is supported by DNA sequence studies (Pryer et al., 2004;
Schuettpelz & Pryer, 2007; Wolf et al., 1994). Although no morphological synapomorphies support he family, an an-
atomical synapomorphy does. The roots have a sclerenchymatous outer cortex combined with an parenchymatous
inner cortical layer six cells wide (Schnieder, 1996). This distinctive anatomy has allowed an Early Cretaceous (Albian)
permineralized fossil to be assigned to the family (Schneider & Kenrick, 2001). This fossil, an epiphyte on a Tempskya
trunk, is the earliest known for the family. Fossils in amber also also known from the Mid-Cretaceous of northern
Myanmar (Regalado et al., 2017).

Four genera are recognized in the Neotropics (Lehtonen et al., 2010). The largest is Lindsaea, which is most spe-
cies-rich in the Guyanan region. It was divided into 23 sections by Kramer (1971), nearly all of these have been
found to be paraphyletic (Lehtonen et al., 2010). Schuettpelz & Pryer (2007) resolved Lindsaea sister to Lonchitis,
Odontosoria, and Sphenomeris; however, Lehtonen et al. (2010) resolved it sister to the Saccolomataceae and all other
Lindsaeaceae. Lonchitis differs from these groups by root anatomy (Smith et al., 2006), two vascular bundles in the
petioles, fleshy, mucilaginous, conspicuously pubescent leaves and sporangia are protected by a false indusium instead
of a true indusium as in other Lindsaeaceae. Also, its petiole bases (and rhizomes?) contain abundant raphides appar-
ently of calcium oxalate (pers. obs). A separate family name, Lonchitidaceae C. Presl ex M. R. Schomb., is available for
the genus. Odontosoria is unusual in the family by indeterminate leaves that scramble on the surrounding vegetation
and use it for support. Sphenomeris consists of a single species (S. clavata) that is circum-Caribbean and grows on
calcareous rocks. The other species previously classified in Sphenomeris have been found to be nested in Odontosoria
(Lehtonen et al., 2010).

Four other genera, entirely Paleotropical, are recognized in the family: Cystodium, Nesolindsaea, Osmolindsaea, Tapeinid-
ium (Lehtonen et al., 2010). Cystodium was formerly placed in the Dicksoniaceae, but petiole and rachis structure
(Croft, 1986) and DNA sequence evidence place it in the lindsaeoids (Korall et al., 2006; Smith et al., 2006; Schuettpelz
& Pyer, 2007). Nesolindsaea and Osmolindsaea, both recently described, are restricted to southeastern Asia and consist
of two species each. They differ from Lindsaea by solenostelic rhizomes. Tapeinidium is characterized by 1(2)-nerved
sori with indusia firm and adnate laterally.

30. Pteridaceae Reichenbach | Maidenhair Family

Plants terrestrial, epipetric, or epiphytic. Rhizomes erect or creeping, scaly or hairy; dictyostelic. Leaves monomorphic
or dimorphic. Petioles with one vascular bundle or several. Blades 1- to 6-pinnate. Veins free or anastomosing. Sori
borne on the lower surface of the lamina, typically spreading along the veins. Indusia present or absent, when present
formed by the revolute margin of the lamina (false indusium), a “true” inner indusium absent. Spores trilete, globose
or globose-tetrahedral. Gametophytes epigeal, green. x=30 (27 and 29 derived in some genera.)

Type species: Pteris longifolia L., of the southeastern United States, Central America, and the West Indies.

Distribution and ecology: Cosmopolitan. In nearly every kind of habitat where plants grow: from mangrove swamps
to páramos, rain forests to deserts, cliff faces to open water.

Genera/species: 50/1100. Subfamilies and their major genera: Subfam. Cryptogrammoideae: Coniogramme
(20), Cryptogramma (3), Llavea (1). Subfam. Ceratopteroideae: Acrostichum (4), Ceratopteris (4). Subfam. Pteri-
doideae: Actiniopteris (5), Anogramma (s), Austrogramme (5), Cosentinia (1), Jamesonia (45), Nephropteris (1), Onychium
(8), Pteris (250), Pterozonium (13), Syngramma (15), Taenitis (15). Subfam. Vittarioideae: Adiantum (200), Anantha-
corus (1), Antrophyopsis (3), Antrophyum (20), Haplopteris (20), Hecistopteris (3), Polytaenium (10), Radiovittaria (8),
Rheopteris (1), Scoliosorus (3), Vaginularia (4); Vittaria (6). Subfam. Cheilanthoideae: Cheilanthes (180), Doryopteris
(30), Pellaea (25), Notholaena (23), Myriopteris (ca. 20), Argyrochosma (17).

96
Economic plants and products: Many species of maidenhair fern (Adiantum) are cultivated as ornamentals (Goud-
ey, 1985).

Discussion: The Pteridaceae is well supported as monophyletic by molecular studies (Pryer et al., 1995; Schneider et
al., 2004d; Zhang et al., 2005; Schuettpelz & Pryer, 2007) and by three apparent synapomorphies: lack of indusia, spo-
rangia spreading along veins (only slightly in some genera, e.g., Cheilanthes), and a base chromosome number of x=30.
A true indusium is completely absent in the family. As a means to protect the developing sporangia, some genera (e.g.,
Cheilanthes, Pellaea) cover the sorus beneath an enrolled leaf margin (false indusium). This characteristic also occurs
in unrelated ferns such as Hypolepis (Dennstaedtiaceae) and Lonchitis (Lindsaeaceae).

The Pteridaceae can be divided into at least five major groups, recognized at the subfamily level. The first group is
the Cryptogrammoideae. It consists of Coniogramme (Asian), Llavea (Texas, southern Mexico and Guatemala), and
Cryptogramma (subcosmopolitan) (Zhang et al., 2005). These genera have sori that follow the veins, often appearing
as lines, and (this needs to be checked) one vascular bundle in the petioles.

A second group is the Ceratopteridoideae. It consists of the aquatic or semiaquatic Acrostichum and Ceratopteris
(Hasebe et al., 1995; Pryer et al., 1995). Its synapomorphies include spores with prominent parallel ridges, 32 or fewer
spores per sporangium, and x=38. The group can be distinguished as the Parkeriaceae or Parkerioideae.

A third group is the Pteridoideae. It is, united by synapomorphies of x=29 and spores with a pronounced equatorial
flange.The group consists of two subclades.The first is composed of Actiniopteris, Anogramma, Austrogramme, Cosentin-
ia, Onychium, Pityrogramma, Pterozonium, Jamesonia (including Eriosorus), Syngramma, and Taenitis (Sánchez-Baracaldo,
2004a,b), and the second of Platyzoma and Pteris (the latter including Neurocallis and Ochropteris). In the first clade, the
sporangia spread along the veins and are usually unprotected by a revolute margin. In the second clade, the sporangia
spread along a vein parallel to the lamina margin and are always protected by a false indusium.

The phylogeny of the first pteriodioid clade was studied by Sánchez-Baracaldo (2004b), who produced a cladogram
showing generic relationships (Figure 55). In another study (2004a), she examined the phylogeny Jamesonia and Eri-
osorus, two high-elevation genera in the Neotropics. Jamesonia, characterized by narrow indeterminate leaves with
small rounded pinnae, was found to be a morphotype or “form genus” that evolved four times from within Eriosorus.
Most of the species of the Jamesonia + Eriosorus clade are Andean, and sister to it are two species endemic to the
mountains of southeastern Brazil. The Jamesonia + Eriosorus clade is sister to a clade containing Pterozonium and the
Old World genera Austrogramme, Syngramma, and Taenitis (Sánchez-Baracaldo, 2004).

As formerly defined (e.g., Tryon & Tryon, 1982), Anogramma is polyphyletic (Nakazato & Gastony, 2003). Anogramma
osteniana, of southeastern South America, belongs in Eriosorus, and A. chaerophylla and A. novogaliciana belong in Pityro-
gramma. With the removal of these species, Anogramma now consists of only two species: A. leptophylla (L.) Link and
A. lorentzii (Hieron.) Diels. These two species are sister to Cosentinia, a monotypic genus of the Mediterranean-Hima-
layan region. This relationship agrees with spore morphology (two ridges on the proximal face above the equatorial
flange) and chromosome number of x=29. Also, the young gametophytes of Anogramma and Cosentinia are spathu-
late and meristematic activity comes from a lateral, not apical, meristem (Pangua & Vega, 1996). The Anogramma +
Cosentinia clade is sister to a clade consisting of Pityrogramma + A. chaerophylla and A. novogaliciana.

The fourth group is the Vittarioideae, which includes Adiantum (Figure 54). If Adiantum is exluded, the group consist of
about 100--130 species worldwide. These were long recognized as a separate family, the Vittariaceae. The 12 genera
recognized as "vittarioid" are Ananthacorus, Antrophyopsis, Antrophyum, Haplopteris, Hecistopteris, Polytaenium, Radiovit-
taria, Rheopteris, Scoliosorus, Vaginularia, and Vittaria. The synapomorphies of this clade are reddish young leaves (pers.
obs.; although a few genera do not exhibit this) and venuloid idioblasts composed of silica in the upper epidermis
(Sundue, 2009).

Adiantum is the only genus of Pteridaceae that bears sporangia on the false indusium instead of the lamina beneath it
(Figure 56 A3). It petioles, rachises, and costae are typically terete and black or castaneous. All species are terrestrial
or epipetric. In contrast, the vittarioids are entirely epiphytic, and some genera have reddish leaves when young (the
distribution of this character needs among vittarioid genera needs to be determined and optimized on a tree). They
are characterized by clathrate rhizome scales, simple, entire leaves, anastomosing veins, one-rowed sporangial stalks
(Wilson, 1959), and a four-celled stomium consisting of two thickened epistomium and two thickened hypostomium

97
cells (Wilson, 1959).

Crane (1997) revised the generic taxonomy of the vittarioids based on rbcL data (Figure 53). He split Vittaria into
three genera: Ananthocorus, Haplopteris, and Radiovittaria. Those species with leaves less than 4 mm wide and with
narrow paraphyses were retained in Vittaria. Those species with obconic terminal cells at the tips of the paraphyses
were placed in either Haplopteris (paleotropical) or Radiovittaria (neotropical) depending on whether their leaves
were distichous or polystichous, respectively. The species formerly placed in Antrophyum (i.e., those with several to
many series of netted veins between the costa and margin) were divided between Antrophyum (paleotropical) and
Scoliosorus (neotropical and African), distinguished by the former having trilete spores and the latter, monolete. Crane
maintained the remaining vittarioid genera (Ananthacorus, Anetium, Hecistopteris, Monogramma, and Polytaenium) in
their former sense. The only vittarioid fern genera lacking paraphyses are Anetium and Polytaenium.) A phylogenetic
study by Schuettpelz et al. (2016) recovered Anetium nested in Polytaenium; also, that it was necessary (to preserve
monophylly) to recognize a new genus Antrophyopsis.

The last group is the Cheilanthoideae, with about 400 species. It is subdivided into the myripteroids, pellaeoids, he-
mionitidoids, and nothlaenoids (Windham et al., 2009). The genera included are Adiantopsis, Aleuritopteris, Allosorus,
Argyrochosma, Astrolepis, Aspidotis, Bommeria, Calciphilopteris, Cheilanthes, Cheiloplecton, Doryopteris, Gaga, Hemionitis,
Myriopteris, Notholaena, Ormopteris, Paragymnopteris, Pellaea (in part), Pentagramma, and Trachypteris. Apparent syn-
apomorphies are spores without an equatorial flange or ridge and x=30 (with 29 derived in some genera, and 27
derived in Argyrochosma; Windham & Yatskievych, 2003). Their sori are usually borne on the vein tips and protected
by a false indusium, but there are significant exceptions such as Hemionitis with sori spread along the veins and indusia
absent. Farina-bearing glandular hairs have evolved has occurred at least twice within the cheilanthoids (Gabriel y
Galán & Prada, 2012). This group was the focus of a phylogenetic study (Figure 54) by Gastony & Rollo (1998).

Figure 53. Phylogeny of the vittarioid ferns. From Crane (1997).

98
Figure 54. Phylogeny of cheilanthoideae (Pteridaceae) based on rbcL. Above the lines are the number of synapomorphies
supporting each clade and autapomorphic branch lengths for terminal taxa. Numbers below lines are bootstrap percentages
based on 100 replicates. From Gastony and Rollo (1998).

99
Figure 55. Phylogeny of subfam. Pteridoideae (Sánchez-Baracaldo, 2004). Parsimony analysis of combined morphological
and molecular data sets. Strict consensus tree of four equally parsimonious trees. Numbers above branches are bootstrap
percentage values, and the number of morphological steps per branch are indicated in parentheses. Numbers of supporting
molecular characters per branch, derived from the coding region of rps4, are written in italic script. Numbers of supporting
molecular character derived from the intergenic spacer rps4-trnS are indicated in roman script. Trees were rooted using
Coniogramme and the two Pteris species as an outgroup.

100
Figure 56. Pteridaceae. A1-3. Adiantum capillus-veneris, A1. Habit. A2. Pinnule. A3. Sporangia on lower surface of indusium.
B1-4. Myriopteris myriophylla. B1. Habit. B2. Stele from rhizome. B3. Pinnule. B4. Segment showing sori protected by enrolled
margin. C1-3. Pellaea sagittata. C1. Habit. C2. Pinnule and sori protected by false indusium. C3. Solenostele of rhizome.

101
31. Dennstaedtiaceae Pic. Serm.
Plants terrestrial, rarely epiphytic. Rhizomes usually long-creeping, solenostelic, unicyclic or (in Pteridium) polycyclic,
hairy or (in Histiopteris) scaly. Fronds monomorphic or slightly dimorphic. Petioles (except in Pteridium) containing an
omega-shaped vascular bundle with the open end of the omega oriented adaxially, usually the dorso-lateral sides of
the base with one to several buds (epipetiolar buds), these extending as soon as produced (sylleptic). Laminae often
large and decompound, sometimes with indeterminate growth or the apex resting while the next pair of pinnae
develop. Stomata polocytic or anomocytic. Sori round and in marginal cups or elongate to linear and covered by false
indusia. Spores tetrahedral-globose and trilete, or reniform and monolete, 64 per sporangium. Gametophytes cordate,
epigeal, green, thin, dorsiventally flattened. Base chromosome number highly variable: x=26, 29, 30, 31, 33, 34, 38, 46,
47, 48.

Type species: Dennstaedtia cicutaria (Sw.) T. Moore (basionym: Dicksonia cicutaria Sw.)

Distribution and ecology: Cosmopolitan, primarily tropical.

Genera/species: 10/170. Major genera: Dennstaedtia (70 spp.), Hypolepis (50), Microlepia (45), Pteridium (15),
Blotiella (15), Paesia (15), Histiopteris (5), Monachosorum (1).

Economic plants and products: Pteridium is the only genus with economic significance. In many parts of the world,
it invades pastures, reduces forage quality, and poisons livestock that eat the young leaves which contain high amounts
of thiaminase, an enzyme that destroys thiamine, or vitamin B (Moran, 2004). The fiddleheads are eaten in the United
Kingdom and Japan and can be found in Asian food stores in plastic pouches containing a sauce. Eating Pteridium fid-
dleheads is discouraged because it is associated with a high incidence of stomach cancer.

Discussion: Morphological synapomorphies for the family are long-creeping rhizomes, an omega-shaped vascular
bundle in the petiole, epipetiolar buds, and submarginal sori. Histiopteris is atypical by pubescent (not scaly) rhizomes,
and Blotiella by erect or ascending rhizomes with approximate leaves. The distinctive epipetiolar buds are described
by Troop and Mickel (1963) for Dennstaedtia, Hypolepis, Paesia, and Histiopteris. Gruber (1981) gives an interesting
account of these buds and their relation to branching pattern of the rhizome in Hypolepis repens. Petiole anatomy is
described by Keating (1968) for the genera in Costa Rica.

A phylogeny of the Dennstaedtiaceae (Perrie et al., 2015) recovered Monachosorum (Asian) as sister to the rest of
the genus and Dennstaedtia paraphyletic with respect to Microlepia. Despite the molecular phylogenetic studies by
Hasebe et al. (1995) and Wolf (1997), sampling of species in the family is still sparse. More species must be added to
the analyses before well supported clades can be confidently determined.

Figure 57. Dennstaedtiaceae, Dennstaedtia. A1. Rhizome with leaf bases. A2. Leaf. A3. Pinnule with marginal, cup-shaped
sori. A4. Sorus.

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 Eupolypods
The Eupolypods consist of 17 families sister to the Pteridaceae. Three synapomorphies support the clade. The first
is monolete spores. All outgroups to the Eupolypods have trilete spores. A few exceptions —such as Sticherus
and Radiovittaria—exist in the outgroup families, but these exceptions represent independent developments of the
monolete condition. Also, within the Eupolypods, there has been a reversion to trilete spores most importantly in the
grammitid clade of the Polypodiaceae.

Second, the perispore loosely covers the exospore and is often broadly folded; it does not tightly conform to the
surface of the exospore, nor does it adhere to it, as in the outgroups (compare Figure 20A,B with F,H,I). Among the
Eupolypods, a reversal in this character occurs in the Davalliaceae and Polypodiaceae, which have the perispore tightly
pressed to the exospore. Finally, the gametophytes are beset minute capitate-glandular hairs. These are most easily
seen along the margins of the prothallus.

The Eupolypods consists of two large clades, Eupolypods I and II, which can generally be distinguished by the vascula-
ture of their petioles (Figure 58). Eupolypods I have petioles containing several vascular bundles with the two adaxial
ones enlarged and several smaller abaxial ones arranged an arc (Figure 588A). The individual vascular bundles are
circular in cross section. In contrast, species belonging to Eupolypods II have only two vascular bundles, these being
elongate in transverse section. The only exception to this helpful aid to identification is the Blechnaceae, which Eu-
polypods II but (frustratingly!) has the petiole vasculature of Eupolypods I. Also, some extremely small ferns (such as
many grammitids and polypods) have only one vascular bundle in their petioles. Nevertheless, because about 70% of
extant species of ferns belong to the Eupolypods (Smith et al., 2006) and most are typical for this character, the two
petiole vasculature types are enormously helpful in fern identification.

Figure 58. Petiole cross sections of Eupolypod ferns. A. Eupolypods I (Dryopteris intermedia; Dryopteridaceae). B.
Eupolypods II (Goniopteris imbricata; Thelypteridaceae).

103
 Eupolypods II
Eupolypods II contains nearly one-third of the extant fern species (Figure 59; Rothfels et al., 2012b). It is characterized
by two vascular bundles in the petioles, each bundle elongate as seen in transverse section (Figure 588; also see peti-
ole vasculatures at end of this book). The recognition of Eupolypod II families below follows Rothfels et al. (2012a, b).

Figure 59. Phylogeny of the families belonging to Eupolypod II (Rothfels et al., 2012).

32. Cystopteridaceae (Payer) Schmakov

Plants terrestrial or epipetric. Rhizomes short- to long-creeping, the scales non-clathrate. Sterile and fertile leaves
small to medium-sized, monomorphic, not articulate. Petioles with 2 vascular bundles at the base, these uniting
distally to form a “U”. Veins free, ending at the margin. Sori round. Indusium absent (Gymnocarpium) or present,
hood-like and attached on the proximal side of the receptacle (Cystopteris). Sporangial stalks 3-celled. Spores mono-
lete, reniform, nongreen. Gametophytes epigeal, green, cordate. x=40 (Gymnocarpium), 42 (Cystopteris, Acystopteris).

Type species: Cystopteris fragilis (L.) Bernh. (basionym: Polypodium fragile L.), cosmopolitan

Distribution and ecology: Mostly northern hemisphere, primarily temperate, and (for Cystopteris) at high elevations
in the tropics, usually in shaded, moist habitats in forests or on rocks.

Genera/species: 3/30. Genera: Cystopteris (21), Gymnocarpium (7), Acystopteris (3).

Economic plants and products: None.

Discussion: The Cystopteridaceae are sister to the rest of Eupolypods II (Fig. 60; Schuettpelz & Pryer, 2007; Rothfels
et al., 2012). A detailed phylogeny of the family was given by Rothfels et al. (2013). Veins that end at the margin (not
before it) are a synapomorphy for the family. Its species usually have creeping rhizomes, monomorphic sterile and
fertile leaves generally less than 40 cm long, and round sori. Gymnocarpium is non-indusiate.The indusia of Acystopteris,
and Cystopteris are unique by being attached to the proximal side of the receptacle and arching over the sporan-
gia. The close relationship between Cystopteris and Gymnocarpium is evidenced by the existence of a hybrid between them,
×Cystocarpium roskumianum Fras.-Jenk., known only from the Pyrenees of Spain (Fraser-Jenkins, 2008). Its parents are C. fragilis
(L.) Bernh. subsp. dickieana (R. Sim) Hyl. and G. robertianum (Hoffm.) Newman. A genus previously recognized in the family,
Cystoathyrium , has now been

104
Figure 60. Phylogeny of the Cystopteridaceae. Maximum likelihood tree based on concatenated chloroplast sequence data.
(From Rothfels et al., 2013).

105
33. Rhachidosoraceae X. C. Zhang

Plants terrestrial; rhizomes creeping, scaly, the scales clathrate; sterile and fertile leaves monomorphic; petioles with 2
vascular bundles, not forming trophopods; laminae 2- to 3-pinnate-pinnatifid, widest basally, the apex non-conform;
rachises, costae, and costules sulcate adaxially, scaly, the scales filiform; veins free, ending before the margin, non-hy-
dathodous; lamina tissue lacking hairs; sori elongate, indusiate, restricted to only one side of the veins (not diplazioid);
sporangial stalks 2 or 3-cells wide at the middle; spores monolete, achlorophyllous; x=41.

Type species: Rhachidosorus, east and southeastern Asia, from Japan to Sumatra to the Philippines.

Distribution and ecology: Southeastern Asia, often in limestone habitats.

Sole Genus: Rhachidosorus (6).

Economic plants and products: none.

Discussion: Rhachidosoraceae can be distinguished by creeping rhizomes, clathrate rhizomes scales, elongate sori
restricted to one side of the vein (i.e., not diplazioid), laminae 2- to 3-pinnate-pinnatifid, and junctures of the costae
and costules with small green awns adaxially.

34. Diplaziopsidaceae X. C. Zhang & Christenh.

Plants terrestrial; rhizomes thick, erect, or decumbent; laminae 1-pinnate; pinnae entire, glabrous, sessile or nearly so;
veins free (Homalosorus) or anastomosing toward the margins, the areoles without included veinlets; sori elongate to
linear; indusia membranaceous; sporangial stalks 2 or 3-cells wide at the middle;. spores monolete, reniform, achloro-
phyllous. Gametophytes epigeal, green, cordate, glabrous or with minute capitate-glandular hairs. x=41 (Diplaziopsis),
40 (Homalosorus).

Type species: Diplaziopsis, southeastern Asia.

Distribution and ecology: Northeastern North America, southeastern Asia.

Genera/species: 2/4. Genus: Diplaziopsis (3), Homalosorus (1).

Economic plants and products: none.

Discussion: The Diplaziopsidaceae are here recognized based on the molecular phylogenetic results of Wei et al.
(2010) and Chunxiang et al. (2011). All of its species have elongate sori and 1-pinnate leaves that lack of hairs, even
in the grooves of the rachises adaxially. The presence of hydathodes might also be a synapomorphy for the family, but
this needs confirmation.

Homalosorus is monotypic and endemic to eastern North America (H. pycnocarpon). Its morphological similarity to
Diplaziopsis flavoviridis, a species of eastern Malesia, was pointed out by Kato & Darnaedi (1988).

106
35. Desmophlebiaceae Mynssen, A. Vasco, R. C. Moran, Sylvestre &
Rouhan.

Plants terrestrial; rhizomes erect or decumbent, the scales lanceolate, entire to dentate, concolorous, brown; petioles
12–82 × 0.2–1.5 cm; laminae ovate to oblong-lanceolate, 1-pinnate, with a conform apical pinna; rachises grooved
adaxially; pinnae adnate or stalked distally, 2–8 pairs per lamina, elliptic or oblong-lanceolate, equilateral, bases trun-
cate, cuneate or obtuse, apices acuminate, margins entire or serrate distally; costae and rachises subglabrous; veins
simple or 2–4 forked, parallel, apices connected by a submarginal vein; sori linear, simple or diplazioid; indusia
persistent, membranaceous, brownish, entire or erose margins; spores ellipsoid, monolete, cristate, non-green; x=?

Type species: Desmophlebium lechleri (Mett.) Mynssen, A. Vasco, Sylvestre, R.C.

Moran and Rouhan, Central and South America.

Distribution and ecology: Neotropics, from Costa Rica to the Brazilian Atlantic Rain Forest. It grows terrestrially in
wet forests from 125–800 m..

Sole Genus: Desmophlebium (2).

Economic plants and products: none.

Discussion: The Desmophlebiaceae are sister to the Hemidictyaceae and Aspleniaceae (Mynssen et al., 2015). For
a long time its two species (D. lechleri and D. longisorum) were classified in Diplazium (Athyriaceae), but molecular
phylogenetic studies resolve it sister to the Hemidictyaceae and Aspleniaceae. The genus is easily recognized by its
imparipinnate laminae, submarginal connecting vein, linear sori.

36. Hemidictyaceae Christenh. & Schneid.

Plants terrestrial; roots proliferous(?); rhizomes erect or nearly so; laminae 1-pinnate, the apices conform (resembling
the lateral pinnae); pinnae entire, glabrous, sessile; veins anastomosing toward the margins, the areoles without includ-
ed veinlets; sori elongate to linear; indusia membranaceous; sporangial stalks 2 or 3-cells wide at the middle;. spores
monolete, reniform, achlorophyllous. Gametophytes epigeal, green, cordate, glabrous or with minute capitate-glandu-
lar hairs. x=31.

Type species: Hemidictyum marginatum (L.) C. Presl, southeastern Asia.

Distribution and ecology: Tropical America, southern Mexico to southern Brazil; low- to middle-elevation forests.

Sole Genus: Hemidictyum (1).

Economic plants and products: none.

Discussion: Hemidictyum is distinctive by leaves large (0.5–1.5 m long), 1-pinnate with a conform terminal segment,
and veins anastomosing toward the margins of the pinnae. The veins endings are thickened and slightly raised. Also,
they are connected at the tips by a continuous submarginal connecting strand.This character distinguished it from the
similar Diplaziopsis (Diplaziopsidaceae).

107
37. Aspleniaceae Newman | Spleenwort Family

Plants terrestrial, epiphytic, or epipetric. Stems dictyostelic, erect, decumbent, or creeping, scaly, the scales clathrate
(Asplenium) or not Leaves monomorphic, rarely subdimorphic with fertile leaves taller and more erect. Petiole bases
with two bundles, or in smaller species with one vascular bundle. Laminae simple to 4-pinnate, rarely pubescent with
acicular hairs, minute (<0.1 mm long) clavate hairs present (always?). Veins free or netted, their tips hydathodous.
Stomata polocytic, staurocytic or paracytic. Sori elongate, usually oriented obliquely to the midrib. Indusium pres-
ent, attached along the vein on one side of the sorus. Sporangia with vertical annulus interrupted by the stalk, the
annulus cells typically 15–28, the stalks with 1 row of cells. Spores monolete, reniform. Gametophytes epigeal,
green, cordate, glabrous or with minute capitate-glandular hairs. x=36 (Asplenium), 38, 39 (sect. Hymenasplenium); 31
(Hemidictyum).

Type species: Asplenium marinum L., primarily of western Europe, the Azores and Canary Islands.

Distribution and ecology: Cosmopolitan, in many habitats.

Genera/species: 2/700. Genus: Asplenium (700).

Economic plants and products: none.

Discussion: The Aspleniaceae is strongly supported by molecular studies (Perrie & Brownsey, 2005; Sano et al., 2000;
Schuettpelz & Pryer, 2007). It is sister to the Diplaziopsidaceae, and morphological synapomorphies that appear to
support their relationship are the lack of hairs on the leaves—even in the grooves of the rachises adaxially—and
elongate sori. The presence of hydathodes might also be a synapomorphy, but this needs confirmation.

Asplenium is the largest genus in the family. Its synapomorphies appear to be clathrate rhizome scales and high number
(20–28) of annulus cells. The larger species tend to have two vascular bundles at the base of the petiole that unite
distally to form a“X”; however, smaller species such as those of the A. trichomanes group, have only a single vascular
bundle (the xylem in this single bundle is X-shaped). This fusion into an X-shape apparently distinguishes the Aspleni-
um from all other families of Eupolypods II.

Sister to Asplenium is Hemidictyum, with which it shares the synapomorphy of hydathodous vein tips. Hemidictyum
differs by large leaves typically more than 1 m long, imparipinnate laminae, the presence of a submarginal connecting
vein, and x=31.

Asplenium resembles Diplazium (Woodsiaceae) and is sometimes confused with it. Both have two vascular bundles
at the base of the petiole and elongate, indusiate sori. Diplazium, however, lacks clathrate rhizome scales, and the two
vascular bundles in its petioles unite distally to form a U, not an X. Another difference is the number of cells in the an-
nulus: Asplenium has 20–28, Diplazium 15–20. Diplazium is always terrestrial, whereas about half the species of Aspleni-
um are epiphytic. Diplazium usually has a single adaxial rachis groove with minute hairs within the groove; in contrast,
Asplenium has two grooves (one on either side of a raised center) and the grooves and adaxial rachis are glabrous.

Many segregate genera of Asplenium have been proposed, such as Camptosorus, Ceterach, Diellia, Holodictyum, Loxos-
caphe, Phyllitis, Pleurosorus, Schaffneria, and Thamnopteris. Their recognition, however, leaves Asplenium paraphyletic
(although some segregate have not yet been included in cladistic analyses) (Schneider et al. 2004).

The recognition of the segregate genus Hymenasplenium, however, does not result in a paraphyletic Asplenium. The
two genera are sister (Murakami, 1995; Schneider et al., 2004). (I prefer to recognize it as a section because of its
morphological similarity to the rest of Asplenium, but either treatment is justified on phylogenetic grounds.) Section
Hymenasplenium, which consists of 50–60 species, differs from Asplenium by the apparently synapomorphic charac-
ters of creeping rhizomes, dorsiventral steles, distichous leaves, and a distinctive rachis-costa architecture (Murakami
& Moran, 1993). The roots differ anatomically from those of Asplenium (Schneider, 1996; Schneider et al., 2004). The
base chromosome number is 39, which differs from 36 in the rest of Asplenium. Hybrids are unknown between sect.

108
Hymenasplenium and Asplenium. Within Hymenasplenium, apogamy has evolved four times: three times in the Old
World and once in the New. The group is excellent to study the evolutionary and ecological significance of sexual
reproduction because it contains several pairs of sexual and apogamous races (Murakami, 1995).

Asplenium is famous for hybridization. In North America, about three-fifths of the sexually reproducing species are of
alloploid origin (Wagner et al., 1993). Autopolyploidy is also occurs in the genus (Lovis, 1979; Moran, 1982; Reichstein,
1981).

The biogeography of Asplenium is unusual. Instead of being divided into the Old and New World clades, the genus is
divided into temperate and tropical ones (Schneider et al., 2004). The temperate clades evolved from the more basal
tropical ones at least six times.

The name Asplenium comes from the Greek, splen, meaning spleen. Dioscorides, a Greek physician in the Roman
army during the first century AD, thought certain species of the genus were useful in treating diseases of the spleen.

38. Woodsiaceae Herter

Plants epipetric or occasionally terrestrial; rhizomes dictyostelic, creeping, decumbent, scaly, the scales non-clathrate.
Sterile and fertile leaves generally small (up to ca. 30 cm long), monomorphic. Petioles with 2 vascular bundles at
the base, these uniting distally to form a “U”. Indument of (often) glandular hairs. Veins free, ending before the lamina
margins, hydathodous. Sori round. Indusia attached around the base of the receptacle, encircling the sorus,
splitting from the apex downward into lobes, filaments, or scale-like segments. Sporangial stalk 3-celled.
Spores monolete, reniform, nongreen. Gametophytes epigeal, green, cordate, usually with minute capitate-glandular
hairs. x=33, 38, 39, 41.

Type species: Woodsia ilvensis (L.) R. Br. (basionym: Acrostichum ilvense L.), of northern North America, Europe, and
Asia.

Distribution and ecology: Mostly north-temperate regions and high elevations in the tropics. Areas of particular
species richness include the mountains of Eurasia and arid regions of the southwestern United States and Mexico.
One speices (W. montevidensis (Spreng.) Hieron. extends southward to the Andes and also occurs in South Africa.
Typically in rocky habitats.

Sole genus: Woodsia (35).

Economic plants and products: None.

Discussion: The most distinctive character of Woodsia is its indusium, which is united around the circumference of
the receptacle and forms a globose cover over the sporangia. At maturity it splits into several into many segments,
the number often being characteristic of a given species. Many species have jointed petioles, the bases of which per-
sist and form an even stubble after the rest of the leaf falls away. These jointed petioles are unique in Eupolypods II.
Woodsia was monographed by Brown (1964).

In a molecular phylogenetic study of the family, Shao et al. (2015) recovered three main clades that they recognized
as subgenera of Woodsia.They provided a key and synopsis for each subgenus (subgen. Woodsia, subgen. Physematium,
and subgen. Cheilanthopsis).They also found that multiple aneuploidy events resulted in the reduction of chromosome
base number from 41 to 33, 37, 38, 39 and 40 during the evolutionary history of the genus.

109
39. Onocleaceae Pic. Serm. | Sensitive Fern Family
Plants terrestrial; rhizomes creeping or erect, dictyostelic, scaly. Sterile and fertile leaves strongly dimorphic, the
fertile turning brown and hard, usually persisting over winter, erect, the pinnae highly reduced, beadlike, with en-
rolled margins enclosing the sori. Petioles with two vascular bundles, often persistent as trophopods. Blades pinna-
tisect to 1-pinnate. Veins areolate or free. Receptacle conic. Indusium present. Annulus cells 24–35. Spores monolete,
green. Gametophytes epigeal, green, cordate, minute glandular hairs present or absent. x=39, 40 or (in Onoclea) 37.

Type species: Onoclea sensibilis L., of eastern North America and eastern Asia.

Distribution and ecology: Mostly north-temperate regions. Forests and wet soils.

Genera/species: 4/5. Genera: Pentarhizidium (2 spp.), Matteuccia (1), Onoclea (1), Onocleopsis (1).

Economic plants and products: The ostrich fern (Matteuccia struthiopteris) is commonly planted outdoors as
ornamental, especially around the foundations of houses. In New England and New Brunswick, its fiddleheads are
harvested and sold as a vegetable. They are the largest export crop of New Brunswick.

Discussion: Two synapomorphies for this family are 24–35 annulus cells and green spores; in contrast, most other
Eupolypod II families have only 11–19 annulus cells and non-green spores (Sundue & Rothfels, 2015). The brown,
indurated dimorphic fertile leaves are highly unusual, as are the sori completely enclosed by the beadlike segments of
the fertile pinnules (Figure 622). The family is sister to the Blechnaceae (Figure 631; Cranfill & Kato, 2003; Gastony &
Ungerer, 1997; Hasebe et al., 1995; Wolf et al., 1994). Gastony and Ungerer (1997) provided a phylogeny of the four
genera and five species (Figure 611).

Each genus has a distinctive geography. Matteuccia is circumboreal; Onoclea is disjunct between eastern North Amer-
ica and eastern Asia; Onocleopsis is endemic to southern Mexico and Guatemala; and Pentarhizidium is limited to
eastern Asia..

Figure 61. Phylogeny of the Onocleaceae. This is a maximum parsimony tree is based on rbcL and using the Blechnaceae (not
shown) as an outgroup. Above the lines are the number of synapomorphies supporting each clade and autapomorphic branch
lengths for terminal taxa. Bootstrap percentages (italic) based on 1000 replicates. Modified from Gastony & Ungerer (1997).

110
Figure 62. Onocleaceae. A. Matteuccia struthiopteris, habit. A2. Fertile pinnule. A3. Sori. A4. Sorus and indusium. A5. Cross-
section of fertile pinnule. B1. Onoclea sensibilis. B2. Sterile pinnule showing venation. B3. Fertile pinnule.

111
40. Blechnaceae Newman | Chain Fern Family; Deer Ferns

Plants terrestrial, rarely climbing epiphytes. Rhizomes dictyostelic, short- to long-creeping, or erect to arborescent,
scaly. Leaves monomorphic or dimorphic, generally more than 30 cm long. Petioles with 4 or more vascular bundles.
Blades simple to 2-pinnate, reddish when young. Rachises usually grooved adaxially. Veins free or netted. Sori elon-
gate to linear, close to and on both sides of the midribs, the indusium opening toward the midribs. Annu-
lus interrupted at the stalk, the stalk with 3 rows of cells. Spores monolete, nongreen. Gametophytes epigeal, green,
cordate, often with a firm midrib, minutely glandular-pubescent. Chromosome numbers various, generally between
28 and 36.

Type species: Blechnum occidentale L., widespread in the Neotropics.

Distribution and ecology: Cosmopolitan, mostly tropical. In many habitats, but absent from dry ones. Rarely epi-
phytic.

Genera/species: 24/270. Major genera: Parablechnum (65 spp.), Austroblechnum (40), Blechnum (25 spp.), Lomario-
cycas (20), Woodwardia (14), Lomaridium (13), Doodia (13), Stenochlaena (8), Sadleria (6), Salpichleana (3).

Economic plants and products: Species of Blechnum, Doodia, Neoblechnum, and Woodwardia are sometimes culti-
vated as greenhouse ornamentals.

Discussion: The Blechnaceae are unique among ferns by elongate sori that are close and parallel to the midribs with
indusia opening toward the midribs, not the margins.The indusia are born along a continuous commisural vein parallel
to the midribs. This vein develops close to the abaxial surface, below the level of the normal veins, after the sporangia
have been initiated (de la Sota & Labouriau, 1961). The indusia are sometimes hidden because, as they mature, they
shrivel and are pushed back and covered by the sporangia. Also distinctive are reddish young leaves that turn green
with age. The family is atypical among Eupolypods II by having petiole vasculature exactly like that of Eupolypods I;
that is, containing 4–12 vascular bundles with the two adaxial ones enlarged (instead of two strap-shaped ones) and
containing hippocampiform xylem.

The Blechnaceae are subcosmopolitan, comprising about 250 species and 24 genera (Gasper et al., 2016; PPG1
2016). Most of the genera and species predominate in the southern hemisphere (Gasper et al., 2017). They occur in
many different habitats, from lowland forests to alpine bogs, from 0–4000 m. Nearly all species are terrestrial, although
Lomaridium is hemiepiphytic. Holoepiphytes are rare.The monophyly of the Blechnaceae is supported by morphology
and DNA sequences, as is its sister relationship to the Onocleaceae (Cranfill & Kato, 2003; Hasebe, 1995; Pryer et al.,
1995; Gasper et al., 2017). The spores in relation to phylogeny were studied by Moran and Labiak (in review), who
found several genera well supported by perine architecture.

Three subgenera are reconized: subg. Stenochlaenoideae, subgen. Woodwardioideae, and subg. Blechnoideae (Gasper
et al., 2017). Subfamily Woodwardiodeae is sister to the other two subfamilies (Fig. 63). It has chain-like sori (the single
linear sorus is broken into several smaller ones), and the veins are often anastomosing. Subfamily Stenochlaenoideae
contains three genera: Salpichlaena, Stenochlaena, and Telmatoblechnum. Their spores are alike (and different from the
rest of the family) by a single perispore layer with spherical deposits (Moran & Lakiak, in review). Salpichlaena is nearly
unique in the plant world by having twining rachises--a character it shares only with Lygodium (Lygodiaceae). Finally,
subf. Blechnoideae is the largest clade, containing most of the diversity in the family, ranging frrom small inconspicuous
plants to trees (Lomariocycas) with trunks to 4 m tall.

112
Figure 63. Phylogeny of the 24 genera of the Blechnaceae, with typical spores at right. Phylogeny after Gasper et al. (2013), and
spores from Moran & Labiak (in review)

113
Plants terrestrial or epipetric. Rhizomes short-creeping or erect, scaly, the scales non-clathrate. Leaves monomorphic.
Petioles with 2 vascular bundles at the base, these uniting distally to form a “U”. Veins free. Sori elongate to
linear, indusiate. Sporangial stalk 3-celled. Spores monolete, reniform, nongreen. Gametophytes epigeal, green, cordate,
usually with minute capitate-glandular hairs. x=40, and in Diplazium 41.

Type species: Athyrium filix-femina (L.) Roth ex Mertens (basionym: Polypodium filix-femina L.), of northern North
America, Europe, and Asia.

Distribution and ecology: Cosmopolitan, usually in shaded, moist habitats in forests, occasionally on rocks.

Genera/species: 6/700. Genera: Diplazium (400 spp.), Athyrium (220), Deparia (70); Cornopteris (9); Anisocampium
(4), Ephemeropteris (3), Pseudoathyrium (1).

Economic plants and products: The fiddleheads of Diplazium esculentum are eaten in Southeast Asia. This species
is also frequently cultivated in greenhouses. Anisocampium niponicum 'Pictum' is one of the most frequently cultivated
outdoor ferns in North America.

Discussion: The family is characterized by the synapomorphy of sori occurring on both sides of the veins in contrast
to other Eupolypod II familes that have fori on only one side (Sundue & Rothfels, 2013). The sori of the family are
typically elongate or linear, but this also occurs in other families of Eupolypods II (e.g., Aspleniaceae). In Athryium and
Anisocampium, the sori and indusia usually extend distally over vein and become C- or J-shaped. In Diplzaium the sori
are paired, back-to-back along the same vein. Cornopteris and Pseudoathyrium are atypical in the family by having round,
non-indusiate sori, a character state shape representing a reversal.

The Athyriaceae are terrestrial and most have x=41, but Athyrium and Deparia have x=40, a synapomorphy for that
genus (Wei et al., 2013). As in other Eupolypods II, two vascular bundles are present in the base of the petiole. These
bundles united distally to form a U-shape—a character helpful in separating the family from the Aspleniaceae, which
has the bundles uniting distally to form an “x”. Many species in the family have reddish leaves, at least when young and/
or short herbaceous green awns on the adaxial surface of the costa-costule junctures (Rothfels et al., 2012). These
awns interrupt the green wing of the axes. An rbcL phylogeny of primarily Asian species showed, among other results,
that Athyrium was polyphyletic and Deparia was monophyletic (Sano et al., 2000).

41. Athyriaceae Alston | Lady ferns [to be done]

42. Thelypteridaceae Ching ex Pichi Sermolli | Beech Ferns, Marsh Fern

Family

Plants terrestrial, more rarely on rocks; stems creeping to erect, scaly, the scales non-clathrate; leaves monomor-
phic or (less commonly) subdimorphic; petioles with 2 vascular bundles at base; laminae most commonly 1-pin-
natifid-pinnatifid, rarely simple or 3-pinnate-pinnatifid, typically lanceolate, elliptic or oblong, but broadly deltate
in the phegopteroids; rachises and costae not or only slightly grooved adaxially, if grooved, then the grooves
not decurrent into those of the next lowest order; veins free or less commonly netted, typically ending at the
segment margins; sori round or rarely elongate; indusia present or absent; sporangial stalks 3-rowed; spores
monolete, nongreen. Gametophytes epigeal, green, cordate, usually minutely glandular-pubescent; antheridia
3-celled; x=27–36.

Type species: Thelypteris palustris Schott, of North America, Europe, and Asia.

Distribution and ecology: Cosmopolitan, mostly tropical; absent from arid regions.

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Genera/species: 30 (14 in New World)/1035. Major genera: Amauropelta (221 spp.), Goniopteris (120 spp.), Menis-
cium (27), Steiropteris (22), Macrothelypteris (10).

Economic plants and products: none. 30 genera.

Discussion: No single character distinguishes the Thelypteridaceae, but it may be easily recognized by a combination
of characters. The petiole bases contain two vascular bundles as is typical of Eupolypods II. In most species (the Phe-
gopteroids being an exception), the laminae are typically1-pinnate pinnatifid with equilateral pinnae (Figure 644). The
veins, if free as they are in most species, tend to be unbrached. Many species have hyaline, acicular hairs (Goniopteris
is a notable exception with stellate or furcate hairs). The acicular hairs are dead and maturity and therefore do not
contain cytoplasm; thus, they tend to be whitish or colorless. The hairs are most commonly unicellular but may also
be multicellular (e.g., Macrothelypteris). Acicular hairs may also occur on the rhizomes, rhizome scales, and sporangial
capsules (e.g., Phegopteris). Sterile-fertile lamina dimorphy is absent or poorly developed in the family, although many
species have fertile leaves with longer more erect petioles compared to the sterile.

As currently circumscribed at least five genera are non-monophyletic, and several others remain untested. Pending
phylogenetic analysis and taxonomic revision, PPG 1 (2016) follows Holttum (1971, 1982).

Some genera of Thelypteridaceae (e.g., Amauropelta and Steiropteris) bear aerophores (a.k.a., pneumatophores) on
the abaxial surfaces of the rachis-costa junctures, whereas other groups lack them (e.g., Goniopteris). Aerophores vary
in shape from low mounds to elongate pegs or spines.They are whitish and contain stomata.The aerophores typically
turn dark and shrivel with age, and when they do so, they can be easily overlooked. In those species with mucilagi-
nous fiddleheads, the aerophores protrude through the mucilage, presumably to aerate the developing leaves. The
mucilage is secreted by minute glandular hairs on the scales (Oliviera et al., 2017). The mucilage harbors a microflora
of diatoms, desmids, and algae (Hennipman, 1968).

The number of genera recognized has varied greatly, even since the advent of molecular systematics. One study based
on three plastid markers and 115 species (representing 27 segregates of the family) recognized seven genera (He &
Zhang, 2012). Almeida et al. (2016) recognized 16 genera, of which eight occur in the Neotropics. Up to 30 mono-
phyletic groups have been proposed as genera within the family by Holttum (1971, 1982) recongized 30 genera, as
has the PPG1 (2016) classification followed here.

The family has two main clades, both recognized as subfamilies. The Phegopteridoideae comprises Macrothelypteris,
Pseudophegopteris, and Phegopteris (Figure 655) These genera have leaf rachises and costae not or only very shallowly
grooved adaxially and laminae typically more than the 1-pinnate-pinnatifid.The second clade is the Thelypteridoideae.
It is predominantly free-veined and1-pinnate-pinnatifid. Within this lineage are the cyclosoroids, which are mostly net-
veined and based on x=36 (e.g., subgenera Cyclosorus, Meniscium, and Goniopteris).

Venation is an important taxonomic character in thelypteroid ferns. Most species have simple, unbranched veins,
and this often distinguishes them from similar ferns (e.g., Dryopteris, Stigmatopteris). In the strict sense, Thelypteris has
branched veins (Figure 644-A2), which distinguishes it from many proposed segregate genera in the family. Also help-
ful is whether the veins meet the laminar margin at the base of the sinus (the cyclosoroid group; Figure 644-A5) or
above it (other groups; Figure 644-A2). Within the cyclosoroids, an important distinction is whether the veins unite
below the sinus with an excurrent vein to the base of the sinus (Figure 644-A5), or whether they are connivent to
the sinus base, that is coming together but not fused. In the phegopteroids, the veins (at least the basal ones) tend
not to reach the margin, ending end in hydathodes just before the margin.

Mucilaginous fiddleheads are common among species of certain genera of Thelypteridaceae, especially Amauropelta
and Steiropteris. The mucilage is secreted by minute glandular hairs on the petioles and scales (Oliveira et al., 2017).
The species having mucilage have long finger-like aerophores that protrude beyond the mucilage. Micro-organisms
such as diatoms and desmids may flourish in the mucilage (Hennipman, 1968).

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Figure 64. Thelypteridaceae. A1. Thelypteris palustris, habit. A2. Segment and veins. A3. Cross-section of the petiole, showing
two vascular bundles, characteristic of Eupolpods II. A4. Phegopteris connectilis. A5. Chingia ferox, segments and venation. A6.
Pseudophegopteris oppositipinna, pinna and non-indusiate sori. A7. Setose sporangial capsule. A8. Christella puberula, indusiate
sorus.

116
Figure 65. Phylogeny of the Thelypteridaceae based on maximum likelihood analysis of rbcL sequence data.Thick branches have
< 95% bootstrap support. From He & Zhang (2012).

117
 Eupolypods I
This clade is sister to Eupolypods II and consists of the Didymochlaenaceae, Hypodematiaceae, Dryopteridaceae,
Lomariopsidaceae, Tectariaceae, Oleandraceae, Davalliaceae, and Polypodiaceae. Helpful in the identification of these
families is the number and arrangement of vascular bundles in the base of the petiole.Typically, there are 4–10 vascular
bundles with the two adaxial ones enlarged (see chart at end of this book). This character can be seen in all mem-
bers of the group unless the plants are small (e.g., grammitids, Cochlidium), in which case they have only one or two
vascular bundles. The enlarged adaxial bundles usually contain a xylem stands that, when viewed in cross-section, are
hooked. This is exactly the situation in the two xylem strands of Eupolypods II. Presumably to two enlarged strands of
Eupolypods I are homologous with those of Eupopolypods II.

43. Didymochlaenaceae Ching ex Li Bing Zhang & Liang Zhang

Plants terrestrial. Rhizomes dictyostelic, erect, massive (to 30 cm wide), scaly. Petioles with 4 or more vascular bundles,
the two adaxial ones enlarged, several smaller ones arranged below (abaxially). Sterile and fertile leaves monomor-
phic, hairs absent but sparse scales present. Laminae 2-pinnate with a conform terminal segment; pinnules dimidiate.
Rachises and costae grooved adaxially, the grooves decurrent into those of the next lower order, with several finger-like
emergences (ca. 1 mm long) at the junctures of the grooves. Veins free, the tips hydathodous. Sori elongate, back to back
along a single vein, indusiate, often slightly sunken abaxially, appearing embossed adaxially; sporangial capsules glabrous.
Gametophytes cordate, epigeal, glabrous. x=41.

Type species: Didymochlaena sinuosa Desv., of India and eastern Asia.

Distribution and ecology: Pantropical but absent from Australia; wet forests.

Genus/species: 1/2(?). Genus: Didymochlaena (2 spp.).

Economic plants and products: Didymochlaena truncatula is widely sold in the horticultural trade as “mahogany
fern,” a name referring to the reddish brown young leaves.

Discussion: The Didymochlaenaceae is sister to the rest of Eupolypods I (Zhang & Zhang, 2015). The family is highly
distinctive morphologically (see italicized characters in the description). The family is nearly unique in Eupolypods I
by elongate sori and the presence of finger-like emergences on the adaxial surfaces of the rachis-costae junctures.
The number of species in the family has not been determined by a comparative study worldwide. The common
widespread species of the Neotropics is Didymochlaena truncatula (Sw.) J. Sm. This species often has an unpleasant,
skunk-like odor. In the field, plants are often first detected by smell instead of sight.

44. Hypodematiaceae Ching

Plants terrestrial or rarely epipetric. Rhizomes dictyostelic, creeping, scaly. PPetioles with 4 or more vascular bundles,
the two adaxial ones enlarged, several smaller ones arranged below (abaxially). Sterile and fertile leaves monomor-
phic. Rachises and costae grooved or terete, the grooves (when present) usually decurrent into those of the next
lower order. Veins free. Sori indusiate, round; sporangial capsules glabrous or (Hypodematium) setose. Gametophytes
cordate, epigeal, glabrous (Didymochlaena) or pubescent (Hypodematium, Leucostegia). x=40, 41.

Type species: Hypodematium crenatum (Forsk.) Kuhn (basionym: Polypodium crenatum Forsk.), of Africa, India, and
Asia.

Distribution and ecology: Pantropical, usually in wet forested habitats.

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Genera/species: 3/9. Major genera: Hypodematium (4), Leucostegia (3), Didymochlaena (2 spp.).

Economic plants and products: None.

Discussion: This family is supported by molecular evidence (Schuettpelz & Pryer, 2007). Previously, its genera were
classified in different families: Luecostegia in the Davalliaceae, and Hypodematium in the Athyriaceae. Both Leucostegia
and Hypodematium have creeping, dorsiventral rhizomes, and this might prove to be a synapomorphy for the family.

45. Dryopteridaceae Herter | Wood Fern Family

Plants terrestrial, epiphytic, epipetric, or hemiepiphytic. Rhizomes dictyostelic, scaly. Petioles with 3 or more vascular
bundles at the base. Leaves monomorphic or (less commonly) dimorphic. Laminae simple to 5-pinnate. Rachises and
costae grooved or terete, the grooves (when present) usually decurrent into those of the next lower order.Veins free
or (less commonly) anastomosing. Sori round or acrostichoid; indusia present or absent. Sporangial stalk 3-rowed.
Spores monolete, reniform, nongreen. Gametophytes cordate, epigeal, green, densely pubescent on margins and both
surfaces, the hairs usually unicellular, papillate. x=41.

Type species: Dryopteris filix-mas (L.) Schott (basionym: Polypodium filix-mas L.), of North America, Europe, and Asia.

Distribution and ecology: Cosmopolitan, usually in forested habitats.

Genera/species: 40–45/1700. Major genera: Elaphoglossum (600 spp.), Dryopteris (275), Polystichum (200), Ctenitis
(100), Arachniodes (50), Bolbitis (45), Megalastrum (92), Polybotrya (37), Parapolystichum (25), Stigmatopteris (23), Lo-
magramma (22), Pleocnemia (19), Lastreopsis (15), Cyrtomium (15), Teratophyllum (11), Mickelia (10), Phanerophlebia
(8), Olfersia (3), Arthrobotrya (3).

Economic plants and products: Dryopteris filix-mas has been used since the Middle Ages as an antihelminthic. Ru-
mohra adiantiformis is widely used in floral arrangements and is commonly called “florist’s fern.”

Discussion: This large and diverse family is strongly supported by molecular data (Schuettpelz & Pryer, 2007; Labiak
et al., 2015; Liu et al., 2015) but lacks morphological synapomorphies. Three subfamilies were recognized by Liu et al.
(2015) for the family: subfam. Dryopteridoideae B. K. Nayer, subfam. Elaphoglossoideae (Pic.Serm.) Crabbe, Jermy &
Mickel, and subfam. Polybotryoideae H.M. Liu & X.C. Zhang.This creation is premature given that branch supports are
weak for the earliest diverging clades in the family.

In general, there apprear to be five main clades in the Dryopteridaceae. The first is a terrestrial group with erect or
short-creeping rhizomes and monomorphic sterile and fertile leaves. It consists of Ctenitis, Dryopteris, Polystichum, Pha-
nerophlebia, Cyrtomium, and Arachniodes. Among the largest genera of Dryopteridaceae is Polystichum, which contains
about 200 species worldwide. It was the subject of a detailed phylogenetic study by Little and Barrington (2003) and,
in the Andes where it is particularly species-rich, by McHenry & Barrington (2014).

The second group is the polybotryoid clade (Moran & Labiak, 2015). It is entirely neotropical except for the mono-
typic Trichoneuron, which is siter to the primarily West Indian Polystichopsis (Liu et al., 2015). It is defined by creeping
rhizomes. Most members of the group are scandent on tree trunks and typically exhibit strong sterile-fertile leaf
dimorphic. The clade consists of Polybotrya, Cyclodium, Maxonia, Olfersia, Pradopteris (gen nov.), Polystichopsis, and Tri-
choneuron.The latter two genera are sister to the others, and atypical from other genera in the clade by thin rhizomes,
terrestrial habit, monomorphic sterile and fertile leaves, and long, straight whitish hairs on the laminae (Liu et al., 2015;
Prado & Moran, 2015).

A third clade is formed by Stigmatopteris, a neotropical genus characterized by terrestrial habit, monomorphic sterile
and fertile leaves, hydathodes, internal punctuate glands in the laminae, and lack of indusia (Moran, 1991). A phylogeny
of this clade was given by Moran & Labiak (2016).

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A fourth group, the “Lastreopsid” clade, is formed by Megalastrum, Lastreopsis, Parapolystichum, and Rumorha (Labiak
et al., 2014). A synapomorphy for this group are colored glandular hairs on the leaves (and on the sporangial stalks
in Lastreopsis, Parapolystichum and Pleocnemia). The plants are terrestrial and generally have monomorphic sterile and
fertile leaves, decompound laminae, basal pinnae prolonged basiscopically, and round sori. All except Megalastrum
have a “Lastreopsis-type” of rachis-costa architecture: the thickened marginal ridge of the costae and costules is sur-
current along the margin of the axis of the next higher order.

The last clade is the bolbitidoid ferns, consisting of Arthrobotrya, Bolbitis, Elaphoglossum, Lomagramma, Mickelia, and
Teratophyllum (Figure 6666). These ferns are characterized by the synapomorphies of dorsiventral rhizomes with an
elongated (in transverse section) ventral meristele, roots borne only from the ventral meristele (not on all sides of
the rhizome), lack of hairs on the leaves, sterile-fertile leaf dimorphy, and acrostichoid sori (Moran et al., 2010). With
further study two other characters will likely prove synapomorphic: the presence of diplodesmic veins and the posi-
tion of the branch buds. Diplodesmic veins are an extra set of veins below (abaxially to) the normal ones that supply
the sporangia. The branch buds are located ventro-laterally in one row on each side of the rhizome and between the
leaf bases. The buds are often inconspicuous and typically do not develop unless the apex of the rhizome is removed
or damaged. If the rhizome has two or more ranks of leaves—that is, if it bears dorsal rows of leaves between the
two lateral ones—branch buds do not develop between the leaves in the dorsal rows. This arrangement of branch
buds is unique among ferns (Moran et al., 2010). The bolbitidoid ferns exhibit a transition series going from terrestrial
(Bolbitis)  scandent on trunks (Arthrobotrya, Lomagramma, Mickelia, and Teratophyllum) epiphytic (Elaphoglossum)
(Moran et al., 2010; Lagomarsino et al., 2012).The Asian Pleocnemia is sister to the bolbitidoids (Liu et al., 2014; Labiak
et al., in review).

One bolbitidoid genus, Elaphoglossum, contains about 600 species and thus constitutes nearly one-third of the
Dryopteridaceae. Its monophyly is supported by trnL-trnF and rps4 (Rouhan et al., 2004; Skog et al., 2004) and by
the synapomorphies of epiphytic habit, distichous leaves, phyllopodia, simple laminae, and free veins (Moran et al.,
2010). A subgeneric classification of Elaphoglossum based on morphology and spores (Mickel and Atehortúa, 1980)
was largely supported by cpDNA sequences (Rouhan et al., 2004). The genus consists of seven main clades (Figure
6767). Section Amygdalifolia is characterized by the unique combination of long-creeping rhizomes, phyllopodia, hy-
dathodes, and reddish young leaves. It is the only species in the genus that is a primary hemiepiphyte (Lagomarsino
et al., 2012). Section Elaphoglossum generally has thick leaves with reduced scales, often appearing glabrous from a
distance. Section Squamipedia is characterized by long-creeping rhizomes, lack of phyllopodia, paired peg-like aero-
phores on the rhizome near the petiole, flaccid tan rhizome scales, leaves less than 15 cm long, and echinulate spores.
The subulate-scaled clade has subulate scales (enrolled throughout their length and often imparting a shaggy or hairy
appearance to the blades). With this clade, section Setosa has hydathodes whereas sect. Polytrichia lacks them. Sec-
tion Lepidoglossa typically has conspicuously scaly laminae, and the body of these scales are flat (non-subulate) with
acicular marginal teeth (the marginal teeth or processes in the other sections are terminated by a slightly swollen,
bulbous cell).

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Figure 66. Phylogeny of the bolbitidoid ferns. 50% majority-rule consensus tree from the Bayesian analyses of a combined
dataset (rps4-trnS + trnL-trnF + coded gaps). Stars indicate values of posterior probabilities (above the branches) and
bootstrap (bellow the branches) equals to 1 and 100%, respectively. (from Moran et al., 2010.)

121
Figure 67. Phylogeny of the sections and subsections of Elaphoglossum based on chloroplast trnL-trnS and rps4 intergenic
spacer regions. Numbers are bootstrap support values; thick horizontal bars are indels. (After Skog et al., 2004 and Rouhan et
al., 2004.)

46. Nephrolepidaceae Pic.Serm.

Type species: Nephrolepis exaltata (L.) Schott (basionym: Polypodium exaltatum L.).

Distribution and ecology: Pantropical and subtropical.

Genera/species: 1/20.

Genera: Economic plants and products: Many species of Nephrolepis are cultivated as ornamentals, with hundreds
of cultivars. The most common, usually grown in hanging baskets, is N. exaltata ‘Bostoniensis’, the Boston Fern. In the
tropics, N. cordifolia is often cultivated as a low hedge or border plant.

Discussion: The rhizome system of Nephrolepis is unique among ferns (Espagnac, 1973; Hovenkamp & Miyamoto,
2005). It consists of two parts: a short, erect, poorly developed leaf-bearing part and a long, horizontal, well developed,
non-leaf-bearing stoloniferous part. The erect rhizomes of Nephrolepis do not produce roots; instead, they produce
stolons, and it is the stolons bear the roots. The stolons are further unusual by not bearing leaves as is typical in other
plants. Also unusual are the rachis scales, which point downward, not upward (distally) as in other ferns (Hovencamp
& Miyamoto, 2005).

An analysis of three plastid sequences, including the maturase matK, resolved Nephrolepis apart from other lomariop-
sids, suggesting that it should be recognized in its own family, distinct frrom the Lomariopsidaceae (Kuo et al., 2011).
A monograph and phylogenetic study are available for Nephrolepis (Hovencamp & Miyamoto, 2005; Hennequin et
al., 2010).

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47. Lomariopsidaceae Alston

Plants terrestrial, epilithic, hemiepiphytic, or epiphytic; rhizomes scaly, long-creeping to erect; petioles with 3 or more
vascular bundles; leaves 1-pinnate; pinnae articulate to the rachis. Rachises and costae grooved or terete. Veins
free. Sori round or acrostichoid. Sporangial stalk 3-seriate. Spores monolete, achlorphyllous or (in Lomariopsis) chlo-
rophyllous; x=41.

Type species: Lomariopsis sorbifolia (L.) Fée (basionym: Acrostichum sorbifolium L.)

Distribution and ecology: Pantropical.

Genera/species: 5/70. Genera: Lomariopsis (45), Cyclopeltis (5), Dracoglossum (1), Thysanosoria (1), Dryopolysticum
(1).

Economic plants and products: None.

Discussion: The apparent morphological synapomorphies of the family are 1-pinnate leaves and pinnae articulate to
the rachis.. The articulation can usually be seen where the pinna joins the rachis as a faint dark line often associated
with a slight swelling. The articulations appear functional only in Nephrolepis, where they fall readily, leaving behind
persistent rachises—a good field character of the genus. Anyone who has cultivated a Boston fern in a hanging basket
knows that the plant is “messy.” The floor beneath needs frequent sweeping to remove fallen pinnae. Although the
pinnae are not normally shed in Cyclopeltis and Lomariopsis, experiments suggest that pinnae are shed in response
to damage or herbivory. The inclusion of Nephrolepis in the family is tentative (Tsutsumi & Kato, 2006; Schuettpelz &
Pryer, 2007; Hennequin et al., 2010). Dracoglossum and Dryopolystichum are atypical in the family by having entire and
1-pinnate-pinnatifid leaves, respectively (Chen et al., 2017).

A monograph and phylogenetic study are available for Lomariopsis (Moran, 2000; Rouhan et al., 2007), and Nephrole-
pis (Hovencamp & Miyamoto, 2005; Hennequin et al., 2010). Thysanosoria consists of a single species (T. pteridiformis
(Cesati) C. Chr., known from a few collections in coastal northwestern New Guinea.

48. Tectariaceae Lellinger

Plants terrestrial. Rhizomes dictyostelic, short-creeping or decumbent, less commonly erect or long-creeping. Petioles
with more than 4 vascular bundles, the adaxial ones enlarged, not jointed to the rhizome. Laminae simple to 4-pin-
nate-pinnatifid, often pubescent with short (<1 mm), jointed hairs, especially on the upper surfaces of the axes. Rachises
and costae not grooved, or if slightly grooved then the grooves not decurrent into those of the next lowest order.
Veins free or anastomosing. Sori typically round; indusia present or absent. Sporangial stalk 3-rowed. Spores monolete,
reniform, nongreen. Gametophytes cordate, epigeal, green, densely puberulent on margins and both surfaces, the hairs
mostly uniseriate, papillate, secretory. x=40 or 41.

Type species: Tectaria trifoliata (L.) Cav.

Distribution and ecology: Mostly tropical, with local extensions into the subtropics. Typically terrestrial in wet for-
ests.

Genera/species: 10/230. Major genera: Tectaria (200), Triplophyllum (22), Arthropteris (12), Pteridrys (8), Hypoderris
(3), Draconopteris (1).

Economic plants and products: none.

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Discussion: The monophyly of the Tectariaceae is supported by DNA sequence evidence (Schuettpelz & Pryer,
2007; Ding et al., 2014). An apparent morphological synapomorphy is the puberulence generally found on the up-
per surfaces of the rachises and costae, and (perhaps) the densely pubescent gametophytes (Nayar & Kaur, 1971).
Arthropteris, mostly a genus of primary hemiepiphytes, is sister the rest of the family, all of which are terrestrial. It is
sister to the rest of the Tectariaceae and has been placed in its own family, the Arthropteridaceae (Liu et al., 2013). Lui
et al. (2007) found Pleocnemia, which had long been classified in the Dryopteridaceae, belonged to the Tectariaceae.
Fa-Guo et al. (2014) and Moran et al. (2014) found that the neotropical Hypoderris is distinct.

Both Arthropteris and Tectaria are in need of monographic work. The neotropical species of Triplophyllum were mo-
nographed by Prado & Moran (2008).

Figure 68. Phylogeny and synapomorphies of three Eupolypods I families.

124
49. Oleandraceae (J. Sm.) Ching ex Pichi Serm.

Plants epiphytic, terrestrial, or epipetric, either creeping and appressed to the substrate, or shrubby with both hor-
izontal terrestrial and erect aerial branches. Roots sometimes extremely long (to 20 cm), branching and producing
root hairs when in contact with the substrate. Stems elongate, dictyostelic with scattered sclerenchyma strands in
the ground tissue, sometimes pruinose, scaly, the scales peltate, typically with the bases overlapping and obscuring
the stem, with marginal, uniseriate cilia. Leaves monomorphic (except one Asian species). Petioles articulate to
phyllopodia, the joint swollen, containing three or four vascular bundles. Laminae simple, entire. Veins free. Sori
round, usually born on the acroscopic branch of a vein cluster, indusiate, the indusia reniform to circular. Sporangia
stalks with glandular hairs distally, just below the capsules; annulus cells 12–14. Spores ellipsoid, monolete, nongreen,
columellate between the exospore and perispore. Gametophytes cordate or rarely strap-shaped, minutely glandular
pubescent. x=41.

Type species: Oleandra neriiformis Cav., type from the Philippines.

Distribution and ecology: Pantropical, most diverse in Asia and Polynesia. Epiphytic in wet forests.

Sole genus: Oleandra (40 spp.)

Economic plants and products: none.

Discussion: The monophylly of Oleandra and its sister relationship to Davalliaceae and Polypodiaceae (Figure 68) is
supported studies of non-coding chloroplast DNA sequences (Hasebe et al., 1995; Schneider et al., 2004; Schuettpelz
& Pryer, 2007; Tsutsumi & Kato, 2006). Oleandra derives its name from the appearance of its shrubby species whose
loose whorls of leaves resemble oleander (Nerium oleander, Apocynaceae).

The family is uniform and distinctive. It is easily recognized by elongate stems, peltate rhizome scales with uniseriate
cilia along the margins (Figure 699A; not the typical dryopteroid tooth composed of two adjacent cells upturned),
petioles with a swollen articulation joint and abscission layer (Phillips & White, 1967), simple and entire leaves, long
and parallel veins between the costa and margin, and round sori usually scattered near the costae, and circular indusia.
The sporangial stalk is two-rowed except just below the sporangium, where there is a secondarily formed third row
of stalk cells, these developing from the capsule. The distal part of the sporangial stalks bear glandular hairs (Figure
699D, E) whose number and number of cells varies between species (this needs to be further documented and
would make an excellent project). The spores are distinctive by being columellate between the exospore and peri-
spore (see SEM images on www.plantsystematics.org). An SEM study of the spores was presented by Liew, 1977).

Besides being long-creeping, the rhizomes have distinctive anatomical characters (Nayar et al., 1968). The ground tis-
sue contains scattered sclerenchyma strands (this character, however, might eventually prove to be be synapomorphic
for Eupolypods I). The stele is dissected into a loose reticulum of slender meristeles resulting from crowded spirally
arranged leaf gaps. In the apparently leafless portions of the rhizomes, these leaf gaps are associated with a solitary
vestigial leaf trace (Figure 699 B,C). Thus, at these leaf gaps the leaves are suppressed; they do not develop outwardly
although their gaps present in the stele. The suppressed leaves can sometimes be seen as low bumps on the rhizome
when the scales are removed. This character might eventually prove to be synapomorphic for the clade consisting of
Oleandraceae, Davalliaceae, and Polypodiaceae.

Although the rhizome scales are peltate—an apparent synapomorphy for the family—those on the costae are bas-
ifixed (Figure 699A). The cilia of the scales resemble those of Gleicheniaceae, Cyatheaceae, or Myriopteris. These cilia
are unlike the teeth of other Eupolypods I, which are composed (ultimately) of the upturned ends of two adjacent
cells. These cilia (either single-celled or multi-cellular) are a synapomorphy for the family.

On the basis of morphology (pers. obs.), Oleandra appears to consist of two clades whose monophyly needs to be
confirmed by molecular phylogenetic studies. The first has rhizomes climbing and appressed to the substrate, widely
spreading apices of the rhizome scale, and leaves produced more or less evenly along its length. The species in this

125
clade (e.g., O. articulata and O. bradei) are holoepiphytic (pers. obs). In contrast, the second clade has a shrubby habit,
with horizontal rhizomes producing few or no leaves but bearing aerial branches with clustered leaves (Hallé et al., fig.
60, 1970).The leaves are clustered along the stems and lack any apparent phyllotaxy. In this second clade the rhizome
scales apices are appressed, not spreading. Some of these aerial rhizomes may become pressed against a tree trunk
and become climbing; at other times the plants appear to be fully epiphytic. The claim by Tsutsumi & Kato (2008,
2010) that these plants are secondary hemiepiphytes (starting on the ground, then growing up a trunk and eventually
loosing the connection with the soil) is unsubstantiated. For some species (e.g., O. costaricensis, it is definitely not true
all the time). Growth habits and field observations of gametophytes need better documentation in this genus. The
two clades might also differ as to whether the cilia of the rhizome scales are unicellular or multicellular and in the
number of paraphyses on the sporangial stalks.

Figure 69. Some characters of Oleandraceae. A. Peltate rhizome scale with cilia (not teeth). B. Dictyostele, dorsal view, showing
suppressed leaves. C. Dictyostele, ventral view. D. Sporangium with two paraphyses below stalk. E. Paraphyses at apex of
sporangial stalk. BT = branch trace; L = Leaf trace; SL = Suppressed leaf trace. A–D. Oleandra wallichii. E. O. pistillaris. (Modified
from Nayar et al., 1968.)

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50. Davalliaceae Mett. ex Frank | Rabbit’s foot Fern Family

Plants epiphytic, rarely epipetric. Rhizomes with a slightly compressed dorsiventral dictyostele, short- to long-creeping,
fleshy, scaly, the scales peltate. Leaves monomorphic, typically glabrous, borne alternately in two rows on the
dorsal surface of the rhizomes, articulate to short phyllopodia. Petioles with several vascular bundles. Laminae
1– to 5-pinnate, rarely simple. Rachises and costae grooved on the upper surface, the grooves with a raised center.
Veins free. Sori terminal on the veins, submarginal or borne well back from the margin, funnelform, cup-shaped, or
lunate. Indusia present. Sporangial stalks 1– or 2-rowed, usually long. Spores ellipsoid, monolete, nongreen, the
perispore thin, appressed to the exospore, not modified into wings or spines. Gametophytes epigeal, cordate, densely
puberulent, the hairs multicellular, papillate, non-glandular; x=40.

Type species: Davallia canariensis (L.) Sm. (basionym: Trichomanes canariensis L.), of the Atlantic Islands, Morocco,
and Iberia.

Distribution and ecology: Tropics and subtropics of the Old World, most species-rich in Asia. Mostly epiphytes in
wet forests.

Genera/species: 4/65. Major genera: Davallia (34 spp.), Araiostegia (11), Davallodes (7), Pachypleuria (2?).

Economic plants and products: Several species of Davallia are cultivated in hanging baskets. They are called “rab-
bit-foot ferns” in allusion to their densely scaly rhizomes.

Discussion: Recent studies have greatly changed the circumscription of the Davalliaceae and the genera within it
(Schuettpelz & Pryer, 2007; Tsutsumi & Kato, 2006; Tsutsumi et al., 2008; Kato & Tsutsumi, 2008). The studies reveal
six major clades within the family, and all of the previously recognized genera are polyphyletic. Gymnogrammitis and
Leucostegia, formerly included in the family on the basis of morphology, are now classified in the Polypodiaceae and
Hypodematiaceae, respectively.

Both Davalliaceae and Polypodiaceae are epiphytes with dorsiventral creeping rhizomes. Their leaves are alternate,
distichous, and articulate to short phyllopodia (Sen et al., 1972; Kato, 1985). Both also show only one or two of rows
of cells in the sporangium stalks, versus the typical 3-rowed stalks in other Eupolypods. A difference between the
two families is chromosome number: x=40 in Davalliaceae, and mostly x=36 or 37 in Polypodiaceae. Chromosome
number also distinguishes the Davalliaceae from the remaining families of Eupolypods II (Dryopteridaceae, Lomariop-
sidaceae, and Tectariaceae), all of which are x=41.

The family was monographed by Nooteboom (1992, 1994). The fine structure of the articulation between the phyl-
lopodium and the petiole was described by Phillips and White (1967). The roots of Davallia are not scattered along
the rhizome but occur only in association with the lateral branch buds, on their ventral and lateral sides (Croxdale,
1976). A review of epiphytism in the group was provided by Kato and Tsutsumi (2013).

51. Polypodiaceae Berchtold & J. Presl | Polypody Family

Plants epiphytic, less often terrestrial or epipetric. Rhizomes dictyostelic, scaly, of two types: creeping and dorsiventral
with the leaves arranged alternately in two rows (Figure 70), or (in grammitids) erect, radial, with leaves polystichous.
Leaves monomorphic or (less commonly) dimorphic. Petioles jointed to the short phyllopodia or not. Laminae simple
to 1-pinnate, rarely more divided. Veins free or anastomosing. Sori nonindusiate, round, rarely elongate to linear.
Sporangial stalk 1- to 3-celled. Spores either yellow and bilateral or (in grammitids) green and tetrahedral, perispore
thin and adhering tightly to endospore, not elaborated into wings or folds. Gametophytes epigeal, green, cordate or
elongate, glabrous or pubescent. Chromosome number variable, most commonly x=35, 36, 37.

Type species: Polypodium vulgare L., of Europe and Asia.

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Distribution and ecology: Cosmopolitan, primarily tropical; epiphytic, rarely saxicolous.

Genera/species: 50/1200. Major genera: Polypods: Polypodium (60 spp.), Selliguea (80), Microsorum (60), Pyrrosia
(55), Pleopeltis (50), Campyloneurum (50), Loxogramme (35 spp.), Loxogramme (30), Pecluma (30), Microgramma (25),
Serpocaulon (40), Phlebodium (4). Grammitids: 30/750: Lellingeria (50), Ceradenia (68), Melpomene (30), Moranopteris
(30), Alansmia (25), Grammitis (22), Stenogrammitis (24), Ascogrammitis (17), Cochlidium (16), Terpsichore (12), Zygo-
phlebia (11), Enterosora (10), Galactodenia (5), Leucotrichum (5), Micropolypodium (3), Lomaphlebia (2).

Economic plants and products: Several genera are cultivated as ornamentals, the most popular being the stag-
horn ferns (Platycerium). In Central America, the rhizomes of several species, collectively called calaguala, are sold in
markets and reportedly good for just about everything.

Discussion: The Polypodiaceae is one of the main families of epiphytic ferns in the tropics. Its monophyly is strongly
supported by cpDNA sequence evidence (Ranker et al, 2004; Schneider et al., 2004). Synapomorphies are round
non-indusiate sori, yellow spores, and phyllopodia (the grammitid clade has green spores and most of its genera lack
phyllopodia). The phyllopodia appear as low enlargements or swellings of the rhizome, not as part of the petiole as in
Elaphoglossum or Oleandra. The color changes abruptly between the phyllopodium and petiole. When the leaves fall,
they do so cleanly, leaving a circular scar on top of the phyllopodia (Phillips & White, 1967).The leaves are arranged in
two ranks on the dorso-lateral side of the rhizome (Figure 7070). This arrangement imparts a distinctive appearance
to the rhizomes, and the plants can usually be identified as belonging to this family by the rhizome alone. The roots
are emitted primarily from the ventral surface of the rhizome.

The family is divided into two clades that correspond neatly to geography: one clade containing the New World
species, and the other the Old World ones. Synammia, a genus of three species in temperate South America, might
be sister to the rest of the family or to the New World clade (Schneider, 2006).

Nested within the New World clade are the grammitid ferns, or dwarf polypodies, previously recognized as the
Grammitidaceae (Ranker et al, 2004; Schneider et al., 2004). They are characterized by reddish setae on the leaves
(these lost in some genera, such as Cochlidium, Grammitis, and Lellingeria), sporangial stalks consisting of a single row
of cells, and green, trilete spores. Scales are always absent from their leaves but present on the rhizomes (could the
reddish setae of the laminae represent highly modified scales?). Gametophytes are elongate, often with an acicular
hair that (when present) is produced at or near the corner of the cell wall. Several large systematic studies have
been done on generic limits within the clade (Labiak et al., 2010a, b; Lehnert et al., 2009; Sundue, 2010; Sundue et al.
2010). The grammitids occur worldwide with little overlap between species in the New World and Old World. The
biogeography of the group was reviewed by Parris (2003), who stated that about 750 species of grammitid ferns are
known in about 30 genera. About 300 species occur in the New World and 450 in the Old. The Old World genera
were reviewed by Parris (2009).

The Polypodiaceae contain the only ferns known to be myrmecophytic. In the Neotropics, there are four myrmeco-
phytic species of Microgramma subgen. Solanopteris, and in the Malesia there are 13 species of Lecanopteris. In both
genera the ants not only defend the plants, but also fertilize them by bringing nutrients into modified rhizomes (Mo-
ran, 2004).

Figure 70. . Typical rhizome of Polypodiaceae, showing creeping habit, two rows of phyllopodia, and ventral roots.

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 Keys to Families and Genera of Neotropical
Ferns and Lycophytes

Key to Families of Lycophytes

1. Plants homosporous; leaves nonligulate; roots without central air canal.......................................................... Lycopodiaceae
1. Plants heterosporous; leaves ligulate; roots with central air canal.
2. Leaves less than 2 cm long, planar, without air chambers; sporangia surficial, not embedded leaf tissue, often
borne in quadrangular strobili that terminate branch tip........................................................................................ Selaginellaceae
2. Leaves longer than 2 cm, terete or nearly so, with 4 air chambers; sporangia embedded in the adaxial face of
the leaf base, not borne in strobili................................................................................................................................................... Isoëtaceae

1. Lycopodiaceae

1. Arial stems isodichotomously branched; elongate horizontal indeterminate stems lacking; sporophylls similar to
trophophylls or, if smaller, green and persisting after spore release; spores foveolate-fossulate
2. Plants erect or decumbent, with lateral gemmae; spores with the equator line concave and the corners trun-
cate, the proximal faces foveolate-fossulate ...................................................................................................................................Huperzia
2. Plants erect to pendulous, without gemmae; spores with the equator line convex between laesureae and with
obtuse corners, proximal faces smooth ................................................................................................................................Phlegmariurus
1. Arial stems isodichotomously branched; elongate horizontal indeterminate stems present; sporophylls dissimilar to
trophophylls, turning straw-colored after spore release; spores reticulate, rugate, or scabrate
3. Strobili pendulous or sessile; creeping stems giving rise to erect branches dorsally; spores rugate
4. Strobili pendulous or nodding; erect stems highly branched, tree-like; sporangia almost enclosed in cavities
formed by the strobilus cortex and coalescent membranous bases of adjacent sporophylls ............... Palhinhaea
4. Strobili erect; erect stems simple or 1-forked, sporangia exposed or enclosed and hidden by free sporo-
phylls
5. Sporangia isovalvate; sporophylls in alternating whorls of 2–5, forming 4–10 longitudinal rows; sporangia
visible between the sporophylls ........................................................................................................................ Pseudolycopodiella
5. Sporangia anisovalvate; sporophylls in alternating whorls of 5 or more, forming 10 or more longitudinal
rows; sporangia hidden by the sporophylls ...............................................................................................................Lycopodiella
3. Strobili erect, sessile or pedunculate; creeping stems giving rise to erect branches laterally; spores reticulate
6. Leafy branches isophyllous, terete
7. Leaf apices with a colorless terminal hair; sporophylls with thin medial wing decurrent onto the axis of
the strobilus; spores reticulate on the proximal faces ..........................................................................................Lycopodium

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 7. Leaf apices without terminal colorless hair; sporophylls without thin medial wing decurrent onto the
axis of the strobilus; spores unornamented on the proximal faces..................................................Austrolycopodium
6. Leafy branches anisophyllous, flattened
8. Dorsolateral leaves alternate; dorsal leaves of flattened branchlets 2 or 3; spores with unornamented
proximal faces ................................................................................................................................................................................Diphasium
8. Dorsolateral leaves decussate; dorsal leaves of flattened branchlets 1; spores with reticulate proximal
faces ............................................................................................................................................................................................Diphasiastrum

2. Isoëtaceae
one genus worldwide: Isoëtes

3. Selaginellaceae
one genus worldwide: Selaginella

4. Equisetaceae
one genus worldwide: Equisetum

5. Psilotaceae
one genus in the Americas: Psilotum

6. Ophioglossaceae

1. Blades simple, lacking a midrib; veins anastomosing; fertile spikes unbranched apically, with sporangia
sunken and in two lateral rows
2. Plants terrestrial; blades entire; roots proliferous Ophioglossum.
2. Plants epiphytic; blades lobed; roots not proliferous Cheiroglossa
1. Blades lobed to 5-pinnate, with a midrib; veins free; fertile spikes branched apically, with sporangia
sessile but not sunken
3. Blades seldom over 7 cm wide, 1-pinnate to 2-pinnate .............................................................................................Botrychium
3. Blades often over 7 cm wide, 2-pinnate-pinnatifid to 5-pinnate
4. Fertile spikes attached above ground at base of blade; lvs deciduous, thin and chartaceous ............. Botrypus
4. Fertile spikes attached at or below ground and well below the blade; lvs wintergreen, leathery
or fleshy...............................................................................................................................................................................................Sceptridium

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 7. Marattiaceae
1. Leaves 1-pinnate or (more rarely) simple or 1-pinnate-pinnatifid; sterile and fertile leaves dimorphic;
synangia fused into a double row of spore-bearing compartments, each compartment opening
by a terminal circular pore.................................................................................................................................................................................. Danaea
1. Leaves 2- to 5-pinnate; sterile and fertile leaves monomorphic; synangium a bivalved structure that
spreads wide apart at maturity, each compartment opening by a terminal elongate slit.
2. Synangia stalked; awns present along the veins adaxially; leaves 1(–2) per plant............................................ Eupodium
2. Synangia sessile; awns absent along the veins adaxially; leaves several per plant................................................. Marattia

8. Osmundaceae

1. Leaves monomorphic; sporangia following veins on abaxial surface of uncontracted, photosynthetic pinnae.
2. Laminae membranaceous, without stomata or mesophyll; sporangia sparsely arranged..........................Leptopteris
2. Laminae herbaceous to subcoriaceous, with stomata and mesophyll; sporangia densely arranged, appearing
confluent.....................................................................................................................................................................................................................Todea
1. Leaves hemidimorphic or dimorphic; sporangia on contracted, nonphotosynthetic pinnae
3. Laminae subcoriaceous, pinnate-pinnatifid, dimoprphic; photosynthetic pinnae with tufts of hairs on abaxial
surface near rachises .................................................................................................................................................................. Osmundastrum
3. Laminae herbaceous or subcoriaceous, 1-pinnate to 1-pinnate-pinnatifid or 2-pinnate
4. Laminae subcoriaceous, evergreen, pinnate....................................................................... Osmunda subgen. Plenasium
4. Laminae herbaceous, deciduous, 1-pinnate-pinnatifid or 2-pinnate
5. Laminae 1-pinnate-pinnatifid, hemidimorphic with fertile pinnae
positioned medially ...................................................................................................... Osmunda subgen. Claytonosmunda
5. Laminae 2-pinnate, hemidimorphic with fertile pinnae
positioned apically, or leaves fully dimorphic ................................................................... Osmunda subgen. Osmunda

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 9. Hymenophyllaceae
(Modified from Ebihara et al., 2006)

1. Rhizomes short-creeping to erect; plants typically terrestrial

2. Involucres bivalvate........................................................................................................................ Hymenophyllum subgen. Fuciformia
2. Involucres tubular
3. Blades venation catadromous
4. Laminae flagellate, proliferous; fertile laminae more than 1 cell thick between veins;
rhizomes erect; roots conspicuous.......................................................................................................................Trichomanes subgen. Feea
4. Laminae non-flagellate, non-proliferous; fertile laminae 1 cell thick between veins;
rhizomes creeping; roots inconspicuous or absent................................................................ Trichomanes subgen. Trichomanes
3. Blade venation anadromous
5. Petioles and rachises pubescent, the hairs bristle-like, light reddish ..................Abrodictyum subgen. Pachychaetum
5. Petioles and rachises glabrous
6. Laminae more than 1 cell thick between veins, usually iridescent.....................Trichomanes subgen. Davalliopsis
6. Laminae 1 cell thick between veins, non-iridescent.....................................................................................................Abrodictyum
1. Rhizomes long-creeping; plants typically epiphytic or hemiepiphytic
7. Involucres bivalvate; rhizomes glabrous, or if sparsely pubescent, the hairs pale brown................... Hymenophyllum
7. Involucres tubular to cylindrical; rhizomes pubescent, the hairs dark
8. False veins present (either between and parallel to main veins or continuous and submarginal)
9. Submarginal false vein absent; lamina margins pubescent,
the hairs dark................................................................................................................................Didymoglossum subgen. Didymoglossum
9. Submarginal false vein present; lamina margins glabrous......................................Didymoglossum subgen. Microgonium
8. False veins absent
10. Rhizomes 0.5–1 mm wide; roots absent or fine and few, usually replaced by short,
lateral, root-like rhizomes
11. Rhizome hairs black; petioles pubescent basally and petiole wings pubescent
with hairs like those on the rhizome; segments with folds parallel to the
veins (these often appearing to be false veins) .............................................................................................................. Crepidomanes
11. Rhizome hairs reddish brown; petioles glabrous; segments without folds
parallel to veins ....................................................................................................................................................................................Polyphlebium
10. Rhizomes > 1 mm wide; roots many and thick
12. Laminae 1-pinnate to bipinnatifid
13. Leaves spreading from substrate; pinnae asymmetric; sori oriented downward (abaxially)
or in same plane as lamina.......................................................................................... Vandenboscshia subgen. Lacosteopsis
13. Leaves appressed to substrate; pinnae symmetric; sori oriented outward (adaxially)
at right angles to substrate........................................................................................................... Trichomanes subgen. Lacostea
12. Laminae 2-pinnate to 5-pinnate
14. Lamina margins pubescent; plants terrestrial,
less commonly epiphytic....................................................................................................... Trichomanes subgen. Trichomanes
14. Lamina margins glabrous; plants hemiepiphytic....................................... Vandenboscshia subgen. Lacosteopsis

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 10. Matoniaceae (Old World)

11. Dipteridaceae (Old World)

12. Gleicheniaceae

1. Rhizomes and buds in forks of the pinnae pubescent; sporangia 8-25 per sorus; veins 2- to 4-forked.
2. Two small pinnae present at the base of each bifurcation; rhizomes protostelic;
spores trilete; x=39. Pantropical..................................................................................................................................................Dicranopteris
2. Two small pinnae absent at the base of each bifurcation; rhizomes solenostelic;
spores monolete; x=43. Tropical America, monotypic.......................................................................................................Gleichenella
1. Rhizomes and buds in forks of the pinnae scaly; sporangia 2-5(-6) per sorus; veins simple or 1-forked.
3. Pinnae not forked; spores trilete................................................................................................................................... Diplopterygium
3. Pinnae forked; spores monolete or trilete.
4. Ultimate segments deltate to oblong to linear; spores monolete; x=34. Pantropical............................. Sticherus
4. Utimate segments rounded, small; spores trilete; x=20, 22. S. Africa, Mascarenes,
Malaysia and Australasia................................................................................................................................................................. Gleichenia

13–15 Schizaeales

1. Leaves climbing by twinning rachises; pinnae short-stalked and bifurcate at the base with a small
bud in the bifurcation; each sporangium covered by a scale-like flap green laminar tissue...... Lygodiaceae (Lygodium)
1. Leaves not climbing; pinnae not bifurcate at the base; sporangia naked
2. Blades 1- to 3-pinnate, lanceolate to ovate to deltate; veins pinnate; spores trilete,
tetrahedral-globose ....................................................................................................................................................... Anemiaceae (Anemia)
2. Blades entire, linear, or if expanded, fan-shaped; veins isodichotomous; spores monolete, ellipsoid.
3. Sporangiophores more than 2 cm long, subdigitate, vertical; sporangia in 2 or more rows
each side of the midrib; spores longitudinally parallel-striate; gametophytes subterranean, tuberous,
............................................................................................................................. Schizaeaceae (Actinostachys)
3. Sporangiophores less than 2 cm long, pinnate, ascending; sporangia in 1 row
on either side of the midrib; spores foveolate to granulate; gametophytes superficial, filamentous
.................................................................................................................................... Schizaeaceae (Schizaea)

16. Salviniaceae

1. Leaves 0.5--3 mm long, alternate; roots present, unbranched..................................................................................................... Azolla

1. Leaves 4—30 mm long, whorled, the third leaf root-like and submerged; roots absent.......................................... Salvinia

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 17. Marsileaceae

1. Laminae absent, fronds consisting solely of filiform petioles; sporocarps single;

sori 2 or 4 per sporocarp.................................................................................................................................................................................. Pilularia
1. Laminae present; sporocarps several or (less commonly) single;
sori several per sporocarp.
2. Pinnae 4; veins usually netted; sporocarps with 1 locule. Cosmopolitan....................................................................Marsilea
2. Pinnae 2; veins free; sporocarps with 2 locules. se. Brazil and Argentina....................................................... Regnellidium

18–25 Cyatheales

1. Rhizomes glabrous; leaves dimorphic.................................................................................................. Plagiogyriaceae (Plagiogyria)

1. Rhizomes hairy, scaly, or bristly; leaves monomorphic.
2. Sori abaxial (borne on the undersurface of the frond)
3. Stems and leaves scaly................................................................................................................................ Cyatheaceae (4 genera)
3. Stems and leaves hairy.
4. Leaves 1-pinnate, not glaucous beneath; plants
usually found below 1000 m................................................................................................................ Metaxyaceae (Metaxya)
4. Leaves 2- to 3-pinnate-pinnatifid, glaucous beneath; plants
usually found above 1000 m.................................................................................................... Dicksoniaceae (Lophosoria)
2. Sori marginal.
5. Rhizomes scaly; plants from southern Venezuela............Cyatheaceae (Cyathea, Hymenophyllopsis clade)
5. Rhizomes hairy; plants from elsewhere.
6. Stems long-creeping, slender, covered by dark bristles; indusia cylindrica.Loxomataceae (Loxomopsis)
6. Stems erect, often massive, sometimes arborescent, covered by golden
to brown hairs; indusia bivalved, clam-like.
7. Laminae gradually reduced toward the base ..........................................................Dicksoniaceae (Dicksonia)
7. Laminae widest at the base.......................................................................................................Cibotiaceae (Cibotium)

25. Cyatheaceae

1. Petiole scales conform (i.e., cells of the same size, shape, and orientation); spores spiny........................ Sphaeropteris
1. Petiole scales marginate (i.e., marginal cells differentiated from the central ones); spores ridged or perforate
2. Petiole scales without a dark apical seta...................................................................................................................................... Cyathea
2. Petiole scales with a dark apical seta
3. Indusia absent; aflebia present at base of petiole; spores 64 per sporangium............................... Gymnosphaera
3. Indusia present; aflebia absent; spores 16 per sporangium..................................................................................... Alsophila

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 26. Saccolomataceae
one genus worldwide: Saccoloma

27. Cystodiaceae (Old World)

28. Lonchitidaceae
one genus worldwide: Lonchitis

29. Lindsaeaceae

1. Sori supplied by two or more veins; ultimate segments dimidiate (midrib along one side)................................ Lindsaea
1. Sori supplied by one vein; ultimate segments not dimidiate
2. Laminae 1-pinnate ................................................................................................................................................ Lindsaea (L. imrayana)
2. Laminae 2- to 4-pinnate
3. Leaves 1-6 m long, scandent, indeterminate..............................................................................................................Odontosoria
3. Leaves to 0.5 m long, erect to spreading, determinate..................................................................................... Sphenomeris.

30. Pteridaceae (5 subfamilies)

1. Plants aquatic, either floating or emergent.......................................................................................... 30a. Subfam. Parkerioideae
1. Plants terrestrial, saxicolous, or epiphytic
2. Plants epiphytic .................................................................................................. 30b. Subfam. Adiantoideae (vittarioid clade)
2. Plants terrestrial or saxicolous
3. Sporangia borne on the indusium...................................................................30b. Subfam. Adiantoideae (Adiantum)
3. Sporangia borne on lamina tissue beneath the indusiu
4. Leaves slightly or strongly holodimorphic or hemidimorphic; segments generally elliptic......................................
.............................................................................................................30c. subfam. Cryptogrammoideae
4. Leaves monomorphous or, if slightly dimorphic, segments various
5. pores without an equatorial flange..........................................................................30e. subfam. Cheilanthoideae
5. pores with an equatorial flange...........................................................................................30d. subfam. Pteridoideae

30a. Subfamily Parkerioideae

1. Mature leaves 1–4 m long, 1-pinnate, coriaceous............................................................................................................... Acrostichum

1. Mature leaves 0.2–0.5 m long, 2–4 pinnate, succulent .....................................................................................................Ceratopteris

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 30b. Subfam. Vittarioideae (excluding Adiantum)

1. Blades forked; veins free....................................................................................................................................................................Hecistopteris

1. Blades entire; veins anastomosing
2. Veins anastomosing to form a single row of areolae; paraphyses present
3. Leaves less than 4 mm wide; apical cell of paraphyses narrow..................................................................................Vittaria
3. Leaves usually more than 4 mm wide; apical cell of paraphyes obconic
4. Rhizomes short-creeping, dorsiventral; paleotropical.....................................................................................Haplopteris
4. Rhizomes sub-erect, radial; neotropical................................................................................................................Radiovittaria
2. Veins anastomosing to form three to many rows of areolae; paraphyses present or absent.
5. Spores reniform, monolete
6. Apical cell of paraphyses narrow; sori in two submarginal lines arranged
parallel to the costa........................................................................................................................................................... Ananthacorus
6. Apical cell of paraphyses round; sori in a reticulating or oblique to the costa................................ Scoliosorus
5. Spores tetrahedral, trilete
7. Costa absent, or if present, not extending to the blade tip; paraphyses present;
paleotropical.............................................................................................................................................................................. Antrophyum
7. Costa present throughout the blade; paraphyses absent; neotropical................................................Polytaenium

30c. Subfamily Cryptogrammoideae

1. Leaves holodimorphic, often succulent; temperate..................................................................................................... Cryptogramma

1. Leaves hemidimorphic (fertile pinnules apical, contracted), often chartaceous; Texas, s. Mexico, Guat. ........... Llavead

30d. Subfamily Pteridoideae

1. Sori born along a vein parallel to the lamina margin; petioles in cross-section generally exhibiting a U-shaped vas-
cular bundle; union of costae and costules often with a short awn adaxially............................................................................. Pteris
1. Sori born along veins running to the lamina margin; petioles in cross-section exhibiting a 1 or 2 terete or elongate
vascular bundles
2. Laminae abaxially usually farinose, the farina usually white or yellow............................................................Pityrogramma
2. Laminae abaxially non-farinose
3. Leaves 2–25 cm long, membranaceous, delicate....................................................................................................Anogramma
3. Leaves 10–35 cm long, chartaceous to coriaceous
4. Segments > 1 cm wide; generally of the tepui region of the Guayanan Sheild............................Pterozonium
4. Segments < 1 cm wide; Central America, Andes, and se. Brazil
7. Rachises stramineous; se. Brazil ...................................................................................................................................Tryonia
7. Rachises castaneous; Central America, Andes, and se. Brazil .................................................................Jamesonia

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 30e. Subfamily Cheilanthoideae
1. Lamina margins with a dark sclerenchymatous spots at the base of the sinuses or (in a few spp.) a dark sclerenchy-
matous band throughout margin.........................................................................................................................................................Doryopteris.
1. Lamina margins without a dark sclerenchymatous spots or bands
2. IAdaxial grooves of petioles and rachises densely pubescent by clavate hairs; fertile segments with blade mar-
gins projecting beyond the scarious false indusial, serrate...................................... Allosorus (Mildella) intramarginalis
2. Adaxial grooves of petioles and rachises glabrous or, if hairs present, these usually flexuous and septate; fertile
segments with blade margins not projecting beyond the scarious false indusia
3. Petioles stramineous to light brown, if atropurpureous, then glabrous or with flexuous septate hairs
0.5–0.6 mm long; axes lacking scales................................................................................................................................................. Pellaea
3. Petioles atropurpureous to castaneous, if lighter, then glandular-viscid; axes often scaly in Cheilanthes and
Myriopteris
4. Blades glabrous (except for 0.1 mm clavate hairs abaxially in Adiantipsis), htn to chartaceous; sori dis-
crete, usually many per ultimate segment.
5. Blades digitate (pinnae radiating from apex of petiole); segments obtuse, oblong; segments abaxially
with minute (< 0.1 mm long) white clavate hairs ..........................................................................................Adiantopsis
5. Blades pinnately divided; segments acuminate, linear, 0.5–1.3 mm wide; segments glabrous abaxially
6. Laminae adaxially faintly striate; segments acute to mucronate .......................................................Aspidotis
6. Laminae adaxially not striate; segments acute to obtuse .............................................................................. Gaga
4. Blades scaly, hairy, or glabrous; sori continuous, or if discrete, blades firm to coriaceous
7. Laminae abaxially farinose; marginal indusial absent.................................................................................Notholeana
7. Laminae abaxially non-farinose, or if farinose, then with conspicuous marginal indusial; marginal indu-
sial usually present although sometimes poorly differentiated..........................Cheilanthes, Cheiloplecton,
Myriopteris

31. Dennstaedtiaceae
1. Sorus supplied by one vein.
2. Indusia scale-like, formed by the recurved margin of the lamina................................................................................Hypolepis
2. Indusia cup-or purse-shaped
3. Sori marginal; lower indusia similar in texture to the above ones, marginal.........................................Dennsteadtia
3. Sori submarginal; lower indusia scarious, submarginal...............................................................................................Microlepia
1. Sorus supplied by two or more veins.
4. Veins anastomosing; pinnae opposite or subopposite
5. Laminae glabrous, often glaucous; rhizomes scaly....................................................................................................Histiopteris
5. Laminae glabrous, often glaucous; rhizomes pubescent.................................................................................................Blotiella
4. Veins free; pinnae all or mostly alternate
6. Rachises flexuous; spores monolete............................................................................................................................................. Paesia
6. Rachises straight; spores trilete............................................................................................................................................... Pteridium

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 32. Cystopteridaceae

1. Sori indusiate; rachis not slightly swollen at junction with pinnae................................................................................. Cystopteris
1. Sori non-indusiate; rachis slightly swollen at junction with pinnae.......................................................................Gymnocarpium

33. Rhachidosoraceae (Old World)

34. Diplaziopsidaceae
only one genus: Homalosorus

35. Desmophlebiaceae
one American genus: Desmophlebium

36. Hemidictyaceae
only one genus: Hemidictyum

37. Aspleniaceae

1. Rhizomes erect, meristeles not elongate; petiole bases not swollen at juncture with rhizome.................... Asplenium
1. Rhizomes creeping, with elongated (in cross-section) ventral meristele; petiole bases swollen at juncture with rhi-
zome.........................................................................................................................................................................................................Hymenasplenium

38. Woodsiaceae
only one genus: Woodsia

39. Onocleaceae

1. Veins free......................................................................................................................................................................................................... Matteuccia

1. Veins anastomosed
2. Laminae widest basally; rhizomes creeping; phyllopodia present............................................................................ Onoclea
2. Laminae narrowed basally; rhizomes erect; phyllopodia absent..................................................................... Onocleopsis

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 40. Blechnaceae
1. Sori several along midrib, chainlike, with several short sori arranged end-to-end (Woodwardioideae)
2. Sterile leaves free-veined; pinnae articulate to the rachis.................................................................... Anchistea (virginica)
2. Sterile leaves areolate-veined; pinnae continuous with the rachis
3. Sterile and fertile leaves dimorphic, pinnatisect; rhizomes long-creeping ...........................Lorinseria (areolata)
3. Sterile and fertile leaves monomorphic, 1-pinnate-pinnatifid; rhizomes short-creeping to suberect......................
..................................................................................................................................................................................................................Woodwardia
1. Veins free; sori single along midrib, linear
4. Fertile leaves 2-pinnate; rachises twining; indusia eventually falling away with sporangia attached.... Salpichaena
4. Fertile leaves simple to 1-pinnate; rachises not twining; indusia persistent
5. Sterile and fertile leaves monomorphic or nearly so
6. Pinnae serrate
7. Lamina apices gradually tapered, pinnatifid; pinnae 8–22 × 1.2–2 cm, broadly adnate; rhizomes erect,
often trunk-like; rhizome scales >10 mm long............................................................ Neoblechnum (brasiliense)
7. Lamina apices conform or nearly so (resembling the lateral pinnae); pinnae 4–15 × 0.5–1.5 cm, ses-
sile but not adnate; rhizomes creeping and becoming erect at their tips when bearing leaves; rhizome
scales <3 mm long............................................................................................................................................ Telmatoblechnum
6. Pinnae entire
8. Rhizomes stoloniferous; leaves <1 m long; rachises lacking scales or nearly so; pinnae adnate or con-
stricted at bases and truncate or rounded, not stalked, entire.................................................................... Blechnum
8. Rhizomes not stoloniferous; leaves >1 m long; rachises with scattered to numerous persistent scales;
pinnae constricted at bases, cordate, stalked, serrulate, especially toward tips........................Parablechnum
5. Sterile and fertile leaves dimorphic
9. Plants arborescent, the trunks to 4 m tall; rhizome scales 2–4 cm long, curved-acicular, with dark central
stripe.............................................................................................................................................................................................Lomariocycas
9. Plants non-arborescent; rhizome scales < 2 cm long, ovate to linear (but not curved), with or without a
dark central stripe
10. Rachises and costae pubescent abaxially........................................................................................................... Cranfillia
10. Rachises and costae glabrous abaxially
11. Proximal pinnae of sterile leaves sessile (but not adnate) or short-stalked
12. Rachises not or only shallowly grooved adaxially...........................................................Parablechnum
12. Rachises deeply grooved adaxially........................................................................................................ Lomaria
11. Proximal pinnae of sterile leaves adnate throught their entire width
13. Plants hemiepiphytic, never stolonferous; rhizome in cross-section with meristeles arranged
in a circle, each surrounded by a dark sclerenchyma sheath.................................................Lomaridium
13. Plants terrestrial, sometimes stoloniferous; rhizome in cross-section with meristeles ar-
ranged indistinctly, not surrounded by a dark sclerenchyma sheath
14. Pinnae serrate to crenate at least distally; tropical and subtropical......Austroblechnum
14. Pinnae entire; aerophores absent; north-temperate........................................... Struthiopteris

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 41. Athyriaceae
1. Rhizomes creeping, protruding beyond current year's growth of leaves; leaves distichous............... Ephemeropteris
1. Rhizomes erect to decumbent, not protruding beyond current year's growth of leaves; leaves polystichous
2. Sori (usually the basal ones) paired back-to-back along the same veins, straight, linear...............................Diplazium
2. Sori (at least some) unpaired, sometimes C- or J-shaped, rarely round
3. Sori round; indusia absent or inconspicuous, often hidden beneath sporangia......................... Pseudoathyrium
3. Sori elongate or linear; indusia present, conspicuous (at least when young)
4. Rachises and costae pubescent adaxially, the grooves of the costules interrupted and not decurrent
into those of the rachis...................................................................................................................................................................Deparia
4. Rachises and costae glabrous adaxially, the grooves of the costules decurrent into those of the rachis
........................................................................................................................................................Athryium

42. Thelypteridaceae

1. Suprabasal pinnae adnate to the rachis

2. Laminae deltate, bipinnate-pinnatifid; sori round; sporangial capsules glabrous............................................ Phegopteris
2. Laminae elliptic-oblong, 1-pinnate basally; sori elliptic; sporangial
capsules setose................................................................................................................................................................................. Stegnogramma
1. Suprabasal pinnae sessile (but not adnate) or stalked
3. Laminae 3- to 4-pinnate; rachis adaxially not grooved.................................................................................. Macrothelypteris
3. Laminae simple to (mostly) 1-pinnate-pinnatisect; rachis adaxially usually grooved
4. At least some of the hairs on the rachis, costae, and/or stem apex scales
forked or stellate............................................................................................................................................................................Goniopteris
4. Hairs on the rachis, costae, and/or stem apex scales simple and unbranched, or rarely absent
5. Laminae 1-pinnate, the pinnae either cut < half way to the costae or entire, undulate or serrate; veins
areolate, formed by united cross veins in 3–25 series between costa and margin
6. Sori mostly discrete, indusiate.............................................................................................................................. Goniopteris
6. Laminae more than 1 cell thick between veins, usually iridescent..................................................... Meniscium
5. Laminae 1-pinnate-pinnatifid or rarely 2-pinnate, the pinnae usually incised half their width or more;
veins meeting margin above the sinus, connivent at the sinus, or with 1 or 2 pairs anastomosing below the
sinus
7. Veins 1- or 2-forked; proximal pinnae not or only slightly shorter than the
longest pairs............................................................................................................................................................................Thelypteris
7. Veins usually simple; proximal pinnae much shorter than the longest or not

140
 8. Lamina with one or usually several pairs of reduced proximal pinnae, the
lowermost vestigial or hastate; veins free, meeting the margin above the sinus
or rarely a the sinus ........................................................................................................................................... Amauropelta
8. Lamina without reduced proximal pinnae, or if reduced pinnae present,
then veins united below the sinus
9. Sori elliptic; sporangial capsules setose..................................................................................... Stegnogramma
9. Sori round; sporangial capsules glabrous
10. Cartilaginous keel or minutely pubescent; false vein running from
sinus toward costa, occasionally lacking; indusia present or absent;
aerophores present at pinna bases abaxially................................................................................Steiropteris
10. Cartilaginous keel lacking; indusia present, persistent; aerophores
absent at pinna bases abaxially............................................................................................................. Cyclosorus

43. Didymochlaenaceae
one genus worldwide: Didymochlaena

44. Hypodematiaceae (Old World)

45. Dryopteridaceae

1. Rhizomes creeping, in cross section with an elongated ventral meristele bearing all the roots; sterile and fertile
leaves dimorphic; sori acrostichoid; laminae generally without hairs (bolbitidoid fern clade)
2. Leaves simple, entire, or rarely (6 species) divided; veins free or rarely (3 species) anastomosing; phyllopodia
often present......................................................................................................................................................................................Elaphoglossum
2. Leaves 1-pinnate or rarely (2 species) simple; veins anastomosing; phyllopodia absent
3. Bulblets absent or, if present, born at the base of the pinna near the rachis; plants scandent hemiepiphytes
or (3 species) terrestrial..........................................................................................................................................................................Mickelia
3. Bulblets present, rarely (2 species) absent, born near the apices of the pinnae or (usually) the terminal seg-
ment; plants terrestrial, rarely low hemiepiphytes......................................................................................................................Bolbitis
1. Rhizomes erect to decumbent, or if creeping then in cross section without an elongated ventral meristele bearing
all the roots; sterile and fertile leaves monomorphic or dimorphic; sori round or (rarely) acrostichoid; laminae with
or without hairs
4. Rhizomes long-creeping (polybotryoid fern clade)
5. Sterile and fertile leaves monomorphic; laminae pubescent by long straight multicellular whitish hairs; plants
terrestrial; West Indies, Nicaragua...................................................................................................................................... Polystichopsis
5. Sterile and fertile leaves strongly dimorphic; laminae glabrous or, if pubescent, the hairs lax and slightly
tortuous; plants terrestrial or scandent on tree trunks; widespread

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 45. Dryopteridaceae (cont.)
6. Vein tips connected by a submarginal connecting strand ..................................................................................... Olfersia
6. Veins tips not connected by a submarginal connecting strand
7. Rhizomes with meristeles surrounded by a dark sclerenchymatous sheath; sori non-indusiate; rachis
groove pubescent within..................................................................................................................................................Polybotrya
7. Rhizomes with meristeles not surrounded by a dark sclerenchymatous sheath; sori indusiate; rachis
groove glabrous withint..........................................................................................................................................................Maxonia
4. Rhizomes short-creeping, decumbent, or erect
8. Lamina margins cartilaginous-spinulose (polystichoid fern clade)
9. Lamina apices gradually tapered, pinnatifid; pinnae and/or pinnules
with a basal acroscopic lobe or auricle........................................................................................................................Polystichum
9. Lamina apices conform, resembling a lateral pinnae; pinnae and/or pinnules
without a basal acroscopic lobe.............................................................................................................................. Phanerophlebia
8. Lamina margins entire to serrate with soft-herbaceous teeth
10. Hydathodes present; medial pinnae with pinnules (if present) arranged catadromically
11. Laminae with internal punctuate glands (best seen in transmitted light); rachises and costae gen-
erally without hairs abaxially; distal pinnules with the basal basiscopic vein springing from the costule,
not the costa............................................................................................................................................................... Stigmatopteris
11. Laminae without internal punctuate glands; rachises and costae generally pubescent abaxially;
distal pinnules with the basal basiscopic vein springing from the costa, not the costule.....Megalastrum
10. Hydathodes absent; medial pinnae with pinnules (if present) usually arranged anadromically
12. Rhizomes decumbent to erect
13. Pinnae or pinnule margins thickened and decurrent onto axis of the next lower order
14. Buds absent distally on rachis ...........................................................................................Parapolystichum
14. Buds present distally on rachis ......................................................................................................Lastreopsis
13. Pinnae or pinnule margins not thickened nor decurrent onto axis of the next lower order
15. Rachises not or only shallowly grooved adaxially;
costa scales typically clathrate .......................................................................................................................... Ctenitis
15. Rachises grooved adaxially; costal scales not clathrate ....................................................Dryopteris
12. Rhizomes creeping
16. Rachises and costal grooves pubescent within
17. Basal pinnae equilateral; spores non-echinulate or only sparsely so......................... Cyclodium
17. Basal pinnae inequilateral, prolonged basiscopically;
spores densely echinulate..........................................................................................................................Arachniodes
16. Rachises and costal grooves glabrous within
18. Basal pinnae equilateral; spores non-echinulate or only sparsely so.............................Rumohra
18. Basal pinnae inequilateral, prolonged basiscopically;
spores densely echinulate ......................................................................................................................Arachniodes

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 46. Nephrolepidaceae
one genus worldwide: Nephrolepis

47. Lomariopsidaceae

1. Plants scandent on tree trunks; sterile and fertile leaves dimorphic; sori acrostichoid.................................. Lomariopsis
1. Plants terrestrial; sterile and fertile leaves monomorphic; sori round.
2. Leaves simple, entire, with apical bud; veins netted..................................................................................................Dracoglossum
2. Leaves 1-pinnate, without apical bud; veins free................................................................................................................ Cyclopeltis

48. Tectariaceae

1. Plants scandent on tree trunks; Juan Fernández Islands.................................................................................................. Arthropteris

1. Plants terrestrial or epipetric; widespread
2. Rhizomes erect or decumbent; leaves arranged radially
3. Rachises not winged; crushed rhizomes and foliage without odor.........................................................................Tectaria
3. Rachises winged; crushed rhizomes and foliage
with odor of toasted almands (cyanogenic) ........................................................................................................... Draconopteris
2. Rhizomes creeping; leaves arranged distichously
4. Spores cristate; veins free................................................................................................................................................. Triplophyllum
4. Spores spiny; veins netted, or if free, then rachis winged.................................................................................... Hypoderris

49. Oleandraceae
one genus worldwide: Oleandra

50. Davalliaceae (Old World)

143
 51. Polypodiaceae

1. Spores yellow, monolete, ellipsoid....................................................................................................................... Polypodioid ferns, Key A

1. Spores green, and (except Loxogramme) trilete, globose-tetrahedral........................................... Grammitid ferns, Key B

Polypodioid ferns, Key A

1. Sori acrostichoid; sterile leaves appressed to substrate, brown at maturity; Peru, Bolivia...... Platycerium andinum
1. Sori round to linear; sterile leaves spreading, green at maturity; widespread
2. Leaves simple, entire
3. Sori linear............................................................................................................................................................................................ Pleopeltis
3. Sori round
4. Sori several between rachis and lamina margin
5. Sori in 2 or more rows between the main lateral veins running from
midrib to margin.................................................................................................................................................. Campyloneurum
5. Sori in 1 row between the main lateral veins running from midrib to margin.......................... Niphidium
4. Sori in 1 row between rachis and lamina margin
6. Veins in a chevron-like pattern with a single excurrent vein in each areole.. Serpocaulon levigatum
6. Veins anastomosed variously, not chevron-like
7. Rhizome scales clathrate; sori covered when young by peltate scales...................................... Pleopeltis
7. Rhizome scales not clathrate; sori not covered when young by peltate scales.......... Microgramma
2. Leaves pinnatisect to (rarely) 2-pinnate-pinnatisect
8. Rachises adaxially puberulent; sporangial capsules often setulose; rhizome scales often with brown hairs at
the point of attachment
9. Laminae pectinate; pinna pairs 20–50............................................................................................................................ Pecluma
9. Laminae pinnate but not pectinate; pinna pairs 1–20................................................... Polypodium dulce Group
8. Rachises adaxially glabrous; sporangial capsules glabrous; rhizome scales glabrous
10. Laminae moderately to densely scaly; sori when young covered by peltate scales...................... Pleopeltis
10. Laminae glabrous or nearly so; sori when young without peltate scales
11. Sori densely paraphysate; s. Chile, Argentina................................................................................................. Synnamia
11. Sori not paraphysate; widespread elsewhere
12. Leaves pinnatifid, the apical segment conform (i.e., resembling the lateral pinnae); sori typically
supplied by two veins............................................................................................................................................. Phlebodium
12. Leaves pinnatifid or 1-pinnate; the apices gradually tapered and pinnatifid; sori supplied by a
single vein
13. Leaves 1-pinnate with 4–7 pairs of pinnae,
these 5–11 cm wide................................................................................................Campyloneurum magnificum
13. Leaves pinnatisect or, if 1-pinnate, with 7–30 pairs of pinnae,
these 0.5–5 cm wide ......................................................................................................................................Serpocaulon

144
Grammitid ferns, Key B
1. Leaves non-setose
2. Laminae entire and black-margined............................................................................................................................................Grammitis
2. Laminae entire to 1-pinnate-pinnatisect, not black-margined
3. Rhizome scales non-clathrate
4. Laminae glandular-pubescent, lax and pendent; rhizome scales orange,
golden-brown or castaneous, shiny............................................................................................................................. Galactodenia
4. Laminae glabrous or nearly so, erect or arching; rhizome scales tan, dull............................................Cochlidium
3. Rhizome scales clathrate, often dark or blackish
5. Leaves 2–5 mm wide; veins and sori one per segment.......................................................................Stenogrammitis
5. Leaves > 10 mm wide; veins and sori several per segment
6. Leaves pinnatifid to 1-pinnate-pinnatifid; sori round, often sunken; spores trilete..................... Lellingeria
6. Leaves simple, entire; sori elongate, superficial, not sunken; spores monolete......................Loxogramme
1. Leaves setose, the setae straight and stiff
7. Hydathodes absent
8. Whitish or brownish spherical glands present among the sporangia and often
on the laminae; veins free .............................................................................................................................................................. Ceradenia
8. Whitish or brownish glands absent; veins anastomosing
9. Leaves pinnatisect to 1-pinnate; sori round, superficial, not surrounded by reddish setae..... Zygophlebia
9. Leaves entire to pinnatisect; sori elongate, deeply or slightly sunken,
often surrounded by reddish setae..................................................................................................................................Enterosora
7. Hydathodes present............................................................................................................................................................................Grammitis
10. Laminae sparsely beset with black, clavate fungi about 1 mm long ...........................................Ascogrammitis
10. Laminae lacking black, clavate fungi
11. Leaves 2–8(–13) mm wide; petioles typically less than 10 mm long; sori one per segment
12. Setae reddish; rhizome scales brightly colored, shiny, non-clathrate,
the cells inflated......................................................................................................................................................Moranopteris
12. Setae whitish; rhizome scales dark brown to blackish, dull, clathrate,
the cells sunken...................................................................................................................................................... Leucotrichum
11. Leaves > 10 mm wide; petioles longer than 10 mm; sori several per segment
13. Leaves long-pendent; setae often paired or stellate
14. Laminar glandular; sporangial capsules glabrous; rhizome scales
brightly colored, shiny.................................................................................................................................... Galactodenia
14. Laminae non-glandular; sporangial capsules setulose; rhizome scales
dull brown or blackish, dull.................................................................................................................................Alansmia
13. Leaves erect or arching
15. Rhizomes scales clathrate, often reddish and slightly iridescent,
non-setulose; sporangial capsules glabrous...........................................................................................Melpomene
15. Rhizomes scales non-clathrate, brown or blackish, not iridescent,
setulose; sporangial capsules setulose..................................................................................................... Terpsichore

145
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146
 American genera of
FERNS and LICOPHYTES
Géneros americanos de
HELECHOS y LICOFITAS
147
HYMENOPHYLLACEAE
Abrodictyum C. Presl
DESCRIPTION: Terrestrial; roots many, robust,
often stilt-like; rhizomes 1–2 mm wide, erect, ascending
or creeping, unbranched, pubescent, the hairs dark,
multicellular; sterile and fertile leaves up to 30 × 18 cm,
monomorphic, polystichous; laminae 1-cell thick between
the veins, 2-pinnate to 4-pinnate-pinnatifid; ultimate
segments narrow, only a few rows of cells along
the veins, the cells large, evident with 10×(ocluidas en
A. rigidum); veins free, anadromous, false veins absent;
sori marginal; indusia tubular to campanulate, the
mouth entire, truncate; receptacles exert; spores green,
tetrahaedral-globose; x=33.
DESCRIPCIÓN: Terrestres; raíces numerosas,
robustas, a menudo como raíces zancudas; rizomas 1–2
mm de ancho, erectos, ascendentes o reptantes, no
ramificados, pubescentes, los pelos oscuros, multicelulares;
hojas estériles y fértiles hasta 30 × 18 cm, monomorfas,
polistícas; láminas una célula de grosor entre las venas,
2-pinnadas a 4-pinnado-pinnatífidas; últimos segmentos
estrechos, solamente unas pocas hileras de células a
lo largo de las venas, las células grandes, evidentes con
10× (ocluidas en A. rigidum); venas libres, anádromas, venas
falsas ausentes; soros marginales; indusios tubulares,
la boca entera, tuncada; receptáculos exertos; esporas
verdes, tetraédricas-globosas; x=33.

SIMILAR GENERA: Trichomanes subgen. Trichomanes GÉNEROS PARECIDOS: Trichomanes subgen.

usually differs by less divided laminae, catadromous
venation, and sori that tend to be clustered toward the
pinna tips. Trichomanes subgen. Davalliopsis differs by
laminae opaque, 3-cells thick between the veins, and
usually iridescent. Hymenophyllum differs by epiphytic
habit, creeping rhizomes, two-valved indusia, and included
receptacles.
Trichomanes difiere usualmente por láminas menos
divididas, nervación catadrómica y soros que tienden a
ser agrupados hacia los ápices de las pinnas. Trichomanes
subgen. Davalliopsis difiere por láminas de 3 células
de grosor entre las venas y usualmente iridiscentes.
Hymenophyllum difiere por hábito epifítico, rizomas
reptantes, indusios bivalvados y receptáculos incluidos.

COMMENTS: Abrodictyum is pantropical and contains COMENTARIOS: Abrodictyum es pantropical y

about 25 species. Four are recognized en the Neotropics:
A. cellulosum, A. sprucei, A. rigidum, and A. windischianum.
These belong to subgen. Pachychaetum. They typically
have short erect rhizomes supported by prop roots and
in aspect resemble a small tree. The large cells along the
narrow margin of veins are easily visible with a 10× lens,
although in T. ridigum (the most widespread neotropical
species) the cells are occluded. Sometimes A. cellulosum
has three-dimensional laminae.The chromosome number
of x=33 is derived within the family (Ebihara et al., 2007).
Abrodictyum is derived from the Greek abros, fine, delicate
+ dictyon, net. The cell walls appear as a net.
consta de 25 especies. Cuatro especies se reconocen
en el Neotrópico: A. cellulosum, A. sprucei, A. rigidum y A.
windischianum. Estas pertenecen al subgén. Pachychaetum.
Típicamente tienen rizomas cortos y erectos, soportados
por raíces zancudas, con el aspecto de un arbolito. Las
células grandes a lo largo de los márgenes estrechos de las
venas son fácilmente visibles con una lupa de 10×, aunque
en A. rigidum (la especie neotropical más difundida) las
células están ocluídas. A veces A. cellulosum tiene laminas
tres-dimensionales. El número cromosómico de x=33
es derivado dentro de la familia (Ebihara et al., 2007).
Abrodictyum deriva del griego abros, fino o delicado +
dictyon, red. Las paredes celulares semejan una red.

LITERATURE: Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of Hymeno-
phyllaceae. Blumea 51: 1–60. Dubuisson, J.-Y., S. Hennequin, L. Bauret, A. Grall, T. Li, A. Ebihara & S. Hennequin. 2016.
Discussion on the taxonomy of African fern Abrodictyum rigidum (Sw.) Ebihara & Dubuisson and description of two new Ab-
rodictyum C. Presl species (Hymenophyllaceae, Polypodiidae) for the Afro-Malagasy region. Phytotaxa 284: 151–168. Ebihara,
A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global molecular phylogeny of the fern genus Trichomanes
(Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of the Linnean Society 155: 1–27. Hébant-Mauri,
R. 1972. Le genre Trichomanes L. (Fougéres Leptosporangiées). Adansonia 12: 469–495.

148
Figure 71. Abrodictyum A–C. A. cellulosum. D–F. A. sprucei. G–H. A. rigidum. (A–F, original, ©Moran, 2010; G, H, from
Mickel & Smith, 2004).

149
PTERIDACEAE
Acrostichum L.
DESCRIPTION: Terrestrial or palustral; roots large,
fleshy; rhizomes erect or ascending, generally clustered;
petioles with many vascular bundles; leaves 2–5 m,
1-pinnate, with a terminal segment similar in shape
to the lateral pinnae, erect or slightly spreading,; pinnae
20–60 pairs, 15–40 × 3–6 cm, the margins entire and
cartilaginously thickened; veins reticulate throughout,
the areoles ca. 1 mm wide, without included veinlets; sori
acrostichoid, paraphysate; spores trilete, whitish to tan;
x=30.
DESCRIPCIÓN: Terrestres o palustres; raíces grandes,
carnosas; rizoma erecto o ascendente, generalmente
amacollado; pecíolos con muchos haces vasculares;
hojas 2–5 m, 1-pinnadas, con una pinna terminal
similar en forma a las pinnas laterales, erectas o un
poco divergentes; pares de pinnas 20 a 60, 15–40 × 3–6
cm, los márgenes enteros y cartilaginosos, engrosados;
nervaduras reticuladas a todo lo largo, las aréolas
ca. 1 mm ancho, sin nervaduras libres incluidas; soros
acrosticoides, parafisados; esporas triletes, blanquecinas
a leonadas; x=30.

SIMILAR GENERA: Pteris differs by non-fleshy roots, GÉNEROS PARECIDOS: Pteris difiere por raíces no
and linear sori protected by a false indusium. carnosas y soros lineales protegidos por un falso indusio.

COMMENTS: Acrostichum is pantropical and has four
species, two of which occur in the Neotropics: A. aureum
L. and A. danaeifolium Langs. & Fischer. The genus is typi-
cally found in brackish or saline habitats near coasts. It can
form dense stands following logging of mangroves. It can
also grow farther inland in wet soils. The ground tissue of
the petioles is composed of aerenchyma, which can easily
be seen in cross section. Aerenchyma is common in many
aquatic plants. The two can be easily distinguished by the
lower surfaces of the pinnae which are puberulent in A.
danaeifolium and glabrous in A. aureum. Also, the peculiar
paraphyses are distinctive, being capitate with a dark and
circular-lobed apex in A. aureum and pale and non-capi-
tate in A. danaeifolium. See Adams & Tomlinson (1979) for
further differences. Within the Pteridaceae, Acrostichum is
sister to Ceratopteris, another aquatic fern genus, but one
quite different in appearance. The genus name is derived
from the Greek akros, summit + stichos, row. In the type
species, the sori are borne only on the distal pinnae.
COMENTARIOS: Acrostichum es pantropical y consta
de cuatro especies, dos de las cuales existen en el
Neotrópico. A. aureum L. y A. danaeifolium Langs. & Fischer.
El género se encuentra típicamente en ambientes salobres
o salinos cerca de las costas. Puede formar poblaciones
densas después de la cosecha de manglares. También
puede ocurrir en tierra firme en suelos húmedos. El
tejido adentro del pecíolo consta de aerénquima, el cual
se puede observar fácilmente en sección transversal.
El aerénquima es común en muchas plantas acuáticas.
Las dos especies pueden ser distinguidas fácilmente
por el envés de las pinnas, el cual es puberulento en A.
danaeifolium y glabro en A. aureum. También, los paráfises
son distintivos, siendo capitados con un ápice oscuro y
circular-lobulados en A. aureum y pálidos y no capitados
en A. danaeifolium.Véase Adams & Tomlinson (1979) para
otras diferencias. Dentro de las Pteridaceae, Acrostichum
es hermano de Ceratopteris, otro género acuático, pero
muy diferente en aspecto. El nombre genérico deriva del
griego akros, cima + stichos, hilera. En la especie tipo, los
soros ocupan solamente las pinnas distales.

LITERATURE: Adams, D. C. & P. B. Tomlinson. 1979. Acrostichum in Florida. American Fern Journal 69: 42–46. García de
López, I. 1978. Revisión del género Acrostichum en la República Dominicana. Moscosoa 1: 64–70. Lloyd, R. M. & T. L. Grergg.
1975. Reproductive biology and gametophyte morphology of Acrostichum danaeifolium from Mexico. American Fern Journal
65: 105–120.

150
Figure 72. Acrostichum A, B. A. aureum. C, D. A. danaeifolium. (From Tryon & Stolze, 1989.)

151
SCHIZAEACEAE
Actinostachys Wall. ex Hook.
DESCRIPTION: Terrestrial; stem short, creeping or
erect, pubescent, the hairs orange or brownish, flexuous;
leaves clumped to spaced; petioles triangular or flattened,
narrowly winged, glabrous, pale green, becoming
brown near the base; laminae linear, grasslike, or
rarely expanded, glabrous; veins free; fertile segments
(sporangiophores) terminal, digitate or subdigitate;
sporangia pear-shaped or oblong, with an apical annulus, in
4–6 rows on the indusiform segments; spores monolete;
gametophytes subterranean, not green, tuberous,
brown hairy; x=134, 140, 168.
DESCRIPCIÓN: Terrestres; rizomas cortos, reptantes o
erectos, pubescentes, los tricomas anaranjados a parduscos,
flexuosos; hojas cespitosas o espaciadas; pecíolos
triangulares o aplanados, estrechamente alados, glabros,
verde pálido a pardos hacia la base; láminas lineares,
graminiformes, escasamente a no foliáceas, glabras;
segmentos fértiles (esporangióforos) terminales,
digitados a subdigitados; esporangios pyriformes u
oblongos, en 4–6 hileras en los segmentos indusiformes;
esporas monoletes; gametofitos subterráneos, no
verdes, tuberosos, pardo-pubescentes; x=134, 140, 168.

SIMILAR GENERA: Schizaea differs by curved, pinnate GÉNEROS PARECIDOS: Schizaea difiere por los
fertile segments, sterile portions of the leaves often segmentos fértiles curvados, pinnados, las partes
dichotomous to flabellate, and sporangia in 2–4 rows. estériles de las hojas a menudo dicotómicas a flabeladas;
esporangios en 2-4 hileras.

COMMENTS: Actinostachys is pantropical and has about
20 species worldwide, two of which are neotropical (A.
pennula and A. subtrigujata). It typically grows on sandy
soil or decaying logs below 600 m. Many species exhibit a
clumped habit that results from repeated branching from
the buds at the base of the petioles. It is often difficult
or find the original stem among the numerous petiole
bases in the clump. In Actinostachys the fertile segments
(sporangiophores) are actually pseudodigitate with a very
short axis. In Schizaea this axis is longer, and the fertile
segments have a feathery appearance. The gametophytes
of Actinostachys are subterranean, tuberous, hairy, and
resemble a short rhizome, whereas those of Schizaea are
superficial, green, and filamentous. Both are mycorrizal.
The species of Actinostachys are often recognized under
Schizaea.The genus name is derived from the Greek aktis,
ray + stachys, spike. The apical segments of the fertile
leaves are digitately arranged.
COMENTARIOS: Actinostachys es pantropical y consta
de casi 20 especies a nivel mundial, dos de las cuales
son neotropicales (A. pennula y A. subtrigujata). Crece
típicamente en suelos arenosos o sobre troncos podridos
por debajo de 600 msnm. Muchas especies exhiben un
hábito amacollado el cual se debe a las ramificaciones
repetidas de brotes en las bases de los pecíolos. Es a
menudo difícil encontrar el tallo original entre las bases
numerosas de los pecíolos. En Actinostachys los segmentos
fértiles (esporangióforos) son en realidad pseudodigitados
con un eje corto. En Schizaea este eje es más largo, y los
segmentos fértiles tienen una apariencia plumosa. Los
gametofitos de Actinostachys son subterráneos, tuberosos,
peludos y se asemejan a un rizoma corto, mientras que
los de Schizaea son superficiales, verdes, y filamentosos.
Los gametofitos en ambos géneros son micorríticos.
A menudo las especies de Actinostachys se tratan bajo
Schizaea. El nombre genérico deriva del griego aktis, raya
+ stachys, espiga. Los segmentos apicales de las hojas
fértiles se disponen digitadamente.

LITERATURE: Lellinger, D. B. 1969. Schizaeaceae (Filicales). In: B. Maguire and collaborators. The botany of the Guayana
Highland—Part VIII. Memoirs of the New York Botanical Garden 18: 2–11. León, B., H. Beltrán & P. Fine. 2005. El género
Schizaea (Schizaeaceae) en el Perú. Revista Peruana de Biología 12: 97–102. Murillo-Pulido, M. T. 1986. Estudio preliminar del
género Schizaea en Colombia. Caldasia 15: 93–101. Takeuchi, M. 1960. O gênero Schizaea no Amazonia. Bol. Mus. Paraense
Emilio Goeldi 5: 1–26.

152
Figure 73. A, B. Actinostachys pennula (Sw.) Hook. (From Mickel & Smith, 2004.)

153
POLYPODIACEAE
Adetogramma T. E. Almeida.
DESCRIPTION: Primarily epiphytic, occasionally
saxicolous; rhizomes long-creeping, scales non-
clathrate, peltate; sterile and fertile leaves monomorphic;
laminae simple, entire, 5–17 x 1–2.5 cm, scaly on
both surfaces, the scales lanceolate, peltate, sessile, non-
clathrate, darkened at point of attachment; veins free,
obscure, 1- or 2-forked, not reaching the lamina margin,
non-hydathodous; sori in one row between the costa
and margin, superficial (not sunken), round or nearly so;
paraphyses resembling the laminar scales but long-stalked,
completely covering the young sori; spores fabiform,
monolete, yellow; x=?
SIMILAR GENERA: Pleopeltis and Microgramma differ
by anstomosing veins. Microgramma further differs by
sessile hairlike or lanceolate paraphyses, and Pleopeltis
further differs by clathrate rhizome scaels and circular,
peltate, pedicellate paraphyses.
COMMENTS: Adetogramma is monotypic, consisting
of a single, Andean species: A. chrysophylla (Hook.) T. E.
Almeida. It occurs from Ecuador to northern Argentina,
from (1800–)3000–4100 m. It is primarily epiphytic but
also occurs occasionally on rocks. Adetogramma is sister
genus to Serpocaulon.The genus name is derived from the
Greek adetos, free + gramme, line. It refers to the free
venation..
LITERATURE: Almeida, T. E., A. Salino, J-Y. Dubuisson & S. Hennequin. 2017. Adetogramma (Polypodiaceae), a new
monotypic fern genus segregated from Polypodium. Phytokeys 78: 109–131.
154
DESCRIPCIÓN: Principalmente epífito, ocasionalmente
saxícola; rizomas largamente rastreros, las escamas no
clatradas, peltadas; hojas estériles y fértiles monomórfas;
láminas simples, enteras, 5–17 x 1–2.5 cm, escamosas
en ambas superficies, las escamas lanceoladas, peltadas,
sésiles, no clatradas, oscuredicas en el punto de inserción;
venas libres, obscuras, 1 o 2 furcadas, no alcanzando
el márgen de la lámina, sin hidatodos; soros en una
sola línea entre la costa y márgen, superficiales (no
hundidos), redondeados o casi así; paráfises se asemejan
las escamas laminares pero largamente pedicelladas,
completamente cubriendo los soros jóvenes; esporas
fabiformes, monletes, amarillas; x=?.
GÉNEROS PARECIDOS: Pleopeltis y Microgramma
difieren por venas anastomosadas. Microgramma difiere
además por los parafisos piliformes o lanceolados, y
Pleopeltis además por las escamas del rizoma clatradas y
los parafises circulares, peltadas, pedicelladas..
COMENTARIOS: Adetogramma es monotípico,
constándose de una sola especie Andina: A. chrysophylla
(Hook.)T. E.Almeida. Existe desde Ecuador hasta Argentina
del norte, de (1800–)3000–4100 m. Es principalmente
epífita pero existe a veces sobre rocas. Adetogramma es
género hermano a Serpocaulon. El nombre genérico se
deriva del griego adetos, libre + gramme, línea. Se refiere
a la venación libre.
Figure 74. A, B. Adetogramma chrysophylla (Hook.) T. E. Almeida. (From Almeida et al, 2017.)

155
PTERIDACEAE
Adiantopsis Fée
DESCRIPTION: Terrestrial or rupestral; rhizomes erect,
decumbent, or short-creeping, scaly, the scales bicolorous
with a dark central stripe; leaves monomorphic, fascic-
ulate, non-farinose; petioles atropurpureous; laminae
pedate, radiate or 1-4-pinnate; pinnules or segments
asymetrical, sessile or short-stalked, glabrous, anadro-
mous; rachises and costa grooved adaxially, with
narrow lateral wings on either side; veins free, simple;
sori marginal, discrete (not in lines), several per segment,
protected by a scarious false indusium; spores echinate or
(2 spp.) cristate, tetrahedral-globose, trilete; x=30.
DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto,
decumbente y cortamente rastrero, escamoso, las escam-
as estrechas, bicolores, con una raya oscura central; hojas
monomorfas, fasciculadas, no farinosas; pecíolos atro-
purpúreos; lámina pedata, radiada o 1 a 4-pinnada; pín-
nulas o segmentos asimétricos, sésiles a cortamente
peciolulados, glabra, anadrómicos; raquis y costas
sulcados o adaxialmente aplanados, con alas estre-
chas y membranáceas en cada lado; nervaduras libres,
simples; soros marginales, discretos (no en líneas), varios
por segmento, protejidos por un indusio falso escarioso;
esporas equinatas o (2 spp.) crestadas, tetraédrico-globo-
sas, triletes; x=30.

SIMILAR GENERA: Adiantum differs by terete petioles
and rachises. Cheilanthes has terete rachises and costae
or, if grooved, then the groove is V-shaped (not flat) and
narrow lateral wings on either side are lacking.
GÉNEROS PARECIDOS: Adiantum difiere por pecío-
los y ráquises teretes. Cheilanthes tiene ráquises y cos-
tas teretes o, si surcados, el surco es en forma de V (no
aplanado) y las alas angostas laterales en cada lado son
ausentes.

COMMENTS: Adiantopsis is completely neotropical and COMENTARIOS: Adiantopsis es completamente neo-

contains ca. 40 species, 9 of which are in the Caribbean.
They occur in wet or seasonally dry forests. It is closely
related to Cheilanthes, especially C. microphylla (Sw.) Sw.
It differs only by the rachis-costa characters. The genus
is notable for containing species with pinnate, pedate, or
digitate lamina architectures. There are ten species with
digitate laminae, with Adiantopsis radiata (L.) Fée being
the most widespread. A key to these ten species may
be found in Link-Pérez et al. (in press). The genus name is
derived from Adiantum and the Greek suffix -opsis, like.
tropical y consta de 40 especies, 9 de las cuales están
en los Caribes. Existe en bosques húmedos o estacio-
nalmente secos. El género es vinculado a Cheilanthes,
especialmente C. microphylla (Sw.) Sw. Difiere solo por
las características del los ráquises y costas. El género es
notable por contener especies con láminas pinnadas, pe-
datas o digitatas. Hay diez especies con láminas digitadas,
con Adiantopsis radiata (L.) Fée siendo la más difundida.
Una clave a estas diez especies se puede encontrar en
Link-Pérez et al. (en prensa). El nombre genérico se deriva
de Adiantum más el sufijo griego, -opsis, parecido.

LITERATURE: Barker, M. S. & R. J. Hickey. 2006. A taxonomic revision of Caribbean Adiantopsis (Pteridaceae). Annals
of the Missouri Botanical Garden 93: 371–401. Hickey, R. J., M. S. Barker & M. Ponce. 2003. An Adiantopsis hybrid from
northeastern Argentina and vicinity. American Fern Journal 93: 42–44. Link-Pérez, M. A. & R. J. Hickey. 2011. Revision of Adi-
antopsis radiata (Pteridaceae) with descriptions of new taxa with palmately compound laminae. Systematic Botany 36: 565–582.
Link-Pérez, M. A., L. E. Watson & R. J. Hickey. 2011. Redefinition of Adiantopsis Fée (Pteridaceae): systematic, diversification,
and biogeography. Taxon 60: 1255–1268. Link-Pérez, M. A., T. G. Ludwig, C. J. Ledford, M. H. Seaboldt & E. B. Sessa.
(en prensa). Lectotypification of Adiantopsis alata (Pteridaceae) and descriptions of new pallmate species in the Guiana Shield.
Systematiic Botany XX: 000–000 Maxon, W. R. 1908. The Cuban species of Adiantopsis. Contributions from the United States
National Herbarium 10: 485–486.

156
Figure 75. A, B. Adiantopsis seemannii (Hook.) Maxon C, D. A. radiata (L.) Fée (From Mickel & Smith, 2004.)

157
PTERIDACEAE
Adiantum L.
DESCRIPTION: Terrestrial or rupestral; rhizome short-
to long-creeping, rarely suberect, scaly; axes of the leaf
dark reddish to black, usually lustrous, terete; lami-
nae simple to 4-pinnate, often reddish when young, elon-
gate spicular cells often present adaxially; ultimate seg-
ments sesile to stalked, never adnate; veins free or rarely
anastomsing; sporangia borne on the inner surface of
the false indusium (not below it); paraphyses absent;
spores globose-tetrahaedral, trilete; x=29, 30.
DESCRIPCIÓN: Terrestres or rupestres; rizoma corta
a largamente rastrero, raramente suberecto, escamoso;
ejes de la hoja pardo-rojizos a negros, lustrosos,
teretes; láminas simples a 4-pinnadas, a menudo ro-
jizas cuando jóvenes, con células espiculares alargadas a
menudo presentes en el haz; últimos segmentos sésiles
a pedicelados, nunca adnatos; nervaduras libres o rara-
mente anastomosadas; esporangios nacen en la cara
interna del falso indusio; parafisos ausentes; esporas
globoso-tetrahédricas, triletas; x=29, 30.

SIMILAR GENERA: Adiantopsis differs by winged peti-
oles and rachises and echinate spores. Lindsaea differs by
its greenish to stramineous, grooved or strongly angled
axes, and submarginal sori with an indusium that open
toward the margen.
GÉNEROS PARECIDOS: Adiantopsis differs by peciío-
los y ráquises alados y esporas equinadas. Lindsaea difiere
por los ejes verdosos o pajizos y sulcados o fuertemente
angulados, y los indusios submarginales con indusios que
abren hacia el margen.

COMMENTS: Adiantum is a cosmopolitan genus of COMENTARIOS: Adiantum es un género cosmopol-

about 200 species, the majority of which are neotropical.
The species grow in a variety of habitats such as shaded
forests and on rocks, but they are always terrestrial. The
autapomorphy for the genus are sporangia borne on the
inner face of the enrolled margin of the segment (the
false indusium). In other Pteridaceae with false indusia, the
sporangia are borne on the lower surface of the lamina,
opposite the recurved margin. Phylogenetic studies show
Adiantum sister to the vittarioid ferns. There are several
well-defined groups within Adiantum. Two of them are
important in the Neotropics and easy to recognize. They
are the group of A. tetraphyllum (laminae 2-pinnate with
a conform apex, pinnules dimidiate) and the group of A.
capillus-veneris (laminae decompound, segments cuneate,
flabellate or nearly round, sori born on the apical side
of the segments). The genus name is derived from the
Greek adiantos, unwettable. An ancient name referring to
the water-repellent leaves.
itano de alrededor de 200 especies, la mayoría de las
cuales son neotropicales. Las especies crecen un una var-
iedad de ambientes como bosques sombreados y rocas,
pero siempre son terrestres. La autapomorfia del género
esporangios nacidos en la cara inferior del margen incur-
vado del segmento (el indusio falso). En otros Pteridace-
ae con inducíos falsos, los esporangios están nacidos en
la cara inferior de la lámina, opuesto el margen reflejo.
Estudios filogenéticos muestran Adiantum hermana a los
helechos vittarioides. Hay varios grupos bien definidos
dentro del género. Dos de ellos, importantes en los Neo-
trópicos y fáciles reconocer, son el grupo de A. tetraphyl-
lum (láminas 2-pinnadas con ápices conformes, pínnulas
demidiadas) y el grupo de A. capillus-venerus (láminas
descompuestas, segmentos cuneados, flabelados, o casi
redondos, los soros nacidos en el lado apical de los seg-
mentos). El nombre genérico se derive del griego adian-
tos, no mojable. Un nombre anciano se refiere a las hojas
que repellen agua.

LITERATURE: Huiet, L., F-W. Li, T-T Kao, J. Prado, A. R. Smith, E. Schuettpelz & K. M. Pryer. 2108. A worldwide
phylogeny of Adiantum (Pteridaceae) reveals remarkable convergent evolution on leaf blade architecture. Taxon 67: 488–502.
Lellinger, D. B. 1991. Common and confusing bipinnate-dimidiate Adiantums of tropical America. American Fern Journal 81:
99–102. Morton, C. V. 1955. Notes on Adiantum in Mexico. American Fern Journal 45: 113–117. Palacios-Rios, M. & R. Riba.
1983. Helechos de Veracruz: Adiantum (Pteridaceae). Boletín de la Sociedad Botánica de México 44: 43–62. Scamman, E. 1960.
The maidenhair ferns (Adiantum) of Costa Rica. Contributions from the Gray Herbarium of Harvard University 187: 3–22.

158
Figure 76. A, B. Adiantum feei. C, D. Adiantum tricholepis. E, F. A. patens. G, H. A. galeottianum. (From Mickel &
Smith, 2004.)

159
POLYPODIACEAE
Alansmia M. Kessler, Moguel, Sundue & Labiak
DESCRIPTION: Epiphytic; rhizomes setose; rhizome
scales usually orange to castaneous, not clathrate, bas-
ifixed (not cordate), hairy on margins and often both
surfaces, or sometimes absent (T. alfarii), the cells tur-
gid; leaves pendulous, indeterminate; blades lacking
black clavate fungi; stipes usually less than 1 cm long, the
pinnae reduced nearly to the base of the leaf; laminae pin-
natisect to 1-pinnate, with tendency for paired or clus-
tered setae; veins simple, free; hydathodes present, often
cretaceous; sori separate, round, superficial (i.e., not sunk-
en in the lamina); sporangial capsules setose; spores
green, trilete and/or monolete; x=37.
DESCRIPCIÓN: Epífitas; rizomas setosos; escamas
del rizoma usualmente anaranjadas a castañas, no clatra-
das, basifijas (no cordadas), peludos en los márgenes y
a menudo los superficies, o a veces ausentes (T. alfarii),
las células turgidas; hojas péndulas, indeterminadas;
láminas se carecen de hongos claviformes negros; pecíolos
usualmente menos de 1 cm de largo, las pinnas reducidas
casi a la base de la hoja; láminas pinnatisectas a 1-pinnadas,
con tendencia tener setas pareadas o agrupadas; venas
simples, libres; hidatodos presentes, a menudo cretásicos;
soros separados, redondos, superficiales (i.e., no hundidos
en la lámina); cápsulas esporangiales setosos; esporas
verdes, triletes y/o monoletes; x=37.

SIMILAR GENERA: Pecluma differs by monolete, yel-
lowish spores and lack of setae on the petioles and blades.
Several species of Pecluma are scaly along the rachis (in
contrast, all grammitids lack laminar scales). Ascogrammitis
is distinguished in part by the presence of small (ca. 0.5
mm long), black, clavate fungi on the lower surface of the
lamina. Leucotrichum differs by rhizome scales blackish,
clathrate, eciliate on both surfaces, leaves determinate,
usually less than 10 cm long, and by the laminar hairs
which are 3-celled, and 1-furcate, the branch cell being
acicular
GÉNEROS PARECIDOS: Pecluma difiere por esporas
monoletes amarillentosas y la falta de setas en los pecío-
los y láminas. Varias especies de Pecluma son escamosas a
lo largo del caquis (en contraste, todos las gramitidoides
carecen de escamas de la lámina). Ascogrammitis se dis-
tingue por la presencia de pequeños hongos negros (ca.
0.5 mm de largo), claviformes, sobre el envés de la lámina.
Leucotrichum difiere por escamas del rizoma negruzcas,
clatradas, eciliadas en ambas superficies, hojas determina-
das, usualmente menos que 10 cm de largo, y pelos de
la lamina 3-celulados, 1-furcados, con una célula acicular
de la rama.

COMMENTS: Alansmia contains 25 species and two va- COMENTARIOS: Alansmia consta de 25 especies y
rieties, all neotropical except for two found in Africa and dos variedades, todos neotropicales excepto dos encon-
the Mascarene Islands. All occur as pendulous epiphytes tradas encontradas en África y las Islas Mascareñas.Todos
in wet shaded forests from low to high elevations. Alans- existen como epífitos péndulos en bosques húmedos
mia is unique among grammitids ferns in having paired sombreados de elevaciones bajas a altas. Alansmia es úni-
hairs on the laminae. It is also unique in producing both co entre helechos grammitoides en tener pelos parea-
trilete and monolete spores; all other grammitids pro- dos en las láminas. También es único en producir esporas
duce only trilete spores. The indeterminate leaves, with triletes y monoletes; todos los otros grammitoides pro-
apices abruptly reduced to s small fiddlehead, are also ducen solamente esporas triletes. Las hojas indetermi-
helpful in identifying the genus. The genus name honors nadas, con ápices abruptamente reducidos, son útiles en
Alan R. Smith (1943– ), American pteridologist. identificar el género. El nombre del género honora Alan
R. Smith (1943– ),.pteridólogo Americano.
LITERATURE: Copeland, E. B. 1956. Ctenopteris in America. Philippine Journal of Science 84: 381–473, t. 1–16. Kessler,
M., A. L. Moguel Velázquez, M. Sundue & P. H. Labiak 2011. Alansmia, a new genus of grammitid ferns (Polypodiaceae)
segregated from Terpsichore. Brittonia. 63: 233–244. Moguel Velázquez, A. L. & M. Kessler. 2009. Taxonomic notes on the
fern species group around Terpsichore lanigera (Polypodiaceae), including the descriptions of three new species and one new
variety. Phytotaxa 2: 35–45. Moguel Velázquez, A. L. & M. Kessler. 2013 Grammitid ferns (Polypodiaceae), III. Alansmia. Flora
Neotropica Monograph 113: 1–168. Smith, A. R. 1993. Terpsichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3:
478–489.

160
Figure 77. A, B. Alansmia cultrata. C–H. A. spathulata. J, K. A. senilis. (From Mickel & Smith, 2004.)

161
PTERIDACEAE
Aleuritopteris Fée
DESCRIPTION: Plants small, usually on rocks; rhizomes
erect or ascending, short; scales (in ours) brown, con-
colorous; petioles and rachises black to reddish brown,
shiny, terete or (in ours) sulcate; laminae abaxially fa-
rinose or (in some Old World species) lacking farina,
farina white (ours) to yellow, adaxially glabrous (ours)
or rarely with hairs or scales; rachises grooved adaxially;
pinnae sessile or subsessile, basal pinnae usually largest;
basal basiscopic segments enlarged, longer than adjacent
acroscopic ones; veins free but often obscure; sori sub-
marginal, consisting of 1–10 sporangia at vein tips, often
confluent at maturity; false indusia present, continuous
or discontinuous, margins (in ours) erose; spores globose
to globose-tetrahedral or tetrahedral, trilete; x = 29, 30.
DESCRIPCIÓN: Plantas pequeñas, usualmente en rocas;
rizomas erectos o ascendentes; escamas (en nuestras)
pardas, concoloras; pecíolos y ráquises negros a pardo
rojizos, lustrosos, teretes o (en nuestros) sulcados; lámi-
nas abaxialmente farinosas o (en algunas especies del
Viejo Mundo) faltando farina, la farina blanqueciña (nues-
tras) a amarillas, adaxialmente glabras o raras veces con
escamas; ráquises surcados adaxialmente; pinnas sésiles o
subsésiles, pinnas basales usualmente lo más grande; seg-
mentos basales basiscópicos agrandados, más largos que
los adyacentes acroscópicos; venas libres pero a menudo
obscuras; soros submarginales, consistiéndose de 1–10
esporangios en los puntos de las venas, a menudos con-
fluentes a la madurez; indusios falsos presentes, contin-
uos o discontinuous, márgenes (en nuestros) erosos; es-
poras globosas a globoso-tetrahédricas, triletes; x=29, 30.

SIMILAR GENERA: Notholaena differs by not or hardly GÉNEROS PARECIDOS: Notholaena difiere por már-
modified margins of the fertile pinnae. Pityrogramma dif- genes de las pinnas fértiles no o casi no modificadas. Pity-
fers by sporangia spreading along the veins. rogramma difiere por esporangios difundiéndose por las
venas.
COMMENTS: Aleuritopteris contains ca. 40 species in COMENTARIOS: Aleuritopteris consta de ca. 40 espe-
the tropics and subtropics of the Old- and New-Worlds cies en los tropicos y subtropicos de los Nuevo y Vie-
It greatest diversity is in China, where 29 species occur, 14 jo Mundos. Su mayor diversidad es en China donde 29
of which are endemic. In the Neotropics it occurs from especies existen, 14 de las cuales son endémicas. En el
Mexico to Costa Rica, and in the Greater Antilles. In the Neotrópico, 1–3 especies han sido segregagadas de la
Neotropics, 1–3 especies have been segregated from the anteriormente ampliamente circunscrita A. farinosa. Ese
formerly broadly circumscribed A. farinosa. That name re- nombre se refiere a un triploide apógamo que es re-
fers to an apogamous triploid that is restricted to eastern stringido a África oriental tropicaland y Oriente Medio
tropical Africa and the adjacent Middle East. The oldest adyyacente. El nombre más viejo para plantas del Nue-
name for New World plants is A. mexicana Fée, but plants vo Mundo es A. mexicana Fée, pero plantas en México
in northern Mexico were called A. chihuahensis by Saiki septentrional fueron llamadas A. chihuahensis por Saiki
(1984b,c) and he described Andean plants as A. peruvi- (1984b,c) y describió plantas Andinas como A. peruviana.
ana. A small group of Mexican/Central American species Un pequeño grupo de especies en México y América
with yellow farina assigned to Aleuritopteris by Pichi Ser- Central con farina amarilla asignado a Aleuritopteris por
molli have been transferred to Notholaena. Our plants Pichi Sermolli ha sido transferido a Notholaena. Nues-
are sexual with 64 spores per sporangium. They grow on tras plantas son sexuales con 64 esporas por esporangio.
rocks, mostly on limestone. The genus name is derived Usualmente existen sobre rocas, mayormente en piedra
from the Greek aleuro- floury or mealy, and pteris, fern. It caliza. El nombre del género se deriva del griego aleuro-
refers to the farina on the abaxial surfaces of the leaves,. harinoso, y pteris, helecho. Se refiere a la farina en las
superfícies abaxiales de las hojas.

LITERATURE: Gangmin, Z., G. Yatskievych & E. A. Hooper. 2013. Aleuritopteris. Pp. 224–235. In: Editorial Committee. Flo-
ra of China (Lycopodiaceae through Polypodiaceae), vols. 2-3: In: C.Y. Wu, P., P. H. Raven & D.Y. Hong (eds.) Fl. China. Science Press
and Missouri Botanical Garden Press, Beijing & St. Louis. Saiki, Y. 1984a. Notes on ferns 2. Asiatic species of the Aleuriopteris
farinosa complex. Journal of Phytogeography and Taxonomy 32: 1–13. Saiki, Y. 1984b. Notes on ferns 3. African and Neotropical
species of the Aleuriopteris farinosa group. Journal of Phytogeography and Taxonomy 32: 81–90. Saiki, Y. 1984c. Notes on ferns
4. Classification of the Aleuriopteris farinosa group. Journal of Phytogeography and Taxonomy 32: 91–98.

162
Figure 78. Aleuritopteris mexicana. (From Mickel & Smith, 2004.)

163
CYATHEACEAE
Alsophila R. Br.
DESCRIPTION: Terrestrial; stems arborescent; sterile
and fertile leaves 1–4.4 m long, monomorphous; petioles
smooth or with stout black spines, scaly, the scales with
black setae (at least apically), concolorous to bicolor-
ous, usually the marginal cells different in shape and orien-
tation from the central ones; laminae 1-pinnate-pinnatifid
to 4-pinnate, the apex pinnatifid or rarely conform; veins
free; axes pubescent adaxially, the hairs thick, strigose, mul-
tiseptate; sori round, the receptacle elevated, subglobose;
indusia present or absent, scalelike or globose, minutely
scaly; spores 16 per sporangium, tetrahedral-globose,
without large equatorial pores; x=69.
DESCRIPCIÓN: Terrestres; tallos arborescentes; ho-
jas estériles y fértiles 1–4.4 m de largo, monomorfas;
pecíolos lisos o con espinas negras fuertes, escamo-
sos, las escamas con setas negras (a lo menos en
el ápice), concoloras a bicoloras, las células marginales
difieren en forma y orientación de las centrales; láminas
1-pinada-pinatífidas a 4-pinadas, el ápice gradualmente
reducido o raras veces conforme; nervaduras libres; ejes
pubescentes adaxialmente, los pelos estrigosos, gruesos,
estrigosos, multiseptados; soros redondos, el receptácu-
lo elevado, subgloboso; indusios presentes o ausentes,
escuamiformes o globosos, diminutamente escamosos;
esporas 16 por esporangio, tetrahédricas-globosas sin
poros ecuatoriales grandes; x=69.

SIMILAR GENERA: Gymnosphaera differs by the pres-
ence of small, skeletonized pinnae (aphlebia) near the
petiole base, and 64 spores per sporangium. Other Cy-
atheaceae differ by lacking a black seta at the apex of the
petiole scale. Sphaeropteris further differs by conform
petiole scales (cells with the same shape and orientation).
GÉNEROS PARECIDOS: Gymnosphaera difiere por la
presencia de pinas pequeñas en esqueletos (aflebias) cer-
ca la base del pecíolo y 64 esporas por esporangio. Otros
Cyatheaceae difieren por la ausencia de una seta negras
en el ápice de la escama del pecíolo. Sphaeropteris difiere
además por escamas peciolares conformes (células con
la misma forma y orientación).

COMMENTS: Alsophila is pantropical and contains COMENTARIOS: Alsophila es pantropical y consta de

about 200 species. It occurs in shaded forests. A name
that has often been used for many species of Alsophi-
la in the American tropics is Nephelea It here placed in
synonymy because there are no constant differences
between it and Alsophila and several hybrids are known
between the two genera. The 2-pinnate-pinnatifid spe-
cies of Alsophila have black spines on the petioles. They
produce narrow lateral branches at the base of the trunk
that grow into the soil where they eventually form erect
stems. Thus the plants form colonies. The genus name is
derived from the Greek alsos, grove + philein, to love. It
refers to the shade-loving habit of the genus.
casi 200 especies. Existe en bosques sombreados. Nephe-
lea, un nombre que ha sido usado a menudo en los trópi-
cos americanos, está ubicado aquí en sinonimia por que
no hay diferencias constantes entre él y Alsophila y varios
híbridos y especies de origen híbrido son conocidos en-
tre los dos. Las especies 2-pinnadas de Alsophila tienen
espinas negras en los pecíolos. Producen ramas laterales
angostas en la base del tronco que crecen hacía el suelo
donde eventualmente forman tallos erectos. Estas espe-
cies forman colonias. El nombre genérico se derive del
griego alsos, arboleda + philein, amar. Se refiere a al hecho
que el género le gusta la sombra.

LITERATURE: Conant, D. S. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. Journal of the Arnold
Arboretum 64: 333–382. Conant, D. S. 1990. Observations on the reproductive biology of Alsophila species and hybrids (Cy-
atheaceae). Annals of the Missouri Botanical Garden 77: 290–296. Gastony, G. J. 1973. A revision of the fern genus Nephelea.
Contributions to the Gray Herbarium of Harvard University 203: 81–148. Korall, P., D. S. Conant, J. S. Metzgar, H. Schnei-
der & K. M. Pryer. 2007. A molecular phylogeny of scaly tree ferns. American Journal of Botany 94: 873–886.

164
Figure 79. A–C. Alsophila cuspidata. F–H, J–L, A. firma. M–R. A. tryoniana (A-C, original ©R. C. Moran 2010; F–R
from Mickel & Smith, 2004.)

165
THELYPTERIDACEAE
Amauropelta Kunze
DESCRIPTION: Terrestrial; rhizomes erect to suberect,
rarely creeping or decumbent; leaves monomorphous;
petioles bases with 2 vascular bundles; laminae
1-pinnate-pinnatifid, reduced toward the base, with
the apex gradually reduced (not conform); aerophores
present or rarely absent; proximal pinnae gradually or
abruptly reduced, sometimes to vestigial pinnae; veins
meeting the margin above the sinus, never united
below the sinus, usually simple (not forked); indumentum
of needle-shaped (rarely hooked) hairs; sori round, rarely
elongate, indusiate or not; sporangia glabrous, rarely
setose; spores finely reticulate; x=29.
DESCRIPCIÓN:Terrestres; rizomas erectas a suberectas,
raras veces reptantes o decumbentes; hojas monomorfas;
las bases de los pecíolos con 2 haces vasculares;
láminas 1-pinnato-pinnatífidas, reducidas hacia la
base, con el ápice gradualmente reducido (no conforme);
aeróforos presentes o raras veces ausentes; pinnas
proximales gradualmente o abruptamente reducidas, a
veces a pinnas vestigiales; venas encontrándose con el
margen por arriba del seno, nunca unido por abajo del
seno, simples (no furcadas); pelos en forma de aguja, raras
veces ganchudo; soros redondos, raras veces elongados,
indusiados o no; esporangios glabros, raramente setosos;
esporas finamente reticuladas; x=29.

SIMILAR GENERA: The other genera of
Thelypteridaceae differ by laminae widest or nearly so
at the base. Cyclosorus, Goniopteris, and Steiropteirs differ
by veins that meet the margin at the base of the sinus,
or either unite or are connivent below it. Thelypteris (s.s.)
differs by long-creeping rhizomes and veins of the sterile
laminae forked.
GÉNEROS PARECIDOS: Los otros géneros de
Thelypteridaceae difieren por láminas más anchas o
casi así en las bases. Cyclosorus, Goniopteris, y Steiropteirs
difieren por venas que encuentran el margen en la base
del sinus, o se unen o están coniventes abajo del sinus.
Thelypteris (s.s.) difiere por rizomas largamente rastreras
y venas bifurcadas de las láminas estériles.

COMMENTS: Amauropelta contains about 220 species,
only a few of which occur outside the Neotropics
(Africa, Madagascar, and Mascarene Islands). Smith (1974)
recognized 9 sections, providing a key to the sections and
a list of their species. In some species the bases of petioles
or even entire croziers are covered with mucilage. This
occurs mostly in extremely wet habitats. The mucilage
is secreted by dense, glandular hairs (Hennipman,
1968). Aerophores usually protrude beyond the layer
of mucilage. The genus name is derived from the Greek
amauro, obscure + pelte, shield. Perhaps referring to the
idusia of the type species
COMENTARIOS: Amauropelta contiene alrededor
de 220 especies, sólo unas de las cuales existen afuera
del Neotrópico (África, Madagascar e Islas Masscarene).
Smith (1974) reconoció 9 secciones, dando una clave a
las secciones y una lista de las especies en cada sección.
En algunas especies las bases de los pecíolos o aún las
prefoliaciones enteras están cubiertos con mucílago. Esto
ocurre a menudo en ambientes extremamente húmedos.
El mucílago está secretado por pelos glandulares densos
(Hennipman, 1968). Aeróforos usualmente se extienden
a fuera la capa de mucílago. El nombre genérico se derive
del griego amauo, obscuro+ pelte, escudo. Tal vez se
refiere a los indusios de la espcie tipo.

LITERATURE: Christensen, C. 1907. Revision of the American species of Dryopteris of the group of D. opposita. Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7,4: 247–336. Christensen, C. 1913. A monograph of the genus Dryopteris,
Part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7,
10:55–282.Hennipman, E. 1968. The mucilage secreting hairs on the young fronds of some leptosporangiates ferns. Blumea
16: 97–103. Ponce, M. M. 1995. Las especies austrobrasileñas de Thelypteris subg. Amauopelta (Thelypteridaceae, Pteridofita).
Darwiniana 33(1–4): 257–283. Salino, A. & S. Semir. 2004. Thelypteris subg. Amauropelta (Kunze) A. R. Sm. (Thelypteridaceae
- Pterophyta) no Estado de São Paulo, Brasil. Lundiana 5: 83–112. Smith, A. R. 1974. A revised classification of Thelypteris
subgenus Amauropelta. American Fern Journal 64: 83–95. Smith, A. R. 1983. 14(4). Polypodiaceae—Thelypteridoideae. In: G.
Harling & B. Sparre, Flora of Ecuador, no. 18. Smith, A. R. 1992. 16. Thelypteridaceae. In: R. M. Tryon & R. G. Stolze, Pteridophyta
of Peru. Fieldiana, Botany, new series 29: iii + 1–80.

166
Figure 80. Amauropelta uncinata. Note in A the gradually reduced lamina base. Note in B dark aerophore at base of pinna. /
Fijase en A la base de la lámina gradualmente reducida. En B, fijase los aeróforos oscuros en la base de la pinna. (From Smith,
Fl. Ecuador, 1983.)

167
THELYPTERIDACEAE
Amblovenatum J. P. Roux
DESCRIPTION: Terrestrial; rhizomes short- to long-
creeping, scales setose on margins and surfaces; leaves
few, mostly 1–2 m long; petioles with 2 vascular bundles;
laminae 1-pinnate-pinnatifid, with proximal pinnae hardly
reduced or with only one greatly reduced pair, the apices
reduced, pinnatifid (not conform); rachises sparsely
pubescent and glandular abaxially, hairs and glands up to
0.1 mm long; pinnae sessile to short-stalked to 2 mm;
aerophores absent; buds lacking; veins 8–14 pairs per
segment, the lowermost often meeting below the sinus
with an excurrent veinlet to the sinus, or veins connivent
near the sinus; indumentum of acicular hairs ca. 0.1 mm
long mixed with a few larger ones ca. 0.5 mm long, also
with many yellow sphaerical glands especially along the
veins and costules abaxially, costal scales absent; sori
round, supramedial, often confined to the apices of the
segments; indusia with marginal glands and a few acicular
hairs; sporangial stalks sometimes glandular; sporangial
capusules glabrous; spores blackish, tuberulate; x=36.
DESCRIPCIÓN: Terrestres; rizomas cortametne a
largamente rastreros, escamas setosas por los márgenes
y superfícies; hojas pocas, principalmente 1–2 m de largo;
pecíolos con 2 haces vasculares; láminas 1-pinnada-
pinnatífida, con pinnas proximales casi no reducidas o con
solo un par muy reducido, los ápices reducidos, pinnatífidos
(no conformes); ráquises esparcidamente pubescentes y
glandulares abaxialmente, los pelos y glándulas hasta 0.1
mm de largo; pinnas séssiles a cortamente pecioluladas
hasta 2 mm; aeróforos ausentes; brotes carentes; venas
8–14 pares por segmento, las más basales a menudo
uniéndose abajo del sinus con una venilla excurrent al
sinus, o venas coniventes cerca del sinus; indumento
de pelos aciculares ca. 0.1 mm de largo mezclados con
unos más grandes ca. 0.5 mm de largo, también con
muchas glándulas esféricas amarillas especialmente
por las venas y cóstulas abaxialmente, escamas costales
ausentes; soros redondeados, supramediales, a menudo
restringidos a los ápices de los segmentos; indusios con
glándulas marginales y unos pelitos acículares; pecíolos de
los esporángios a veces glandulares; cápsulas esporangiales
glabras; esporas negruscas; x=36.

SIMILAR GENERA: All other genera ofThelypteridaceae GÉNEROS PARECIDOS: Todos los otros géneros
differ by lacking spherical yellow glands along the abaxial de Thelypteridaceae difieren por las glandulas esféricas
surfaces of the costules and veins. amarillas por las superficies abaxiales de las cóstulas y
venas.

COMMENTS: Amblovenatum contains ca. 12 species, COMENTARIOS: Amblovenatum consta de ca. 12

and is native to the East Africa, tropical Asia, Queensland
especies y es indígena a África oriental, Asia tropical.,
, and in the Pacific to Tahiti. One species, A. opulentumQueenslandia y en el Pacífico hasta Tahiti. Una especie,
(Kaufuss) J. P. Roux, is naturalized in the New World. Since
A. opulentum (Kaufuss) J. P. Roux, es naturalizada en el
it was first collected in the 1950s, it has spread rapidly,
Nuevo Mundo. Desde fue colectada por primera vez en
now occuring from Florida to southern Brazil, in both los 1950, ha difundo rápidamente y ahora existe desde
open-disturbed and shaded-forest habitats, from 0–700 Florida hasta al sur de Brasil, en ambientes arbolados y
m. In the Neotropics, the spherical yellow glands along abiertos perturbados, a 0–700 m. En el Neotrópico, las
the abaxial surfaces of the veins are absolutely diagnostic
glandulas esféricas amarillas por las superficies abaxiales
of the species.The genus name is derived from the Greek de las venas son absolutamente diagnósticas de la especie.
amblo-, abortion + venatum, vein. It is unclear what thisEl nobre genérico se deriva del griego amblo-, aborción
refers to. + venatum, venas. No se pone en claro a lo que se
refiere.
LITERATURE: Fosberg, F. R. & M.-H. Sachet. 1972. Three Indo-Pacific Thelypteris species reinterpreted and a new African
species described. Smithsonian Contributions to Botany 8: 1–10. He L-J, Zhang X-C. 2012. Exploring generic delimitation within
the fern family Thelypteridaceae. Molecular Phylogenetics and Evolution 65: 757–764. Holttum, R. E. 1977. Studies in the family
Thelypteridaceae. XII. The genus Amphineuron Holttum. Blumea 23, 205–218. Holttum, R. E. 1981. Thelypteridaceae. In: C. G.
G. J. van Steenis & R. E. Holttum (gen. eds.). Flora Malesiana, series II - Pteridophyta, Ferns and Fern Allies, 1(5): 1–599. Roux, J.
P. 2009. Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23. South
African National Biodiversity Institute, Pretoria

168
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Figure 81. Amblovenatum opulentum

169
PTERIDACEAE
Ananthocorus Maxon & Underwood
DESCRIPTION: Epiphytic or epipetric; rhizomes
short- creeping solenostelic, short-creeping, dorsiven-
tral; rhizome scales clathrate; leaves 15–45 × 0.9–1.5
cm, simple, entire, glabrous, coriaceous, distichous,
clumped; petioles absent or very short and flattened;
costae present; veins areolate, 2 or 3 rows of areoles
between the costa and margin; so r i submarginal
on each side of the costa, nearly continuous; indusia
absent; paraphyses abundant, slender, without a swollen
apical cell; spores monolete, nongreen; gametophytes
lacking gemmae; x=120.
DESCRIPCIÓN: Epífitas o epipétricas; rizomas corta-
mente rastreros, solenostélicos, dorsiventrales; escam-
as del rizoma clatradas; hojas 15–45 × 0.9–1.5 cm,
simples, enteras, glabras, coriáceas, dísticas, amacolla-
das; pecíolos ausentes o muy cortos y achatados; costas
presentes; nervaduras areoladas, en 2 o 3 hileras de
areolas entre la costa y el margen; so r o s submargi-
nales en cada lado de la costa, casi continuous; indusios
ausentes; parafisos abundantes, delgados, sin una célula
apical hinchada; esporas monoletes, no verdes; gameto-
fitos sin yemas; x=120.

SIMILAR GENERA: Radiovittaria and Vittaria differ by GÉNEROS PARECIDOS: Radiovittaria y Vittaria difie-
having only one row of areoles between the costa and ren por tener una sola hilera de areolas entre la costa
margin (vs. 2 or 3) and by the presence of gemmae y el margen (vs. 2 o 3) y por la presencia de gemas en
on the gametophytes. el gametofito.

COMMENTS: Consists of only one species, Anantha-
corus angustifolius (Sw.) Underw. & Maxon, widespread
from Mexico to Bolivia. It occurs as an epiphyte in wet
or semi-deciduous forests from 50–1500 m. Anantha-
corus is the only vittarioid fern whose gametophytes
lack gemmae. Its gametophytes bear antheridia scat-
tered over the surface instead of being predominantly
on small plants or on germinating gemmae as in other
vittarioid ferns (Farrar, 1974). The rows of aeroles be-
tween the midrib and margin are often hard to see
because of the thick lamina. The genus name is derived
from the Greek an, without + anthos, flower + Acorus,
Sweetflag. Resembling Acorus but without flowers.
COMENTARIOS: Consta de una sola especie, Anan-
thacorus angustifolius (Sw.) Underw. & Maxon, difundi-
do desde México hasta Bolivia. Ocurre como epífito
en bosques húmedos o semi-deciduous de 50–1500
m. Ananthacorus es el único helecho vittarioide cuy-
os gametofitos faltan de yemas. Sus gametofitos llevan
anterídios salpicados por la superficie en lugar de estar
principalmente en plantas pequeñas o en yemas germi-
nantes como en otros helechos vittarioides (Farrar,
1974). Las hileras de areolas entre la costa y el margen
son a menudo difícil ver a causa de la lámina gruesa. El
nombre genérico se derive del griego an, sin + anthos,
flor + Acorus. Se asemeja Acorus per sin flores.

LITERATURE: Benedict, R. C. 1911. The genera of the fern tribe Vittarieae: Their external morphology, venation, and re-
lationships. Bulletin of the Torrey Botanical Club 38: 153–190. Crane, E. H. 1997. A revised circumscription of the genera of
the fern family Vittariaceae. Systematic Botany 22: 509–517. Crane, E. H., D. R. Farrar & J. F. Wendel. 1995. Convergent
simplification leads to a polyphyletic Vittaria. American Fern Journal 85: 283–305. Farrar, D. R. 1974. Gemmiferous fern
gametophytes—Vittariaceae. American Journal of Botany 61: 146–155. Maxon, W. R. 1908. Studies of tropical American
ferns—No. 1. A new genus allied to Vittaria. Contributions from the United States National Herbarium 10: 486–487.

170
Figure 82. A–D. Ananthacorus angustifolius. (From Mickel & Smith, 2004.)

171
BLECHNACEAE
Anchistea C. Presl
DESCRIPTION: Terrestrial; rhizomes long-creep-
ing, scaly; sterile and fertile lvs monomorphic, decidu-
ous, 1-pinnate-pinnatifid; petioles with several vascu-
lar bundles, the two adaxial ones larager than the others;
pinnae of the sterile leaves 15–20 pairs; veins forming
one row of areoles between the costa and margin; sori
elongate, forming a chain parallel to the costa, su-
perficial, not sunken in the lamina tissue; x=35.
SIMILAR GENERA: Osmunda cinnamomea dif-
fers by holodimorphic sterile and fertile leaves that are
clumped. Woodwardia differs by clumped leaves. Lorinse-
ria differs by sterile and fertile leaves dimorphic, com-
pletely areolate venation, and sori sunken in the lamina
tissue.
COMMENTS: Consists of only one species, Anchistea
virginica (L.) C. Presl, which occurs primarily on the costal
plain in the eastern North America. It grows in swamps,
marshes, bogs, roadside ditches, generally in moist plac-
es on acid soils. The common name "chain fern" refers
to the short, elongate sori in a series that resemble a
chain. The genus name is derived from the Greek an-
chistos, a relation, alluding to this fern’s relationship to
Woodwardia, another genus of Blechnaceae..
LITERATURE: Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodi-
ales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227.
172
DESCRIPTION: Terrestres; rizomas largamente ra-
streros, escamosos; hojas estériles y fértiles monomór-
fas, decíduas, 1-pinnada-pinnatifida; pecíolos con vari-
os haces vasculares, los dos adaxiales más grandes de los
otros; pinnas de las hojas estériles 15–20 pares; venas
formando una hilera de areolas entre la costa y el mar-
gen; soros alargados, formando una cadena para-
lela a la costa, superficiales, no hundidos en el tejido
láminar; x=35.
SIMILAR GENERA: Osmunda cinnamomea difiere
por sus hojas estériles y fértiles holodimorfas que son
amacollados. Woodwardia difiere por hojas agrupados.
Lorinseria difiere por hojas estériles y fértiles dimorfas,
nervación areolada y soros hundidos en el tejido láminar.
COMMENTS: Consta de sola una especie, Anchistea
virginica (L.) C. Presl, la cual existe principalmente en la
planicie costal de América del Norte oriental. Habita
pantanos, humedales, turberas, zanjas por los lados de
los caminos, y generalmente lugares húmedos en suelos
ácidos. El nombre común "helecho de cadena" refiere a
los soros cortos, alrgados en una série que asemeja una
cadena. El nombre genérico se deriva del griego, an-
chistos, una relación, aludiendo a la relación de este
helecho a Woodwardia, otro género de Blechnaceae..
Figure 83. Anchistea virginica. Note chain-like sori in upper right. (From Small, 1935.)

173
ANEMIACEAE
Anemia Sw. subgen. Anemia
DESCRIPTION: Terrestrials or saxicolous; rhizomes
compact, short-creeping or ascending, dictyostel-
ic , densely pubescent, the hairs brown or orange;
l e a v e s p o l y s t i c h o u s ; sterile and fertile leaves
hemidimorphic, rarely holodimorphic; laminae pinnatifid
to 4-pinnate; basal pinnae typically modified into 2
erect, long-stalked fertile spikes; veins free or areo-
late; sporangia cylindrical or pear- shaped with an apical
annulus; spores tetrahedral- globose, striate with paral-
lel smooth or spiny ridges; x = 38.
DESCRIPCIÓN: Terrestres o saxícolas; rizomas
compactos, cortamente rastreros o ascendentes, dic-
tiostélicos, densamente pubescentes, los pelos pardos
o naranjados; hojas p o l i s t élicos, estériles y fértiles
hemidimórfas, raras veces holodimórfas; láminas pin-
natífidas a 4-pinnadas; pinnas basales típicamente
modificadas en 2 espigas fértiles lárgamente pe-
dunculadas, erectas, (la hoja raras veces es comple-
tamente fértile); nervaduras libres o areoladas; espo-
rangios cilíndricos o piriformes con un anillo apical;
esporas tetraédricas-globosas, estriadas con filas para-
lelas lisas o espinosas; x = 38.

SIMILAR GENERA: Botrychium differs by erect rhi-
zomes, fleshy leaves, a single fertile spike, and glo-
bose sporangia. Anemia subgen. Anemiorrhiza differs by
longer-creeping, solenostelic rhizomes, blackish rhizome
hairs, and distichous leaves.
GÉNEROS PARECIDOS: Botrychium difiere por rizo-
mas erectos, hojas carnosas, única espiga fértil e espor-
angios globosos. Anemia subgén. Anemiorrhiza difiere por
rizomas más largamente rastreros, solenostélicos, pelos
de los rizomas negruzcos y hojas dísticas.

COMMENTS: Anemia contains about about 115 living
species, of which 85 are neotropical, 16 African-Mad-
agascan, and one is Indian It is most diverse in Brazil
(80 species) and Mexico (20 species). Nearly all species
occur in dry, open or semi-open habitats. They often
hybridize when growing together. Ploidy levels in the
genus range from 2x to 14x. Mohria, an African genus
with scales instead of hairs, is sister to Anemia (Skog et
al., 2002). Anemia subgen. Anemiorrhiza (see next page)
is sometimes recognized as Ornithopteris Bernh. It is sis-
ter to the rest of the genus. The genus name is derived
from the Greek aneimon, naked, and alludes to the
exposed sporangia.
COMENTARIOS: Anemia consta de alrededor de
115 especies vivientes, de las cuales 85 son neotrópico
a l e s , 1 6 Áfricanas-Madagascarenas y una de India. Es
más diverso en Brasil (80 especies) y México (20 espe-
cies). Casi todas las especies existen en habitats secos,
abiertos, o semi-abiertos. A menudo se hibridizan
cuando en poblaciones mixtas. Niveles de poliplodea
en el género varian desde 2x hasta 14x. Mohria, un
género Africano con escamas en lugar de pelos, está
hermano a Anemia subgén. Anemia (Skog et al., 2002).
Anemia subgén. Anemiorrhiza (vea la próxima página) se
reconoce a veces como Ornithopteris. Es hermano al res-
to del género. El nombre genérico se deriva del griego
aneimon, desnudo, y refiere a los esporangios expues-
tos.

LITERATURE: de la Sota, E. R. & J. T. Mickel. 1968. Sinopsis de las especies Argentinas del género Anemia Sw. (Schi-
zaeaceae). Revista del Museo de la Plata, n.s., Sección Botánica 11: 133–152. Labiak, P., J. T. Mickel & J. Garrison Hanks.
2015. Molecular phylogeny and character evolution of Anemiaceae (Schizaeales). Taxon 64: 1141–1158. Mickel, J. T.
1962 A monographic study of the fern genus Anemia, subgenus Coptophyllum. Iowa State College Journal of Science 36:
349–482. Mickel, J. T. 1981. Revision of Anemia subgenus Anemiorrhiza (Schizaeaceae). Brittonia 33: 413–429. Mickel,
J. T. 1982. The genus Anemia (Schizaeaceae) in Mexico. Brittonia 34: 388–413. Mickel, J. T. 2016. Anemia (Anemiaceae).
Flora Neotropica Monograph 118: 1–182.

174
Figure 84. A, B. Anemia munchii. C, D. A. phyllitidis. E, F. A. hirsuta × A. phyllitidis. (From Mickel & Smith, 2004.)

175
ANEMIACEAE
Anemia subgen. Anemiorrhiza Prantl
DESCRIPTION: Calciphiles, usually saxicolous; rhi-
zomes creeping, solenostelic, densely pubescent, the
hairs blackish; leaves distichous; sterile and fertile
leaves holodimorphic or hemidimorphic; laminae
pinnatifid to 4-pinnate; basal pinnae typically modified
into 2 erect, long-stalked fertile spikes; veins free or are-
olate; sporangia cylindrical or pear-shaped with an apical
annulus; spores tetrahedral-globose, striate with parallel
smooth or spiny ridges; x = 38.
DESCRIPCIÓN: Calcifílas, usualmente saxícolas; ri-
zomas reptantes, solenostélicos, densamente pubes-
centes, los pelos negruzcos; hojas dísticas; hojas
esteriles y fértiles holodimórfas o hemidimorfas;
láminas pinnatífidas a 4-pinnadas; pinnas basales típi-
camente modificadas en 2 espigas fértiles largamente
pedunculadas, erectas; nervaduras libres o areoladas; es-
porangios cilíndricos o piriformes con un anillo apical;
esporas tetraédricas-globosas, estriadas con filas parale-
las lisas o espinosas; x = 38.

SIMILAR GENERA: Anemia subgen. Anemia differs by
compact, dictyostelic rhizomes, orange to castaneous
rhizome hairs, and polystichous leaves. Botrychium differs
by erect, glabrous rhizomes, fleshy leaves, a single fertile
spike, and globose sporangia.
GÉNERO PARECIDO: Anemia subgen. Anemia difiere
por rizomas compactos, dictiostélicos, pelos de los rizo-
mas anaranjados a castaños y hojas polisticas. Botrychi-
um difiere por rizomas erectos glabros, hojas carnosas,
única espiga fértil e esporangios globosos.

COMMENTS: Anemia subgén. Anemiorrhiza contains COMENTARIOS: Anemia subgén. Anemiorrhiza con-
13 species that occur primarily on limestone in the sta de 13 especies que existen principalmente sobre
Caribbean Basin. Most of the species have small rang- piedra caliza en la Cuenca del Caribeo. La mayoría de
es, except for A. adiantifolia (L.) Sw., which is wide- las especies tienen rangos pequeños, con excepción
spread. Nearly all occur in dry, open or semi-open de A. adiantifolia (L.) Sw., la cual es muy difundida. Casi
habitats. They often hybridize when growing together. todas existen en hábitats secos, abiertos o semi-abi-
A key to the species can be found in Mickel (1981, ertos. A menudo se hibridizan cuando en poblaciones
2016). The subgenus, which is sister to subgen. Ane- mixtas. Una clave a las especies se puede encontrar
mia, is sometimes recognized at generic rank as Or- en Mickel (1981, 2016). El subgénero, lo que es her-
nithopteris Bernh. The African Mohria is nested with mano al subgén. Anemia, se reconoce a veces al nivel
Anemia. Besides the characters given above, Anemia genérico como Ornithopteris Bernh. El género Africa-
subgén. Anemiorrhiza tends to have more coriaceous no Mohria está anidado dentro de Anemia. Además
laminae and the terminal cell of the laminar hairs are de las características dadas arriba, Anemia subgén.
acute, not swollen. About half of the species have ho- Anemiorrhiza suele tener láminas más coriaceas y la
lodimorphic sterile and fertile leaves. Most species of célula terminal de los pelos foliares son agudos, no
subgén. Anemiorrhiza are diploid, whereas those of hinchados. Casi una mitad de las especies tienen ho-
subgen. Anemia, are tetraploid, hexaploid, or higher. jas estériles y fértiles holodimorfas. La mayoría de
The genus name is derived from the Greek ornis, bird, las especies de subgén. Anemiorrhiza son diploides,
and pteris, fern. The allusion is unknown. mientras que las de subgén Anemia son tetraploides,
hexaploides o más alto. El nombre genérico se deriva
del griego ornis, pájaro, y pteris, helecho. La alusión se
desconoce.
LITERATURE: de la Sota, E. R. & J. T. Mickel. 1968. Sinopsis de las especies Argentinas del género Anemia Sw. (Schi-
zaeaceae). Revista del Museo de la Plata, n.s., Sección Botánica 11: 133–152. Labiak, P., J. T. Mickel & J. Garrison Hanks.
2015. Molecular phylogeny and character evolution of Anemiaceae (Schizaeales). Taxon 64: 1141–1158. Mickel, J. T.
1962 A monographic study of the fern genus Anemia, subgenus Coptophyllum. Iowa State College Journal of Science 36:
349–482. Mickel, J. T. 1981. Revision of Anemia subgenus Anemiorrhiza (Schizaeaceae). Brittonia 33: 413–429. Mickel, J.
T. 1982. The genus Anemia (Schizaeaceae) in Mexico. Brittonia 34: 388–413. Mickel, J. T. 2016. Anemia (Anemiaceae).
Flor a Neotropica Monogr aphs 00: 000–000.

176
Figure 85. Anemia subgen. Anemiorrrhiza. A, B. Anemia munchii. C, D. A. phyllitidis. E, F. A. hirsuta × A. phyllitidis.
(From Mickel & Smith, 2004.)

177
PTERIDACEAE
Anogramma Link
DESCRIPTION: Terrestrial; rhizomes inconspicuous,
erect, hairy (not scaly); leaves 5–25 cm long, fascicu-
late, annual; petioles ca. 1 mm wide, as long or slightly
longer than the lamina, green to stramineous; laminae
elongate-triangular, 2- to 4-pinnate, not farinose, anad-
romic, thin; terminal segments 0.5–2 mm wide; rachises
grooved adaxially; veins free; sori spreading along the
veins, without parafises; indusia absent; spores tetrahe-
dral-globose, trilete, dark brown, with an equatorial ridge
and two parallel flanges; x=29.
SIMILAR GENERA: Gastoniella differs by scaly rhizomes,
narrower ultimate segments, and whitish or tan spores, .
Pityrogramma differs by scaly rhizomes and laminae usual-
ly farinose beneath. Cystopteris has two vascular bundles
in the petiole indusiate sori.
DESCRIPCIÓN: Terrestres; rizomas inconspicuos, erec-
tos, pelosos (no escamosos); hojas 5–25 cm de lar-
go, fasciculadas, anuales; pecíolos c. 1 mm de ancho, tan
largo como las láminas o escasamente más largos que
ellas, verdes o pajizos; láminas alargado-triangulares, 2- a
4-pinnadas, no farinosas, anádromas, delgadas, glabras;
segmentos terminales 0.5–2 mm de ancho; ráquises sur-
cados adaxialmente; nervaduras libres; soros difundidos
a lo largo de las nervaduras, sin parafisos; indusios
ausentes; esporas tetraédrico-globosas, triletes, pardas
oscuras, con una fila ecuatorial y dos costillas paralelas;
x=29.
GÉNEROS PARECIDOS: Gastoniella difiere por rizo-
mas escamosos, segmentos últimos más estrechos y es-
poras blanquecinas o leonadas. Pityrogramma difiere por
rizomas escamosos y las láminas usualmente farinosas en
el envés. Cystopteris y tiene dos haces vasculares en el
pecíolo y soros indusiados.

COMMENTS: Anogramma consists of two pantropical COMMENTARIOS: Anogramma consta de dos es-
species: A. leptophylla (L.) Link and A. lorentzii (Hieron.) pecies pantropicales: A. leptophylla (L.) Link y A. lorentzii
Diels. Both grow on moist banks in semi-open areas (Hieron.) Diels. Ambas ocurren en bancos húmedos en
from 1000–2800 m. Two species formerly in Anogram- lugares semi-abiertas de 1000–2800 m. Dos especies
ma are now placed in Gastoniella (i.e., G. chaerophylloides previamente en Anogramma ahora están ubicadas en
(Desv.) Li Bing Zhang & Liang Zhang and G. novogaliciana Gastoniella (i.e., G. chaerophylloides (Desv.) Li Bing Zhang
(Mickel) Li Bing Zhang & Liang Zhang). Anogramma is sis- & Liang Zhang y G. novogaliciana (Mickel) Li Bing Zhang
& Liang Zhang.). Anogramma es hermana a Cosentinia,
ter to Cosentinia, a monotypic genus of the Mediterra-
un género monotípico de la región Mediterráneo a las
nean-Himalayan region.The gametophytes of Anogramma Amalhayas. Los gametofitos tienen varias características
have several unusal characteristics. The young ones are inusuales. Los jóvenes son espatulados, no cordados y la
spathulate, not cordate, and meristematic activity is from actividad meristamática es de un meristemo lateral, no
a lateral meristem, not an apical one. These characters apical. Estos rasgos se compartan con Cosentinia. Si no
are shared with Cosentinia. If fertilization or sporophyte toma lugar la fecundación ni producción del esporofito,
production does not occur, the gametophytes can survive los gametofitos pueden sobrevivir por una temporada
an unfavorable growing season by means of a tubercle desfavorable por medio de un tubérculo desarrolla-
formed by elaborating the thickened archegonial region do por elaborar la región aquegonial o “cojín”. Cuando
or cushion. When favorable growth conditions return, condiciones favorables regresan, el tubérculo regenera
lóbulos nuevos gametofíticos. Los gametofitos no pueden
the tubercle regenerates new lobes. The gametophytes
desarrollarse más si un nuevo esporofito se produce. Por
cannot further develop if a new sporophyte is produced. eso, los gametofitos perenan solo en un sentido limitado.
Thus, the gametophytes perennate only in a limited sense. El nombre genérico se deriva del griego ano, hacia arriba
The genus name is derived from the Greek ano, upward + gramme, línea. Refiere a los soros alargados en los seg-
+ gramme, line. It refers to the elongate sori on the ter- mentos terminales.
minal segments.
LITERATURE: Nakazato, T. & G. J. Gastony. 2003. Molecular phylogenetics of Anogramma species and related genera
(Pteridacae: Taenitidoideae). Systematic Botany 28: 490–502. Pangua, E. & B. Vega. 1996. Comparative study of gametophyte
development in Cosentinia and Anogramma (Hemionitidaceae) and Cheilanthes (Sinopteridaceae). Pages 497–508. In: J. M.
Camus, M. Gibby & R. J. Johns (eds.). Pteridology in perspecitive. Royal Botanic Garden, Kew. Tryon, R. M. 1962. Taxonomic fern
notes. II. Pityrogramma (including Trismeria) and Anogramma. Contributions from the Gray Herbarium 189: 52–76. Zhang, L.-
B. & L. Zhang. 2017. Phylogeny of the fern subfamily Pteridoideae (Pteridaceae; Pteridophyta), with the description of a new
genus: Gastoniella. Molecular Phylogenetics and Evolution 109: 59–72.

178
Figure 86. A–C. Anogramma leptophylla. D, E. A. lorentzii. (A–C from Mickel & Smith, 2004; D, E, © R. C. Moran, 2010.)

179
DRYOPTERIDACEAE
Arachniodes Adans.
DESCRIPTION: Terrestrial; rhizomes creeping; leaves
monomorphic; petioles with more than three vascu-
lar bundles; laminae widely deltate or pentagonal, 2- to
5-pinnate; medial pinnae prolonged acroscopically, the
basal basiscopic side cuneate; basal pinnules anadrom-
ically arranged, stalked; rachises and costae with
grooves glabrous, without proliferous buds; veins free;
sori round; indusium orbicular-reniform; spores mono-
lete; x=41.
SIMILAR GENERA: Dryopteris differs by rhizomes
erect or ascending, pinnules catadromic, and (in the
Neotropics) leaves often capitate-glandular. Polystichum
differs by the basal acroscopic pinnules slightly elongate,
and generally the margins of the pinnules are spinulose.
Lastreopsis and Rumohra differ by a thickened marginal
ridge of the costa or costules surcurrent along the green
laminar margin of the segment of the next highest order
(not adaxially along the midrib of the costule). Saccoloma
differs by marginal sori and a single, continuous (not divid-
ed) omega-shaped vascular bundle in the petiole. Pradoia
differs by adaxial grooves of rachises and costae pubes-
cent, as least at the junctures.
COMMENTS: Arachniodes is pantropical with about 50
species, most of which are Asian. In the Neotropics it con-
tains about 4 species, and these occur mostly above 2000
m. The most widespresd species is A. denticulata (Sw.)
Ching), which highly variable. Christensen (1920) listed
numerous forms and varieties of this species, and many of
these probably represent distinct taxa, but biosystemat-
ic work is needed. The genus name is derived from the
Greek arachnion, spider’s web + odes, having the form or
nature of. What it refers to is unknown.
LITERATURE. Christensen, C. 1920. The a monograph to the genus Dryopteris. Part II. The tropical American bipinnate-de-
compound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 8, 6: 101–120. Proctor, G. R. 1985. Ferns of
Jamaica. British Museum (Natural History), London.
180
DESCRIPCIÓN: Terrestres; rizomas rastreros esca-
mosos; hojas monomorfas; pecíolo con tres o más hac-
es vasculares; lámina ampliamente deltada o pentagonal,
2-5-pinnada; pinnas medias prolongadas acroscópica-
mente, el lado basal basiscópico cuneado; pínnulas dis-
puestas anadrómicamente, pediculadas; raquis y cos-
tas con surcos glabros, sin yemas prolíferas; nervaduras
libres; soros redondos; indusio orbicular-reniforme; es-
poras monoletes; x=41.
GÉNEROS PARECIDOS: Dryopteris difiere por rizo-
mas erectos o ascendentes, pínnulas catadrómicas y (en
el Neotrópico) hojas a menudo glanduloso-capitadas.
Polystichum difiere por las pínnulas basales basiscópicos
alargados, y generalmente los márgenes de las pínnulas
son espinulosos. Lastreopsis y Rumohra difieren por una
fila marginal engrosada de la costa o cóstula surcurrente
por el margen del tejido laminar verde del segmento del
próximo orden más alto (no adaxialmente por la cóstula
del segmento). Saccoloma difiere por soros marginales y
un haz vascular continuo (no dividido) en forma de ome-
ga en el pecíolo. Pradoia difiere por los surcos adaxiales
de los raquises y costas pubescentes, al menos en las
uniones.
COMENTARIOS: Arachniodes es pantropical con casi
50 especies, la mayoría de las que son Asiáticas. En el
Neotrópico contiene alrededor de 4 especies, e estas
existen principalmente arriba de 2000 m. La especie
más difundida es A. denticulata (Sw.) Ching), la cual es
muy variable. Christensen (1920) listó numerosas formas
y variedades de él, y muchos de estos probablemente
representan taxones distintos, pero se necesita trabajo
biosistemático. El nombre genérico se deriva del griego
arachnion, tela de araña + odes, con la forma o la carac-
terística de algo. Se desconoce a lo que se refiere.
Figure 87. A, B. Arachniodes denticulata. (From Mickel & Smith, 2004.)

181
PTERIDACEAE
Argyrochosma (J. Sm.) Windham
DESCRIPTION: Terrestrial or rupestral; rhizomes
creeping or erect, scaly, the scales glandular, commonly
not sclerotic, concolorous or slightly bicolorous; leaves
2-4 pinnate, with separate segments all the way to
the apex (not becoming confluent in a pinnatifid
apical portion), monomorphic; petiole with a single vas-
cular bundle; pinnules short-stalked or narrowed toward
the base, typically white or yellow farinose beneath;
veins free, the tips not or only weakly thickened; sori sub-
marginal along the distal 1/4-2/3 of each vein, partially
hidden by the weakly to strongly revolute margins of the
pinnules; spores trilete; x=27.
SIMILAR GENERA: Pellaea differs by nonglandular rhi-
zome scales and nonfarinose laminae. Cheilanthes also
differs by nonfarinose lamina. Notholaena differs by bi-
colorous (centrally sclerotic) rhizome scales.
COMMENTS: Argyrochosma is exclusively American
genus of about 20 species. It center of diversity is the
Chihuahuan desert region of northern Mexico and ad-
jacent Texas, where it has 8 species. Most of the species
are apogamous and grow on or among rocks. The base
chromosome number of x=27 is unique among cheilan-
thoid ferns. Argyrochosma is sister to Pellaea and Astrolepis.
The gametophytes of Argyrochosma lack farina, where as
those of Notholeana have it (Windham, 1987). The gener-
ic name comes from the Greek argyros, silver, + chosma,
poweder. It refers to the white (silvery) powder on the
abaxial surface of the laminae.
LITERATURE: Sigel, E. M., M. D. Windham, L. Huiet, G. Tayskievych, K. M. Pryer. 2011. Species relationships and farina
evolution in the cheilanthoid fern genus Argyrochosma (Pteridaceae). Systematic Botany 36: 554–564. Tryon, R. M. 1956. A
revision of the American species of Nothlaena. Contributions from the Gray Herbarium of Harvard.
182
DESCRIPCIÓN: Terrestres o rupícolas; rizomas cor-
tamente rastreros o erectos, escamosos, las escamas
glandulares, comúnmente no escleróticas, concoloras o
ligeramente bicoloras; hojas 2-4-pinnadas, 2-4-pinnadas,
con segmentos separatas hacia el ápice (no llegan-
do a ser confluentes en una parte apical pinnatífi-
da), monomorfas; pecíolo con un haz vascular; pínnulas
cortamente pediculadas o basalmente constrictas, típica-
mente blancas o amarillo-cerosas en el envés; ner-
vaduras libres, las puntas débilmente o no engrosadas;
soros submarginales a lo largo del 1/4-2/3 apical de cada
nervadura, ocultos parcialmente por los márgenes débil
a fuertemente reflejos de las pínnulas; esporas triletes;
x=27.
GÉNEROS PARECIDOS: Pellaea difiere por escamas
del rizoma no glandulares. Cheilanthes difiere también por
láminas nofarinosas. Notholaena difiere por escamas del
rizoma bicoloras (escleróticas en la parte central).
COMENTARIOS: Argyrochosma es género exclusiv-
amente americano de casi 20 especies. Su centro de
diversidad es la región del Desierto Chihuahuense del
norte de México y el área adyacente de Texas, donde
se encuentra 8 especies. La mayoría de las especies son
apógamas y crecen sobre o entre rocas. El número cro-
mosómico básico de x=27 es único entre los helechos
cheilantoides. Argyrochosma es hermana a Pellaea y Astro-
lepis. Los gametófitos de Argyrochosma carecen de una fa-
rina, mientras que los de Notholaena la tienen (Windham,
1987). El nombre genérico se deriva del griego argyros,
plata, + chosma, polvo. Se refiere al polvo blanco (platea-
do) en el envés de las láminas.
Figure 88. A-C. Argyrochosma incana. D-G. A. delicatula. E-G. A. delicatula. H, J. A. incana. K, L. A. fendleri. (From Mickel
& Smith, 2004.)

183
POLYPODIACEAE
Ascogrammitis M. Sundue
DESCRIPTION: Plants epiphytic, rarely saxicolous;
rhizomes short-creeping, dorsiventral, unbranched; rhi-
zome scales usually clathrate, the cells flat (not turgid
or inflated), margins usually reddish setose, sometimes
entire or provided only with a single apical seta; leaves
pubescent and setose, the fertile portions more densely
setose than sterile portions, usually bearing a black,
clavate fungus ca. 1 mm long; laminae 1-pinnatisect
to 1-pinnate; veins greenish (not dark); hydathodes
present, usually cretaceous or not; sori slightly eliptic,
sometimes provided with receptacular setae; paraphyses
absent; sporangial capsules glabrous; spores trilete, green;
x=37?
DESCRIPCIÓN: Plantas epífitas, raras veces saxícolas;
rizomas cortamente rastreros, dorsiventrales, no ram-
ificados; escamas del rizoma usualmente clatradas,
las células planas (no túrgidas o hinchadas), márgenes
usualmente setosos-rojizos, a veces enteros o con
una seta apical; hojas pubescentes y setosas, las partes
fértiles más setosas que las estériles, usualmente por-
tando un hongo negro clavado pequeño (ca. 1 mm
long); láminas 1-pinnatisectas a 1-pinnadas; venas
verdosas (no oscuros); hidatodos presentes, usual-
mente cretásicos; soros levemente elípticos, a veces con
setas receptaculares; cápusulas esporangiales glabras es-
poras triletes, verdes; x=37?

SIMILAR GENERA: Lellingeria differs by rhizome scales
with hyaline marginal setae, fronds that lack reddish se-
tae, costae not visible on the abaxial side of the blade,
and hairs with an acicular rather than setiform branch.
Mycopteris differs by concolorous, orange to castaneous
rhizome scales with turgid (swollen) cells, leaves not more
densely setose in the fertile parts, and blackish veins.
Melpomene differs by reddish-brown scales with entire
margins and frequently with multiple apical papillate cells.
Dried specimens have a sweet, spicy odor. Pecluma differs
by monolete yellowish spores and lack of foliar setae.
GÉNEROS PARECIDOS: Lellingeria difiere por las es-
camas del rizoma con setas marginales hialinas, hojas que
faltan setas rojizas, costas no visibles en el envés de la lámi-
na y pelos con una rama acicular en lugar de setiforme.
Mycopteris difiere por escamas de rizoma concoloras,
anaranjadas a castaneas con células túrgidas (hichadas),
hojas no más densamente setosas en las partes fértiles
y venas negruzcas. Melpómene difiere por escamas ro-
jizas-pardas enteras y frequentamente con varias células
apicales papilosas. Ejemplares secas tienen un olor dulce
especiosa. Pecluma difiere por esporas monoletes amaril-
lentosas y la falta de setas foliares.

COMMENTS: Ascogrammitis contains about 17 species.
It is entirely neotropical, occurring in the Greater Antil-
les, Guadeloupe, and Mexico to Bolivia. The species occur
in cloud forests usually from 1800–3200 m. The fungus
Acrospermum maxonii is a nearly constant associate of the
genus. Its mycelia are whitish and the spore cases (as-
comes) are black, clavate, and ca. 1 mm long. The fungus
does not appear to harm the fern in any way. The genus
name refers to the ascomes (Greek, asco-, bag or blad-
der) of the fungal associate.
COMENTARIOS: Ascogrammitis consta de ca. 17 epe-
cies. Es completamente neotropical y existe en las Antillas
Mayores, Guadeloupe, México hasta Bolivia. Las especies
inhabitan bosques nublados usualmente desde 1800–
3200 m. El hongo Acrospermum maxonii es casi constan-
temente asociado con el género. Sus micelias son blan-
quecinas y las esporangios (ascomas) son negras, clavadas
y ca. 1 mm de largo. El hongo no parece dañar el helecho
en ninguna manera. El nombre genérico refiere a las as-
comas (Griego, asco-, bolsa o vejiga) del hongo asociado.

LITERATURE: Smith, A. R. 1993. Terpsichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3: 478–489. Sundue,
M. 2008. A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae). Brittonia 62: 357–399. Sundue, M.,
I. Olivares & M. Kessler. 2018. Ascogrammitis lehnertii (Polypodiaceae): A new and dominant understory-species from a Diverse
Community of Grammitid Ferns in the Andes of Ecuador. Systematic Botany, 43:709–716 [contains key to all species].

184
Figure 89. Ascogrammitis david-smithii A. Habit. B. Rhizome scale. C. Detail of abaxial lamina surface, note ascomes of
Acrospermum. Ascogrammitis colombiensis D. Habit. E. Rhizome scale. F. Detail of abaxial lamina surface including ascomes of
Acrospermum. Ascogrammitis cuencanana G. Habit. H. Detail of adaxial lamina surface. J. Detail of abaxial lamina surface, note
ascome of Acrospermum. (From Sundue, 2008.)

185
ASPLENIACEAE
Asplenium L.
DESCRIPTION: Epiphytic, terrestrial or saxicolous;
stems erect to creeping, scaly, the scales clathrate; sterile
and fertile leaves monomorphous, rarely subdimorphous;
petioles with 2 vascular bundles basally which unite to-
ward the distally to form a single X-shaped vascular bun-
dle; laminae simple to 4-pinnate, anadromic; axes of the
lamina grooved, the grooves decurrent, glabrous; veins
free; sori elongate to linear, borne singly (not double)
per vein indusiate; annulus of the sporangia consisting
of 20-28 cells; spores bilateral, brown; x=36.
DESCRIPCIÓN: Epífitas, terrestres o saxícolas; tallos
erectos a rastreros, escamosos, las escamas clatradas;
hojas estériles y fértiles monomorfas, raras veces subdi-
morfas; pecíolos con 2 haces vasculares en la base los
cuales se unen distalmente formando un solo haz vas-
cular en forma de “X”; láminas simples a 4-pinnadas,
anádromas; ejes de la lámina surcados, los surcos decur-
rentes, glabros; venas libres; soros alargados a lineares,
nacen uno por vena (no doblados) indusios; anillo del
esporangio consiste de 20-28 células; esporas bilate-
rales, pardas; x=36.

SIMILAR GENERA: Asplenium and Diplazium can be
difficult to distinguish; nevertheless, three characteristics
consistently separate them.The first is the number of cells
in the annulus: Asplenium has 20-28, Diplazium has 15-20.
The second is the number of rows of cells in the stalk
of the sporangium: Asplenium has only 1, Diplazium has
2 or 3. The third is the form of the vascular bundle(s) in
the distal part of the petiole: in Asplenium it is X-shaped,
in Diplazium the two strands are separate and elliptic.
Unfortunately, the characteristics cited above are often
difficult to observe. There are two other characteristics
that, although not always as constant as the above, are
often helpful. The rhizome scales of Asplenium are usu-
ally clathrate; those of Diplazium are never clathrate. The
sori of Asplenium are almost always borne singly along
a single vein; those of Diplazium are often paired back
to back along a single vein. See Diplazium for additional
comments.
GÉNEROS PARECIDOS: Asplenium y Diplazium
pueden ser difíciles de distinguir; sin embargo, tres car-
acterísticas los separan consistentemente. La primera
es el número de células en el anillo: Aplenium tiene 20-
28, Diplazium tiene 15-20. La segunda es el número de
células en el pecíolo del esporangio: Asplenium tiene 1,
Diplazium tiene 2 o 3. La tercera es la forma del haz
vascular en la parte distal del pecíolo: en Asplenium forma
una ”X”, en Diplazium forma dos haces distintos, elípti-
cos. Lamentablemente, las características arribas citadas
son a menudo difíciles de observar. Hay otras dos car-
acterísticas que aunque no son tan constantes (como las
anteriores), a menudo son útiles. Las escamas del rizoma
de Asplenium usualmente son clatradas; las de Diplazium
nunca son clatradas. Los soros de Asplenium casi siempre
nacen únicamente por una sóla vena; las de Diplazium
a menudo nacen pareados, en par en par, por una sóla
vena. Véase Diplazium para comentarios adicionales.

COMMENTS: Asplenium is cosmopolitan and contains COMENTARIOS: Asplenium es cosmopolito y consta

approximately 700 species, making it is one of the larg-
est fern genera in the world. Section Hymenasplenium is
sometimes segregated as distinct because it is sister to
the rest of Asplenium. The genus name comes from the
Greek splen, spleen. Dioscorides thought a fern with this
name was useful in treating diseases of the spleen.
de aproximadamente 700 especies, haciéndolo uno de
los más grandes géneros de helechos en el mundo. A vec-
es la sección Hymenasplenium se reconoce como distinto
por que es hermana el resto de Asplenium. El nombre
genérico se deriva del griego splen, esplín. Dioscorides
pensó un helechos con este nombre era útil para enfer-
medades del esplín

LITERATURE: Morton, C. V. & D. B. Lellinger. 1966. The Polypodiaceae subfamily Asplenioideae in Venezuela. Memoirs of
the New York Botanical Garden 15: 1–49. Schneider, H., S. J. Russell, C. J. Cox, F. Bakker, S. Henderson, F. Rumsey, J.
Barrett, M. Gibby & J. C. Vogel. 2004a. Chloroplast phylogeny of asplenioid ferns based on rbcL and trnL-F spacer sequences
(Polypodiidae, Aspleniaceae) and its implications for biogeography. Systematic Botany 29: 260–274. Stolze, R. G. 1986. Polypo-
diaceae – Asplenioideae. In: G. Harling and B. Sparre (eds.). Flora of Ecuador, no. 23.

186
Figure 90. Asplenium. Note linear sori and clathrate rhizome scales. Fijase en los soros lineares y escamas del rizoma clatradas.
(Original, ©Robbin Moran, 2009.)

187
ASPLENIACEAE
Asplenium (Loxoscaphoid clade)
DESCRIPTION: Epiphytic; rhizomes short-creeping
to suberect, the scales clathrate; leaves 5–15 cm
long, clumped, monomorphic; petioles scaly, with two
vascular bundles at the base; blades finely dissected,
2-4-pinnate; segments slender, linear, fleshy; veins free;
sori near the margin, protected by a pouch-like indusia
that opens toward the margin; spores bilateral, brown;
x=35.
DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros
a suberectos, las escamas clatradas; hojas 5–15 cm de
largo, amacolladas, monomorfas; pecíolos escamosos,
con 2 haces vasculares en la base; láminas finamente
disectadas, 2-4-pinnadas; segmentos delgados, lineares,
carnosos; margen de la lámina engrosado y decurrente
por el raquis; venas libres; soros cerca del margen,
protegidos por indusios como bolsa que abra hacia
el margin; esporas bilaterales, pardas; x=35.

SIMILAR GENERA: Asplenium (s.s.) differs only by the GÉNEROS PARECIDOS: Asplenium (s.s.) difiere
shape of the sorus: elongate to linear and opening toward solamente por la forma del soro: alargado a lineal y
the midrib or vein. abriéndose hacia la costa o vena.

COMMENTS: The "loxoscaphoid" ferns are nested
in Asplenium. They are primarily African and Asian, with
the exact number of species being unknown. Because
the group includes A. theciferum (Kunth) Mett., the type
of Loxoscaphe T. Moore, the term "loxoscaphoid" is
used for the clade (the name Asplenium subg. Daraea
T. Moore, also applies to the clade). The loxoscaphoid
clade is characterized by short cup-shaped sori with
the appearance of a Davallia sorus. In fact, the type, A.
theciferum, was first described in Davallia (as Davallia
thecifera Kunth), Typically, each ultimate segment bears
only a single vein and single sorus. The lateral sides of the
indusium sometimes fused to the lamina and occasionally
form a pocket-like structure. The base chromosome
number of x=35 also differs from Asplenium, which
has x=36. Asplenium theciferum is the only Neotropical
species in the clade. It is an epiphyte in wet forests
from 1200–1900 m. It also occurs in Africa, where it is
represented by var. concinnum (Schrad.) Schelpe. The
only difference between the two varieties is the short or
absent apical lobe that extends beyond the sorus in the
American plants; this lobe is absent or very short in the
African plants. The name is derived from the Greek loxos,
oblique + skaphe, a small boat or bowl. The bowl-shaped
sori are oblique to the costae.
COMENTARIOS: Los helechos "loxoscaphoides" son
anidados en Asplenium. Son principalmente Áfricanos y
Asiáticos, con el númber exacto de especies no conocido.
A causa del grupo incluye A. theciferum (Kunth) Mett., , el
tipo de Loxoscaphe, el termino "loxoscaphoide" se aplica
al clado (el nombre Asplenium subg. Daraea T. Moore,
tambien se aplica al clado). El clado loxoscaphoide se
caracteriza por soros cortos en forma de copa con la
apariencia de un soro de Davallia. De hecho, el tipo,
A. theciferum, fue descrito por primera vez en Davallia
(como D. thecifera Kunth). Típicamente, cada segmento
ultimo lleva una sola vena y un solo soro. Los lados
laterals del indusio a veces se fusionan a la lámina y
ocasionalmente forman una estructura como una bosilla.
El número cromosómico basal de x=35 también difiere
de Asplenium, lo cual tiene x=36. Asplenium theciferum es
la única especie Neotrópical. Es una epífita en bosques
húmdos de 1200–1900 m. La especie está representada
en África por la var. concinnum (Schrad.) Schelpe. La única
diferencia entre las dos variedades es el lóbulo apical
corto o ausente que extiende más alla el soro en las
plantas Americanas; este lóbulo es ausente o muy corto
en las plantas Africanas. El nombre se deriva del griego
loxos, oblicuo + skaphe, una barca pequeña o escudilla.
Los soros en forma de escudilla son oblicuos a la costa.

LITERATURE: Bellefroid, E., S. Khadijah Rambe, O. Leroux1 & R. L. L. Viane. 2010. The base number of ‘loxoscaphoid’
Asplenium species and its implication for cytoevolution in Aspleniaceae. Annals of Botany 106: 157–171. Morton, C. V. & D. B.
Lellinger. 1966. The Polypodiaceae subfamily Asplenioideae in Venezuela. Memoirs of the New York Botanical Garden 15: 1–49.

188
Figure 91. A. Asplenium (Loxoscaphe) theciferum. (From Mickel & Beitel, 2004.)

189
ASPLENIACEAE
Hymenasplenium Hayata
DESCRIPTION: Epiphytic, terrestrial or saxicolous; rhi- DESCRIPCIÓN: Epífitas, terrestres o saxícolas; rizomas
zomes creeping, dorsiventral, with 2 rows of leaves erectos a rastreros, dorsiventrales, con dos hileras de
on the dorsolateral surfaces, scaly toward the apices, hojas en la superficie dorsolateral, escamosos hacia
the scales clathrate; sterile and fertile leaves monomor- los ápices, las escamas clatradas; hojas estériles y fértiles
phous; petioles slightly swollen at their junction with monomorfas; pecíolos levemente hinchados en su
the rhizome, with 2 vascular bundles basally, these uniting unión con el rizoma, con 2 haces vasculares en la base,
distally to form a single X-shaped strand; laminae 1-pin- estos uniéndose distalmente en forma de “X”; láminas
nate to 1-pinnate-pinnatifid, anadromic; axes of the lam- simples a 1-pinnadas a 1-pinnado-pinnatífidas, anádromas;
ina grooved, the grooves decurrent, glabrous; veins free; ejes de la lámina surcados, los surcos decurrentes, glabros;
sori elongate to linear, borne singly (rarely double) per venas libres; soros alargados a lineares, nacen uno por
vein, indusiate; annulus of the sporangia consisting of vena (raras veces doblados), indusiados; anillo del espo-
20-28 cells; spores bilateral, brown; x=38, 39. rangio consiste de 20-28 células; esporas bilaterales,
pardas; x=38, 39.
SIMILAR GENERA: Diplazium and sect. Asplenium differ GÉNEROS PARECIDOS: Diplazium y la secc. Aspleni-
by polystichous leaves, slender (non-swollen) petiole bas- um difieren por hojas polísticas, las bases de los pecíolos
es, and chromosome number. delgados (no hinchados) y número cromosómico.

COMMENTS: Hymenasplenium is pantropical and con- COMENTARIOS: Hymenasplenium es pantropical y

tains ca. 50 species, of which 11 are neotropical. In the
Americas it extends from Mexico and the Antilles to
southern Brazil. It occurs in wet forests, generally below
2000 m, and is remarkable for the variety of growth habits
among its species (terrestrial, saxicoulous, hemiepiphytic)
and diversity of spore types. The leaves resemble those
typical of sect. Asplenium, but the creeping rhizomes with
distichous leaves and swollen petiole bases are strikingly
different. The chromosome number of the genus is x=38
or 39, in contrast to Asplenium which is uniformly x=36.
The New World species fall into two clades. The first, the
H. laetum clade (6 spp.), is characterized by nonconform
and pinnatifid leaf apex, cristate spores, and brown or
blackish rhizomes in living plants The second, the H. ripar-
ium clade (5 spp.), is characterized by greenish rhizomes
when living, swollen petiole bases, conform, subconform
or hastate leaf apices, and spiny or papillose spores. The
genus name comes from the Greek hymen, membrane +
Asplenium. The type species has membranous leaves.
consta de ca. 50 especies, de las cuales 11 son neotrop-
icales. En las Américas se extiende desde México y las
Antillas hasta Brasil del sur. Existe en bosques húmedos,
generalmente abajo de 2000 m, y es destacado por la
variedad de hábitos de crecimiento entre sus especies
(terrestre, saxícola, hemiepífita) y diversidad de tipos de
las esporas. Las hojas se asemejan a ellas que son típicas
de secc. Asplenium, pero obviamente diferente son los
rizomas reptantes con hojas dísticas y bases hinchados de
los pecíolos. El número cromosómico el género es x=38
o 39, mientras que es uniformemente x=36 en Aspleni-
um. Las especies del Nuevo Mundo cayen en dos clados.
El primero, el clado de H. laetum (6 spp.), se caracteriza
por rizomas en plantas vivas pardos o negruzcos, ápices
foliares no conformes y pinatifidos y esporas cristadas. El
segundo, el clado de H. riparium (5 spp.), se characteriza
por rizomas en plantas vivas verduzcos, bases hinchados
de los pecíolos, ápices foliares conformes, subconformes
o hastatos y esporas espinosas o papilosas. El nombre del
género se deriva del griego hymen, membrana + Aspleni-
um. La especie tipo tiene hojas membranáceas.

LITERATURE: Moran, R. C. & M. Sundue. 2005. Asplenium (sect. Hymenasplenium) basiscopicum, a new species from Bolivia.
Brittonia 56: 124–127. Murakami, N. 1995. Systematics and evolutionary biology of Hymenasplenium. Journal of Plant Research
108: 257–268. Murakami, N. & R. C. Moran. 1993. Monograph of the neotropical species of Asplenium sect. Hymenasplenium
(Aspleniaceae). Annals of the Missouri Botanical Garden 80: 1–38. Regalado, L. & C. Prada. The genus Hymenasplenium (As-
pleniaceae) in Cuba, including new combinations for the neotropical species. American Fern Journal 101: 265–281. Smith, A. R.
1976. Diplazium delitescens and the neotropical species of Asplenium sect. Hymenasplenium. American Fern Journal 66: 116–120.
Xu, K-W. et al. 2018. A global plastid phylogeny uncovers extensive cryptic speciation in the fern genus Hymenasplenium (As-
pleniaceae). Molecular Phylogenetics and Evolution 127: 203–216.

190
Figure 92. Hymenasplenium. A. H. repandulum. B. H. delitescens. C. H. obtusifolium. D. Vasculature of dorsiventral rhizome; note
distichous leaves with swollen bases. E. H. ortegae. F. H. laetum. (From Murakami & Moran, 1993.)

191
PTERIDACEAE
Astrolepis D. M. Benham & Windham
DESCRIPTION: Rhizomes compact, horizontal to erect,
occasionally branched; rhizome scales stramineous to cas-
taneous, concolorous to weakly bicolorous, linear, ciliate
to entire; leaves monomorphic, clumped, 7-80 cm long;
petioles with 2 vascular bundles, 1/6–1/3 the leaf
length, terete; blades linear to linear-oblong, pinnate
to pinnate-pinnatifid, coriaceous, abaxially clothed with
imbricate, ciliate scales with underlying scurf of substel-
late scales, adaxially with substellate to long-ciliate
scales, often glabrescent, dull; pinnae 15–42 pairs,
short-stalked to subsessile, articulate, ovate to oblong or
narrowly deltate, cordate to truncate or rounded at base,
coriaceous; pinna margins undifferentiated, not recurved;
veins obscure, free; sporangia along terminal 1/3 of veins;
spores dark brown, tetrahedral-globose, rugose; x=29.
DESCRIPCIÓN: Rizomas compactos, horizontales a
erectos, ocasionalmente ramificados; escamas del rhi-
zoma estramineas a castañas, concoloras a debilmente
bicoloras, lineares, ciliadas a enteras; hojas monomorfas,
amacolladas, 7–80 cm de largo; pecíolos con 2 haces
vasculares, 1/6–1/3 el largo de la hoja, teretes; láminas
lineares a lineares-oblongas, pinnadas a pinnada-pi-
natífidas, coriáceas, con escamas imbricadas ciliadas en
el envés arriba de una caspilla de escamas subesteladas,
en el haz con escamas subesteladas a largo-ciliadas,
a mendo glabrescentes, opaco; pinnas 15–42 pares,
corte-pedicelado a subsessiles, articuladas, ovadas a ob-
longas o estrechamente deltadas, cordatas a truncatas o
redondeadas; margenes de las pinnas no diferenciadas,
no recurvados; venas obscuras, libres; esporangios en la
tercera parte terminal de las venas; esporas pardas oscu-
ras, globoso-tetrahédricas, rugosas; x=29.

SIMILAR GENERA: Notholeana differs by a powder GÉNEROS PARECIDOS: Notholeana difiere por un
(usually whitish) on the lower surface of the pinnae. polvo (usualmente blanco) en el envés de las pinnas.

COMMENTS: All are epipetric or terrestrial in rela-
tively dry regions. The genus occurs from California and
Oklahoma south to Argentina, and is disjunct in Alabama,
Georgia, and Hispaniola. All species occur in Mexico. The
genus consists of five diploid species which form auto-
polyploids (6 taxa) and allopolyploids (13). The diploids
are sexual, with 64 spores per sporangium, the spores
less than 50 mμ long. All polyploids are apomictic, with
32 spores per sporangium. Triploids have spores 50–70
mμ long, tetraploids more than 71 mμ long. Astrolepis to
be monophyletic with a chromosome base number of
29, fronds pinnate to pinnate-pinnatifid with many pinna
pairs, two vascular bundles in the petiole, imbricate scaly
abaxially, and substellate to ciliate scales on adaxial blade
surface. The genus name is derived from the Greek astro,
star + lepis, scale. Star-shaped scales occur on the upper
surface of the pinnules.
COMENTARIOS: Todas son epipétricas o terrestres en
regiones secas. El género existe en California y Oklahoma
al sur hasta Argentina, y es disjunto en Alabama, Geor-
gia y Española. Todas las especies existen en México. El
género consta de cinco especies diploides que forman
autopolyploideas (6 taxones) y allopolyploideas (13). Los
diploides son sexuales con 64 esporas por esporangio,
las esporas menos de 50 mμ de largo. Todos los polip-
loideas son apomícticas, con 32 esporas por esporangio.
Los triploides tienen esporas 50–70 mμ de largo, tetra-
ploides más de 71 mμ de largo. Astrolepis es un grupo
monofilético con un número cromosómico base de 29,
hojas pinadas a pinada-pinatífidas con muchos pares de
pinas, dos haces vasculares en el pecíolo, escamas imbri-
cadas en el envés, y escamas subesteladas, ciliadas en el
haz. El nombre genérico se deriva del griego astro, estrella
+ lepis, escama. Escamas en form de estrellas existen el
haz de las pínnulas.

LITERATURE: Beck, J. B., M. D. Windham & K. M. Pryer. 2011. Do asexual polyploidy lineages lead short evolutionary
lives? A case study from the fern genus Astrolepis. Evolution 65-11: 3217–3229. Benham, D. M. 1992. Additional combinations in
Astrolepis. American Fern Journal 82: 59–62. Benham, D. M. & M. D. Windham. 1992. Generic affinities of the star-scaled cloak
ferns. American Fern Journal 82: 47–58. Hevly, R. H. 1965. Studies of the sinuous cloak fern (Notholaena sinuata) complex.
Journal of the Arizona Academy of Science 3: 205–208.

192
Figure 93. Astrolepis. In F and K, which show the adaxial surface of the pinnae, note the characteristic stellate scales. En F y
K, que muestran la superficie adaxial de las pinnas, fijase las escamas esteladas características. A–G. Astrolepis sinuata. H–L. A.
crassifolia. M–Q. A. laevis. (From Mickel & Smith, 2004.)

193
ATHYRIACEAE
Athyrium Roth.
DESCRIPTION: Terrestrial; rhizomes erect to creep-
ing, scaly, the scales not clathrate; sterile and fertile leaves
monomorphous; petioles with 2 vascular bundles ba-
sally, these uniting distally to form a “U”, bases swollen
with a uniseriate row of tuberculate aerophores; lam-
inae 1-pinnate to 4-pinnate; veins free; rachis and costa
grooved, the grooves decurrent; sori shortly linear or
J-shaped or horseshoe-shaped, with the distal por-
tion hooked around the tip of the vein; indusia mem-
branous, the margins ciliate or lacerate; sporangial stalks
with 3 rows of cells; spores bilateral, brown; x=40.
SIMILAR GENERA: Diplazium differs by having straight
(not J-shaped) sori. Asplenium differs by having straight
(not J-shaped) sori, and the 2 vascular bundles in the pet-
iole uniting distally to form an “X.” Athyrium is always ter-
restrial, whereas Asplenium can be terrestrial, epiphytic or
saxicolous. Ephemeropteris differs by creeping rhizomes
and distichous leaves.
COMMENTS: Athyrium contains about 100 species and
is cosmopolitan, but most species occur in eastern Asia.
It has few species in the Neotropics. Athyrium filix-femina
(L.) Roth, the type species of the genus, is represented in
the Neotropics by a complex of species, probably none
of which are conspecific with the European type. Names
that apply to this complex in Central America are A. arcua-
tum Liebm. and A. bourgaei Fourn., and in South America
the plants are often called A. dombeyi Desv. These species
occur above 1500 m. They both have creeping rhizomes,
in contrast to the previous ones that have erect rhizomes.
The genus name is derived from the Greek athyros, door-
less. It apparently refers to the sporangia only tardily
pushing back the outer edge of the indusium.
LITERATURE: Stolze, R. G. 1994. Athyrium. In: Harling, G. & L. Andersson, editors. Flora of Ecuador, 14 (5B). Polypodiace-
ae-Dryopteridoideae-Physematieae 49: 1–108.
194
DESCRIPCIÓN: Terrestres; tallos erectos o rastreros,
escamosos, las escamas no clatradas; hojas estériles y
fértiles monomorfas; pecíolos con dos haces vascu-
lares en la base, estos uniéndose distalmente en forma
de “U”, las bases hinchados con una hilera uniserada
de aróforos tuberculados; láminas 1-pinnada a 3-pinna-
do-pinnatífidas; nervaduras libres; raquis y costas surcados,
los surcos decurentes; soros cortamente lineares o en
forma de “J” a hipocrepiformes con la porción dis-
tal uncinada alrededor de la punta de la nervadura;
indusio membranáceo, los márgenes glandulosos, ciliados
o lacerados; pedículos de los esporangio con 3 hileras de
células; esporas reniformes, pardas; x=40.
GÉNEROS PARECIDOS: Diplazium difiere por nunca
tener soros rectos (no en forma de J), Asplenium difiere
por tener soros rectos (no en forma de J), y los dos haces
vasculares en el pecíolo unen distalmente para fomar un
“X”. Athyrium es siempre terrestre, mientras que Aspleni-
um puede ser terrestre, epífita, o saxícolo. Ephemeropteris
difiere por rhizomas reptantes con hojas dísticas.
COMENTARIOS: Athyrium consta de casi 100 especies
y es cosmopolito, pero la mayoría de las especies existen
en Asia oriental. Tiene pocas especies en el Neotrópico.
Athyrium filix-femina (L.) Roth, la especie tipa del género,
está representada en el Neotrópico por un complejo de
especies, probablemente ninguna es conespecífica con el
tipo de Europa. Nombres que aplican a este complejo
en America Central son A. arcuatum Liebm. y A. bour-
gaei Fourn., y en America del Sur las plantas se llaman A.
dombeyi Desv. En el sentido amplio, esto es la única espe-
cies que existe arriba de 1500 m. El nombre genérico se
deriva del griego athyros, sin una puerta. Aparentamente
refiere a los esporángios que empujan atrás tardiamente
el márgen exterior del indusio.
Figure 94. A. Athyrium bourgeaui. B. Pinnule with J-shaped sori. C. Athyrium arcuatum. (From Mickel & Smith, 2004.)

195
BLECHNACEAE
Austroblechnum Gasper & V. A. O. Dittrich
DESCRIPTION: Terrestrial or epipetric; rhizomes erect,
ascending, or creeping, stoloniferous or not, scaly, the
scales concolorous or rarely bicolorous with pale mar-
gins, ovate to lanceolate, entire; sterile and fertile leaves
dimorphic; petioles usually longer in fertile leaves; laminae
reduced proximally (rarely truncate) sometimes with
vestigial pinnae, narrowly elliptic or ovate, pinnatisect
to pinnate, rarely entire, apices pinnatifid or pinnatisect;
rachises glabrous or sparsely scaly; buds rarely present;
aerophores absent; pinnae partially or totally adnate to
the rachis, entire or crenate to serrate, plane to slightly
revolute; veins free, hydathodous; sori linear, indusia entire
to erose or fimbriate; x=33.
DESCRIPCIÓN: Terrestres o epífitos; rizomas erectos,
ascendentes o rastreros, estoloníferos o no; escamosos,
las escamas concoloras o raras veces bicoloras con már-
genes palidos, ovadas; hojas estériles y fértiles dimorfas;
pecíolos usualmente más largos en hojas fertiles; lámi-
nas reducidas proximalmente (raras veces truncadas),
a veces con pinnas vestigiales, angostamente elípticas o
ovadas, pinnatisectas a pinnadas, raras veces enteras, ápic-
es pinnatífidos o pinnatisectos; ráquises y costas glabros o
esparcidamente escamosos; yemas raras veces presentes;
aeroforos ausentes; pinnas parcialmente o totalmente ad-
nadas al ráquis, enteras o crenadas a serradas, planos o
levamente revolutas; venas libres, con hídadotos; soros
lineares; indusios enteros o erosos o fimbriados; x=33.

SIMILAR GENERA: Blechnum differs by sterile and fer-
tile leaves monomorphous. Parablechnum differs by having
proximal and medial pinnae sessile (not adnate) and the
presence of aerophores. Lomaridium differs by climbing
on trunks and bicolorous rhizome scales; Lomariocycas
differs by arborescent habit and petiole scales bicolorous
with a dark central stripe.
GÉNEROS PARECIDOS: Blechnum difiere por hojas
estériles y fértiles monomorfas. Parablechnum difiere por
tener las pinnas proximales y mediales sésiles (no adna-
tas) y la presencia de aeroforos. Lomaridium difiere por
subir truncos y escamas bicoloras del rizoma rectos; Lom-
ariocycas difiere por hábito arborescente y escameas del
pecíolo bicoloras con una raya oscura central.

COMMENTS: Austroblechnum contains about 40 in the
tropics and, in the southern hemisphere, subtropics. Most
species occur from middle to high elevations, usually in
forests and along streams. The genus name is derived
from the Latin australis, southern, and the Blechnum, an
ancient Greek name for a fern. It refers to the distribution
of the genus, which is principally in the Southern Hemi-
sphere..
COMENTARIOS: Austroblechnum consta de casi 40 es-
pecies en los tropicos y, en el Hemisferio Sur, los subtrópi-
cos. La mayoría de las especies existen desde elevaciones
medias a altas, usualmente en bosques y por riochuelos.
El nombre genérico se deriva del Latín australis, sur, y
Blechnum, un nombre anciano Griego de un helecho. Se
refiere a la distribucion del género, que es principalmente
en el Hemisferio Sur.

LITERATURE: Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodi-
ales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227. Rojas-Alvarado, A. F. 2008.
Notes in the Blechnum lherminieri complex (Blechnaceae) from the Neotropics. Métodos en Ecología y Sistemática 3: 8–29.
Rolleri, C. H. & C. Prada. 2006. Revisión de los grupos de especies del género Blechnum (Blechnaceae-Pteridophyta): el grupo
B. penna-marina. Acta Botánica Malacitana 31: 7–50.

196
Figure 95. Austroblechnum. A, B . A. wardiae. C. A. stoloniferum. D. A. lehmannii (From Mickel & Smith, 2004.)

197
LYCOPODIACEAE
Austrolycopodium Holub
DESCRIPTION: Terrestrial; main horizontal stems
creeping, subterranean or superficial, giving rise to two
dorsolateral ranks of determinate erect shoots ultimate
branchlets terete (not flattened), isophyllous, with leaves
polystichous, lacking hyaline hair-like apicies; strobili
erect, pedunculate or sessile; sporophylls peltate,
without a decurrent medial wing, lacking mucilage
cavities; sporangia isovalvate, reniform, the epidermal
cells with sinuate anticlinal walls; spores smooth on the
proximal face, reticulate on the distal face; gametophytes
subterranean, broadly obconic (carrot-shaped); x=31.
DESCRIPCIÓN:Terrestres; tallos principales horizantales
reptantes, subterraneos o superficiales, dando lugar a dos
hileras dorsolaterales de vastagos erectas determinadas;
ramitas últimas teretes (no achatados), isofílicas, con
hojas polísticas, carente de ápices filiformes, hialinos;
estróbilos erectos, pedunculados o sésiles; esporofilos
peltatos, sin una ala decurrente medial, carente
de cavidades que contiene mucílago; esporangios
isovalvados, reniformes, brevamente estípitatas, con
células epidérmicas sinuosas; esporas lisas en la cara
proximal, reticuladas en la cara distal; gametofitos
subterráneos, anchamente obconicos; x=31.

SIMILAR GENERA: All other Lycopodiaceae with
distinct strobili differ by sporophylls with medial wings
decurrent on the axis of the strobilus. Dendrolycopodium
and Spinulum further differ by their occurrence only in
the Northern Hemisphere. Lycopodium differs by leaves
with colorless, hair-like tips, sporophylls with decurrent
medial wings, and spores reticulate on both faces, and
x=34.
GÉNEROS PARECIDOS: Todos los otros
Lycopodiaceae con estróbilos distinctos difieren por
esporofilos con alas mediales decurrentes en el eje del
estróbilo. Dendrolycopodium y Spinulum difieren además
por su ocurréncia solamente en el Hemisfério Norte.
Lycopodium difiere además por las hojas con ápices
piliformes y hialinos, y esporas reticuladas en ambas
superficies y x=34.

COMMENTS: Austrolycopodium consists of 5 species COMENTARIOS: Austrolycopodium consta de 5

worldwide and occurs in Australia,Tasmania, New Zealand,
Juan Fernandez, Costa Rica, Hispaniola, Andes from
Venezuela to Tierra del Fuego, Malvinas, southern Brazil,
South Georgia, Kerguelen, and Mt. Aberdare in Uganda.
Four species occur in South America, usually above
2000 m in open or semi-open habitats or sometimes
lower elevations in high southern latitudes. The most
widespread species is A. magellanicum (P. Beauv.) Holub.,
which is common in dry páramos. In southeastern Brazil,
Chile, Argentina and Bolivia, A. erectum (Philippi) Holub is
common found at high elevations. In Chile and Argentina,
L. assurgens (Fee) Holub is found. Worldwide the species
of Austrolycopodium are poorly defined. The genus needs
monographic revision. The genus name is derived from
the Latin austro-, southern + Lycopodium. It refers to the
predominantly southern distribution of genus.
especies y existe Australia, Tasmania, Nueva Zealanda,
Juan Fernández, Costa Rica, Hispaniola, los Andes desde
Venezuela a Tierra del Fuego, las Malvinas, sur de Brazil,
Georgia Austral, Kerguelen y Mt. Aberdare en Uganda.
Cuatro especies existen en América del Sur, usualmente
arriba de 2000 m en ambientes abiertos o semiabiertos
o a veces elevaciones más bajo en latitudes altas al sur. La
especie más difundida es A. magellanicum (P. Beauv.) Holub.,
la cual es común en los páramos secos. En el sureste
de Brasil, Chile, Argentina y Bolivia, A. erectum (Philippi)
Holub. se encuentra comunmente en elevaciones altas. En
Chile y Argentina se encuentra L. assurgens (Fee) Holub.
Mundial, las species de Austrolycopodium son pobremente
definidas. El género necesita revisión monográfica. El
nombre genérico se deriva del Latín austro-, sur +
Lycopodium. Se refiere a la distribución sur predominante
del género.

LITERATURE: Holub, J. 1991. Some taxonomic changes within Lycopodiales. Folia Geobotanica et Phytotaxonomica 26:
81–80. Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Øllgaard, B. 1988.
Lycopodiaceae. In: G. Harling & L. Andersson, editors. Flora of Ecuador, no. 33. 1992. Øllgaard, B. 1992. Neotropical Lycopodia-
ceae—an overview. Annals of the Missouri Botanical Garden 79: 687–717. Øllgaard, B. & P. G. Windisch. 2014. Lycopodiaceae
in Brazil. Conspectus of the family I. The genera Lycopodium, Austrolycopodium, Diphasium, and Diphasiastrum. Rodriguésia
62(2): 2014.. Øllgaard, B. & P. G. Windisch. 2016. Lycopodiaceae in Brazil. Conspectus of the family II. The genera Ly-
copodiella, Palhinhaea, and Pseudolycopodiella. Rodriguésia 67: 691–719.

198
Figure 96. Austrolycopodium erectum. A, C. Habit. B. Branchlet apex, showing acute (not piliform) leaf apices. D. Strobilus
with sporophylls. E. Peltate sporophylls; note lack of decurrent medial wing at base. (From Øllgaard & Windisch, 2014.)

199
SALVINIACEAE
Azolla Lam.
DESCRIPTION: Floating aquatics; plants typically 0.5–
2 cm long, more or less triangular; roots present, slender,
unbranched; stems hidden by the imbricate leaves; leaves
0.4–2 mm, distichous, appressed, sessile, with an upper
and lower lobe, the lower lobe translucent and floating,
the upper lobe green or reddish, arched-erect, papillose;
sori generally paired, one megasporangiate (smaller) and
one microsporangiate (larger), borne on the underside of
the first leaf of a lateral branch, covered by a membrana-
ceous indusium; x=22.
SIMILAR GENERA: None.
DESCRIPCIÓN: Acuáticas flotantes; plantas típi-
camente 0.5–2 cm de largo, más o menos triangulares;
raíces presentes, delgadas, no ramificadas; tallos ocultos
por las hojas imbricadas; hojas 0.4–2 mm, dísticas, ad-
presas, sésiles, con un lóbulo superior e inferior, el lóbulo
inferior translúcido y flotante, el superior verde o rojizo,
arqueado-erecto, papiloso; soros generalmente pareados,
uno megasprangiodo (más pequeño) y el otro microspo-
rangiado (más grande), portados sobre el envés de la pri-
mera hoja de una rama lateral, envueltos por un indusio
membranáceo; x=22.
GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Azolla consists of abut 7 species world-
wide, mostly in tropical and subtropical regions of the
world.They grow on the surface of stagnant or slow-mov-
ing water but may become stranded on mud. In the tem-
perate zones, the plants turn reddish in the fall. Azolla
is well known for its symbiosis with the nitrogen-fixing
cyanobacteria Anabaena azollae Strasb. The cyanobacte-
ria can be observed by crushing a leaf on a microscope
slide and examining it with a compound microscope. The
plants are a source of nitrogen, and are used as fertilizer
in the rice paddies of southeast Asia. Four species have
long been recognized in the New World, but according to
Evrard & Van Hove (2004), there are only two: A. cristata
Kaulf. (leaf hairs 2(-3)-celled, glochidia mostly 2 to pluri-
septate; megaspores variously granular) and A. filiculoides
Lam. (leaf hairs unicellular, glochidia mostly septate; mega-
spores warty). A molecular phylogenetic analysis suggests
there are three species in the New World (Reid et al.,
2006). The genus name is derived from the Greek azo, to
dry + olluo, to kill. The fern is presumably killed by drying.
COMENTARIOS: Azolla consta de casi 7 especies
mundial, principalmente en las zonas tropicales y sub-
tropicales del mundo. Crecen sobre la superficie de agua
estancada o en arroyos lentos pero pueden estar sobre
lodo. En las zonas templadas, las plantas se tornan rojizas
en el otoño. Azolla es bien conocida por su simbiosis con
la cianobacteria fijadora de nitrógeno Anabaena azollae
Strasb. La cianobacteria puede ser observada por trit-
urar una hoja sobre un portaobjetos y examinándolas
bajo un microscopio compuesto. Las plantas son una rica
fuente de nitrógeno y se emplean como fertilizante en
los campos de arroz de sureste Asia. Cuatro especies ha
sido reconocidas en el Nuevo Mundo, pero conforme a
Evrard & Van Hove (2004), hay solamente dos especies: A.
cristata Kaulf. (pelitos de la hoja 2(-3) celulares, glochidias
2 a pluriseptadas, megasporas granulares variosamente)
y A. filiculoides Lam. (pelitos de la hoja unicelulares, las
glochidias septadas; megaesporas verrugosas). Un análisis
filogenético molecular sugiere que hay tres especies en
el Nuevo Mundo (Reid et al., 2006). El nombre genérico
se deriva del griego azo, secar + olluo, matar. Presumible-
mente el helecho se muere al secar.

LITERATURE: Evrard, C. & C. Van Hove. 2004. Taxonomy of the American Azolla species (Azollaceae). Syst. Geogr. Pl. 74:
301–318. Perkins, S. K., G. A. Peters, T. A. Lumpkin & H. E. Calvert. 1985. Scanning electron microscopy of perine archi-
tecture as a taxonomic tool in the genus Azolla. Scanning Electron Microscopy 4: 1719–1734. Saunders, R. M. K. & K. Fowl-
er. 1993. The supraspecific taxonomy and evolution of the fern genus Azolla (Azollaceae). Plant Systematics & Evolution 184:
175–193. Reid, J. D., G. M. Plunkett & G. A. Peters. 2006. Phylogenetic relationships in the heterosporous fern genus Azolla
(Azollaceae) based on DNA sequence data from three noncoding regions. International Journal of Plant Science 167: 529538.
Svenson, H. K. 1944. The New World species of Azolla. American Fern Journal 34: 69–84.

200
Figure 97. Azolla sp. B. Anabaena azollae Strasb. C. Leaf cut lengthwise showing the upper and lower lobes, note the
chamber containing Anabaena in the upper lobe. D. Microsporangiate sorus. E. Two sori, the upper microsporangiate, the
lower megasporangiate. F, I. Mature megaspores. G. Massula with microspores and septate glochidia. H. Massula attached to
megaspore. (D–I, original ©Moran, 2016; D–I from Eames, 1936.)

201
BLECHNACEAE
Blechnum L.
DESCRIPTION: Terrestrial, rarely hemiepiphytic or ep-
iphytic; stems scaly, creeping or erect, in some species
massive and arborescent; sterile and fertile leaves mono-
morphous or dimorphous; petioles with several vascular
bundles, the two adaxial ones enlarged; laminae pinna-
tisect to 1-pinnate, rarely simple; pinnae or segments
entire or serrate, never lobed; aerophores absent or
present at the base of the pinnae abaxially, sometimes
also along the petioles; veins free, bifurcate; sori linear,
parallel to the costae; indusia opening toward the cos-
tae (not towards the margin); spores bilateral, nongreen;
2n=28, 29, 31, 32, 33, 34, 36.
DESCRIPCIÓN: Terrestres, hemiepífitas, o raras veces
epífitas; tallos escamosos, rastreros a erectos, en algunas
especies masivos y arborescentes; hojas estériles y fér-
tiles monomorfas o dimorfas; pecíolos con varios haces
vasculares, los dos adaxiales agrandodos; láminas pinna-
tisectas o 1-pinnadas, raras veces simples; pinnas o
segmentos enteros a serrados, nunca lobados; aeróforos
ausentes o presentes en las bases de las pinnas abaxial-
mente y algunas veces a lo largo del pecíolo; venas libres,
bifurcadas; soros lineares, paralelos a las costas; indu-
sios abriéndose hacia la costa (no hacia el margen); es-
poras bilaterales, no verdes; 2n=28, 29, 31, 32, 33, 34, 36.

SIMILAR GENERA: Plagiogyria differs by three vascular GÉNEROS PARECIDOS: Plagiogyria difiere por tres
bundles in the petiole base and pinnae adnate. Polypodium haces vasculares en la base del pecíolo y pinnas adna-
differs by having round, nonindusiate sori and leaves artic- tas.Polypodium difiere por tener soros redondos sin indu-
ulate to the rhizomes. sios y hojas articuladas al rizoma.

COMMENTS: Blechnum consists of about 25 species
and is pantropical. In the future it will probably be divid-
ed into smaller monophyletic genera. In the dimorphic
species of Blechnum with narrow fertile pinnae, it is often
difficult to interpret the soral condition. The sporangia,
when mature, cover the entire abaxial surface of the pin-
nae, obscuring the costae and indusia. This imparts the
false impression of an acrostichoid sorus. The only solu-
tion to the correct interpretation is careful observation.
Usually a vestige of the indusium can be seen near the
margin of the pinna. In the Blechnum occidentale group
(those species with stolons and monomorphous sterile
and fertile leaves) hybrids are frequent. Although they
have abortive spores, the hybrids can become abundant
locally through reproduction by stolons. The genus name
is derived from the Greek blechnon, an ancient name for
ferns in general.
COMENTARIOS: Blechnum consta de casi 25 especies
y es pantropical. En el futuro se espera que será dividido
en géneros más pequeños monofiléticos. En las espe-
cies dimorfas de Blechnum con pinnas fértiles angostas,
a menudo es difícil interpretar la condición soral. Los
esporangios, cuando maduros, cubren toda la superficie
abaxial de las pinnas fértiles, obscureciéndo las costas y
los indusios. Da la impresión falsa de un soro acrosti-
coide. La sola solución para la interpretación correcta es
la observación cuidadosa. Usualmente se puede ver un
véstigio del indusio cerca del margen de la pinna. En el
grupo de Blechnum occidentale (las especies con hojas
estériles y fértiles monomorfas) híbridos son frequentes.
Aunque tienen esporas abortivas, los híbridos pueden
ser abundantes localmente via reproducción por estolo-
nes. El nombre genérico se deriva del griego blechnon, un
nombre antiguo para helechos en general.

LITERATURE: Broadhurst, J. 1912. The genus Struthiopteris and its representatives in North America. Bulletin of the Torrey
Botanical Club 39: 257–278, 357–385. Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for
Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227. Mu-
rillo, M. T. 1968. Blechnum subgénero Blechnum en Sur América, con especial referencia a las especies de Colombia. Nova
Hedwigia 16: 329–366, t. 110–147. Rojas-Alvarado, A. F. 2008. Notes in Blechnum lherminieri complex (Blechnaceae) from
the Neotropics. Métodos en Ecología y Sistemática 3(Supl. 1): 8–29. Rolleri, C. & C. Prada. 2006. Catálogo comentado de
las especies de Blechnum L. (Blechnaceae, Pteridophyta) en Mesoamérica y Sudamérica. Annales Jardín Botánico de Madrid 63:
67–106. Rolleri, C., C. Prada, L. Passarelli, J. M. Gabriel y Galán, & M. M. Ciciarelli. 2012. Revisión de especies monomor-
fas y subdimorfas del género Blechnum (Blechnaceae-Polypodiophyta). Botánica Complutense 36: 51–77.

202
Figure 98. A. Blechnum appendiculatum. B, C. B. occidentale. D. B. polypodioides. E, F. B. serrulatum. G. B. gracile.
(From Mickel & Beitel, 1988.)

203
DENNSTAEDTIACEAE
Blotiella R. M. Tryon
DESCRIPTION: Terrestrial; rhizomes decumbent to
erect, pubescent; leaves up to 3 m long, distant, pendu-
lous to subscandent; petiole bases in cross-section with
a vascular bundle in the form of a omega (Ω); laminae
pubescent on both sufaces, 1- o 2-pinnate-pinnatifid,;
pinnae subopposite; pinnules sessile, mostly adnate; ra-
chises and costae moderately to densely pubescent, the
hairs 3–10 mm long, septate, spreading; veins netted,
the areoles polygonal, without included veinlets; sori
marginal, continuos at the base of the sinus, supplied by
more than one vein, blade margin recurved and differen-
tiated as an indusium (false indusium); inner indusium
absent; spores bilateral, monolete; x=38.
SIMILAR GENERA: Histiopteris differs by laminae gla-
brous and glaucous, with stipule-like pinnules at the base
of the pinnae. Hypolepis differs by sori supplied by a single
vein. Lonchitis differs by free veins.
COMMENTS: Blotiella contains about 15 species, all in
Africa and Madagascar except for one American species,
B. lindeniana (Hook.) R. M. Tryon. This species grows in
wet forests from 1000–2000 m in the Greater Antilles
and from Costa Rica to eastern Brazil. Its leaves, soft and
pliable, are often supported on the surrounding vegeta-
tion. Blotiella is sister to Histiopteris. Both have laminae
with intermittent apical growth, opposite pinnae, and
areolate veins. The genus name honors Marie-Louise Tar-
dieu-Blot, (1902–), French pteridologist
LITERATURE: Tryon, R. M. 1962. Taxonomic fern notes. III. Contributions from the Gray Herbarium 191: 91–107.
204
DESCRIPCIÓN: Terrestres; rizomas ecumbentes a
erectos, pubescentes; hojas hasta 3 m de largo, distan-
tes, péndulas a subescandentes; la base del pecíolo en
sección cruz con un haz vascular en forma de omega
(Ω); láminas pubescentes en ambas superficies, 1- a
2-pinnada-pinnatífidas; pinnas subopuestas; pínnulas
sésiles, a menudo adnatas; requises y costas moderada
a densamente pubescentes, los pelos 3–10 mm de largo,
septados, divergentes; venas anastomosadas, las are-
olas poligonales, sin venas incluídas; soros marginales,
continuos por las bases de los senos, suministrados por
más de una vena, margen de la lámina recurvado y difer-
enciado como indusio (indusio falso); un indusio interior
ausente; esporas bilaterales, monoletes; x=38.
GÉNEROS PARECIDOS: Histiopteris difiere por lámi-
nas glabras y glaucas, con pínnulas basales que asemejan
como estípulas en la base de las pinnas. Lonchitis difiere
por venas libres.
COMENTARIOS: Blotiella consta de alrededor de 15
especies, todas en África y Madagascar con excepción de
una especie Americana, B. lindeniana (Hook.) R. M. Tryon.
Esta especie inhabita bosques húmedos de 1000–2000
m en las Antillas Mayores y desde Costa Rica a Brasil
oriental. Sus hojas, suaves y flojas, están a menudo apoy-
adas por la vegetación circundante. Blotiella es hermana
a Histiopteris. Los dos tienen láminas con crecimiento
apical intermitente, pinnas opuestas y venas areoladas.
El nombre genérico honora a Marie-Louise Tardieu-Blot,
(1902–), pteridólogo francés.
Figure 99. . Blotiella lindeniana. A. Leaf and rhizome. B. Adaxial surface of lamina. C. Abaxial surface of lamina. D. Sorus in
sinus. E. Detail of pinna rachis abaxially. (original, © R. C. Moran, 2010.)

205
DRYOPTERIDACEAE
Bolbitis Schott
DESCRIPTION: Terrestrial, saxicolous, or hemi-epiphyt-
ic; rhizome creeping, scaly, in cross section with a sin-
gle, wide, ventral vascular bundle and 3 or 4 dorsal ones;
sterile and fertile leaves dimorphous, the fertile ones
more erect, longer petiolate, and with narrower pinnae;
laminae simple and entire to 1-pinnate-pinnatifid, glabrous
or sparsely scaly, often with buds at or near the apex;
pinnae continuous (not articulate to) the rachis; rachis
and costae above without decurrent grooves, glabrous;
veins reticulate (free in some Old World species), with
or without included veinlets; sori acrostichoid; indusia
absent; spores monolete; x=41.
DESCRIPCIÓN: Terrestres, saxícolas, o hemiepífitas; ri-
zomas reptantes, escamosos, en corte transversal con
un sólo haz vascular ancho ventral y con 3 o 4 haces
dorsales; hojas estériles y fértiles dimorfas, las hojas
fértiles más erectas, con pecíolos más largos y pinnas más
angostas; láminas simples y enteras a 1-pinnado-pinnatí-
fidas, glabras o escasamente escamosas, a menudo con
brotes en o cerca del ápice; pinnas continuas (no artic-
uladas) al raquis; raquis y costas en la superficie superior
sin surcos decurrentes, glabros; venas reticulate (libres
en algunas especies del Viejo Mundo), con o sin venillas
incluidas; soros acrosticoides; indusios ausentes; espo-
ras monoletes; x=41.

SIMILAR GENERA: Mickelia differs by (usually) scan-
dent habit, laminae imparipinnate (not pinnatifid at the
apices), and buds (when present) at the base of the medi-
al pinnae. Tectaria differs by the pubescent upper surfaces
of its axes and non-acrostichoid (usually round) sori.
GÉNEROS PARECIDOS: Mickelia difiere por ser
(usualmente) hábito escandente, láminas imparipinadas
(no pinatífidas en los apices) y brotes (si presentes) en la
base de las pinnas mediales. Tectaria difiere por la pubes-
cencia en la superficie superior de los ejes y usualmente
soros no acrostichoides (usualmente redondos).

COMMENTS: Bolbitis contains about 50 species, with
about 12 in the Neotropics. It is primarily a genus of low-
to middle-elevation forests. Most of the species grow on
the forest floor or among on rocks in streams. Bolbitis is
sister to the rest of the bolbitidoid ferns; that is, Arthro-
botrya, Elaphoglossum, Lomagramma, Mickelia, and Terato-
phyllum, It greatly resembles Mickelia, with which many of
its species were formerly classified, and no single char-
acter distinguishes them. There are, however, tendencies
to differ. Most species of Bolbitis bear buds at the lamina
apices, whereas only two species of Mickelia have buds,
and these occur at the bases of the medial pinnae. The
lamina apex of Bolbitis is usually pinnatifid, whereas that of
Mickelia is conform, similar to the lateral pinnae. The ge-
nus name is derived from the Greek bolbition, diminutive
of bolbos, bulb. It alludes to the small buds formed on the
leaves of some species.
COMENTARIOS: Bolbitis consta de casi 50 especies,
con casi 12 en el Neotrópico. Es principalmente un géne-
ro de bosques de elevaciones bajas a medias. La mayoría
de las plantas existen en el sotobosque o entre rocas de
riachuelos. Bolbitis es hermana al resto de los helechos
bolbitidoides; i.e., Arthrobotrya, Elaphoglossum, Lomagram-
ma, Mickelia y Teratophyllum. Se parece mucho a Mickelia,
con que muchas de sus especies fueron clasificadas ante-
riormente, y no carácter por su mismo distingue los dos.
Pero hay, no obstante, tendencias diferir. La mayoría de
las especies de Bolbitis llevan brotes en los ápices de las
láminas, mientras que solo dos especies de Mickelia llevan
brotes, estos en las bases de las pinnas mediales. El ápice
de la lámina de Bolbitis es usualmente pinnatífido, mien-
tras que lo de Mickelia es conforme, similar a las pinnas
laterales. El nombre genérico se derive del griego bolbi-
tion, diminutivo de bolbos, bulbo. Se refiere a los brotes
pequeños encontrados en las hojas de algunas especies.

LITERATURE: Hennipman, B. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Botanical Series 2: xiii
+ 331 pp., 12 pl, 87 figs. Moran, R. C., P. Labiak & M. Sundue. 2010. Phylogeny and character evolution in bolbitidoid ferns
(Dryopteridaceae). International Journal of Plant Sciences 171: 547–559. Moran, R. C., P. Labiak & M. Sundue. 2010. Synopsis
of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae). Brittonia 62: 337–356.

206
Figure 100. A–C. Bolbitis aliena (Sw.) Alston. D, E. B. serratifolia (Mert. ex Kaulf.) Schott. F–H. B. umbrosa (Liebm.) Ching. (From
Mickel & Smith, 2004.)

207
PTERIDACEAE
Bommeria E. Fourn.
DESCRIPTION: Terrestrial; rhizomes short- to
long-creeping, scaly and sometimes hairy with unicellular
hairs; leaves monomorphous; petioles castaneous to
atropurpureous, terete or adaxially grooved; laminae
1-16 cm, generally as long as wide, pentagonal, pe-
date, deeply lobed, the indument of the upper sur-
face consisting of unicellular acicular hairs, on the
lower surface of scales and unicellular acicular hairs or
occasionally enrolled, occasionally with 2-cell hairs with
a glandular head; basal segments generally connected to
the apical ones by a wing along the rachis. the basiscopic
lobe elongated or bipinnatifid; veins free or netted, the
areoles without included veinlets; sporangia spreading
along the veins, covering 2/3-3/4 the distance from the
margin to the costa, occasionally restricted toward the
margins; indusium absent; paraphyses absent; spores
globose; x = 30.
DESCRIPCIÓN: Terrestres; rizomas corto a largamente
rastreros, escamosos y a veces pelosos con tricomas
unicelulares; hojas monomorfas; pecíolos castaño a at-
ropurpúreos, teretes a sulcados adaxialmente; lámina
1-16 cm, generalmente tan larga como ancha, pen-
tagonal, pedata, profundamente lobada, el indumento
del haz de tricomas unicelulares, aciculares, el indu-
mento del envés de escamas y tricomas unicelulares, acic-
ulares o a veces enrollados, ocasionalmente con tricomas
2-celulares con la punta glandulosa; segmentos basales
conectados generalmente a los apicales por un ala angos-
ta que corre a lo largo del raquis, el lobo basiscópico alar-
gado a bipinnatífido; nervaduras libres o anastomosadas,
las aréolas sin nérvulos incluidos; esporangios difundi-
dos a lo largo de las nervaduras, que cubren 2/3-3/4
de la distancia del margen a la costa, ocasionalmente se
restringien hacia los márgenes; indusio ausente; paraf-
isos ausentes; esporas globosas; x = 30.

SIMILAR GENERA: Doryopteris differs by glabrous lam- GÉNEROS PARECIDOS: Doryopteris difiere por sus
inae y anastomosing ones (except for two species). Hemi- láminas glabras y nervaduras anastomosadas (con excep-
onitis differs by recurved indusia. ción de dos especies). Hemionitis difiere por su indusios
reflexos.

COMMENTS: Bommeria has 5 species and occurs from
the southwestern United States to Costa Rica. The ge-
nus typically grows in dry, rocky habitats. The genus name
honors Joseph Edouard Bommer (1829–1895), Belgian
pteridologist.
COMENTARIOS: Bommeria tiene 5 especies y ocurre
desde el suroeste de los Estados Unidos hacia Costa Rica.
El género crece típicamente en hábitats secas, rocosas.
El nombre genérico honora Joseph Edouard Bommer
(1829–1895), pteridólogo Bélgico.

LITERATURE: Gastony, G. J. and C. H. Haufler. 1976. Chromosome numbers and apomixis in the fern genus Bommeria
(Gymnogrammaceae) Biotropica 8: 1–11. Haufler, C. H. 1979. A biosystematic revision of Bommeria. Journal of the Arnold
Arboretum 60: 445–476. Haufler, C. H. & G. J. Gastony. 1978. Systematic implications of spore morphology in Bommeria and
related fern genera. Systematic Botany 3: 241–256. Maxon, W. R. 1913. Studies of tropical American ferns—No. 4. Contributions
from the United States National Herbarium 17: 133–179. Ranker, T. A. 1989. Spore morphology and generic delimitation of
New World Hemionitis, Gymnopteris, and Bommeria (Adiantaceae). American Journal of Botany 76: 297–306. Ranker, T. A. &
C. H. Haufler. 1990. A new combination in Bommeria (Adiantaceae). American Fern Journal 80: 1–3.

208
Figure 101. A–C. Bommeria pedata (Sw.) E. Fourn. D–G. B. subpaleacea Maxon. (From Mickel & Smith, 2004.)

209
OPHIOGLOSSACEAE
Botrychium Sw..

DESCRIPTION: Terrestrial; plants small and inconspic-
uous, fleshy to herbaceous; roots not proliferous; leaves
(in temperate regions) dying in late summer or fall; leaf
sheaths closed; leaf primordia glabrous; trophophores
usually less than 7 cm wide, 1- to 2-pinnate, oblong to
broadly triangular, sessile to stalked; pinnae entire to pin-
natifid, glabrous; veins free, forked; sporophores typically
present on juvenile plants, joined to trophophore well
above ground; x=45.
DESCRIPCIÓN: Terrestres; plantas pequeñas y incon-
spículas, carnosas a herbáceas; raíces no prolíferas; hojas
(en las zonas templadas) muriéndose en el verano tarde
o el otño; vainas de las hojas cerradas; primórdios de las
hojas glabros; trofoforos usualmente menos de 7 cm de
ancho, 1- a 2-pinadas, oblongos a anchamented deltados,
sésiles o peciolulados; pinnas enteras a pinatífidas, glabras;
venas libres, furcados; sporoforos típicamente presentes
en plantas juveniles, únidos al trofoforo bien arriba del
suelo; x=45.

SIMILAR GENERA: Sceptridium differs by sporophores GÉNEROS PARECIDOS: Sceptridium difiere por spo-
united to common stalk below ground. Botrypus differs roforos únidos a un pecíolo común abajo del suelo. Botry-
by larger size and more highly divided trophophores. pus difiere por su tomaño más grande y trofoforos más
divididos.

COMMENTS: Botrychium has about 30 species world-
wide. It is most diverse in north-temperate and boreal
North America, but also occurs in southern South Amer-
ica and Eurasia. They are small plants, usually less than 15
cm tall, and for that reason easily overlooked. Like all mem-
bers of the Ophioglossaceae, Botrychium lacks root hairs
and depends on endomycorrizal fungi (genus Glomus) for
uptake of water and nutrients. Quite likely because of
their strong reliance on mycorrizal nutrition, Botrychium
species have become reduced in size and morphologi-
cal complexity relative to other ferns. The genus name
is derived from the Greek botrys, cluster, refering. to the
grape-like clusters of sporangia.
COMENTARIOS: Botrychium tiene casi 30 especies
mundial. Es mas diverso en América del Norte boreal y
templado, pero existe también en el America del Sur aus-
tral y Eurasia. Son plantas pequeñas, usualmente menos
de 15 cm de altura, y por eso fácilmente sobrepasado.
Como otros miembros de Ophioglossaceae, Botrychi-
um carece de pelos de raíces y depende de hongos en-
domicorrizales (género Glomus) para absorbar agua y
nutrientes. Es probable dado su dependencia fuerte en
alimentación micorrizal, que especies de Botrychium lle-
garon a ser reducidas en tomaño y complexidad mor-
fológico relativo a otros helechos. El nombre genérico se
deriva del griego botrys, grupo, referiéndose a los espor-
angios que se asemejan a un grupo de uvas. .

LITERATURE: Clausen, R. T. 1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19(2): 1–177,
figs. 1–33. Farrar, D. R. 2011. Moonwort (Botrychium) systematics. http://www.herbarium.iastate.edu/botrychium.html Hauk,
W. D., C. R. Parks & M. W. Chase. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL and trnL–F plastid
DNA sequences and morphology. Molecular Phylogenetics and Evolution 28: 131–151. Hauk, W. D., L. Kennedy & H. M.
Hawke. 2012. A phylogenetic investigation of Botrychium s.s. (Ophioglossaceae): evidence from three plastid DNA sequence
datasets. Systematic Botany 37: 320–330.

210
Figure 102. A, B. Botrychium tolucaense. C–E. B. minganense. F. B. matricariifolium. G. B. simplex. H. B. lunaria. J. B. mormo.
(A–D from Mickel & Smith, 2004; F–J, original.)

211
OPHIOGLOSSACEAE
Botrypus Sw..

DESCRIPTION: Terrestrial; plants small to large; roots
not proliferous; leaves herbaceous, deciduous, in the tem-
perate zones dying in fall; leaf sheaths open; leaf primordia
pubescent; trophophores often more than 7 cm wide,
3- to 5-pinnate, broadly deltate, sessile, herbaceous,
with a midrib; veins free, forked; sporophores attached
above ground at base of trophophore, often absent
or aborted; x=46.
DESCRIPCIÓN: Terrestres; plantas pequeñas y grandes;
raíces no prolíferas; hojas herbáceas, decíduas, en las zo-
nas templadas muriéndose en el otño; vainas de las hojas
abiertas; primórdios de las hojas pubescentes; trofoforos
usualmente más de 7 cm de ancho, 3- a 5-pinadas, an-
chamented deltados, sésiles, herbáceos, con una costa;
venas libres, furcados; sporoforos únidos al trofoforo
arriba del suelo en la base del trofoforo, típicamente
ausentes o abortados; x=46.

SIMILAR GENERA: Sceptridium differs by sporophores GÉNEROS PARECIDOS: Sceptridium difiere por spo-
united to common stalk below ground. Botrychium differs roforos únidos a un pecíolo común abajo del suelo. Botry-
by smaller size and less highly divided trophophores. Ane- chium difiere por su tomaño más pequeño y trofoforos
mia differs by having two erect fertile stalks. menos divididos. Anemia difiere por dos espigas fértiles.

COMMENTS: Botrypus has 2 or 3 species worldwide.
The only species in the Americas is B. virginianus (L.)
Michx., which occurs from Canada to Bolivia, typically in
wet woods and (in the tropics) cloud forests. Unlike oth-
er Ophioglossaceae (Botrychium, Cheiroglossa, and Scep-
tridium), its leaf sheaths are open, and the leaf primordia
are pubescent. Gametophytes, which are potato-shaped
and about 1 cm wide, can usually be found still attached
to young sporophytes by digging up plants with leaves less
than 5 cm long. A trophophore may not always develop,
and when this happens, an small (2–3 mm long) aborted
bud can be found on the adaxial surface of the common
stalk a few cm below the trophophore. The genus name
is derived from the Greek botrys, cluster, and pous, foot,
refering. to the clusters of sporangia are borne on a foot
or stalk.
COMENTARIOS: Botrypus tiene 2 o 3 especies mun-
dial. La única especie en las Américas es B. virginianus
(L.) Michx., la cual existe desde Canadá hasta Bolivia,
típicamente en bosques húmedos y (en el Neotrópico)
bosques nubosos. Diferente de otras Ophioglossaceae
(Botrychium, Cheiroglossa y Sceptridium), sus vainas de las
hojas son abiertas, y los primórdios de las hojas son pu-
bescentes. Gametofitos, la cuales son casi 1 cm de ancho y
tienen la forma de una papa, se puede encontrar undios a
esporofitos jóvenes por excavar plantas con hojas menos
de 5 cm de largo. Un trofoforo no siempre se desarolla,
y cuando esto occure, un brote abortado pequeño (2–3
mm de largo) ocurre en la superficie adaxial del pecíolo
común. El nombre genérico se deriva del griego botrys,
grupo, y pous, pie, referiéndose a los esporangios que se
nacen en un pie o pecíolo.

LITERATURE: Butters, F. K. 1917. Botrychium virginianum and its American varieties. Rhodora 19: 209. Clausen, R. T.
1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19(2): 1–177, figs. 1–33. Hauk, W. D., C. R.
Parks & M. W. Chase. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL and trnL–F plastid DNA sequences
and morphology. Molecular Phylogenetics and Evolution 28: 131–151. Hauk, W. D., L. Kennedy & H. M. Hawke. 2012. A
phylogenetic investigation of Botrychium s.s. (Ophioglossaceae): evidence from three plastid DNA sequence datasets. Systematic
Botany 37: 320–330.

212
Figure 103. A. Botrypus virginianus. (Mickel & Smith, 2004.)

213
POLYPODIACEAE
Campyloneurum C. Presl
DESCRIPTION: Epiphytic, occasionally terrestrial or
rupestral; rhizomes short- to long-creeping scaly, the
scales usually lanceolate, clathrate or nonclathrate, gla-
brous; sterile and fertile leaves monomorphous;
petioles articulate to the stem or to short phyllopodia;
laminae simple, entire (but 1-pinnate in 2 species), ob-
long to narrowly elliptic, glabrous or with a few scales
on the rachis; venation areolate with included veinlets,
these usually between prominent, lateral veins; sori in
2–4 rows between the main lateral veins, borne on
the tips of included veinlets, round, without indusia,
and several rows between the rachis and margin of the
lamina; sporangia glabrous; paraphyses absent; spores bi-
lateral, yellow; x=37.
SIMILAR GENERA: Niphidium differs by one row of
sori between the lateral veins. Other genera of Polypo-
diaceae with simple, entire leaves, such as Microgramma
and Pleopeltis, are easily distinguished by having only one
row of sori between the rachis and margin of the lamina.
Elaphoglossum differs by acrostichoid sori and (almost al-
ways) free veins.
COMMENTS: Campyloneurum consists of about 50 spe-
cies and is entirely neotropical. Its species grow primarily
in wet forests.Two species are atypical by having 1-pinnate
laminae: C. decurrens (Raddi) C. Presl and C. magnificum T.
Moore. They are included in the genus because they have
venation like that of the simple-leaved species. The genus
name is derived from the Greek kampylos, arched + neu-
ron, vein. It refers to the secondary veins that are arched
between the main lateral ones.
LITERATURE: Lellinger, D. B. 1988. Some new species of Campyloneurum and a provisional key to the genus. American
Fern Journal 78: 14-35. León, B. 1992. A taxonomic revision of the fern genus Campyloneurum (Polypodiaceae). Ph.D. Disser-
tation. Aarhus University, Denmark. Rojas, A. F. 2005. El complejo de Campyloneurum angustifolium (Sw.) Fée en Costa Rica.
Lankesteriana 5: 41–48. Vasques, D. T. & J. Prado. 2011. Campyloneurum C. Presl (Polypodiaceae) in São Paulo State, Brazil.
Hoehnea 38: 147–163.
214
DESCRIPCIÓN: Epífitas, ocasionalmente terrestres o
saxícolas; rizomas cortamente a largamente rastreros, es-
camosos, las escamas usualmente lanceoladas, clatradas o
no clatradas, glabras; hojas estériles y fértiles mono-
morfas; pecíolos articulados al tallo o a las filopódios
cortos; láminas simples, enteras, (pero 1-pinnadas en
2 especies), oblongas a angostamente elípticas, glabras
o con pocas escamas sobre el raquis; venación areo-
lada, con venillas incluídas, estas usualmente entre venas
laterales prominentes; soros en 2–4 hileras entre las
venas laterales principales, se nacen en los ápices
de venillas incluídas, redondos, sin indusios, en varias
hileras entre el raquis y margen de la lámina; esporangios
glabros; parafises ausentes; esporas bilaterales, amarillas;
x=37.
GÉNEROS PARECIDOS: Niphidium difiere por una
hilera de soros entre las venas laterales. Otros géneros
de Polypodiaceae con hojas simples y enteras, tales como
Microgramma y Pleopeltis, son distinguidos facilmente por
tener una sola hilera de soros entre el raquis y margen de
la lámina. Elaphoglossum difiere por soros acrostichoides
y (casi siempre) venas libres.
COMENTARIOS: Campyloneurum consta de alrededor
de 50 especies y es completamente neotropical. Sus es-
pecies se encuentra principalmente en bosques húmedos.
Dos especies son atípicas por tener láminas 1-pinnadas:
C. decurrens (Raddi) C. Presl y C. magnificum T. Moore. Es-
tán incluídas dentro del género porque tienen venación
parecida a éso en las especies con hojas simples. El nom-
bre genérico se deriva del griego kampylos, arqueado +
neuron, vena. Se refiere a las venas secundarias que son
arcuadas entre las venas laterales principales.
Figure 104. A, B. Campyloneurum amphostenon. C, D. C. angustifolium. E–G. C. tenuipes. H. C. xalapense. J, K. C. repens.
L–N. C. phyllitidis. (From Mickel & Beitel, 1988.)

215
POLYPODIACEAE
Ceradenia L.E. Bishop
DESCRIPTION: Typically epiphytic; rhizome scales non-
clathrate, often ciliate and with whitish, marginal glands
when young; petiole often articulate; laminae simple,
pinnatifid or peripinnate, often pectinate; veins often
1-forked, sometimes simple; hydathodes absent; setae
reddish; trichomes simple or branched, sometimes glan-
dular; sori round or oblong, superficial or subimpressed;
paraphyses glandlike, waxy, globose or pyriform,
whitish, brown-yellow to yellowish, multicellular; spores
green, tetrahedral; x=37.
DESCRIPCIÓN: Típicamente epífitas; escamas del
rizoma no clatradas, a menudo ciliadas y con glándulas
marginales blanquecinas cuando jóvenes; pecíolo a menu-
do articulado; láminas simples, pinnatífidas o peripinnadas,
a menudo pectinadas; nervaduras a menudo 1-bifurcadas,
a veces simples; hidátodos ausentes; setas rojizas; tri-
comas simples o ramificados, a veces glandulares; soros
redondeados u oblongos, superficiales o subimpresos;
paráfisis como glándulas, cerosos, globosos o piri-
formes, blanquecinos, pardo-amarillentos a amarillentos,
multicelulares; esporas verdes, tetraédricas; x=37.

SIMILAR GENERA: Zygophlebia and Enterosora differ
by anastomosing veins, but this is difficult to see because
of the thickness of the lamina. They lack glands in the sori,
or if present (Zygophlebia), then they are brown and not
waxy or whitish. Many other genera of Grammitidace-
ae, such as Lellingeria, Melpomene y Terpsichore, can be
distinguished by the presence of hydathodes. Cochlidium
Grammitis, Lellingeria, and Luisma differ by the absence of
reddish setae.
GÉNEROS PARECIDOS: Zygophlebia e Enterosora di-
fieren por venas anastomosadas, pero esto es difícil de
distinguir debido al grosor de la lámina. Faltan glándulas
en los soros, y si son presentes (Zygophlebia), entonces
son pardos y no cerosos blanquecinos. Se puede distin-
guir muchos otros géneros de las Grammitidaceae, tales
como Lellingeria, Melpomene y Terpsichore, por la presen-
cia de hidatodos. Cochlidium Grammitis, Lellingeria y Luis-
ma difieren por la ausencia de setas rojizas.

COMMENTS: Ceradenia is a primarily neotropical with COMENTARIOS: Ceradenia es principalmente neo-

about 60 species, with 2 in the Azores and Liberia, and 6
in Africa-Madagascar. It is the largest genus of Neotropi-
cal grammitid ferns. Nearly all grow in cloud forests. The
most distinctive characteristic of this genus is the whitish
glands in the sorus, from which the genus takes its name.
These can be so numerous that they obscure the young
sporangia and can be mistaken for spores in more mature
specimens. Some species also have branched, gland-bear-
ing hairs on the lamina, axes, and margins of the rhizome
scales. In these locations, the hairs are often branched,
with 2–5 glands per hair. Sometimes the glands are diffi-
cult to observe in specimens collected in alcohol or dried
with high heat. Ceradenia is monophyletic and sister to
Enterosora (Ranker et al., 2004). The genus name is de-
rived from the Greek keras, wax, and aden, gland, referring
to the distinctive glands in the sorus.
tropical con casi 60 especies, con 2 en los Azores y Libe-
ria, y 6 en África-Madagascar. Es el género más grande de
helechos neotropicales. Casi todos existen en bosques
nublados. El carácter más distintivo del género es las
glándulas blanquecinas en el soro, de las cuales el género
toma su nombre. Estas pueden san tan numerosas que
esconden los esporangios jóvenes, y pueden ser malin-
terpretados como esporas en ejemplares más maduros.
Algunas especies también tienen pelitos glandulíferos por
las laminas, ejes y márgenes de las escamas del rizoma.
En estas locaciones, las pelitos son a menudo ramificados
con 2–5 glándulas por pelito. A veces las glándulas son
difíciles de observar en ejemplares colectados en alcohol
o secados con alto calor. Ceradenia es monofilético y her-
mano a Enterosora (Ranker et al., 2004). Ceradenia deriva
del griego keras, cera, e aden, glándula, refiriéndose a las
glándulas distintivos en el soro.

LITERATURE: Bishop, L. E. 1988. Ceradenia, a new genus of Grammitidaceae. American Fern Journal 78: 1–5. Bishop, L.
E. 1989. New species of Ceradenia subgen. Ceradenia. American Fern Journal 79: 14–25. Bishop, L. E. 1989. New species of
Ceradenia subg. Ceradenia. American Fern Journal 79: 14–25. Labiak, P. H. & J. Prado. 2003. Grammitidaceae (Pteridophyta)
no Brasil com ênfasis nos gêneros Ceradenia, Cochlidium, e Grammitis. Hoehnea 30: 243–283. León-Parra, A. 2012. Ceradenia
(Polypodiaceae). Flora of Colombia 29: 7–83. Smith, A. R. 1993. New species and combinations of Ceradenia (Grammitidace-
ae). Novon 3: 182–185.

216
Figure 105. Ceradenia. In G and K, note whitish glands among sporangia. En G y K, fijase las glándulas blanquecinas entre los
esporangios (From Smith, 1993.)

217
PTERIDACEAE
Ceratopteris Brongn.
DESCRIPTION: Plants free-floating or rooting
in the mud; rhizomes inconspicuous, short-creeping,
sparsely scaly; sterile and fertile fronds dimorphic,
the fertile ones contracted and more erect compared to
the sterile; sterile fronds pinnatifid to 3-pinnate, glabrous;
veins areolate; buds present in the pinna sinuses; sori
marginal, with the margin recurved over the sori; annulus
weak and ill-defined; spores tetrahedral-globose, 16 or 32
per sporangium; x=39.
SIMILAR GENERA: None.
COMMENTS: Ceratopteris is pantropical and subtropical
usually occurs in standing or slow-moving water. The ge-
nus was monographed by Lloyd (1974), who recognized
three species in the Neotropics: C. pteridoides (Hook.)
Hieron., C. richardii Brongn. and C. thalictroides (L.) Brongn.
The spores, which are 70–150 micrometers long, are
among the largest of any homosporous fern. They bear
have parallel ridges that can easily be seen under a dis-
secting microscope at 30X. The chromosome number of
x=39 is unique in the family. The genus is frequently used
for genetic research because it has the shortest life cycle
of any homosporous fern (three months), and the plants
grow easily in the laboratory. Reproduction by buds in
the sinuses of the pinnae is extremely common. The spe-
cies are frequently sold for use in aquaria. Ceratopteris is
the sister genus to Acrostichum, which looks quite differ-
ent but also grows in wet soils. The two are classified in
the subfamily Parkerioideae. The generic name is derived
from the Greek keras, horn + pteris, fern. It alludes to the
antler-like fertile leaves.
LITERATURE: Gastony, G. J. & D. R. Rollo. 1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern
United States and adjacent Mexico--a molecular phylogenetic reassessment of generic lines. Aliso 17: 131–144. Lloyd, R. M.
1974. Systematics of the genus Ceratopteris Brongn. (Parkeriaceae) II. Taxonomy. Brittonia 26: 139–160.
218
DESCRIPCIÓN: Plantas flotantes o echando raíces
en el lodo; rizomas inconspícuos, cortamente rasteros,
escamosos; las hojas estériles y fértiles dimorfas, las
fértiles contraídas y más erectas en comparación con las
estériles; láminas estériles pinatifídas a 3-pinnadas, glabras;
venas areoladas; brotes presentes en los sinuses de las
pinnas; soros marginales, con los márgenes recurvados
por los soros; anillos débiles y mal definidos, 16 o 32 por
esporangio; esporas tetraédricas-globosas; x=39.
GÉNEROS PARECIDOS: Ninguno.
COMENTARIOS: Ceratopteris es pantropical y sub-
tropical y ocurre usualmente en aguas estancadas o len-
tas. El género fue monografiado por Lloyd (1974) quién
reconoció tres especies en el Neotrópico: C. pteridoides
(Hook.) Hieron., C. richardii Brongn. y C. thalictroides (L.)
Brongn. Las esporas, cuales son 70–150 micrometros
de largo, son unas de los más grandes entre helechos
homospóricos. Tienen crestas paralelas fácilmente vistas
con un microscopio de disección a 30X. El número cro-
mosómico de x=39 es único en la familia. El género se
usa frecuentemente para investigacíon genética porque
tiene el ciclo de vida más corto de cualquier helecho
homospórico (tres meses), y las plantas crecen fácilmente
en el laboratorio. La reproducción por brotes en los si-
nuses de las pinnas es muy común. Se venden las especies
con frecuencia para uso en los acuarios. Ceratopteris es el
género hermana a Acrostichum, que parece muy diferente
pero también inhabita suelos mojados. Los dos se clasif-
ican en la subfamilia Parkerioideae. El nombre genérico
se deriva del griego keras, cuerno + pteris, helecho. Se
refiere a las hojas fértiles como cuernos.
Figure 106. A–D. Ceratopteris pteridoides. D–G. C. thalictroides (From Mickel & Smith, 2004.)

219
PTERIDACEAE
Cheilanthes Sw.
DESCRIPTION: Terrestrial or rupicolous; rhizomes
prostrate or ascending, scaly; plants generally less than
30 cm long; leaves monomorphic or subdimorphic, her-
baceous to coriaceous; petioles with 1 vascular bun-
dle, often castaneous to blackish y shiny, terete or sulcate
adaxially; laminae 1-5-pinnate, not farinose; veins free,
the tips not claviform; indusium reflexed, scarious;
paraphyses absent; spores tetrahedral-globose to glo-
bose, trilete, 64 or, more commonly, 32 per sporangium.
x=29, 30.
DESCRIPCIÓN: Terrestres o rupícolas; rizoma postra-
do o ascendente, escamoso; plantas generalmente
menores de 30 cm de largo; hojas monomorfas o
subdimorfas, herbáceas a coriáceas; pecíolo con un haz
vascular, a menudo castaño a negruzco y lustroso, terete
o sulcado adaxialmente; lámina 1-5-pinnada, no farinosa;
nervaduras libres, los ápices no claviformes; esporangios
dispuestos sobre o cerca de los ápices de las nervaduras;
indusio generalmente reflexo, escarioso; parafisos
ausentes; esporas tetraédrico-globosas a globosas, triletes,
64 o, más comúnmente, 32 por esporangio; x=29, 30.

SIMILAR GENERA: Notholaena and Argyrochosma dif- GÉNEROS PARECIDOS: Notholaena y Argyrochosma
fer by farinose laminae. Notholeana differs furthermore difieren por láminas farinosas. Notholeana difiere además
by sori not covered by the reflexed margin of the lamina. por los soros no cubiertos por el margen reflexo de la
lámina.

COMMENTS: Cheilanthes is cosmopolitan and contains COMENTARIOS: Cheilanthes es cosmopolito y consta

about 150 species. It occurs mainly in dry habitats, and
most of its species are apogamous with 32 spores per
sporangium. In the New World, its center of diversity and
endemism is Mexico. Even with the removal of Adian-
topsis, Aleuritopteris, Argyrochosma, Astrolepis, Cheiloplecton,
Mildella, and Notholaena, Cheilanthes remains a large and
heterogeneous group. Future studies will probably result
in the recognition of additional segregate genera, such as
Physapteris. The genus name is derived from the Greek
cheilos, lip + anthos, flower. It alludes to the indusia and
the sori.
de casi 150 especies. Existe principalmente en hábitats
secas, y la mayoría de sus especies son apógamas con 32
esporas por esporangio. Su centro de diversificación y
endemismo en el Nuevo Mundo es en México. Aun con
la exclusión de Adiantopsis, Aleuritopteris, Argyrochosma,
Astrolepis, Cheiloplecton, Mildella y Notholaena, Cheilanthes
permanece como un grupo numeroso y heterogéneo. Es
probable que estudios futuros resulten en la designación
de géneros adicionales segregados, tales como Physapter-
is.El nombre genérico se deriva del griego cheilos, labio +
anthos, flor. Alude al indusio y los soros.

LITERATURE: Mickel, J. T. 1979.The fern genus Cheilanthes in the continental United States. Phytologia 41: 431–437. Reeves,
T. A. 1979. Monograph of the fern genus Cheilanthes Subgenus Physapteris (Adiantaceae). Ph.D. dissertation. Arizona State
University, Tempe. Tryon, R. M. 1956. A revision of the American species of Notholaena. Contributions to the Gray Herbarium
of Harvard University 179: 1–106.

220
Figure 107. A–C. Cheilanthes kaulfussii. D, E. C. longipila. F–H. C. lendigera. J–P. C. myriophylla. (From Mickel & Smith, 2004.)

221
OPHIOGLOSSACEAE
Cheiroglossa C. Presl.
DESCRIPTION: Epiphytes; roots dichotomously divid-
ed, smooth (without root hairs), proliferous; leaves pen-
dent; trophophores arching or pendent, laminae stalked,
palmately lobed, broadly fan-shaped, to 45 × 30 cm;
margins entire; veins anastomosing, major areoles 35 × 8
mm; sporophores numerous, to 10, arising on each side
of base of trophophore; sporangial clusters with sporan-
gia in 2 rows embedded in compact, linear spike, apicu-
lum absent; gametophytes subterranean, 1–4 mm thick,
brown to white, cylindric, repeatedly branched, branch-
es 2–20, and the whole ± stellate structure sometimes
reaching 20 mm. x=?
DESCRIPCIÓN: Epífitas; raíces divididas dicotómica-
mente, lisas (sin pelitos de raices), prolíferas; hojas péndu-
las, trofoforos arcuadas o pendentes, láminas pecioluladas,
lobuladas palmadamente, anchamente flabelliforme, has-
ta 45 × 30 cm; márgenes enteros; venas anastomosadas,
areolas más grandes 35 × 8 mm; sporoforos numero-
sos, hasta 10, producidos en cada lado de la base del
trofoforo; esporangios en 2 hileras enterrado en espigas
compactas lineares, apículas ausentes; gametofitos suber-
raneos, 1–4 mm de grosor, pardos a blancos, cilíndricos,
ramificadas repetidamente, las ramas 2–20, la estructura
entera ± esteliforme, a veces hasta 20 mm; x=?

SIMILAR GENERA: Cheiroglossa differs by terrestrial GÉNEROS PARECIDOS: Cheiroglossa difier por habito
habit, simple, entire trophophores, and sporophores lon- terrestre, trofoforos simples y enteros, y esporoforos lar-
ger-stalked and only one per leaf. gamente peciolulados y sola una por hoja.

COMMENTS: Cheiroglossa consists of a single species,
O. palmata (L.) C. Presl, which occurs in the Neotropics,
Zaire and Madagascar, Réunion, Seychelles, and Vietnam. It
is a rare epiphyte, found from 0–2500 m. It often occurs
(like Psilotum) in the persistent, sheathing leaf bases of
palm where humus accumulates. It also occurs on mossy
trunks, especially in the crotches of trees.The genus name
is derived from the Greek cheiros, hand + glossa, tongue,
alluding to the trophophore that resembles a hand and
the sporophores, tongues.
COMENTARIOS: Cheiroglossa consta de una sola espe-
cie, O. palmata (L.) C. Presl, que occur en el Neotrópico
y Zaire, Madagascar, Réunion, Seychelles y Vietnam. Es un
epífita rara, se encuentra de 0–2500 m. Muchas veces ex-
iste (como Psilotum) en las bases vainadores persistentes
de palmas, donde humus se acumula. También existe en
troncos musgosos y las horquetas. El nombre genérico se
deriva del griego cheiros, mano + glossa, lengua, aludién-
dose trofoforo que parece como un mano y los esporo-
foros como lenguas.

LITERATURE: Hauk, W. D., C. R. Parks & M. W. Chase. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL
and trnL–F plastid DNA sequences and morphology. Molecular Phylogenetics and Evolution 28: 131–151.

222
Figure 108. Cheiroglossa palmata. (original ©R. C. Moran, 1992.)

223
CIBOTIACEAE
Cibotium Kaulf.
DESCRIPTION: Terrestrial; trunks erect, to 1.5 m tall
(to 15 m in Asia and Pacific), densely lanose, the hairs soft,
golden; leaves 2–4 m long; petioles densely lanose like the
trunk; laminae 2-pinnate-pinnatifid to 3-pinnate, glabrous,
glaucous or green abaxially; veins free; sori marginal; in-
dusial bivalvate, the lower valve narrower and fitting
into the recurved upper one, both valves differing from
the lamina tissue in color; spores non-green, tetrahedral,
with a prominent equatorial flange. x=68.
SMILAR GENERA: Culcita differs by non-arborescent
trunks, broadly deltate laminae that are green abaxially
(never glaucous), and valves of the indusial of equal size.
Lophosoria differs by abaxial, non-indusiate sori.
COMMENTS: Culcita consists of 9 species, of which 2
occur in northern Central America, 3 in Asia, and 4 in
Hawaii. Our species typically grow in forests or thickets
from (–750)1000–2500(–3000) m. The two American
species are C. schiedei Schldtl. & Cham. (Veracruz and
Oaxaca) and C. regale Verschaff. & Lem. (s. Mexico to
Honduras and El Salvador). The genus name is derived
from the Greek kibotos, a small chest or casket, alluding
to the shape of the indusium.
LITERATURE: Maxon, W. R. 1912. Studies of tropical American ferns. No. 3. Contributions from the United States National
Herbarium 16: 54–58. Geiger, J. M. O., P. Korall, T. A. Ranker, A. C. Kleist & C. L. Nelson. 2014. Molecular phylogenetic
relationships of Cibotium and the origin of the Hawaiian endemics. American Fern Journal 103: 141–152.
224
DESCRIPCIÓN: Terrestres; troncos erectos, hasta 1.5
m de altura (hasta 15 m en Asia y el Pacífico), densa-
mente lanosos; los pelos suaves, dorados; hojas 2–4 m
de longitud; pecíolos densamente lanosos como el tron-
co; láminas 2-pinnado-pinnatífidas a 3-pinnadas, glabras,
glaucas o verdes abaxialmente; nervaduras libres; soros
marginales; indusios bivalvados, la valva inferior más
angosta y cabe a dentro lo de arriba recurvada, las
dos valvas difieren del tejido de la lámina en color; espo-
ras no verdes, tetrahédricas, con un cíngulo prominen-
te ecuatorial. x=68.
GÉNEROS PARECIDOS: Culcita difiere por troncos
no arborescentes, láminas anchamente deltadas que son
verdes abaxialmente (nunca glaucas) y valvas del indusio
de tamaño igual. Lophosoria difiere por soros abaxiales
no indusiados.
COMENTARIOS: Culcita consta de 9 especies, de las
cuales 2 existen en América Central septentrional, 3 en
Asia y 4 en Hawaii. Nuestras especies crecen típicamente
en bosques o matorrales desde (–750)1000–2500(–
3000) m. Las dos especies Americanas son C. schiedei
Schldtl. & Cham. (Veracruz and Oaxaca) and C. regale
Verschaff. & Lem. (s. Mexico to Honduras and El Salva-
dor).El nombre genérico se deriva del griego kibotos, una
caja pequeña o arquita.
Figure 109. Cibotium. A–C. Cibotium schiedei. D. C. regale. (From Mickel & Smith, 2004.)

225
POLYPODIACEAE
Cochlidium Kaulf.
DESCRIPTION: Epiphytic or saxícolous; rhizome scales
not clathrate, eciliate, dull or somewhat lustrous, brown-yel-
low to brown; leaves less than 25 cm long, typically simple,
linear, entire or deeply serrate, glabrous or with simple,
2–8-celled hairs (without dark setae); veins free or con-
nivent basally en a commissural vein, sterile veins simple
or 1-forked; hydathodes present; sori grouped in the
distal part of the lamina; separate or linearly confluen;
paraphyses absent; spores green, tetrahedral. x=33, 34?,
35.
DESCRIPCIÓN: Epífitas o rupícolas; escamas del rizoma
no clatradas, sin cilias, opacas o algo lustrosas, pardo-am-
arillentas a pardas; hojas menos de 25 cm de longitud,
típicamente simples, lineares, enteras o profundamente
serradas, glabras o con pelos simples 2–8-celulares
(sin setas oscuras); nervaduras libres o conniventes
basalmente en una nervadura comisural, nervaduras
estériles simples o 1-bifurcadas; hidátodos presentes; so-
ros agrupados en la parte distal de la lámina, separa-
dos o linearmente confluentes; paráfisis ausentes; esporas
verdes, tetrahédricas. x=33, 34?, 35.

SMILAR GENERA: Grammitis differs by having a dark
laminar margin. Luisma differs by two vascular bundles
in the petiole. Several species of Lellingeria resemble C.
serrulatum (Sw.) L. E. Bishop, but can be distinguished by
clathrate rhizome scales and at least a few small hairs
(especially on the petiole and raquis) unequally forked.
GÉNEROS PARECIDOS: Grammitis difiere por un
margen laminar oscuro. Luisma difiere por dos haces
vasculares en el pecíolo. Varias especies de Lellingeria se
parecen a C. serrulatum (Sw.) L. E. Bishop, pero pueden
distinguírseles por escamas del rizoma clatradas y al
menos algunos pelitos (especialmente en el pecíolo y ra-
quis) desigualmente furcados.

COMMENTS: Cochlidium consists of 16 species primarily COMENTARIOS: Cochlidium consta de 16 especies

in the Neotropics, with one species (C. serrulatum) rang- principalmente en el Neotrópico, con una especie (C.
ing from Africa to islands of the Indian Ocean (Madagscar, serrulatum) extendiéndose a África y las islas del Oceano
Mauritius, Amsterdam Island). The genus is unique among Indico (Madagascar, Mauricio, Isla Amsterdam). El géne-
grammitid ferns by the sori restricted to the apex of the ro único entre los helechos grammitoides por los soros
blades. In many species the sori are fused (coensori) into restingidos a la parte distal de la lámina. En muchas espe-
a central line.Two common and widespread species in cies los soros son fusionados (coenosoros) en una línea
the Neotropics are C. linearifolium (Desv.) Maxon ex C. central. Dos especies comunes y muy difundidas en el
Chr. and C. rostratum (Hook.) Maxon ex C. Chr. The two Neotrópico son C. linearifolium (Desv.) Maxon ex C. Chr.
have simple and entire laminae and look very much alike, y C. rostratum (Hook.) Maxon ex C. Chr. Las dos tienen
but it is often possible to separate them on elevation: C. láminas simples y enteras y se parecen mucho, pero a
linearifolium occurs between 0–800 m, whereas C. rostra- menudo es posible separarlas por elevación: C. linearifoli-
tum occurs between 600–2600 m. Only one species of um existe entre los 0–800 m, mientras tanto C. rostratum
Cochlidium has a serrate lamina: C. serrulatum. Ranker et al. existe entre los 600–2600 m. Solamente una especie de
(2004) found that Cochlidium is monophyletic and proba- Cochlidium tiene lámina serrada: C. serrulatum. Ranker et
bly sister to the black-margined, simple-leaved species of al. (2004) encontraron que Cochlidium es monofilético
Grammitis (Grammitis s.str.). The genus name is derived y probablemente hermana a las especies de Grammitis
from the Greek cochlea, spoon + eidos, like. The distal con hojas simples y bordes negros (Grammitis s. str.). El
part of the leaf is spoon-like. nombre genérico deriva del griego cochlea, cuchara +
eidos, parecido. La parte distal de la hoja se asemeja a
una cuchara.
LITERATURE: Bishop, L. E. 1978. Revision of the genus Cochlidium (Grammitidaceae). American Fern Journal 68: 76–94.
Maxon, W. R. 1914. Notes upon Polypodium duale and its allies. Contributions from the United States National Herbarium 17:
398–406. Labiak, P. H. & J. Prado. 2003. Grammitidaceae (Pteridophyta) no Brasil com ênfasis nos gêneros Ceradenia, Coch-
lidium, e Grammitis. Hoehnea 30: 243–283. Ranker, T. A., A. R. Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K.
Staub & H. Schneider. 2004. Phylogeny and evolution and grammitid ferns (Grammitidaceae): a case of rampant morphological
homoplasy. Taxon 53: 415–428.

226
Figure 110. C–E. Cochlidium serrulatum (Sw.) L. E. Bishop. G. Cochlidium rostratum (Hook.) Maxon ex C. Chr.; note sorus
sunken in the central groove (From Mickel & Beitel, 1988.)

227
THELYPTERIDACEAE
Coryphopteris Holttum.
DESCRIPTION: [Applies to the American species
only] Rhizomes long-creeping, 1.5–3 mm wide; leaves
monomorphic, dying back in winter, often 1 cm or more
apart, fertile leaves often somewhat longer than sterile
leaves, 25–80 cm. Petioles 12–45 cm × 1–3 mm, at
base sparsely set with tan, ovate, glabrous scales; blades
lanceolate, 14–35 cm, proximal pinnae largest, or lowest
pair slightly smaller, blades tapered gradually to pinnatifid
apex; pinnae deeply pinnatifid to ca. 1 mm from costa,
3–10 × 1–2 cm; segments oblong, somewhat oblique,
entire; proximal pair of veins from adjacent segments
meeting margin above sinus; indument abaxially of
sparsely set hairs 0.2–0.4 mm primarily on costae, also
of yellowish short-stalked glands less than 0.1 mm, blade
tissue with numerous reddish to orangish, resinous,
shiny, sessile, hemispheric glands; blades adaxially with
scattered hairs on veins; sori round, medial; indusia tan,
often glandular; sporangia glabrous; x = 64.
DESCRIPCIÓN: [Se aplica a la especie americana
solamente] Rhizomas largamente rastreros, 1.5–3 mm
de ancho; hojas monomorfas, muriéndose en el invierno,
a menudo 1 cm o más distante; hojas fértiles a munudo
un pocito más largas que las estériles, 25–80 cm; pecíolos
12–45 cm × 1–3 mm, en la base con escamas esparsas
leonadas, ovadas, glabras; láminas lanceoladas, 14–35
cm, pinnas proximales las más largas, o par mas basal un
pocito mas pequeño; lámina gradualmente estrechada a
un ápice pinatifido; pinnas profundamente pinnatífidas
hasta ca. 1 mm from costa, 3–10 × 1–2 cm; segmentos
oblongos, un pocito obliquo, enteros; pares proximales de
venas de segmentos adjacentes llegando arriba del sinus;
indumento abaxialmente de pelos esparcidos, 0.2–0.4
mm principalmente en las costas, también de glándulas
corto-peciolulados amarillentos menos de 0.1 mm, tejido
laminar con muchas glándulas rojizas a naranjadas,
resinosas, sésiles, hemisféricas; láminas abaxialmente
con pelos dispersas por las venas; soros redondeados,
mediales; indusios leonados, a menudo glandulares;
esporangios glabros; x=64.

SIMILAR GENERA: Coryphopteris greatly resembles GÉNEROS PARECIDOS: Coryphopteris e parece

Amauropelta y Parathelypteris, y las distinciones mucho a Amauropelta y Parathelypteris, y las distinciones
morfologicas no son claras. morfologicas no son claras.

COMMENTS: Coryphopteris has ca. 50 species, mostly
Asian., with one species in the Americas: C. simulata
(Davenp.) S. E. Fawc., comb. ined. This species occurs
primarily in the northeastern United States in low-nutrient,
acidic soils–a habitat preference shared with many Old
World species of Coryphopteris. The species is closely
related to P. nipponica (Franch. & Sav.) Ching of Japan,
providing yet another example of a floristic relationship
between eastern North America and eastern Asia. The
morphological distinctions between Coryphopteris and it
sister genus Parathelypteris (H. Itô) Ching are not clear.The
genus name is derived from the Greek corypha, summit,
and pteris, fern. It alludes to the type species, which grows
only on high ridges.
COMENTARIOS: Coryphopteris consta de ca. 50
especies, mayormente asiáticas, con una especie en las
américas: C. simulata (Davenp.) S. E. Fawc., comb. ined.
Esta especie existe principalmente en los Estados Unidos
nororiental en suelos acídicos y de bajo nutrientes--un
habitat compartido con muchas especies de Coryphopteris
en el Viejo Mundo. Esta especie es emparentada
ceranamente a P. nipponica (Franch. & Sav.) Ching de
Japón, suministrado aun otro ejemplo de una relacion
floristica entre America del Norte oriental y Asia oriental.
Las distinctiones morfologicos entre Coryphopteris y su
genero hermano Parathelypteris (H. Itô) Ching no son
claras. El nombre genérico e deriva del griego corypha,
cumbre, y pteris, helechos. Alude a el especies tipo, que
existe solo en filas altas.

LITERATURE: Almeida, T. E., S. Hennequin, H. Schneider, A. R. Smith, J. A. N. Bastista, A. J. Ramalho, K. Proite &
A. Salino. 2016. Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of
neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688–700. Holttum, R.
E. 1976. Studies in the family Thelypteridaceae X. The genus Coryphopteris. Blumea 23:18–47. He,L. J. & X. C. Zhang. 2012.
Exploring generic delimitation within the fern family Thelypteridaceae. Molecular Phylogenetics and Evolution 65:757–764.

228
Figure 111. C–E. Coryphopteris simulata (Davenp.) S. Fawcett, comb. ined. (From Ogden, 1981.)

229
BLECHNACEAE
Cranfillia Gasper & V. A. O. Dittrich.
DESCRIPTION: Plants terrestrial; rhizomes short-
creeping to usually suberect or erect, stoloniferous or
not, the scales reddish brown to blackish, sometimes
bicolorous, lanceolate or oblong-attenuate, entire
scales; sterile and fertile leaves dimorphic; petioles
stramineous to dark brown, scaly proximally, often
abundantly hairy, or hairs sometimes sparse to absent;
blades linear-oblong to deltate, pinnate or deeply
pinnatifid proximally, pinnatifid distally, proximally truncate
or with gradually to subabruptly reduced pinnae; rachises
scaly and often pilose, the hairs uniseriate, septate,
sometimes sparse; buds absent or (in C. caudata) present;
aerophores absent; pinnae subpetiolulate proximally
or often becoming fully adnate distally, entire to crenate
or dentate along margins; veins free, hydathodous; sori
linear, indusia more or less entire, sometimes pubescent;
x = 33.
DESCRIPCIÓN: Plantas terrestres; rizomas cortamente
rastreros a usualmente suberectos o erectos,
estoloníferos o no, las escamas pardas a negruzcas, a veces
bicoloras, enteras; hojas estériles y fértiles dimorfas;
pecíolos estramíneos a pardo oscuros, escamosos
proximalmente, a menudo muy peludo, o pelos esparsas
a ausentes; láminas lineares-oblongas a deltadas,
pinnadas o profundamente pinnatífidas proximalmente,
pinnatífidas distalmente, proximalmente truncadas o
gradualmente a subabruptamente con pinnas reducidas;
ráquises escamosos y a menudo pilosos, los pelos
uniseriados, septados, a veces escasos; brotes ausentes
o (en C. caudata) presentes; aeróforos ausentes; pinnas
subpecioluladas hacia la base de la lámina, tornandose
adnadas distalmente en la lámina, enteras a crenatas
o dentadas; venas libres, con hidatodos; soros lineales;
indusios lineares, a veces pubescentes; x=33.

SIMILAR GENERA: Blechnum differs by stolons, GÉNEROS PARECIDOS: Blechnum difiere por
monomorphic sterile and leaves, and lack of hydathodes. estolones, hojas esteriles y fertiles monomofas y la falta
Austroblechnum differs by all pinnae adnate to the rachis. de hidatodos. Austroblechnum difiere por todas las pinnas
adnatas al raquis.

COMMENTS: Cranfillia has ca. 12 species, with three in
the Neotropics and the remainder in Oceania. It is a genus
of wet, shaded forest floors. The Neotropical species are
C. sampaioana (Brade) Gasper & V.A.O. Dittrich, C. caudata
(Baker) V.A.O. Dittrich & Gasper, and C. mucronata (Fée)
V.A.O. Dittrich & Gasper. Cranfillia is characterized by pilose
rachises and costae, with exception of C. sampaioana,
where hairs are very tiny and scarce. Cranfilia caudata is
distinctive by flagelliform, radicant leaf apices. A detailled
morphological and anatomical description of the species
(as Blechnum sprucei) was given by Gabriel y Galán et al.
(2008).The genus is sister to Austroblechnum + Blechnum.
The genus name honors Raymond Cranfill (ca. 1956– ),
American pteridologist who first studied Blechnaceae using
molecular phylogenetic methods.
COMENTARIOS: Cranfilia consta de 12 especies, con
tres en el Neotrópico y las demás en Oceania. Es un
género de sotobosques sombreados húmedos. Las
especies Neotropicales son C. sampaioana (Brade)
Gasper & V.A.O. Dittrich, C. caudata (Baker) V.A.O.
Dittrich & Gasper y C. mucronata (Fée) V.A.O. Dittrich
& Gasper. Cranfillia se caracteriza por ráquises y costas
pilosos, con la excepción de C. sampaioana donde los
pelos son muy pequeños y esparsas. Cranfilia caudata es
distinctiva por ápices foliares flageliformes y radicantes.
Una descripción morfológica y anatómica detallada de
la especie fue dada por Gabriel y Galán et al. (2008).
El género es hermano a Austroblechnum + Blechnum.
El nombre genérico honora a Raymond Cranfill (ca.
1956– ), pteridólogo estadounidense, quien estudió por
primera vez Blechnaceae usando métodos moleculares
filogenéticos.

LITERATURE: V. A. O. Dittrich, A. L. Gasper, A. Salino, R. Monteiro & A. L. de Gasper. 2018. The family Blechnaceae
(Polypodiopsida) in Brazil: key to the genera and taxonomic treatment of Austroblechnum, Cranfillia, Lomaridium, Neoblechnum
and Telmatoblechnum for southern and southeastern Brazil. Phytotaxa 303: 1–33. Gabriel y Galán, J. M., L. M. Passarelli,
C. Prada & C. H. Rolleri. 2008. Sporophyte morphology and gametophyte development of the fern Blechnum sprucei
(Pteridophyta: Blechnaceae). Revista de Biologia Tropical Vol. 56(4): 2027–2040 Gasper, A. L. & V. A. O. Dittrich, A. R.
Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and
combinations. Phytotaxa 275: 191–227. .

230
Figure 112. C–E. Cranfillia caudata (From Gabriel y Galan et al., 2008.)

231
HYMENOPHYLLACEAE
Crepidomanes (C. Presl) C. Presl subgen. Crepidomanes
DESCRIPTION: Mostly epiphytes; roots absent, but
root-like short shoots often present; rhizomes 0.1–
0.8 mm wide, long-creeping, frequently branched,
densely pubescent, the hairs dark brown or (in
ours) black; sterile and fertile monomorphic; petioles
to 8 cm long, pubescent basally and along margin of
wing, the hairs like those on the rhizomes; sterile
laminae 1-cell thick between the veins, 1-pinnate to 4-pin-
nate, the apices not proliferous; veins anadromous, free,
false veins often present; receptacles exert; spores green,
tetrahaedral-globose; x=36.
DESCRIPCIÓN: Mayormente epífitas; raíces ausentes,
pero vástagos cortos que parecen a raíces a menudo
presentes; rizomas 0.1–0.8 mm de ancho, largamente
rastreros, ramificados con frecuencia, densamente
pubescentes, los pelos pardo-oscuro o (en nuestra)
neguzcos; hojas estériles y fértiles monomorfas; pecíolos
hasta 8 cm de largo; pubescentes en la base y por el
margen de la ala del pecíolo, los pelos como los del
rizoma; láminas estériles una célula de grosor entre las
venas, 1-pinnadas a 4-pinnadas, los ápices no prolíferos;
venas anádromas, libres, falsas venas a menudo pre-
sentes; receptáculos exertos; esporas verdes, tetrahédr
cas-globosas; x=36.

SIMILAR GENERA: Polyphlebium differs by reddish
brown rhizome hairs, presence of roots, glabrous peti-
ole bases and petiole wing margins, and lack of folds in
the lamina parallel the veins. Didymoglossum differs by the
presence of false veins in the laminae and smaller leaves
(< 6 cm).
GÉNEROS PARECIDOS: Polyphlebium difiere por pe-
los de los rizomas pardo-rojizos y presencia de raíces,
bases de los pecíolos y márgenes de las alas de los pecío-
los glabros, y falta de pliegues en la lámina paralelos a las
venas. Didymoglossum difiere por la presencia de venas
falsas en las láminas y hojas más cortas (< 6 cm).

COMMENTS: Crepidomanes consists of about 30 spe- COMENTARIOS: Crepidomanes consta de casi 30

cies of wet forests. All of its species are paleotropical ex-
cept for C. pyxidiferum (L.) Dubuisson & Ebihara, which is
widespread in the Neotropics. It is distinctive by bearing
folds in the lamina parallel to the veins. Perhaps it is not
distinct from C. melanotrichum of Africa and Madagascar. lámina paralelas a las veins. Tal vez es lo mismo como C.
It resembles Polyphlebium diaphanum and P. hymenophyl-
loides and is often misidentified as these species. Crepi-
domanes pyxidiferum can be distinguished from these
species by black (not red-brown) rhizome hairs, petiole
bases pubescent on all sizes (the hairs like those of the
rhizome), and wings of the petioles pubescent on mar-
gins. An unusual character it shares with Didymoglossum
and Polyphlebium is rhizomes branched to produce short
shoots resembling roots (Schneider, 2000). The genus
name is derived from the Greek crepis, slipper, and manes,
cup. It alludes to the shape of the sorus.
especies de bosques húmedos. Todos sus especies son
paleotropicales con excepción de C. pyxidiferum (L.)
Dubuisson & Ebihara, la cual está muy difundida en el
Neotrópico. Es distinctiva por llevando pliegues en la
melanotrichum de África y Madagascar. Se asemeja a Poly-
phlebium diaphanum y P. hymenophylloides, y es a menu-
do mal identificada con estas especies. Crepidomanes
pyxidiferum se puede distinguir por los pelos negruzcos
de los rizomas (no pardo-rojizos), bases de los pecíolos
pubescentes en todos lados (los pelos como los en el
rizoma) y alas de los pecíolos pubescentes por el margen.
Un carácter inusual que se comparta con Didymoglos-
sum y Polyphlebium es rizomas ramificados produciendo
vástagos cortos que parecen a raíces (Schneider, 2000).
El nombre genérico se deriva del griego crepis, zapatilla, y
manes, copa. Se refiere a la forma del soro.

LITERATURE: Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of Hymeno-
phyllaceae. Blumea 51: 1–60. Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global molecular
phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of
the Linnean Society 155: 1–27. Schneider, H. 2000. Morphology and anatomy of roots in the filmy fern tribe Trichomaneae H.
Schneider (Hymenophyllaceae, Filicatae) and the evolution of rootless taxa. Botanical Journal of the Linnean Society 132: 29–46.
Slosson, M. 1916. Notes on Trichomanes—I. The identity of Trichomanes pyxidiferum L. Bulletin of the Torrey Botanical Club
42: 651–658.

232
Figure 113. Crepidomanes pyxidiferum. Note dark hairs at petiole base and along wing and folds in the lamina parallel to the
segment midribs. (From Mickel & Smith, 2004.)

233
PTERIDACEAE
Cryptogramma R. Br.
DESCRIPTION: Saxicolous; stems short-creeping ,de-
cumbent to erect, scaly; sterile and fertile leaves di-
morphic, with the fertile leaves longer-petiolate, taller and
more erect than the sterile; sterile leaves 5-20 cm long;
petioles grooved on the upper surface, with 1 vascular
bundle; laminae 2-4-pinnate, glabrous on both surfaces,
not farinose beneath, scales rare or absent, the apex pin-
natifid, tapered; ultimate segments sessile or short-stalked;
veins free; false indusia present; sporangia borne along
the veins, often intermixed with farina-producing glands,
64-spored; spores yellow, trilete, verrucate, without
equatorial flange; x=30.
DESCRIPCIÓN: Saxícolas; tallos cortamente rastreros,
decumbentes a erectos, escamosos; hojas estériles y
fértiles dimorfas, con la fértil más largamente pecioladas,
más largas y erectas que las estériles; hojas estériles 5-20
cm de largo; pecíolos surcados en la superficie superior,
con 1 haz vascular; láminas 2-4-pinnado, glabras en am-
bas superficies, no farinosas en el envés, escamas raras
o ausentes, el ápice pinnatífido, reducido gradualmente;
segmentos últimos sésiles o cortamente peciolulados;
nervaduras libres; inducíos falsos presentes; esporan-
gios nacen a lo largo de las venas, a menudo mezcladas
con glándulas farinosas, con 64 esporas; esporas amarillas,
triletes, verrucatas, sin un borde ecuatorial; x=30.

SIMILAR GENERA: Cystopteris differs by round sori,
presence of a true indusium (not marginal), and two vas-
cular bundles in the petiole. Anogramma differs by sori not
protected by a reflexed marginal indusium.
GÉNEROS PARECIDOS: Cystopteris difiere por soros
redondeados y un indusio verdadero (no marginal) y dos
haces vasculares en el pecíolo. Anogramma difiere por no
tener los soros no protegidos por un indusio marginal
reflexo.

COMMENTS: Cryptogramma is circumboreal with one COMMENTARIOS: Cryptogramma es circumboreal

species in southern South America. It consists of 9 spe- con una especie en el sur de América del Sur. Consta
cies, 5 of which occur in the Americas. All species are sax- de 11 especies, 5 de las cuales existen en las Américas.
icolous. Cryptogramma differs from most genera of Pteri- Todas las especies son saxícolas. Cryptogramma difiere de
daceae by dimorphic leaves, false indusial, and verrucose la mayoría de géneros de Pteridaceae por hojas dimór-
spores that lack an equatorial flange. It is closely related to fas, indusios falsos y esporas verrucosas que carecen de
Llavea and the paleotropical Coniogramme. Two sections un reborde ecuatorial. Es cercanamente relacionado a
are recognized: sect. Homopteris (Rupr.) C. Chr., found in Llavea y el paleotropical Coniogramme. Se reconocen dos
northern North America, Asia, and extreme northeast- secciones: secc. Homopteris (Rupr.) C. Chr. de regiones
ern Europe. It has creeping rhizomes, membranous leaves, septentrionales de América del Norte y Asia y noreste
and occurrence on calcareous rocks. It consists of one Europa. Tiene rizomas rastreros, hojas membranáceas, y
species, C. stelleri (S. G. Gmel.) Prantl. Section Cryptogram- ocurre en rocas calcáreas. Consta de una sola especie,
ma is found in boreal and temperate North America and C. stelleri (S. G. Gmel.) Prantl. La secc. Cryptogramma se
Eurasia, with one species (C. fumariifolia (Baker) H. Christ) encuentra en las zonas boreales y templadas de América
in southern Chile and Argentina. The section consists of del Norte y Eurasia, con una especie (C. fumariifolia (Bak-
8 species characterized by erect rhizomes, thicker leaves, er) H. Christ) en el sur de Chile y Argentina. La sección
and occurrence on acidic rocks. Cryptogramma is derived consta de 8 especies y se caracteriza por rizomas erectos,
from the Greek cryptos, hidden, and gramme, line, refer- hojas más gruesas, y una existencia en rocas ácidas. Cryp-
ring to the lines of sori along the veins that are hidden by togramma se deriva del griego cryptos, ocultos, y gramme,
the false indusium. línea, referiéndose a las líneas de soros por las venas que
son ocultos por el indusio falso.
LITERATURE; Alverson, E. R. 1989. Cryptogramma cascadensis, a new parsely fern from western North America. American
Fern Journal 79: 95–102. Alverson, E. R. 1995. Cryptogramma. Pages 137–139. In: Flora of North America Editorial Commit-
tee. Flora of North America, Volume2, Pteridophytes and Gymnosperms. Oxford Univ. Press, New York. Metzgar, J. S., E. R.
Alverson, S. Chen, A. V. Vaganov & S. M. Ickert-Bond. 2013. Diversification and reticulation in the circumboreal fern genus
Cryptogramma. Molecular Phylogenetics and Evolution 67: 589–599. Zhang, G-M. & X-C. Zhang. 2003. Taxonomic revision of
the genus Cryptogramma R. Br. from China. Acta Phytotaxonomica Sinica 41: 475–482.

234
Figure 114. A, B. Cryptogramma (sect. Cryptogramma) acrostichoides. C, D. C. (sect. Homopteris) stelleri. (Original, ©R. C.
Moran, 2011.)

235
DRYOPTERIDACEAE
Ctenitis (C. Chr.) C. Chr.
DESCRIPTION: Terrestrial or rarely rupicolous; rhi-
zomes erect to obliquely ascending; petioles with more
than three vascular bundles; leaves monomorphous; lam-
inae 1-pinnate-pinnatifid to 4-pinnate-pinnatifid, cata-
dromic, often glandular, the glands cylindrical, appressed;
rachis and costae flat to rounded (not grooved) on
the upper surface, puberulent, often scaly beneath,
the scales typically clathrate; veins free, the tips narrow
(not hydathodous), ending at the margin; sori round; in-
dusia present or absent; sporangial stalks with unicellular
cylindrical glands; x=41.
DESCRIPCIÓN: Terrestres o raramente rupícolas;
pecíolos con más de tres haces vasculares; rizomas erec-
tos a oblicuamente ascendentes; láminas 1-pinnado-pin-
natífidas a 4-pinnado-pinnatífidas, catádromas, a menudo
glandulosas, las glándulas cilíndricas, adpresas; raquis y
costas aplanados a redondeados (no surcados) en la
superficie superior, puberulentos, a menudo escamo-
sos abaxialmente, las escamas típicamente clatradas;
nervaduras libres, las puntas delgadas (no como un hid-
atodo), terminando en el margen; soros redondeados;
indusios presentes o ausentes; pedículos esporangiales
con glandulos unicelulares, cilíndricos; x=41.

SIMILAR GENERA: Megalastrum differs by strigose,
whitish hairs on the upper surface of the axes, and hy-
dathodes on the upper surface of the lamina. Lastreopsis
has a completely different kind of architecture on the up-
per surface of the rachis and costas.
GÉNEROS PARECIDOS: Megalastrum difiere por pe-
los estrigosos, blanquecinos en la superficie superior de
los ejes y hidatodos en la superficie superior de la lámina.
Lastreopsis tiene un tipo diferente de arquitectura en el
haz de la los ráquises y costas.

COMMENTS: Ctenitis is pantropical and contains about
125 species, of which about 50 are Neotropical. It grows
on wet forest floors mostly from 0–2000 m. It appears
to contain two species groups: one with laminae lanceo-
late and 1-pinnate-pinnatifid (e.g., C. subincisa (Langsd. &
Fischer) Ching), the other with laminae deltate to ovate
and 2-pinnate-pinnatifid or more finely divided (e.g., C. ex-
celsa (Desv.) Proctor). Ctenitis has distinctive hairs, called
appropriately “Ctenitis hairs,” on the upper surface of its
lamina axes. These hairs are short (0.2–0.5 mm long), of-
ten reddish, several-celled, and often appear jointed be-
cause the cells twist at right angles to each other upon
drying. These hairs are useful in identifying the genus, but
they are also found in other genera such as Cyclopeltis,
Lastreopsis, and Triplophyllum. Ctenitis is derived from the
Greek kteis, comb, and refers to the pectinate pinnae of
certain species.
COMENTARIOS: Ctenitis es pantropical y consta de
alrededor de 125 especies, de las cuales casi 50 son Neo-
tropicales. Crece sobre los suelos de bosques húmedos
principalmente de 0–2000 m. Parece contener dos gru-
pos de especies: uno con láminas lanceoladas y 1-pinna-
do-pinnatífidas (e.g., C. subincisa (Langsd. & Fischer) Ching),
el otro con láminas deltadas a ovatas y 2-pinnado-pinnatí-
fidas (e.g., C. excelsa (Desv.) Proctor). Ctenitis tiene pelos
distinctivos, llamados apropriadamente “pelos de Ctenitis,
en la superficie superior de los ejes de las láminas. Estos
pelos son cortos (0.2–0.5 mm de largo), a menudo ro-
jizos, multicelulares, y a menudo aparecen artículados a
causa de las células torcen en angulos rectos a cada uno
cuando se secan. Estos pelos son útiles en identificar el
género, pero ocurren en otros géneros como Cyclopel-
tis, Lastreopsis y Triplophyllum. Ctenitis se deriva del griego
kteis, peine, y se refiere a las pinnas pectinadas a ciertas
especies.

LITERATURE: Christensen, C. 1913. A monograph of the genus Dryopteris. Part I. The tropical American pinnatifid-bipin-
natifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 7, 10: 55–282. Christensen, C. 1920. A mono-
graph of the genus Dryopteris. Part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr.,
Naturvidensk. Math. Afd. ser. 8, 6: 1–132. Hennequin, S. et al. 2017. Global phylogeny and biogeography of the fern genus
Ctenitis (Dryopteridaceae), with a focus on the Indian Ocean region. Molecular Phylogenetics and Evolution doi: http://dx.doi.
org/10.1016/j.ympev.2017.04.012. Viveros, R. S., G. Rouhan & A. Salino. 2018. A taxonomic monograph of the fern genus
Ctenitis (Dryopteridaceae) in South America. Phytotaxa 385(1): 1–83.

236
Figure 115. A–C. Ctenitis submarginalis. D, E. C. nigrovenia. F–H. C. melanosticta. J–M. C. excelsa. (From Mickel & Smith,
2004.)

237
CULCITACEAE
Culcita C. Presl
DESCRIPTION: Terrestrial; rhizome subterranean,
stout, short, not forming a trunk, the apex with dense,
long, golden hairs. Leaves monomorphic; stipes hairy, with
an omega-shaped vascular bundle; blades 4- or 5-pin-
nate, deltate, glabrous to slightly hairy, coriaceous; rachis-
es and costae grooved on the upper surface; veins
free; sori marginal; indusia bivalvate, the lower valve
fitting inside the outer (upper), outer valve green like the
laminar tissue, inner valve whitish to tan, differentiated;
spores trilete, nongreen; x = 66.
DESCRIPCIÓN: Terrestres; rizoma subterráneo, cor-
to y grueso, no formando un tronco, ápice con pelos
densos, largos, y dorados. Hojas monomorfas; pecíolos
pelados, con una haz vascular en forma de una omega;
láminas 4- o 5-pinnadas, deltadas, glabras a ligeramente
peludas, coriáceas; ráquises y costas surcados en el
haz; venas libres; soros marginales; indusios bivalvos,
la valva inferior cabe adentro del superior, la valva supe-
rior verde como el tejido láminar, la inferior blanceciña a
leonada, diferenciada; esporas triletes, no verdes; x = 66.

SIMILAR GENERA: Dicksonia and Cibotium differ by
arborescent trunks, laminae to only 3-pinnate-pinnatifid,
and non-grooved rachises and costae adaxially. Dicksonia
further differs by laminae strongly tapered toward the
base. Dennsteadtia also has marginal sori, but the indusia
are cup-shaped, not bivalvate. It lacks long, golden hairs
on the rhizome and petiole bases.
GÉNEROS PARECIDOS: Dicksonia y Cibotium difieren
por troncos aborescentes, láminas hasta 3-pinadas-pin-
natífidas, y ráquises y costas no surcados adaxialmente.
Dennsteadtia tiene también los soros marginales, pero los
indusios son en forma de una copa, no bivalvos. Carece
de pelos largos dorados en el rizoma y bases de los
pecíolos.

COMMENTS: Culcita consists of two species: C. coniifo-
lia (Hook.) Maxon of the Neotropics, and C. macrocarpa
C. Presl of Spain, the Azores, Madeira, and Tenerife The
neotropical species grows above 2000 m in cloud forests,
generally in semi-open places. Previously, Calochleana, a
genus of 5 species from southeastern Asia and the South
Pacific, was classified as a subgenus of Culcita, but molecu-
lar phylogenetic studies have resolved it in the Dicksonia-
ceae (Korall et al., 2006). Culcita as been resolved sister to
Plagiogyria (Plagiogyriaceae), with which it seems to have
little in common. The name Culcita means bed or cushion
is Latin. It might refer to the arched, cushion-like indusia,
or perhaps the soft, hairy rhizome apex, or the use of the
hairs to stuff pillows.
COMENTARIOS: Culcita consta de dos especies: C.
coniifolia (Hook.) Maxon en el Neotrópico, y C. macrocar-
pa C. Presl en España, los Azores, Madeira, y Tenerife. La
especie neotropical existe arriba de 2000 m en bosques
nubosos, generalmente en lugares semi-abiertos. Previa-
mente, Calochleana, un género de 5 especies del sureste
de Asia y el Sur Pacífico, fue clasificado como un subgéne-
ro de Culcita, pero estudios filogenéticos moleculares o
resuelvan en Dicksoniaceae (Korall et al., 2006) . Culcita
ha sido resuelto hermano a Plagiogyria (Plagiogyriaceae),
con que parece tener poco en común morfológicamente
o anatómicamente. El nombre Culcita significa cama o
cojín en Latín. Tal vez se refiere a los inducíos arqueados,
como cojín, o el ápice del rizoma suave y peludo, o el uso
de los pelos en almohadas.

LITERATURE: Gastony, G. J. 1981. Spore morphology in the Dicksoniaceae. I. The genera Cystodium, Thyrsopteris, and Cul-
cita. American Journal of Botany 68: 808–819. Korall, P. K. M. Pryer, J. S. Metzgar, H. Schneider & D. S. Conant. 2006.
Tree ferns: monophyletic groups and their relationships as revealed by four protein-coding platid loci. Molecular Phylogenetics
and Evolution 39: 830–845. Maxon, W. R. 1922.The genus Culcita. Journal of the Washington Academy of Science 12: 454–460.
Sen, U. 1967. Anatomy of Culcita macrocarpa. Canadian Journal of Botany 46: 43–46. White, R. A. & M. D. Turner.
1988. Calochleana, a new genus of dicksonioid ferns. American Fern Journal 78: 86: 95.

238
Figure 116. A–D. Culcita coniifolia. (From Mickel & Smith, 2004.)

239
CYATHEACEAE
Cyathea J. E. Sm.
DESCRIPTION: Usually terrestrial; stems typically ar-
borescent; petioles scales bicolorous or concolorous,
lacking long black setae, the borders with cells of
different size, shape, and orientation from those
in the medial part; sterile and fertile leaves 1–4(–6)
m long, monomorphous; laminae 1-pinnate to 4-pinnate
(most species 2-pinnate-pinnatisect), the apex pinnatifid
or rarely conform; veins free or anastomosing; sori round,
receptacles elevated, usually paraphysate; indusia present
or absent, globose or cup-shaped or scalelike; spores tet-
rahedral, 64 per sporangium; x=69.
DESCRIPCIÓN: Usualmente terrestres; tallos típica-
mente arborescentes; escamas de los pecíolos bicolor-
as o concoloras, sin setas largas negras, los bordes
con células de tamaño, forma y orientación de las
en la parte medial; hojas estériles y fértiles 1–4(–6) m
de largo, monomorfas; láminas 1–pinnadas a 4-pinnadas
(la mayoría de las especies son 2-pinnada-pinnatisectas),
el ápice pinnatífido o raras veces conforme; nervaduras
libres o anastomosadas; soros redondos, receptáculos el-
evados, usualmente con paráfises; indusios presentes o
ausentes, globosos o con forma de copa o escamiformes;
esporas tetrahédricas, 64 por esporangio; x=69.

`SIMILAR GENERA: See Alsophila and Sphaeropteris for GÉNEROS PARECIDOS: Véase Alsophila y Sphaerop-
comparison. Dicksonia differs by marginal sori and hairs teris para comparación. Dicksonia difiere por soros mar-
instead of scales on the trunks and petioles. ginales y pelos en lugar de escamas sobre los troncos y
los pecíolos.
COMMENTS: Cyathea is pantropical and contains about COMENTARIOS: Cyathea es pantropical y consta de
235 species in the Neotropics. It is sister to Alsophila, the alrededor de 235 especies en el Neotrópico. Es hermana
two sharing the synapomorphy of marginate petiole scales a Alsophila, los dos compartiéndo la sinapomorfia de es-
(i.e., scales with cells of the borders differentiated from camas peciolares marginadas (i.e., escamas con las células
those of the medial portions. Cnemidaria, long recognized por los bordes diferentes de las de la parte mediana de
as a genus, is nested within Cyathea. It is herein treated as la escama).Cnemidaria, reconocido por mucho tiempo
Cyathea, Cnemidaria group (which see). Trichipteris differs como género, es anidado dentro de Cyathea. Se trata aquí
from Cyathea only by the absence of indusia, a structure como el grupo de Cyathea, Cnemidaria (vease). Trichipteris
that has been lost independently several times in Cyathea. difiere de Cyathea sólo por la ausencia del indusio, una
Lehnert revised the species of Cyathea with globose in- estructura que ha sido perdido varias veces en Cyathea.
dusia and white petiole scurf (Lehnert, 2008) those with Lehnert revisó las especies de Cyathea con indusios glo-
globose indusia and dark petiole scurf (Lehnert, 2009). bosos y caspilla peciolar blanquecina (Lehnert, 2008) y las
The genus name is derived from the Greek kyathos, wine con indusios globosos y caspilla peciolar oscura (Lehnert,
cup. It alludes to the shape of the indusium. 2009). El nombre genérico se deriva del griego kyathos,
copa para vino. Alude a la forma del indusio.

LITERATURE: Barrington, D. S. 1978. A revision of the genus Trichipteris. Contributions of the Gray Herbarium of Harvard
University 208: 3–91. Lehnert, M. 2006. The Cyatheaceae and Dicksoniaceae (Pteridophyta) of Bolivia. Brittonia 58: 229–244.
Lehnert, M. 2008. On the identification of Cyathea pallescens (Sodiro) Domin (Cyatheaceae): typifications, reinstatements and
new descriptions of common Neotropical tree ferns. Botanical Journal of the Linnean Society 158: 621–649. Lehnert, M. 2009.
Resolving the Cyathea caracasana complex (Polypodiopsida: Cyatheaceae). Beiträge zur Naturkunde A, Neue Serie 2: 409–445.
Lehnert, M. 2011. Species of Cyathea in America related to the western Pacific species C. decurrens. Phytotaxa 26: 39–59.
Lehnert, M. 2011. The Cyatheaceae (Polypodiopsida) of Peru. Brittonia 63: 11–45. Lehnert, M. 2012. A synopsis of the species
of Cyathea (Cyatheaceae – Polypodiopsida) with pinnate to pinnate-pinnatifid fronds. Phytotaxa 61: 17–36. Lehnert, M. 2015. A
synopsis of the exindusiate species of Cyathea (Cyatheaceae-Polypodiopsida) with bipinnate-pinnatifid or more complex fronds,
with a revision of the C. lasiosora complex. Phytotaxa 243(1): 1–53. Lellinger, D. B. 1988. The disposition of Trichopteris (Cy-
atheaceae). American Fern Journal 78: 90–94. Riba, R. 1969 [“1967”]. Revisión monográfica del complejo Alsophila swartziana
Mart. (Cyatheaeceae). Anales del Instituto de Biología de la Universidad Nacional de México, sér. bot. 38: 61–100. Tryon, R. M.
1976. A revision of the genus Cyathea. Contributions from the Gray Herbarium of Harvard University 206: 19–98. Windisch, P.
G. 1977. Synopsis of the genus Sphaeropteris (Cyatheaceae) with a revision of the neotropical exindusiate species. Botanische
Jahrbücher für Systematik 98: 176–198.

240
Figure 117. Cyathea caracasana var. boliviensis. A. Pinna. B. Sphaeropteroid (globose) indusia. C. Marginate petiole scale; note
cells not all of same shape as in Sphaeropteris. (From Tryon & Stolze, 1989.)

241
CYATHEACEAE
Cyathea (Cnemidaria C. Presl, clade)
DESCRIPTION: Terrestrial; stems non-arborescent,
or if so (rarely), up to 1.5 m tall; sterile and fertile leaves
1-3.5 m long, monomorphous; petioles smooth or spiny,
scaly, the scales bicolorous with a dark brown center
and white margins, lacking setae; laminae 1-pinnate to
1-pinnate-pinnatisect (rarely 2-pinnate), with con-
form or pinnatifid apices; pinnae equilateral, sessile or
short-stalked; veins free or anastomosing; rachis and
costae glabrous adaxially (except in C. suprastrigosa
R.C. Moran); sori round, the receptacle elevated, subglo-
bose; indusia scalelike, attached to the base of the recep-
tacle, either semicircular and attached on the costular side
of the receptacle, or circular and completely encircling
the receptacle; spores trilete, with three large equa-
torial pores and often smaller ones; x=69.
SIMILAR GENERA: Cyathea (s.s.) may have laminae
1-pinnate to 1-pinnate-pinnatisect; however, the axes are
always pubescent adaxially with thick strigose hairs. Most
species of Cyathea differ by arborescent trunks.
DESCRIPCIÓN: Terrestres; tallos no arborescentes,
o si (raras veces), hasta 1.5 m de alto; hojas estériles y
fértiles 1-3.5 m de largo, monomorfas; pecíolos lisos o
espinosos, escamosos, las escamas bicoloras con un cen-
tro oscuro y márgenes blanquecinos, carecente de setas;
láminas 1-pinnadas a 1-pinnada-pinnatisectas (raras
veces 2-pinnadas), con los ápices conformes o pinnatí-
fidos; pinnas equiláteras, sésiles o cortamente peciolula-
das; nervaduras libres o anastomosadas; raquis y costas
glabros adaxialmente (con excepción de C. suprastrigo-
sa R.C. Moran); soros redondos, con receptáculos eleva-
dos, subglobosos; indusios escamiformes, únidos a la base
del receptáculo, semicirculares y únidos al lado costular
del receptáculo, o circular y completamente rodeando
el receptáculo; esporas tetrahédricas-globosas, con
tres poros grandes ecuatoriales y a menudo unos
más pequeños; x=69.
GÉNEROS PARECIDOS: Cyathea (s.s.) peude tener
láminas 1-pinnadas a 1-pinnada-pinnatisectas; no obstan-
te, los ejes siempre son pubescentes adaxialmente con
pelos gruesos estrigosos. La mayoría de especies de Cy-
athea difiere por troncos arborescentes.

COMMENTS: The Cnemidaria clade is entirely neotrop-
ical and contains about 27 species. It inhabits wet forests
from low to middle elevations. Cnemidaria is monophy-
letic, nested within Cyathea (Korall et al., 2006).Besides
the characteristics in boldface in the description, many
species of Cnemidaria have anastomosing veins, conform
or subconform lamina apices, and short stems. It is rarely
arborescent. When a trunk is present, it is usually less than
one meter tall.The genus name is derived from the Greek
knemis, spoke of a wheel + aris, resembling. The lowest
veins radiate like spokes of a wheel.
COMENTARIOS: El clado de Cnemidaria es completa-
mente neotropical y consta de casi 27 especies. Inhabita
bosques húmedos de elevaciones bajas a medias. Cnemi-
daria es monofiletico y anidado dentro de Cyathea (Korall
et al., 2006). Además de las características en negritas da-
das la descripción, muchas especies de Cnemidaria tienen
venas anastomosadas, ápices de las hojas conformes o
subconformes y tallos cortos. Raras veces es arborescen-
te, pero cuando si, el tronco es usualmente menos de un
metro de altura. El nombre genérico se deriva del griego
knemis, un rayo de una rueda + aris, semejante. Las venas
basales se asemejan a los rayos de una rueda.

LITERATURE: Korall, P., K. M. Pryer, J. S. Metzgar, H. Scheider & D. S. Conant. 2006. Tree ferns: monophyletic groups
and their relationships as revealed by four protein-coding plastid loci. Molecular Phylogenetics & Evolution 39: 830–845. Leh-
nert, M. 2011. A synopsis of the species of Cyathea (Cyatheaceae–Polypodiopsida) with pinnate to pinnate-pinnatifid fronds.
Phytotaxa 61: 17–36. Lehnert, M. & A. Weigand. 2017. A synopsis of the Neotropical species of Cyathea (Cyatheaceae;
Polypodiopsida) with bipinnate fronds. Brittonia 69: 71–90. Stolze, R. G. 1974. A taxonomic revision of the genus Cnemidaria
(Cyatheaceae). Fieldiana, Botany 37: 1–98.

242
Figure 118. A–E. Cyathea (Cnemidaria) grandifolia. F–H. C. horrida. (Original, ©Moran & Mickel, 2010.)

243
CYATHEACEAE
Cyathea (Hymenophyllopsis Goebel, clade)
DESCRIPTION: Terrestrial or rupestral; rhizomes
ascending, short-creeping, or erect, scaly; leaves 10–30(–
40) cm long, monomorphic, fasciculate; petioles terete
or sulcate, brown to black; blades 1-pinnate-pinnatifid
to 3-pinnate-pinnatifid, several cells thick between
the veins, lacking stomata, glabrous to sparsely scaly
on both surfaces; pinnules or ultimate segments narrow,
often linear; veins usually one per ultimate segment;
sori marginal or submarginal; indusium bivalved,
entire, repand or broadly lacerate; paraphyses absent;
spores tetrahedral-globose, trilete, brown; x=?
DESCRIPCIÓN: Terrestres o rupestres; rizomas
ascendentes, cortamente rastreros, o erectos, escamosos;
hojas 10–30(–40) cm de longitud, monomorfas,
fasciculatadas; pecíolos teretes o sulcados, pardos o
negros; láminas 1-pinnado-pinnatífidas, de varias
cellulas de grosor entre las venas, careciendo de
estomas, glabras en ambas superficies; pínnulas
o segmentos últimos estrechos, a menudo lineares;
nervaduras usualmente una por segmento último; soros
marginales o submarginales; indusios bivalvados,
enteros, repandos o anchamente lacerados; parafises
ausentes; esporas tetrahédricas-globosas, triletes; x=?

SIMILAR GENERA: Hymenophyllum differs by GÉNEROS PARECIDOS: Hymenophyllum difiere por

pubescent rhizomes (not scaly), laminae only one cell rizomas pubescentes (no escamosos), láminas solamente
layer thick between the veins, and green spores. With low una célula de grosor entre las venas, y esporas verdes.
magnification the cells of its laminae are evident, whereas Con magnificación baja las células de sus láminas son
those of Hymenophyllopsis are obscure. evidentes, mientras las de Hymenophyllopsis son obscuras.

COMMENTS: Hymenophyllopsis consists of 8 species, all COMENTARIOS: Hymenophyllopsis consta de 8

endemic to the Guyana Shield of southern Venezuela and
adjacent Guyana and Brazil.The species are usually epipetric
on sandstone cliff faces and ledges or terrestrial at the
base of boulders from 1100–2700 m. Hymenophyllopsis
has recently been shown to be nested within Cyathea,
and its species will eventually be transferred to that genus.
Hymenophyllum + the Greek suffix, opsis, like. The plants
resemble Hymenophyllum by their small size, thin laminae,
absence of stomata, and marginal sori.
especies, todas endémicas al Escudo Guyanés en
Venezuela del sur y partes adyacentes de Guyana y Brasil.
Las especies son usualmente epipétricas sobre farallones
de piedra arenosa o terrestres en la base de peñas desde
1100–2700 m. Hymenophyllopsis se ha demonstrado
ser anidado dentro de Cyathea, e sus epecies deben
ser transferido eventualmente a ese género para.
Hymenophyllum + el sufijo griego, opsis, similar. Las plantas
se asemejan Hymenophyllum por su tamaño pequeño,
láminas delgadas, ausencia de estomas, y soros marginales.

LITERATURE: Korall, P. K. M. Pryer, J. S. Metzgar, H. Schneider & D. S. Conant. 2006. Tree ferns: monophyletic groups
and their relationships as revealed by four protein-coding plastid loci. Molecular Phylogenetics and Evolution 39: 830–845.
Lellinger, D. B. 1984. Hymenophyllopsidaceae (Filicales). In: B. Maguire and collaborators, The Botany of the Guayana Highland
Part XII. Memoirs of the New York Botanical Garden 38: 2–9. Wolf, P. G., S. D. Sipes, M. R. White, M. L. Martines, K. M.
Pryer, A. R. Smith, & K. Ueda. 1999. Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and
Lophosoriaceae: evidence from rbcL nucleotide sequences. Plant Systematics and Evolution 219: 263–270.

244
Figure 119. Cyathea (Hymenophyllopsis) hymenophylloides. (Original, ©R. Moran, 2016.)

245
DRYOPTERIDACEAE
Cyclodium C. Presl
DESCRIPTION: Terrestrial, hemiepiphytic, or rheoph-
ytic; rhizomes creeping, scaly; leaves monomorphic or
slightly dimorphic; petioles with 4 or more vascular bun-
dles; laminae pinnate to 2-pinnate-pinnatifid, glabrous or
nearly so, anadromous, chartaceous to coriaceious, the
apex pinnatifid, abruptly tapered or similar in form to the
lateral pinnae; longest segments on the acroscopic side
of the pinnae; rachis and costae grooved adaxially, the
grooves more or less continuous with those of the lower
order, puberulent within the groove; costae abaxially with
uniseriate trichomes, reddish or brownish, appressed;
veins free or anastomosing, ending at the margins, not en-
larged; sori round; indusium round, peltate or attached
at a narrow sinus, often deciduous before the sporangia
mature; spores monolete; x=41.
DESCRIPCIÓN: Terrestres, hemiepífitas o reofíticas; ri-
zomas reptantes, escamosos; hojas monomorfas a algo
dimorfas; pecíolos con 4 o más haces vasculares; lámina
pinnada a 2-pinnado-pinnatífida, glabra o casi así, anádro-
ma, cartácea a coriácea, el ápice abruptamente reducido
o pinnatífido similar en forma a las pinnas laterales; seg-
mentos más largos en el lado acroscópico de las pinnas;
raquis y costas sulcados adaxialmente, los surcos más o
menos confluentes de un eje al siguiente, pelosos por
dentro; costas abaxialmente con tricomas uniseriados,
rojizos oparduscos, adpresos; nervaduras libres o anas-
tomosadas, terminándose antes del margen, no hichados;
soros redondeados; indusio redondeado, peltado o
unido a un seno estrecho, a menudo deciduo antes de la
madurez del esporangio; esporas monoletes; x=41.

SIMILAR GENERA: Arachniodes differs by more finely
divided blades. Polybotrya differs by (mostly) climbing rhi-
zomes and strongly dimorphic sterile and fertile leaves.
Dryopteris differs by erect or decument rhizomes. Stig-
matopteris differs by thinner blades with internal punctu-
ate glands, and veins ending before the margins in clavate
hydathodes.
COMMENTS: Cyclodium is entirely neotropical and con-
tains about 10 species, these mostly in the Guianas. It is
primarily a lowland genus, growing only up to 1500 m.
It forms a clade with other dryopteroid ferns that have
creeping rhizomes and (often) dimorphic sterile and
fertile leaves, such as Maxonia, Olfersia, Polybotrya, and
Polystichopsis. Hybrids are known between Cyclodium and
Polybotrya. The genus name is derived from the Greek
kyklodes, ring-like. It alludes to the circular indusia.
GÉNEROS PARECIDOS: Arachniodes difiere por lámi-
nas anadrómicas y rizomas reptantes. Polybotrya es (prin-
cipalmente) un helecho trepador y tiene hojas estériles y
fértiles dimorfas. Dryopteris difiere por rizomas erectos o
decumentes. Stigmatopteris difiere por laminas más delga-
das con glándulas internas punctatas y venas terminando
ante del margen en hidatodos clavados.
COMENTARIOS: Cyclodium es completatmente neo-
tropical y consta de casi 10 especies, estas principalmente
en las Guianas. Es un género de las tierras bajas, alca-
ncando solo hasta 1500 m. Forma un clado con otros
helechos dryopteroides que tienen rizomas reptantes y
(a menudo) hojas estériles y fértiles dimorfas, tales como
Maxonia, Olfersia, Polybotrya, and Polystichopsis. Se con-
ocen híbridos entre Cyclodium y Polybotrya. El nombre
genérico se deriva del griego kyklodes, como un anillo.
Alude a los indusios circulares.

LITERATURE: Engels, M. E. & B. K. Canestraro. 2017. XCyclobotrya: a new hybrid genus between Cyclodium and Poly-
botrya (Dryopteridaceae) from the Brazilian Amazon. Brittonia 69: 307–312. Smith, A.R. 1986. Revision of the neotropical fern
genus Cyclodium. American Fern Journal 76: 56–98.

246
Figure 120. Cyclodium meniscioides. A. Habit. B. Venation. C. Fertile pinnae. D. Detail of sori. (Original, ©Robbin Moran,
2009.)

247
LOMARIOPSIDACEAE
Cyclopeltis J. Sm.
DESCRIPTION: Terrestrial; rhizomes erect or decum-
bent, scaly; sterile and fertile leaves monomorphous;
laminae 1-pinnate with a subconform apical pinna;
pinnae articulate to the rachis, entire or nearly so,
the basiscopic side auriculate and overlapping the
upper surface of the rachis; rachis and costae round-
ed or flattened above (not grooved), puberulent adaxially,
these 0.1–0.2 mm long, reddish, articulate; veins free; sori
round, in 2 or 3 rows between the costa and mar-
gin; indusia peltate. Spores monolete.
SIMILAR GENERA: Nephrolepis differs by having sto-
lons, pinnatifid (not conform) lamina apices, and only one
row of sori between the costa and margin. Lomariopsis
differs by epiphytic or hemiepiphytic habit, long-creeping
rhizomes, sterile-fertile leaf dimorphy, and acrostichoid
sori.
COMMENTS: Cyclopeltis has one species in the New
World (C. semicordata (Sw.) J. Sm.) and five in southeast-
ern Asia and the western Pacific. The American species
is common and widespread in wet forests mostly below
600 m. The articulations of the pinnae can be seen as
dark line around the base of the costa where it joins the
rachis.They appear to be nonfunctional; that is, the pinnae
do not disarticulate. The presence of articulations is the
only morphological synapomorphy known for the Lom-
ariopsidaceae.The genus name is derived from the Greek
kyklos, circle + peltis, shield, and refers to the indusium.
LITERATURE: Lu, J. M. & D. Z. Li. 2006. A study on systematic position of Cyclopeltis. Acta Botanica Yunnan 28(4): 337–340.
[in Chinese]
248
DESCRIPCIÓN: Terrestres; rizomas erectos o decum-
bentes, escamosos; hojas estériles y fértiles monomórfi-
cas; láminas 1-pinnadas con una pinna apical subcon-
forme; pinnas articuladas al raquis, enteras o casi
enteras, el lado basiscópico auriculado y traslapa
la superficie superior del raquis; raquis y costas en la
superficie superior redondeados o planos (no surcados),
puberulentos adaxialmente, estos 0.1–0.2 mm de largo,
rojizos, articulados; nervaduras libres; soros redondea-
dos, dispuestos en 2 o 3 hileras entre la costa y el
margen; indusios peltatos. Spores monolete.
GÉNEROS PARECIDOS: Nephrolepis difiere por ten-
er estolones, ápices de la láminas pinnatífidos (no con-
formes) y solamente una hilera de soros entre la cos-
ta y el margen. Lomariopsis difiere por hábito epifítico
o hemiepifítico, rizomas largamente rastrero, dimorfismo
entre las hojas estériles y fértiles y soros acrosticoides.
COMENTARIOS: Cyclopeltis tiene una especie en el
Nuevo Mundo (C. semicordata (Sw.) J. Sm.) y cinco en
el sureste de Asia y el Pacífico occidental. La especie
americana es común y difundida y occure en bosques
húmedos principalmente bajo los 600 m. Las articula-
ciones de las pinnas puede ser observadas como líneas
oscuras alrededor de la base de la costa donde se une
al raquis. Aparecen ser no funcional; o sea, las pinnas no
se desprenden. La presencia de las articulaciones es la
única sinapomorfia conocida para las Lomariopsidaceae.
El nombre genérico se deriva del griego kyklos, círculo +
peltis, shield, a se refiere al indusio.
Figure 121. Cyclopeltis semicordata (Sw.) J. Sm. Note basal basiscopic lobe of pinna overlapping the rachis. (From Rovirosa,
1909)

249
THELYPTERIDACEAE
Cyclosorus Link
DESCRIPTION: Rhizomes long-creeping, blackish,
glabrous or nearly so, internodes generally about 3
cm long; petiole bases with 2 vascular bundles; laminae
1-pinnate-pinnatifid, not or hardly reduced at the base,
gradually pinnatifid to the tip; rachises and costae
glabrous to pubescent, sparsely scaly, the scales broadly
ovate to lanceolate, thin, with marginal hairs; costae
grooved adaxially; basal veins from adjacent segments
anastomosing and producing an excurrent vein that
ends in the base of the sinus, the suprabasal veins free,
reaching the margin, all veins sparsely glandular, the glands
spherical, red; aerophores and buds absent; hairs (when
present) acicular; sori round, indusiate; sporangial stalks
with reddish glands, capsules glabrous; x=36.
DESCRIPCIÓN: Rizomas largamente rastreros,
negruscos, glabros o casi asi, entrenudos
generalmente ca. 3 cm de largo; bases de los pecíolos
con 2 haces vasculares; láminas 1-pinnate-pinnatifidas, no
o poco reducidas en la base, gradualmente pinnatifidas
hacia el ápice; ráquises y costas glabras a pubescentes,
esparcidamente escamosos, las escamas anchamente
ovadas a lanceoladas, delgadas, con pelos marginales;
costas surcadas adaxialmente; venas basales desde
segmentos adyacentes anastomosándose y
produciendo una vena excurrente que termina en la
base del sinus, las venas suprabasales libres, alcanzando
el margen, todos las venas esparciadmente glandular, las
glándulas esféricas rojizas; aeróforos y brotes ausetnes;
pelos (cuando presentes) aciculares; soros redondos,
indusiados; pecíolos de los esporángios con glándulas
rojizas, cápsulas glabras; x=36.

SIMILAR GENERA: Amauropelta typically differs by GÉNEROS PARECIDOS: Amauropelta típicamente

erect rhizomes, laminae reduced basally, and absence of
reddish glands. Amblovenatum differs by short-creeping
rhizomes and yellow glands on veins abaxially and
sporangial stalks. Parathelypteris and Thelypteris differ by
smaller size (pinnae rarely more > 8 cm long), free veins,
and glabrous sporangial stalks.
difiere por rizomas erectos, láminas reducidas basalmente,
y la ausencia de glándulas rojizas. Amblovenatum difiere
por rizomas cortamente rastreros y glándulas amarillas
esféricas en las venas abaxialmente y los pecíolos
esporangiales. Parathelypteris y Thelypteris difieren por
tamaño mas pequeño (pinnas raras veces más de > 8 cm
de largo), venas libres y pecíolos esporangiales glabros.

COMMENTS: Cyclosorus contains 2 generally recognized COMENTARIOS: Cyclosorus consta de 2 especies

species, with only one in the New World It is pantropical generalmente reconocidas, con solamente una en el
and subtropical. The species typically occur from 0–1700 Nueva Mundo.. Es pantropical y subtropical. Las especies
m in freshwater swamps, marshes, lake margins, and mats típicamente existen desde 0–1700 m en pantanos de agua
of floating vegetation.The most common and widespread dulce, humedales, márgenes de lagos, alfombras flotantes
species is C. interruptus (Willd.) H. Ito. It represents a de vegetación. La especie más común y difundida es C.
complex badly in need of study. See Holttum (1981) and interruptus (Willd.) H. Ito. Representa un complejo que
Smith (1971) for the many names that apply. Plants in necesita estudio. Véase Holttum (1981) y Smith (1971)
Jamaica are tetraploids, whereas ones from South Africa para los varios que se aplican. nombres for the many
and Asia are diploids.The genus name is derived from the names that apply. Las plantas de Jamaica son tetraploides,
Greek kyklos, circle + soros, sorus, referring to the round mientras que las de Sur África y Ásia son diploides. El
sori. nombre genérico se deriva del griego kyklos, círculo
+ soros, sorus, a se refiere al los soros redondeados.
LITERATURE: Fosberg, F. R. & M.-H. Sachet. 1972. Three Indo-Pacific Thelypteris species reinterpreted and a new African
species described. Smithsonian Contributions to Botany 8: 1–10. Holttum, R. E. 1971. Studies in the family Thelypteridaceae
III. A new system of genera in the Old World. Blumea 19: 17–52. Holttum, R. E. 1981. Thelypteridaceae. In: C. G. G. J. van
Steenis & R. E. Holttum (gen. eds.), Flora Malesiana, series 2 - Pteridophytea, ferns and fern allies 1(5): 331–599. Holttum, R. E.
1970. Studies in the family Thelypteridaceae I. A comparative study of the type-species of Thelypteris Schmidel, Cyclosorus Link,
and Ampelopteris Kunze. Blumea 18: 195–215. Ponce, M. 1987. Revisión de las Thelypteridaceae (Pteridophyta) Argentinas.
Darwiniana 28: 317-390. Small, J. K. 1931. Ferns of Florida. The Science Press, New York. Smith, A. R. 1971. Systematics of the
neotropical species of Thelypteris section Cyclosorus. University of Calififornia Publications in Botany. 59: 1–143.

250
Figure 122. Cyclosorus interruptus. (Original, ©R. C. Moran, 2019.)

251
CYSTOPTERIDACEAE
Cystopteris Bernh.
DESCRIPTION: Terrestrial or saxicolous; rhizomes
creeping, scaly, the scales thin, translucent; sterile and fer-
tile leaves dimorphous, thin-textured; petioles with 2
vascular bundles; laminae usually 15–40 cm long, 2-pin-
nate-pinnatifid to 4-pinnate, anadromic; axes grooved
on the upper surface, the grooves decurrent; veins free,
the tips slender (not enlarged); sori round, medial or in-
framedial; indusia hyaline, scalelike, attached to the
proximal side of the receptacle and (when young)
arching over the sporangia, evanescent, often hidden
by the mature sporangia; spores monolete, bilateral; x=42.
SIMILAR GENERA: Woodsia resembles Cystopteris but
differs by having the indusium attached completly around
the base of the receptacle. At maturity, the indusium
splits into lobes or hairs that form a loose cup around
the sporangia. In sterile specimens, the veins tips are diag-
nostic: those of Woodsia are enlarged, whereas those of
Cystopteris are slender.
COMMENTS: Cystopteris contains about 10 species
and is primarily temperate. The common and widespread
species in the neotropics is C. fragilis (L.) Bernh. It is usually
found above 2500 m. Throughout Latin America this spe-
cies has diploid, tetraploid, and hexaploid cytotypes. Some
specimens have misshapen spores, suggesting hybridiza-
tion.Two notes about observing identifying the genus:The
indusium is often pushed back and hidden by the sporan-
gia. Therefore, it is difficult to see in mature sori. The two
vascular bundles in the petioles can usually be seen in
specimens where the petiole has been broken.The genus
name is derived from the Greek kystos, bladder + pteris,
fern. It refers to the inflated indusium.
LITERATURE: Blasdell, R. F. 1963. A monographic study of the fern genus Cystopteris. Memoirs of the Torrey Botanical Club
21(4): 1–102. Pearman, R. W. 1976. A scanning electron microscopic investigation of the spores of the genus Cystopteris. British
Fern Gazette 11: 221–230.
252
DESCRIPCIÓN: Terrestres o saxícolas; rhizomas rep-
tantes, escamosos, las escamas delgadas, translúcidas;
hojas estériles y fértiles dimorfas, de textura delgada;
pecíolos con 2 haces vasculares; láminas usualmente
15–40 cm de largo, 2-pinnado-pinnatífidas a 4-pinnadas,
anadrómicas; ejes surcados en la superficie superior, los
surcos decurrentes; nervaduras libres, los ápices delgados
(no engrosados); soros redondeados, mediales o infra-
mediales; indusios hialinos, como una escama, unidos
al lado proximal del receptáculo y (cuando jóvenes)
arqueando sobre los esporangios, evanescentes, a
menudo escondido por los esporangios maduros; espo-
ras monoletes, bilaterales; x=42.
GÉNEROS PARECIDOS: Woodsia se parece a
Cystopteris pero difiere por tener el indusio unido com-
pletamente alrededor de la base del receptáculo. Al mad-
urar, el indusio se hende en lobulos o pelos que forman
una copa suelta alrededor de los esporangios. En ejem-
plares estériles, las venas son diagnósticas: las de Woodsia
son engrosadas, mientras que las de Cystopteris son del-
gadas.
COMENTARIOS: Cystopteris consta de alrededor de
10 especies y es principalmente templado. La especie
común y muy difundida en el neotrópico es C. fragilis (L.)
Bernh. Se encuentra usualmente sobre los 2500 m. Por
todo América Latín esta especie tiene citotipos diploides,
tetraploides y hexaploides. Algunos ejemplares tienen es-
poras malformadas, sugiriendo hibridación. Dos notas so-
bre identificar el género: El indusio es a menudo empuja
hacia atrás y escondido por los esporangios. Por eso es
difícil verlo en soros maduros. Los dos haces vasculares
en el pecíolo se pueden ver usualmente en ejemplares
donde el pecíolo ha sido quebrado. El nombre genérico
se deriva del griego kystos, vejiga + pteris, helecho. Se
refiere al indusio inflado.
Figure 123. A–D. Cystopteris fragilis. G, H, J. C. membranifolia. (From Mickel & Smith, 2004.)

253
MARATTIACEAE
Danaea Sm.
DESCRIPTION: Terrestrials (rarely saxicolous), fleshy;
rhizomes creeping or erect; sterile and fertile leaves di-
morphic, 1-pinnate (rarely simple), with a conform ter-
minal pinna or this is replaced by a bud; petioles with
or without swollen nodes, basally with rounded, earlike,
fleshy stipules; pinnae opposite, entire or with serrate
apices; rachis swollen-nodose at the insertion of the
pinnae; veins free, simple or 1-forked near the base,
parallel to each other; fertile leaves long-petiolate and
therefore taller than the sterile, with narrower pinnae;
sporangia fused into an elongate, double-rowed synan-
gium, with 20–100 locules, opening by terminal pores;
synangia covering most of the distance between the costa
and margin; x=40.
DESCRIPCIÓN: Terrestres (raras veces saxícolas), car-
nosas; rizomas reptantes o rectos; hojas estériles y fértiles
dimorfas, 1-pinnadas (raras veces simples), con una pinna
terminal conforme o en su lugar un brote; pecíolos con o
sin nudos hinchados, basalmente con estípulas redondea-
das parecidas a orejas; pinnas opuestas, enteras o con
ápices serrados; raquis nodoso-hinchado en la unión de
las pinnas; nervaduras libres, simples o 1-furcadas cerca
de la base, paralela entre sí; hojas fértiles lárgamente pe-
cioladas y por eso más largas que las estériles, con pinnas
más angostas; esporangios fusionados en sinangios elon-
gados de doble hilera, con 20–100 lóculos, abriéndose
por poros terminales; sinángios cubren la mayoría de la
distancia entre la costa y el margen; x=40.

SIMILAR GENERA: Lomariopsis differs by its long-creep-
ing and climbing stems, alternate pinnae, and lack of swol-
len nodes along the rachis. Metaxya differs by hairs at
the base of the petioles, the absence of swollen nodes at
the rachis-pinna junctures, and round sori. Once-pinnate
species of Pteris differ by areolate veins and sori along the
margins.
GÉNEROS PARECIDOS: Lomariopsis difiere por sus
tallos largamente reptantes y rastreros, pinnas alternas
y la ausencia de nudos hichados por el raquis. Metaxya
difiere por pelos en la base de los pecíolos, la aucencia
de nudos hinchados en la unión del raquis y pinnas, y so-
ros redondos. Especies de Pteris 1-pinnadas difieren por
venas areoladas y soros por los márgenes.

COMMENTS: Danaea is a completely American genus COMENTARIOS: Danaea es un género completa-

with about 50 species. They grow in wet forests at low to mente americano con alrededor de 50 especies. Se en-
middle elevations. When dried, the swollen nodes on the cuentran en bosques húmedos en elevaciones bajas a
petioles and rachises shrink and darken. In several species, medias. Cuando es secado, los nudos hinchados de los
the apical pinna is often or always replaced by a bud. After pecíolos y ráquises se encogen y oscuren. En varias espe-
the leaf has died, the petiole bases and stipules persist cies, la pinna apical es a menudo o siempre replazado por
on the stem and contribute substantially to its diameter. un brote. Después de que la hoja se muere, las bases de
The genus totally lacks hairs; the only indument consists los pecíolos persisten sobre el tallo y contribuyen sub-
of peltate, irregular scales that provide few distinctions stancialmente a su diámetro. El género carece totalmente
between the species. When collecting Danaea, it is im- de pelos; el único indumento consiste de escamas pelta-
portant to note whether the rhizome is creeping or erect. das iregulares que provienen pocas distinciones entre las
The very young leaves of some species are iridescent. especies. Cuando se colecta a Danaea, hay que notar si
Three subgenera have been recognized (Christenhusz et el rizoma es reptante or erecto. Las hojas muy jóvenes
al., 2010). The genus is named for Giovanni Pietro Maria de algunas especies son iridescentes.Tres subgéneros han
Dana (1736–1801), professor of botany and friend of J. sido reconocidos (Christenhusz et al., 2010). El género se
E. Smith. nombra para Giovanni Pietro Maria Dana (1736–1801),
profesor de la botánica y amigo de J. E. Smith.
LITERATURE: Christenhusz, M. J. M., H. Tuomisto, J. S. Metzgar & K. M. Pryer. 2008. Evolutionary relationships within
the Neotropical, eusporangiate fern genus Danaea (Marattiaceae). Molecular Phylogenetics & Evolution 46: 34–48. Christen-
husz, M. J. 2010. Danaea (Marattiaceae) revisited: biodiversity, a new classification and ten new speies of a neotropical fern
genus. Botanical Journal of the Linnean Society 163: 360–385. Moran, R. C. & M. H. Grayum. 2017. The hybrid nature of
Danaea plicata (Marattiaceae), a Costa Rican endemic. Brittonia 00: 000–000. Rolleri, C. 2004. Revisión del género Danaea
(Marattiaceae - Pteridophyta). Darwiniana 42(1–4): 217–301. Tuomisto, H. & R. C. Moran. 2001. Marattiaceae. In: G. Harling
(editor). Flora of Ecuador 66: 23–68.

254
Figure 124. Danaea elliptica. Note in C the rounded stiples, and in B the poricidal synangia. Fijase en C las estipulas
redondeadas y en B los sinángios poricidales. (from Proctor, 1985; drawn by Peter J. Edwards.)

255
LYCOPODIACEAE
Dendrolycopodium A. Haines.
DESCRIPTION: Terrestrial; main horizontal shoots
subterranean, long-creeping, giving rise laterally to 2
alternate rows of erect dendroid shoots; trophophylls
isophyllous or weakly anisophyllous, arranged in
alternating pseudowhorls of 3, collectively forming 6
total ranks, lacking spine-like or hair-like tips; strobili
sessile, with subpeltate sporophylls, these with a medial
wing decurrent on the strobilus axis; sporangia isovalvate,
with epidermal cells with sinuate, pitted sidewalls; spores
reticulate on the distal hemisphere, with isolated ridges
on the proximal hemisphere; gametophytes subterranean,
holosprophytic, disk-shaped; x=34.
DESCRIPCIÓN: Terrestres; vástagos principales
horizantales subterráneos, largamente rastreros,
dando lugar lateralmente a 2 hileras alternas de
vástagos erectos dendroides; trofofilios isofilicos o
debilmente anisophyllous, arreglados en pseudoverticelos
alternando de 3, colectivamente formando 6 hileras
totales, faltando un ápice espiniforme o piliforme;
estróbilos séssiles, con esporofílios subpeltatos, estos
con una ala medial decurrente por el eje del estróbilo;
esporángios isovalvados, las células epidérmicas con las
paredes anticlinales sínuas picadas; esporas reticuladas
en hemisfério distal, con filas aislados en el hemisfério
proximal; gametofitos subterráneos, holosaprofíticos,
disciformes; x=34.

SIMILAR GENERA: Austrolycopodium differs by strobili
pedunculate and sporophylls peltate, without a decurrent
medial wing on the strobilus axis. Lycopodium differs by
non-dendroid habit of the erect shoots, piliform leaf
apices, and usually pedunculate strobili. Spinulum differs by
the erect shoots lacking horizontal branchlets, and 8–10
rows of trophophylls, these terminating in a minute spine.
GÉNEROS PARECIDOS: Austrolycopodium difiere por
estróbilos pedunculados y esporofilios peltados, sin una
ala medial decurrente en el eje del estróbilo. Lycopodium
difiere por hábito no dendroide de los vástagos erectos,
ápices de los trofofílios piliformes y estróbilos usualmente
pedunculados. Spinulum difiere por los vástagos erectos
faltando ramitas horizontales y con 8–10 hileras de
trofofílios, estos terminándose en una espinita.

COMMENTS: Dendrolycopodium consists of four species COMENTARIOS: Dendrolycopodium consta de cuatro

and is primarily northern temperate and circumboreal. especies y es principalmente en las zonas templadas del
Three species occur in North America: D. dendroideum, D. norte y circunboreal. Tres especies existen en América
hickeyi, and D. obscurum (the fourth species, D. juniperoidium, del Norte: D. dendroideum, D. hickeyi, and D. obscurum
is Asian). They typically grow on shaded forest floors (the fourth species, D. juniperoidium, es Asiático). Existen
but may also grow in sunny habitats where their lateral típicamente en sotobosques sombreados pero pueden
branchlets tend to become strongly ascending. Each year, ocurrir en ambientes soleados donde sus ramitas laterales
the subterranean stem produces and erect shoot and a son más ascendentes. Cada ano el tallo suberráneo
small, weak horizontal shoot. The latter does not develop produce un vástago erecto y una rama debil pequeño
unless the main stem dies or is injured. A key to the three horizontal. Este último no se desarrolla a menos que
species in the Americas may be found in Haines (2003) el tallo principal muere o se daña. Una clave a las tres
and Hickey (1977). The genus name is derived from the especies en las Américas se puede encontrar en Haines
Greek dendro, tree, and Lycopodium. It refers to the tree- (2003) y Hickey (1977). El nombre genérico es derivado
like growth form of the arial shoots.. del griego dendro, arbol, y Lycopodium. Se refiere al forma
de crecimiento dendroide de las vástagos aéreos.
LITERATURE: Haines, A. 2003. The families Huperziaceae and Lycopodiaceae of New England, a taxonomic and ecological
reference. V. F. Thomas Co., Bowdin, Maine. Hickey, R. J. 1977. The Lycopodium obscurum complex in North America. American
Fern Journal 67: 45–48. Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178.
Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the Missouri Botanical Garden 79: 687–717. Øllgaard,
B. & K. Tind. 1993. Scandinavian ferns. Rhodos, Copenhagen. Small, J.K.1935. Ferns of the vicinity of New York. The Science
Press, Lancaster, Pennsylvania. Primack, R. B. 1973. Growth patterns of five species of Lycopodium. American Fern Journal 63:
3–7. Whittier, D. P. 1977. Gametophytes of Lycopodium obscurum as grown in axxenic culture. Canadian Journal of Botany
55: 563–567.

256
Figure 125. Dendrolycopodium obscurum (from Small, 1935). Lower left compares the phyllotaxy of the ultimate horizontal
branchlets of the three species in North America: 1–3 D. obscurum, 5–6 D. hickeyi, and 7–9 D. dendroideum (from Hickey,
1977). Top row is dorsal view, middle row is ventral view, and lower row is front view..

257
DENNSTAEDTIACEAE
Dennstaedtia Bernh.
DESCRIPTION: Terrestrial or epiphytic; rhizomes
creeping, pubescent, without scales; leaves 1–4(-7)
m long, erect or supported by surrounding vegetation;
petiole bases frequently with buds that give rise to
new stems, in transverse section with a vascular bundle
en the form of an omega; laminae 2-pinnate-pinnatifid
to 4-pinnate, rarely (in D. wercklei (H. Christ) R.M. Tryon)
1-pinnate, some species with buds in the axils of the distal
pinnae; pinnules arranged anadromically; veins free, end-
ing behind the margin in slender or clavate apices; sori
marginal, deflexed, terminal on the vein apices, in most
species borne in the sinuses between lobes; indusia cup-
shaped; spores tetrahaedral-globose, trilete, nongreen.
x=30, 33, 34, 46, 47.
SIMILAR GENERA: Hypolepis differs by false indusia.
Paesia and Pteridium differs by elongate sori supplied by
several to many veins. Saccoloma differs by persistant pet-
iole bases around a short, thick rhizome.
COMMENTS: Dennstaedtia is mostly pantropical with
about 20 species in the New World. With the excep-
tion of D. bipinnata and D. globulifera, all other species in
the genus have epipetiolar buds. These are borne on the
sides of the petiole near the base and may number more
than one. They are an integral part of the branching sys-
tem of the plant. Epipetiolar branches are found in other
dennstaedtioid genera such as Histiopteris, Hypolepis, and
Pteridium. The genus name is named for August Wilhelm
Dennstaedt (1776–1826), German botanist and author.
DESCRIPCIÓN: Terrestres o epífitas; rizomas rastre-
ros, pubescentes, sin escamas; hojas 1–4(-7) m de largo,
erectas o apoyándose sobre la vegetación circundante;
bases de los pecíolos frecuentemente portando
yemas que dan lugar a nuevos tallos, en corte trans-
versal con un haz vascular en forma de omega; láminas
2-pinnado-pinnatífidas a 4-pinnadas, raras veces (en D.
wercklei (H. Christ) R.M. Tryon) 1-pinnatas, algunas espe-
cies con yemas en las axilas de las pinnas distales; pínnulas
arregladas anadrómicamente; venas libres, terminando
antes del margen, con las puntas delgadas o claviformes;
soros marginales, deflexos, terminales en las puntas de
las venas, naciendo en la mayoría de las especies en los
senos entre los lobulos; indusios en forma de copa;
esporas tetrahédricas-globosas, triletes, no verdes. x=30,
33, 34, 46, 47.
GÉNEROS PARECIDOS: Hypolepis difiere por false
indusia. Paesia y Pteridium tienen soros alargados y sum-
inistrados con varias a muchas venas. Saccoloma difiere
por los bases de los pecíolos persistentes alrededor de
un rizoma corto y grueso.
COMENTARIOS: Dennstaedtia es mayormente pan-
tropical con alrededor de 20 especies en el Nuevo
Mundo. Con excepción de D. bipinnata y D. globulifera,
todas las otras especies del género tienen ramas epipe-
ciolares. Estas provienen de los lados del pecíolo cerca
de la base y pueden ser varias en número. Son una parte
integral de la sistema de ramificación de la planta. Ramas
epipeciolares se encuentran en otros géneros de Denn-
staedtiaceae como Histiopteris, Hypolepis y Pteridium. El
nombre genérico honra August Wilhelm Dennstaedt
(1776–1826), botánico Alemán y autor.

LITERATURE: Navarrete, H. & B. Øllgaard. 2000. The fern genus Dennstaedtia (Dennstaedtiaceae) in Ecuador, - new
characters, new species and a new combination. Nordic Journal Botany 319–346. Perrie, L. R., R. K. Wilson, L. D. Shepherd, D. J.
Ohlsen, E. L. Batty, P. J. Brownsey & M. J. Bayly. 2014. Molecular phylogenetics and generic taxonomy of Blechnaceae ferns. Taxon
63: 745–758. Tryon, R. M. 1960. A review of the genus Dennstaedtia in America. Contributions from the Gray Herbarium of
Harvard University 187: 23–52. Yanez, M., D. Arana, G. J. Marquez & A. Oggero. 2014. The genus Dennstaedtia Bernh.
(Dennstaedtiaceae) in Argentina. Phytotaxa 174: 69–81.

258
Figure 126. Dennstaedtia. A–J. D. cornuta. Q–K. D. bipinnata. R–U. D. globulifera. (From Mickel & Smith, 2004.)

259
ATHYRIACEAE
Deparia.
DESCRIPTION: Plants terrestrial; rhizomes short-
creeping; sterile and fertile leaves monomorphic, dying
back in winter; petioles 1/3–2/3 length of lamina, bases
swollen, persisting as trophopod over winter, the sides
dentate (ours) or not; vascular bundles 2; laminae
elliptic to ovate-lanceolate, 1-pinnate-pinnatifid
[pinnatifid to 3-pinnate-pinnatifid]; pinnae not articulate
to rachis, segment margins entire, crenulate, or serrate;
proximal pinnae (several pairs) reduced or not, sessile,
equilateral; costae adaxially shallowly grooved, grooves
not continuous with that of rachis; indument on
rachis and costae (both sides) of multicellular hairs;
veins free; sori elongate, ± straight (ours) or hooked
at distal end; indusia linear, laterally attached, persistent;
spores tuberculate; x=40.
SIMILAR GENERA: Athyrium differs by 2-pinnate or
more divided laminae, adaxial surfaces of the leaf axes
glabrous, and J-shaped sori. Homalosorus differs by smooth
(not dentate) sides of the petiole bases, 1-pinnate laminae,
and leaf axes glabrous adaxially. Thelypteridaceae differs
by typically having round sori.
COMMENTS: Deparia consists of ca. 50 species
worldwide, occurring in eastern North America, eastern
Asia, tropical Africa, the Pacific Islands, and Australia. Only
one species is American: D. acrostichoides (Sw.) M. Kato. It
grows primarily on the floors of shaded, mesic forests.The
hairs of the leaf axes adaxially are said to be homologous
with scales. The genus name is derived from the Greek
depas, saucer, referring to the suacer-shaped indusium of
the type species, D. prolifera (Kaulfuss) Hook. & Grev. This
character is very unusual in the genus..
LITERATURE: Kato, M. 1984. A taxonomic study of the athyroid fern genus Deparia with main reference to the Pacific spe-
cies. Journal of the Faculty of Science, University of Tokyo, Sect. 3, Botany 13: 375–340. Li-Yaung Kuo, L-Y., A. Ebihara, T-C.
Hsu, G. Rouhan, Y-M Huang, C-N. Wang, W-L. Chiou & M. Kato. 2018. Infrageneric revision of the fern genus Deparia
(Athyriaceae, Aspleniineae, Polypodiales). Systematic Botany 43: 645–655. Ogden, E. C. 1981. Field guide to northeastern ferns.
New York State Museum Bulletin, no. 444. Albany, New York.
260
DESCRIPCIÓN: Plantas terrestres; rizomas cortamente
reptantes; hojas estériles y fértiles monomórfas,
muriéndose en el invierno; pecíolos 1/3–2/3 la longitud de
las láminas, bases hinchados, persistiéndose por el invierno,
los lados dentados (nuestro) o no; haces vasculares
2; láminas elípticas a ovado-lanceoladas, 1-pinnado-
pinnatífidas; pinnas no articuladas a los ráquises, márgenes
de los segmentos enteros, crenulados o serrados; pinna
proximal (o varias) reducidas o no, sésilies, equiláteras;
costas adaxialmente surcadas levemente, los surcos
no contínuos con los de los ráquises; indumento
en los ráquises y costas (ambos lados) de pelos
multicelulares; venas libres; soros alargados, ± rectos
(nuestro) o enganchados distalmente; indusios lineales,
unidos lateralmente, persistentes; esporas tuberculadas;
x=40.
GÉNEROS PARECIDOS: Athyrium difiere por las
laminas 2-pinnadas o mas divididas, superficies adaxiales
de los ejes de las laminas glabros y soros en forma de J.
Homalosorus difiere por los lados lisos (no dentados) de
las bases de los peciolos, laminas 1-pinnadas y ejes de
las laminas glabras adaxialmente. Thelypteridaceae difiere
por tipicamente tener soros redondeados.
COMENTARIOS: Deparia consta de 50 especies
mundial, distribuido en América del Norte oriental,
Asia oriental, África tropical, las Islas Pacíficas y Australia.
Solamente una especies es Americana, D. acrostichoides
(Sw.) M. Kato. Existe principalmente en los sotobosques
mésicos sombreados. Los pelos por los ejes de la lámina
se dicen que son homólogos con escamas. El nombre
genérico es derivado del griego depas, platillo, y se
refiere al forma del indusio en la especie tipa D. prolifera
(Kaulfuss) Hook. & Grev. Este carácter es muy inusual
dentro del género.
Figure 127. Deparia acrostichoides. (From Ogden, 1981.)

261
DESMOPHLEBIACEAE
Desmophlebium Mynssen, A. Vasco, Sylvestre, R. C. Moran & Rouhan
DESCRIPTION: Plants terrestrial; rhizomes erect or
decumbent, the scales lanceolate, entire to dentate, con-
colorous, brown; petioles 12–82 × 0.2–1.5 cm; laminae
ovate to oblong-lanceolate, 1-pinnate, with a conform
apical pinna; rachises grooved adaxially; pinnae adnate or
stalked distally, 2–8 pairs per lamina, elliptic or oblong-lan-
ceolate, equilateral, bases truncate, cuneate or obtuse,
apices acuminate, margins entire or serrate distally; costae
and rachises subglabrous; veins simple or 2–4 forked, par-
allel, apices connected by a submarginal vein; sori
linear, simple or diplazioid; indusia persistent, membrana-
ceous, brownish, entire or erose margins; spores ellipsoid,
monolete, cristate, non-green; x=?
SIMILAR GENERA: Diplazium differs by lacking a sub-
marginal connecting vein. Also, many species of Diplazium
are more than 1-pinnate. Olfersia differs by typical eupoly-
pods II petiole vasculature, 2-pinnate fertile leaves, and
acrostichoid sori.
COMMENTS: Desmophlebium contains two species: (D.
lechleri Mynssen et al. y D. longisorum (Baker) Mynssen
et al.)) and is entirely Neotropical, occurring from Costa
Rica to the Brazilian Atlantic Rain Forest. It grows terres-
trially in wet forests from 125–800 m. For a long time
its two species were classified in Diplazium (Athyriaceae),
but molecular phylogenetic studies resolved it sister to
the Hemidictyaceae and Aspleniaceae. The genus is eas-
ily recognized by its imparipinnate laminae, submarginal
connecting vein, linear sori. The generic name is derived
from the Greek, desmos, band or halter, and phlebos, vein.
It refers to the distinctive submarginal connecting vein.
LITERATURE: Mynssen, C. M., A. Vasco, R. C. Moran, L. S. Sylvestre & G. Rouhan. 2016. Desmophlebiaceae and Des-
mophlebium: a new family and genus of Eupolypods II ferns. Taxon 65: 19–34.
262
DESCRIPCIÓN: Plantas terrestres; rizomas erectos o
decumbentes, las escamas lanceoladas, enteras dentatas,
concoloras; pecíolos 12–82 × 0.2–1.5 cm; láminas ova-
das a oblongo-lanceolatas; 1-pinnadas, con ápice con-
forme; ráquises surcados adaxialmente; pinnas adnatas o
pecioluladas distalmente, 2–8 pares por lámina, elípticas a
oblongo-lanceolatas, equiláteras, bases truncadas cunea-
das o obtusas, ápices acuminadas, márgenes enteras o
serradas distalmente; costas y ráquises subglabros; venas
simples o 2–4 furcados, paralelas, los ápices conecta-
dos por una vena submarginal; soros lineares, sim-
ples o diplazioides; indusios persistentes, membranaceos,
parduscos, enteras o erosos; esporas elipsoides, mono-
letes, cristadas, no verdes; x=?
GÉNEROS PARECIDOS: Diplazium difiere por la falta
de una vena submarginal conectante. También, muchas
especies de Diplazium son más de 1-pinnada. Olfersia di-
fiere por la nervadura de los pecíolos de tipo de eupoly-
pods II, hojas fértiles 2-pinnadas y soros acrosticoides.
COMENTARIOS: Desmophlebium consta de dos es-
pecies (D. lechleri Mynssen et al. y D. longisorum (Baker)
Mynssen et al.) y está restringido al Neotrópico, desde
Costa Rica hasta el bosque lluvioso de Bresil Atlántico.
Crece terestrialmente en bosques mojados de 125–800
m. Por mucho tiempo sus dos especies fueron clasificadas
en Diplazium (Athyriaceae), pero estudio filogenéticos
moleculares lo resuelvaran hermano a Hemidictyaceae
y Aspleniaceae. El género se reconoce fácilmente por las
láminas imparipinnadas, vena submarginal conectante y
soros lineales. El nombre genérico se deriva del griego,
desmos, banda, y phlebos, vena. Se refiere a la distinctiva
vena submarginal conectante.
Figure 128. Desmophlebium lechleri. (From Mynssen et al., 2016.)

263
DICKSONIACEAE
Dicksonia L’Hér.
DESCRIPTION: Terrestrial; stems arborescent, to 10
m tall, densely covered by adventitious roots; petioles
with dense, wool-like, golden brown hair, lacking
spines; sterile and fertile leaves 1–3.5 m long, monomor-
phous; laminae 2-pinnate-pinnatifid to 4-pinnate, cori-
aceous, gradually reduced basally and apically; axes
adaxially not grooved, pubescent; veins free; sori mar-
ginal, reflexed; indusia bivalved, with a scarious inner
(abaxial) lobe and a thicker green outer (adaxial) lobe,
the two lobes appearing clam-shaped; spores tetrahe-
dral-globose; x = 65.
SIMILAR GENERA: Alsophila, Cyathea, and Sphaerop-
teris differ by scaly petioles, abaxial sori (not marginal),
and varied shapes of indusia but never bivalved (in some
species the indusia are lacking). Culcita differs by lacking
a trunk, deltate laminae, and rachises and costa grooved
adaxially.
DESCRIPCIÓN: Terrestres; tallos arborescentes, has-
ta 10 m de longitud, densamente cubierto por raíces ad-
venticias; pecíolos cubiertos por pelos como lana, densos,
largos, pardo-dorados, espinas ausesntes; hojas estériles
y fértiles 1–3.5 m de largo, monomorfas; láminas 2-pin-
nado-pinnatífidas a 4-pinnadas, coriáceas, gradualmente
reducidas basalmente y apicalmente; ejes adaxial-
mente no surcados, pubescentes; nervaduras libres; so-
ros marginales, reflejados; indusios bivalvados, con una
valva interior (abaxial) escariosa y una valva exterior
(adaxial) más gruesa y verde, las dos valvas semejan a una
almeja; esporas tetrahédricas-globosas; x = 65.
GÉNEROS PARECIDOS: Alsophila, Cyathea y Sphaer-
opteris difieren por pecíolos escamosos, soros abaxiales
(no marginales), e indusios variados pero nunca bivalva-
dos (en algunas especies los indusios están ausentes). Cul-
cita difiere por faltar un tronco, láminas deltatas y ráquises
y costas surcados adaxialmente.

COMMENTS: Dicksonia contains about 20 species, pri-
marily in the Malesian-Australian region. Three species
occur in the New World: D. berteroana (Colla) C. Chr.
(endemic to Juan Fernández Islands), D. sellowiana Hook.
and D. spruceana Kuhn. (Noben et al. (2018) recognized
8 taxa, but this needs verification). Only D. sellowiana
is common and widespread. It grows above 1500 m in
cloud forests. It can usually be recognized from a distance
by its basally tapering laminae, short petioles, and dead
leaves that persist around the stem to form a “skirt.” Dick-
sonia is named for James Dickson (1738–1822), Scottish
physician and botanist.
COMENTARIOS: Dicksonia consta de alrededor de
20 especies principalmente en la región Malesian-Aus-
traliana. Three especies existen en el Nuevo Mundo:
D. berteroana (Colla) C. Chr. (endémica a las islas Juan
Fernández), D. sellowiana Hook. and D. spruceana Kuhn.
(Noben et al. (2018) reconocieron 8 taxones, pero esto
se necesita verificaión). Solamente D. sellowiana es común
y difundida. Existe arriba de 1500 m en bosques nubo-
sos. Usualmente se puede reconocer de una distancia
por sus láminas reducidas basalmente, pecíolos cortos, y
hojas muertas que persisten alrededor del tallo forman-
do una “falda”. Dicksonia se nombra para James Dickson
(1738–1822), médico escocés y botánico.

LITERATURE: Korall, P. K. M. Pryer, J. S. Metzgar, H. Schneider & D. S. Conant. 2006. Tree ferns: monophyletic groups
and their relationships as revealed by four protein-coding platid loci. Molecular Phylogenetics and Evolution 39: 830–845. Maxon,
W. R. 1913. The North American ferns of the genus Dicksonia. Contributions from the United States National Herbarium 17:
153–156. Noben S, Kessler M, Quandt D, et al. 2017. Biogeography of the Gondwanan tree fern family Dicksoniaceae – a
tale of vicariance, dispersal and extinction. Journal of Biogeography 41: 402–413. Noben, S., M. Kessler, A. Weigand, et al.
2018. A taxonomic and biogeographic reappraisal of the genus Dicksonia (Dicksoniaceae) in the Neotropics. Systematic Botany
43: 389–357. Wolf, P. G., S. D. Sipes, M. R. White, M. L. Martines, K. M. Pryer, A. R. Smith, & K. Ueda. 1999. Phyloge-
netic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: evidence from rbcL nucleotide
sequences. Plant Systematics and Evolution 219: 263–270.

264
Figure 129. A–C. Dicksonia sellowiana. (From Mickel & Smith, 2004.)

265
GLEICHENIACEAE
Dicranopteris Bernh.
DESCRIPTION: Terrestrial; rhizomes long-creeping,
pubescent, the hairs stiff, multicellular, bearing leaves in
one row on the dorsal surface; leaves indeterminate, of
periodic growth; pinnae opposite, forked pseudodichoto-
mously, the bud in the angle of each fork pubescent
by dark bristles; penultimate segments pectinate; veins
2–4-forked; sori round, without indusia; sporangia usu-
ally 8–15 per sorus, sessile; spores trilete; x=39.
SIMILAR GENERA: Sticherus differs by scaly buds in the
pinna forks, scales (never bristles) present on the costae
and costules, veins 1-forked, and 3–6 sporangia per sorus.
Diplopterygium differs by unforked pinnae and the pres-
ence of scales (not bristles) on the leaf.
COMMENTS: Dicranopteris is pantropical and consists
of about 10 species. It is found in open or semi-open hab-
itats, especially roadsides. Their leaves grow periodically,
forming a dormant bud on the tip of the rachis while the
two opposite pinnae develop beneath. When the pinnae
finish their development, the bud at the tip of the rachis
continues growth. This process may repeat itself many
times, resulting in leaves up to 20 m long. The pinnae are
also forked and have a dormant bud in the axil(s). These
buds usually do not develop unless the rachis bud is dam-
aged or destroyed. Two species are common and wide-
spread in the Neotropics: Dicranopteris flexuosa (Schrad.)
Underw. and D. pectinata (Willd.) Underw. The genus
name is derived from the Greek dikranos, two-pronged +
pteris, fern. The leaves and pinnae are repeatedly forked.
LITERATURE: see Sticherus; véase Sticherus.
266
DESCRIPCIÓN: Terrestre; rizomas largamente rastre-
ros, pubescentes, los pelos tiesos multicelulares, llevando
hojas en una hilera en la superficie dorsal; hojas indeter-
minadas, de crecimiento periódico; pinnas opuestas, fur-
cadas pseudodicotómicamente, el brote en el ángulo
de cada bifurcación pubescente por cerdas oscuros;
segmentos penúltimos pectinados; venas 2-4-furcadas;
soros redondos, sin indusios; esporangios usualmente
8–15 por soro, sésiles; esporas triletes; x=39.
GÉNEROS PARECIDOS: Sticherus difiere por brotes
escamosos en las furcaciones de las pinnas, y escamas
(nunca cerdas) usualmente presentes en las costas y
cóstulas. Diplopterygium difiere por pinnas no bifurcadas
y la presencia de escamas (no cerdas) en la hoja.
COMENTARIOS: Dicranopteris es pantropical y consta
de ca. 10 especies. Se encuentra en habitats abiertos o
semi-abiertos, especialmente en taludes. Sus hojas crecen
periódicamente y forman un brote latente en el ápice
del raquis mientras las dos pinnas opuestas se desarrol-
lan abajo. Cuando las pinnas terminan su desarrollo, el
brote latente apical del raquis continua su crecimiento.
Este proceso puede repetirse muchas veces resultando
en hojas de hasta 20 cm de largo. Las pinnas son tam-
bién furcadas y tienen un brote latente en la(s) axila(s).
Estos brotes usualmente no se desarrollan a menos que
el brote del raquis sea dañado o destruido. Dos especies
son comunes y muy difundias en el Neotrópico: Dicran-
opteris flexuosa (Schrad.) Underw. y D. pectinata (Willd.)
Underw.El nombre genérico se deriva del griego dikranos,
dos-cuernos + pteris, helecho. Las hojas y pinnas se bifur-
can repetidamente.
Figure 130. A–D: Dicranopteris flexuosa. (From Mickel & Smith, 2004.)

267
DIDYMOCHLAENACEAE (EUPOLYPODS I)
Didymochlaena Desv.
DESCRIPTION: Terrestrial; leaves 1–2 m long, the ster-
ile and fertile monomorphous; laminae 2-pinnate, im-
paripinnate; pinnae 15–20 pares, sessile; pinnules 1–2.5
× 0.5–1 cm, subdimidiate, sessile, obtuse, entire or cren-
ate, straight along the basiscopic side; basal acroscopic
pinnule overlapping the rachis; ridges of the axes with
spine-like projections on the upper surface of the junc-
tures of the rachis, costae, and costules; axes scaly; veins
free, hydathodous; sori elongate, along both sides of
the vein and curved around its apex, embossed adaxially;
indusia round distally, entire; spores monolete; x=41.
SIMILAR GENERA: Lindsaea differs by larger size and GÉNEROS PARECIDOS: Lindsaea difiere por tamaño
sori close and parallel to the margin. más grande y soros parallelos y cercanos al margen.
COMMENTS: A pantropical genus consisting of a single
species D. truncatula (Sw.) J. Sm. (a second species might
occur in Madagascar). It grows in wet forests at low to
mid elevations and is common and widespread through-
out the Neotropics. Didymochlaena has two characters
unique among the Eupolpods I: elongate sori and minute
spine-like projections on the upper surface of the costa
where it joins the costule. The young unfurling leaves are
reddish brown, giving rise to the common name of the
plant in horticulture: mahogany fern. The leaves emit a
skunk-like odor when cut or crushed, similar to that of
Phanerophlebium. Plants are often detected first in the
field by their odor. The genus name is derived from the
Greek didymos, double + chlaena, cloak. It refers to the
indusia that bends around the vein apex and covers both
sori, thus appearing double.
LITERATURE: Zhang, L-B. & L. Zhang. 2015. Didymochlaenaceae: A new fern family of eupolypods I (Polypodiales). Taxon 64:
27–38.
268
DESCRIPCIÓN: Terrestres; hojas 1–2 m de largo, las es-
tériles y fértiles monomorfas; láminas 2-pinnadas, im-
paripinnadas; pinnas 15–20 pares, sésiles; pínnulas 1–2.5
× 0.5–1 cm, subdimidiadas, sésiles, obtusas, enteras o
crenuladas, rectas a lo largo del lado basiscópico; pín-
nulas basales basiscopicas sobrelapando el raquis;
crestas de los ejes con proyecciones espiniformes en
la superficie adaxial en las uniones del raquis, costas y
cóstulas; ejes escamosos; nervaduras libres, hydatodos
presentes; soros alargados, a lo largo de ambos lados
de la nervadura y curvados alrededor de su ápice; adax-
ialmente un poquito elevado; indusios redondeados en el
extremo distal, enteros; esporas monoletes; x=41.
COMENTARIOS: Un género pantropical que consiste
de una sola especie D. truncatula (Sw.) J. Sm. (una segunda
especie posiblemente existe en Madagascar). Crece en
bosques húmedos en elevaciones bajas o medias, es es
común y muy difundida por todo el Neotrópico. Didy-
mochlaena tiene dos características únicas entre los Eu-
polypods I: soros alargados y proyecciones espiniformes
minutas en la superficie superior de las costa donde se
une con las pínnulas. Las hojas jóvenes en el proceso de
desplagarse son pardo rojizas, dando lugar al nombre
común en la horticultura de helecho de mahogany. Cuan-
do cortadas o estrujadas, las hojas emiten un olor como
zorrillo, similar a eso en Phanerophlebium. Las plantas se
detectan primero en al campo por su olor. El nombre
genérico se deriva del griego didymos, double + chlaena,
capa. Se refiere al los indusios que doblan alrededor del
ápice de la vena para cubrir los dos soros, por eso apa-
reciéndose doble.
Figure 131. Didymochlaena truncatula (From Tryon & Stolze, 1991.)

269
HYMENOPHYLLACEAE
Didymoglossum sect. Didymoglossum
DESCRIPTION: Epiphytic; roots absent; rhizomes ca.
1mm wide, long-creeping, branched, densely pubescent,
the hairs blackish; leaves to 6 cm long, entire to bipinnat-
ifid, margins pubescent, the hairs dark, simple or paired
or arranged in stellate groups of 3 or 4; petioles short or
absent, often pubescent with dark hairs like those on the
rhizomes; costae present or absent, if absent; veins cata-
dromous, often flabellate; false veins present as short,
faint streaks between the true veins, continuous sub-
marginal false vein absent; sori one to several, margin-
al in the distal part of the leaf; indusia funnel-shaped or
tubular, bilabiate, the lobes often with dark margins;
receptacles exert; spores green, trilete; x=34.
SIMILAR GENERA: Didymoglossum sect. Microgonium
differs by the presence of a continuous submarginal false
vein, marginal hairs absent, indusia truncate (not bival-
vate) without a dark margin. Polyphlebium differs by false
veins absent and larger leaves.
COMMENTS: Section Didymoglossum is mainly neo-
tropical and contains about 20 species. Wessels Boer
(1962) recognized 15 species in his monograph of the
neotropical taxa. They grow in wet forests up to ca. 1500
m, usually as low-trunk epiphytes and sometimes on
moss-covered rocks. Often they are overlooked because
of their small size. An unusual character of Didymoglos-
sum (both subgenera) is that the young laminae are flat;
they do not form a fiddlehead. Another unusual char-
acter is false veins. These are elongate collenchymatous
cells that appear as faint streaks between and parallel to
the main veins. Section Microgonium also has a continuous
submarginal false vein parallel to the margin (this lacking
in sect. Didymoglossum). The dark hairs on the rhizome
are specialized for adhesion to the substrate, branching
when they touch the surface (Schneider, 2000).The genus
name is derived from the Greek didymos, double + glossa,
tongue. The mouth of the indusium typically consists of
two widely spreading lobes.
LITERATURE: Schneider, H. 2000. Morphology and anatomy of roots in the filmy fern tribe Trichomaneae H. Schneider
(Hymenophyllaceae, Filicatae) and the evolution of rootless taxa. Botanical Journal of the Linnean Society 132: 29–46. Wessels
Boer, J. G. 1962. The New World species of Trichomanes sect. Didymoglossum and Microgonium. Acta Botanica Neerlandica
11: 277–330.
270
DESCRIPCIÓN: Epífitas; raíces ausentes; rizomas a 1
mm de ancho, largamente rastreros, ramificados, densa-
mente pubescentes, los pelos negruzcos; hojas a 6 cm de
largo, enteras a bipinnatífidas, márgenes pubescentes;
pecíolos cortos o ausentes, a menudo pubescentes con
pelos oscuros como los en el rizoma; costas presentes o
ausentes, si ausentes; venas catádromas, a menudo flabel-
ladas; venas falsas presentes como rayos cortos débiles
entre las venas verdaderas, falsa vena continua sub-
marginal ausente; soros uno a varios, marginales en la
parte apical de la hoja; indusios infundibuliformes o tubu-
lares, bilabiados, los lóbulos con márgenes oscuros;
receptáculos exertos; esporas verdes, triletes; x=34.
GÉNEROS PARECIDOS: Didymoglossum sect. Microgo-
nium difiere por la precencia de una falsa vena continua
submarginal, pelos marginales ausentes, indusios trunca-
dos (no bilobados) sin un margen oscuro. Polyphlebium
difiere por venas falsas ausentes y hojas más grandes.
COMENTARIOS: La secc. Didymoglossum es principal-
mente neotropical y consta de ca. 20 especies. Wessels
Boer (1962) reconoció 15 especies en su monografía de
los taxones neotropicales. Crecen en bosques húmedos
hasta 1500 m, usualmente como epífitos en las partes bas-
al de troncos y a veces sobre rocas cubiertas con musgos.
Usualmente están pasados por alto a causa de su tamaño
pequeño. Una característica poco común de Didymoglos-
sum (ambos subgéneros) es que las láminas jóvenes son
planas; no forman un cayado. Otra característica inusual
es falsas venas. Estos son células colenquímatas alarga-
das y se aparecen como rayas finas entre (y paralelas a)
las venas principales. La sección Microgonium tiene tam-
bién una falsa vena continua submarginal (esto ausente
en secc. Didymoglossum). Los pelos oscuros en el rizoma
son especializados para adherir al sustrato, ramificando
en ápice cuando tocan la superficie (Schneider, 2000). El
nombre genérico se deriva del griego didymos, doble +
glossa, lengua. La boca del indusio típicamente consta de
dos lóbulos ampliamente divergentes.
Figure 132. A. Didymoglossum kraussii. B. D. membranaceum. C, E. D. bucinatum. D. D. ovale. F. D. hymenoides. (Modified
from Mickel & Smith, 2004.)

271
HYMENOPHYLLACEAE
Didymoglossum sect. Microgonium
DESCRIPTION: Epiphytic; roots absent; rhizomes ca.
1mm wide, long-creeping, branched, densely pubescent,
the hairs blackish; leaves to 6 cm long, entire to bipin-
natifid, margins glabrous; petioles short or absent, of-
ten pubescent with dark hairs like those on the rhizomes;
costae present or absent, if absent; veins catadromous,
often flabellate; false veins present as short, faint streaks
between the true veins, continuous submarginal false
vein present; sori one to several, marginal in the distal
part of the leaf, usually completely immersed in the lami-
na; indusia funnel-shaped or tubular, truncate (not bila-
biate), without dark margins; receptacles exert; spores
green, trilete; x=34.
SIMILAR GENERA: Didymoglossum sect. Didymoglos-
sum differs by the absence of a continuous submarginal
false vein, the indusia bilabiate (not truncate) with dark
margins, and marginal hairs present. Polyphlebium differs
by false veins absent and larger leaves.
COMMENTS: Section Microgonium is mainly paleotrop-
ical and consists of about 10 species. They occur as ep-
iphytes or on mossy rocks in wet tropical forests. In his
monograph of the neotropical taxa, Wessels Boer (1962)
recognized 4 species: T. ekmanii, T. godmanii, T. hookeri, and
T. kapplerianum. Often they are overlooked because of
their small size. An unusual character of Didymoglossum
(both subgenera) is that the young laminae are flat; they
do not form a fiddlehead. Both subgenera of Didymoglos-
sum lack roots but have root-like rhizomes (Schneider,
2000). These are short, leafless rhizomes that are densely
pubescent with adhesive hairs.They help anchor the plant
to the substrate. (For comments about adhesive hairs and
false veins, see subgen. Didymoglossum.) The chromosome
number x=34 is considered derived within the family.The
name Microgonium is derived from the Greek mikros,
small + gonos, progeny, referring to the size of these ferns.
LITERATURE: Schneider, H. 2000. Morphology and anatomy of roots in the filmy fern tribe Trichomaneae H. Schneider (Hy-
menophyllaceae, Filicatae) and the evolution of rootless taxa. Bot. J. Linn. Soc. 132:29—46. Wessels Boer, J. G. 1962. The New
World species of Trichomanes sect. Didymoglossum and Microgonium. Acta Botanica Neerlandica 11: 277-330.
272
DESCRIPCIÓN: Epífitas; raíces ausentes; rizomas a 1
mm de ancho, largamente rastreros, ramificados, densa-
mente pubescentes, los pelos negruzcos; hojas a 6 cm
de largo, enteras a bipinnatífidas, márgenes glabros;
pecíolos cortos o ausentes, a menudo pubescentes con
pelos oscuros como los en el rizoma; costas presentes o
ausentes, si ausentes; venas catádromas, a menudo flabel-
ladas; venas falsas presentes como rayos cortos débiles
entre las venas verdaderas, falsa vena continua sub-
marginal presente; soros uno a varios, marginales en
la parte distal de la hoja, usualmente inmersados en la
lámina; indusios infundibuliformes o tubulares, truncados
(no bilabiados), sin márgenes oscuros; receptáculos
exertos; esporas verdes, triletes; x=34.
GÉNEROS PARECIDOS: Didymoglossum sect. Didymo-
glossum difiere por la ausencia de una falsa vena conti-
nua submarginale, indusios bilabiados (no truncados) con
bordes oscuros, y pelos marginales presentes. Polyphlebi-
um difiere por venas falsas ausentes y hojas más grandes.
COMENTARIOS: La sect. Microgonium es principal-
mente paleotropical y consta de casi 10 especies. Existen
como epífitas o sobre rocas musgosos en bosques húme-
dos. En su monografía de taxones neotropicales, Wessels
Boer (1962) reconoció 4 especies: T. ekmanii, T. godmanii, T.
hookeri y T. kapplerianum. Muchas veces están pasados por
alto a causa de su tamaño pequeño. Una característica
poco común de Didymoglossum (ambos subgéneros) es
que las láminas jóvenes son planas; no forman un cayado.
Los dos subgéneros de Didymoglossum faltan de raíces
pero tienen rizomas que asemejan a raíces (Schneider,
2000). Estos son rizomas cortas sin hojas que son dens-
amente pubescentes con pelos adhesivos. Ayudan anclar
la planta al sustrato. (Para comentarios sobre pelos ad-
hesivos y falsas venas, vease subgén. Didymoglossum.) El
número cromosómico x=34 se considera derivado den-
tro de la familia. El nombre Microgonium se deriva del
griego mikros, pequeño + gonos, progenie, refiriéndose al
tamaño de estos helechos.
Figure 133. A. Didymoglossum godmanii. B. D. ekmanii. (Modified from Mickel & Smith, 2004.)

273
LYCOPODIACEAE
Diphasiastrum Holub
DESCRIPTION: Terrestrial; main horizontal stems
subterranean or creeping; ultimate branchlets
dorsiventral, anisophyllous, with decussate
trimorphic leaves, one dorsal and one ventral rank
of narrow leaves, and two lateral ranks of broad
laterally compressed leaves, a few species with radially
arranged, isophyllous leaves; branchlet leaves herbaceous
throughout; strobili pedunculate; sporophylls subpeltate;
sporangia isovalvate, the epidermal cells with sinuate
anticlinal walls; spores reticulate on all faces; gametophytes
obconic, carrot-shaped; x=23.
DESCRIPCIÓN: Terrestres; tallos principales
horizantales subterráneos o reptantes; ramitas últimas
dorsiventrales, anisofílicas, con hojas decusadas
trimórfas, una hilera dorsal y una ventral de
hojas angostas, y dos hileras dorsolaterales de
hojas anchas comprimidas, unas especies con hojas
arregladas radialmente y isofílicas; hojas de las ramitas
herbaceas por todo parte; estróbilos pedunculados;
esporofilios subpeltatos; esporangios isovalvados, con
células epidérmicas sinuosas, las paredes anticlinales
sinuadas; esporas reticuladas en ambas caras; gametofitos
obcónicas, con forma de una zanahoria; x=23.

SIMILAR GENERA: Diphasium differs by the flattened GÉNEROS PARECIDOS: Diphasium difiere por las
ultimate branchlets having two or three rows of ventral achatadas ramitas últimas con dos o tres hileras de hojas
leaves. ventrales.

COMMENTS: Diphasiastrum consists of about 20 species COMENTARIOS: Diphasiastrum consta de casi 20

and is primarily northern temperate and circumboreal. especies y es principalmente en las zonas templadas del
It grows in shaded forests to (more abundantly) in norte y circunboreal. Existe en bosques sombreados
open sunny or partially shaded habitats. The genus was hasta (más abundantamente) ambientes abiertos o
monographed by Wilce (1965), and Holub (1975) parcialmente sombreados. El género fue monographiado
discussed the relationships to other groups. The flattened por Wilce (1965) y Holub (1975) discutió las relaciones
four-ranked ultimate branches, which occur in nearly a otros grupos. Las ramitas últimas achatadas con
all species of the genus, are distinctive. These branches cuatro hileras de hojas, las cuales ocurren en casi todas
often exhibit constrictions where growth has slowed or las especies, son distinctivas. Estas ramitas a menudo
ceased completely during the winter or (in the tropics) exhiben constricciones donde el crecimiento ha llegado
unfavorable growing season. Diphasiastrum is remarkable a ser más lento or parado completamente durante el
for its ability to form fertile interspecific hybrids with invierno o (en el trópico) estación de crecimiento no
viable spores capable of producing normal, functional favorable. Diphasiastrum es notable por su habilidad de
gametophytes. In Finland, a clone of D. complanatum was formar híbridos interespecíficos fértiles con esporas
250 m in diameter and estimated at more than 800 years viables capáces de producir gametofitos normal
old (Oinonen, 1967). There are five species in the United funcionales. En Finlandia, un clono de D. complanatum
States and Canada, and two (D. thyoides (Willd.) Holub fue 250 m en diámetro y estimado tener más que 800
[widespread] and D. falcatum B. Øllg. & P. G. Windisch años de edad (Oinonen, 1967). Hay cinco especies en
[s.e. Brazil]) in the Neotropics. The genus name is derived los Estados Unidos y Canadá y dos (D. thyoides (Willd.)
from Diphasium, another genus of Lycopodiaceae, and Holub [difundida] y D. falcatum B. Øllg. & P. G. Windish
the Greek suffix -astrum, false. The genus resembles [s.e. Brasil]) en el Neotrópico. El nombre genérico es
Diphasium. derivado de Diphasium, otro género de Lycopodiaceae,
y el sufijo griego –astrum, falso. El género se asemeja a
Diphasium.
LITERATURE: Aagaard, S. M. D., J. Greilhuber, X-C. Zhang & N. Wikström. 2009. Occurrence and evolutionaryy origins
of polyploids in the clubmoss genus Diphasiastrum (Lycopodiacae. Molecular Phylogenetics and Evolution 52: 746–754. Holub,
J. 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47: 97–110. Oinonen, E. 1967. Sporal regeneration of ground
pine (Lycopodium complanatum) in southern Finland in light of the size and age of its clones. Acta Forestalia Fennica 83: 76–85.
Øllgaard, B. & K. Tind. 1993. Scandinavian ferns. Rhodos, Copenhagen. Øllgaard, B. & P. G. Windisch. 2014. Lycopodiaceae
in Brazil. Conspectus of the family I. The genera Lycopodium, Austrolycopodium, Diphasium, and Diphasiastrum. Rodriguésia 62(2):
2014. Wilce, J. H. 1965. Section Complanata of the genus Lycopodium. Beihefte zur Nova Hedwigia 19: i–ix, 1–233.

274
Figure 134. Diphasiastrum. A. Spores of D. complanatum, proximal hemisphere (above) and distal (below). B. D. thyoides,
habit. C. Branchlests of D. complanatum showing seasonal growth contrictions; left, dorsal, right, ventral. D. Ultimate branchlet,
dorsal view. E. Ultimate branchlet, ventral surface; note medial row of reduced leaves. F. Carrot-shaped gametophyte of D.
digitatum with young sporophyte attached. G. D. complanatum. H. D. alpinum. I. D. tristachyum. (A, B, F from Øllgaard and
Tind, 1993; G, H, I from Wilce, 1965; B, D, E from Mickel and Smith, 2004.)

275
LYCOPODIACEAE
Diphasium C. Presl ex Rothm.
DESCRIPTION: Terrestrial; main horizontal stems
subterranean, creeping to subscandent; ultimate
branchlets dorsiventral, anisophyllous, with
dimorphic leaves, two dorsolateral ranks of broad
alternate leaves, two or three ventral ranks
of narrow membranous-tipped leaves; strobili
pedunculate or sessile; sprophylls subpeltate; sporangia
with cells sinuate, finely curled side walls; spores reticulate
with large meshes and unornamented proximal faces;
gametophytes obconic; x=34–36, ca. 90.
DESCRIPCIÓN: Terrestres; tallos principales
horizantales subterráneos, reptantes a subescandentes;
ramitas últimas dorsiventrales, anisofílicos, con
hojas dimórfas, 2 hileras dorsolaterales de hojas
alternas anchas, 2 o 3 hileras ventrales de hojas,
cada hojas con apices membranáceos; estróbilos
pedunculados o sésilies; esporofilios subpeltatos;
esporangios con células sinuosas, las paredes laterales
finamente curvadas; esporas reticuladas con grandes
redes, las caras proximales no ornamentadas; gametofitos
obcónicas; x=34–36, ca. 90.

SIMILAR GENUS: Diphasiastrum differs by ultimate GÉNERO PARECIDO: Diphasiastrum difiere por las
branchlets with four ranks of leaves: two lateral, one ramitas últimas con cuatro hileras de hojas: dos laterales,
dorsal, and one ventral. una dorsal y una ventral.

COMMENTS: Diphasium consists of five species,
with two in the Neotropics: D. jussiaei (Desv. Ex Poir.)
C. Presl ex Rothm. and D. gayanum (Rémy) Holub. The
first is widespread from Costa Rica to southern Brazil
and Argentina, the second is restricted to Chile, Juan
Fernandez, and Argentina. A third species occurs from
Indonesia to New Zealand: D. scariosum (G. Forst.)
Rothm. The plants are often creeping, trailing, or hanging
over banks, and occur from 1700—3700 m.The flattened
ultimate branches with 2 or 3 ranks of scarious leaves
below are distinctive. Unlike Diphasiastrum, there is no
medial, dorsal row of leaves. The genus name is derived
from the Greek diphasios, two-fold. It is unknown what
this refers to, but perhaps it is the two ranks of wide
lateral leaves.
COMENTARIOS: Diphasium consta de cinco especies
con dos en el Neotrópico: D. jussiaei (Desv. Ex Poir.) C.
Presl ex Rothm. y D. gayanum (Rémy) Holub. La primera
es muy difundida desde Costa Rica hasta el sur de Brasil
y Argentina, y la segunda es restringida a Chile, Juan
Fernández y Argentina. Una tercera especie existe desde
Indonesia hasta Nueva Zelandia: D. scariosum (G. Forst.)
Rothm. Las plantas son a menudos reptantes o colgantes
sobre taludes, y ocurren 1700–3700 m. Las ramitas
últimas achatadas con 2 o 3 hileras de hojas escariosas
en el envés son distinctivas. A diferente de Diphasiastrum,
no hay una hilera medial de hojas. El nombre genérico se
deriva del griego diphasios, duplo. No se conoce a que
refiere, pero tal vez es los dos hileras laterals de hojas
anchas.

LITERATURE: Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the Missouri Botanical Garden 79:
687–717. Øllgaard, B. & P. G. Windisch. 2014. Lycopodiaceae in Brazil. Conspectus of the family I. The genera Lycopodium,
Austrolycopodium, Diphasium, and Diphasiastrum. Rodriguésia 62(2): 2014 (on-line only).

276
Figure 135. Diphasium jussiaei. A. Ultimate branchlet, ventral surface. B. Ultimate branchlet, dorsal surface. C. Strobilus with
caudate apex. D. Habit. (Original, ©Robbin C. Moran, 2016.)

277
ATHYRIACEAE (EUPOLYPODS II)
Diplazium Sw.
]DESCRIPTION: Terrestrial; rhizomes creeping to erect,
scaly, the scales not clathrate; sterile and fertile leaves
monomorphous; petioles with 2 vascular bundles ba-
sally, these uniting distally to form a “U”; laminae
simple to 4-pinnate; veins free or areolate, the areoles
without included veinlets; sori elliptic to linear, usually
paired back to back along a single vein, indusiate,; an-
nulus consisting of 15–20 cells; spores bilateral, brown;
x=41.
SIMILAR GENERA: Asplenium differs by more than 20
cells per annulus and, typically, unpaired sori. Whereas
Diplazium is always terrestrial, Asplenium can be terrestri-
al, epiphytic or saxicolous, Another difference is that the
annulus of Asplenium generally remains extended after
dehiscence, and often the distal part of the capsule splits
divaricately, resembling two small wings. The annulus of
Diplazium returns to its original position after dehiscence,
forming an elongate “C,” and the distal part of the cap-
sule walls do not split apart. Callipteris differs by rhi-
zome (and often laminar) scales with black margins and
black, bifid teeth.
COMMENTS: Diplazium consists of about 350 species,
of which about 150 are neotropical. It grows primarily
in wet forests at low to middle elevations. In the Neo-
tropics, few fern genera have received as little study as
Diplazium. Consequently, there are numerous taxonomic
and nomenclatural problems, and these require mono-
graphic study to solve. The sori arranged back-to-back
along the same vein are characteristic of the genus and
are called diplazioid sori. Many species have distinctive
hairs on the grooves of the rachises, and a peculiar flange
of tissue where the costules join the pinna rachises. The
genus name is derived from the Greek diplazios, double.
The indusia are often double, or back-to-back along the
same vein.
LITERATURE: Stolze, R. G. & L. Pachecho. 1994. Diplazium. In: Harling, G. & L. Andersson. Flora of Ecuador, 14 (5B). Polypo-
diaceae-Dryopteridoideae-Physematieae 49: 1–108. Testo, W. L., M. Sundue & A. Vasco. 2017. Diplaziium hybrids involving
D. plantaginifolium and D. ternatum from Mexico and Central America. Brittonia 69: 295–306.
278
DESCRIPCIÓN: Terrestres; rizomas rastreros a erec-
tos, escamosos, las escamas no clatradas; hojas estériles
y fértiles monomorfas; pecíolos con 2 haces vascu-
lares basalmente, éstos uniéndose distalmente
para formar un haz vascular en forma de “U”;
láminas simples a 4-pinnadas; venas libres o areoladas
sin venillas incluidas; soros elípticos o lineares, usual-
mente pareados de par en par por una sóla vena,
con indusios,; anillo consiste de 15–20 células; esporas
bilaterales, pardas; x=41.
GÉNEROS PARECIDOS: Asplenium difiere por más
de 20 células por anillo y, típicamente, soros no pareados.
Diplazium es siempre terrestre, pero Asplenium puede
puede ser terrestre, epífita, o saxícola. Otra diferencia es
que el anillo de Asplenium generalmente se queda exten-
dido después de dehiscencia, y a menudo la parte distal
de la cápsula se hunde divaracatamente, pareciendo dos
pequeñas alas. El anillo en Diplazium regresa a su posición
original después de la dehiscencia, formando una “C” alar-
gada, y la parte distal de la cápsula no se hunde. Callipteris
difiere por escamas del rizoma (y a menudo las de las
láminas) con márgenes negros y dientes bífidos negros.
COMENTARIOS: Diplazium consta de casi 350 espe-
cies, de las cuales casi 150 son neotropicales. Crece prin-
cipalmente en bosques húmedos en elevaciones bajas a
medias. En el Neotrópico, pocos géneros de los helechos
han recibidos tan poco estudio como Diplazium. En con-
secuencia, hay numerosos problemas taxonómicos y
nomenclaturales, y requieren estudio monográfico para
resolverlos. Los soros arreglados espalda a espalda por
la misma vena son característicos del género y se llaman
soros diplazioides. Muchas especies tienen pelitos ca. 0.1
mm de largo en los surcos de los ráquises, y una pliega de
tejido donde las cóstulas se unen con los ráquises de las
pinnas. El nombre genérico se deriva del griego diplazios,
doble. A menudo los inducíos son dobles, o espalda a
espalda por la misma vena.
Figure 136. Diplazium lonchophyllum. A. Leaf (hoja). B. Pinna with sori (pinna con soros). C. Diplazioid sori (soros
diplazioides). (Original, ©Robbin C. Moran, 2016.)

279
ATHYRIACEAE (EUPOLYPODS II)
Diplazium (Ceratolepis clade)
DESCRIPTION: Terrestrial; rhizomes erect or decum-
bent, the scales with black borders and black, bifid
marginal teeth; sterile and fertile leaves monomor-
phous; petioles with 2 vascular bundles basally, these
uniting distally to form a U-shaped strand; laminae simple
to 1-pinnate-pinnatifid, broadly cut, never finely divided;
veins areolate or rarely free, the areoles without includ-
ed veinlets; sori linear, indusiate, usually paired back to
back along a single vein; annulus consisting of 15–20 cells;
spores bilateral, brown; x=41.
SIMILAR GENERA: The other species of Diplazium are
distinguished by rhizome scales without dark borders.
Also, most species of Diplazium have free veins, where-
as the Callipteris group almost always has anastomosing
veins.
COMMENTS: The Callipteris group consists of about 20
species in the New World and 70 in the Old. It grows
on wet, shaded, forest floors from 0–2000 m. The group
appears to be monophyletic but nested within Diplazium.
The laminae are often fleshy, and the teeth of the rhizome
scales are dark and bifid. The name Callipteris is derived
from the Greek callos, beautiful, + pteris, fern.
LITERATURE: Pacheco, L. & R. C. Moran. 1999. Monograph of the neotropical species of Callipteris (Woodsiaceae) with
anastomosing veins. Brittonia 51: 343–388.
280
DESCRIPCIÓN: Terrestres; rizomas erectos a decum-
bentes, las escamas del rizoma con márgenes negros
y dientes bífidos negros; hojas estériles y fértiles mono-
morfas; pecíolos con 2 haces vasculares basalmente,
éstos uniéndose distalmente para formar un haz vascular
en forma de “U”; láminas simples a 1-pinnada-pinnatífidas,
cortadas ampliamente (nunca finamente divididas); venas
areoladas or raras veces libres, las areolas sin venillas in-
cluidas; soros lineares, con indusios, usualmente parea-
dos de par en par por una sóla vena; anillo consiste de
15–20 células; esporas bilaterales, pardas; x=41.
GÉNEROS PARECIDOS: Los otros especies de Dipla-
zium se distinguen por las escamas del rizoma sin bordes
oscuras. También, la mayoría de las especies de Diplazium
tienen venas libres, mientras que los en el grupo de Cal-
lipteris usualmente son anastomosantes.
COMENTARIOS: El grupo Callipteris consta de casi 20
especies en el Nuevo Mundo y 70 en el Viejo. Existe en
sotobosques húmedos y sombreados de 0–2000 m. El
grupo se parece ser monofilético pero anidado dentro
de Diplazium. Las láminas a menudo son carnosas, y los
dientes de las escamas del rizoma son oscuros y bífidos.
El nombre Callipteris se deriva del griego callos, hermoso,
+ pteris, helecho.
Figure 137. Diplazium (Ceratolepis clade) chimborazense (From Pacheco & Moran, 1999.)

281
GLEICHENIACEAE
Diplopterygium (Diels) Nakai
DESCRIPTION: Terrestrial; rhizomes long-creeping,
scaly; leaves indeterminate, of periodic growth, bearing
pairs of opposite pinnae, the distalmost pair with a scaly
resting bud between them; pinnae not forked, pin-
nate-pinnatisect or 2-pinnate; penultimate segments
pectinate; veins once-forked; sori round, not indusiate;
sporangia 2–5 per sorus; spores trilete; x=56.
DESCRIPCIÓN: Terrestres; rizomas largamente ras-
treros, escamosos; hojas indeterminadas, de crecimiento
periódico, llevando pares de pinnas opuestas; el par más
distal con una yema latente escamosa entre sí; pin-
nas no surcadas, pinadas-pinnatisectas o 2-pinnadas;
segmentos penúltimos pectinados; venas una vez fur-
cadas; soros redondos, no indusiados; esporangios 2–5
por soro; esporas triletes; x=56.

SIMILAR GENERA: Sticherus and Dicranopteris differ GÉNEROS PARECIDOS: Sticherus y Dicranopteris difi-
by forked pinnae. Dicranopteris and Gleichenella differ by eren por pinnas furcadas. Dicranopteris y Gleichenella di-
pubescent axillary buds, veins 2–4 times forked, and spo- fieren por yemas axilares pubescentes, venas 2–4 veces
rangia 8–15 per sorus. furcadas, y esporangios 8–15 por soro.

COMMENTS: Diplopterygium has about 10 spe-
cies worldwide, all in Asia (India to Japan and Malesia).
Only one species occurs in the Neotropics: D. bancroftii
(Hook.) A. R. Sm. This species generally grows in road-
sides and steep banks at middle elevations (1000–2600
m). This genus is sometimes treated in Gleichenia, but it is
maintained here as distinct because of its differences in
pinna architecture and chromosome number. The genus
name is derived from the Greek diploos, double + pterygi-
um, little wing. The opposite pinnae resemble wings.
COMENTARIOS: Diplopterygium tiene casi 10 especies
mundialmente, todos en Asia (India a Japón y Malesia).
Solamente una especie existe en el Neotrópico: D. ban-
croftii (Hook.) A. R. Sm. Esta especie se encuentra usual-
mente en taludes y al lado de los caminos en elevaciones
medias (1000–2600 m). Algunas veces este género se
trata en Gleichenia, pero aquí se mantiene distinto debido
a sus diferencias en la arquitectura de las pinnas y en el
número cromosómico. El nombre genérico se deriva del
griego diploos, doble + pterygium, alita. Las pinnas opues-
tas se asemejan alas.

LITERATURE: Ching, R. C. 1940. On the genus Gleichenia Smith. Sunyatsenia 5: 269–288. Holttum, R. E. 1957. Florae Male-
sianae Precursores XVI. On the taxonomic subdivision of the Gleicheniaceae with descriptions of new Malaysian species and
varieties. Reinwardtia 4: 257–280. Holttum, R. E. 1957. Morphology, growth habit, and classification in the family Gleicheniaceae.
Phytomorphology 7: 168–184. Maxon, W. R. 1909. Gleicheniaceae. North American Flora 16: 53-63. Østergaard Andersen,
E. & B. Øllgaard. 1996. A note on some morphological terms of the leaf in the Gleicheniaceae. Amererican Fern Journal 86:
52–57. Østergaard Andersen, E. & B. Øllgaard. 2001. Gleicheniaceae. In: G. Harling & L. Andersson, editors. Flora of Ecuador
66: 105–170. Underwood, L. M. 1907. American ferns VIII. A preliminary review of the North American Gleicheniaceae. Bulletin
of the Torrey Botanical Club 34: 243–262.

282
Figure 138. A–F. Diplopterygium bancroftii (Hook.) A. R. Sm. (From Mickel & Smith, 2004.)

283
PTERIDACEAE
Doryopteris J. Sm.
DESCRIPTION: Terrestrial or rupestral; rhizomes erect,
scaly, the scales bicolorous with a dark central strip
and wide pale borders; leaves monomorphic or slightly
dimorphic; petioles brown or black, with 1 or 2 vascular
bundles, terete or flattened adaxially, if flattened then with
narrow wings on either side; laminae glabrous, with
black sclerotic band, this sometimes restricted to
small sinuses, rarely cordiform, deltate or reniform; veins
netted or (in 8 species) free, often hard to see because
of the thick lamina, the areoles without included veinlets;
sori continuous along the margin; indusia formed by
the recurved margin; x=30.
SIMILAR GENERA: Bommeria differs by pubescent
laminae and free veins (only two species of Doryopteris
have free veins). Hemionitis differs by sori spreading be-
tween the midrib and margin.
DESCRIPCIÓN: Terrestres o rupícolas; rizomas erectos,
escamosos, las escamas bicoloras estrechamente lance-
oladas, con un centro oscuro y ancho, los bordes pardo
claro; hojas monomorfas a casi dimorfas; pecíolos pardos
a negros, con 1 o 2 haces vasculares, terete o aplanado
adaxialmente, en este caso acompañado por bordes o
emarginaciones laterales estrechas; láminas glabras con
una banda esclerótica, esto a veces restingida a es-
cotaduras pequeñas, raramente cordiformes, deltadas
o reniformes; nervaduras generalmente oscurecidas por
la gruesa lámina; areoladas o (en 8 especies) libres, las
aréolas sin nérvulos incluidos; soros continuos a lo lar-
go del margen; indusios formados por el margen foliar
reflexo; x=30.
GÉNEROS PARECIDOS: Bommeria difiere por lámi-
nas pubescentes y venas libres (solo dos especies de Do-
ryopteris tienen venas libres). Hemionitis difiere por los
soros difundidos entre la costa y el margen.

COMMENTS: Doryopteris has 39 species and occurs
in the Neotropics, South Africa, and Hawaii. In the Neo-
tropics, 34 species occur, mostly occur in the Planalto re-
gion of southern Brazil. The genus typically grows in dry,
rocky habitats. The most constant character to distinghuis
the genus is the black marginal sclerenchymatous band,
this sometimes restricted to small sinuses. Phylogenetic
studies have shown that Doryopteris is most closely re-
lated to Trachypteris and Adiantopsis of South America
and Pellaea calomelanos of Africa (Gastony & Rollo, 1995,
1998). As defined here, the genus includes Ormopteris J.
Sm. and Lytoneuron (Klotzsch) J. C.Yesilyurt, both of which
are monophyletic. The genus name is derived from the
Greek dory, lance + pteris, fern. It alludes to the shape of
the blade in some species.
COMENTARIOS: Doryopteris tiene 39 especies e ex-
iste en los Neotropics, Sur de África y Hawaii. En los
Neotropics, 34 especies existen, la mayoría en la región
Planalto del sur de Brasil. El género crece típicamente
en ambientes secas y rocosas. El carácter más constante
distinguirlo es la banda negra marginal esclerótica, esto a
veces restingida a los sinuses pequeñas. Estudios molec-
ulares han demostrado que Doryopteris es más cercana-
mente relacionado a Trachypteris y Adiantopsis de Améri-
ca del Sur y Pellaea calomelanos de África (Gastony &
Rollo, 1995, 1998). Como definido aquí, el género incluye
Ormopteris J. Sm. y Lytoneuron (Klotzsch) J. C. Yesilyurt., los
cuales son monofiléticos. El nombre genérico se deriva
del griego dory, lanza + pteris, helecho. Refiere a la forma
de las láminas en algunas especies.

LITERATURE: Gastony, G. J. & D. R. Rollo. 1995. Phylogeny and generic circumscriptions of cheilanthoid ferns (Pteridaceae:
Cheilanthoideae) inferred from rbcL nucleotide sequences. American Fern Journal 85: 341–360. Gastony, G. J. & D. R. Rollo.
1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern United States and adjacent Mexico–a molecular
phylogenetic reassessment of generic lines. Aliso 17: 131–144. Tryon, R. M. 1942. A revision of the genus Doryopteris. Contri-
butions to the Gray Herbarium 143: 1–80. Tryon, R. M. 1944. Dynamic phytogeography of Doryopteris. American Journal of
Botany 31: 471–473. Tryon, R. M. 1962. The fern genus Doryopteris in Santa Catarina and Rio Grande do Sul, Brazil. Sellowia
14: 51–59. Yesilyurt, J. C., T. Barbará, H. Schneider, S. Russell, A. Culham & M. Gibby. 2015. Identifying the generic limits
of the Cheilanthoid genus Doryopteris. Phytotaxa 221(2): 101–122.

284
Figure 139. Doryopteris. A–C. Doryopteris palmata. D, E. Doryopteris concolor. (From Mickel & Smith, 2004.)

285
LOMARIOPSIDACEAE (EUPOLYPODS II)
Dracoglossum Christenh.
DESCRIPTION: Terrestrial; Rhizomes short-creeping,
polystichous, scaly; sterile and fertile leaves monorphic,
simple, entire or slightly sinuate, typically with an apical
bud; venation obscure, reticulate, with 4 or 5 rows of
areoles between the midrib and margin; sori round,
placed regularly in two rows between the main lateral
veins; indusia present or absent, when present circular,
thick, peltate; spores brown. x=?.
DESCRIPCIÓN: Terrestres; rizomas cortamente
rastreros, polísticos, escamosos; hojas estériles y fértiles
monomorfas, simples, enteras o levemente sinuadas,
típicamente con un brote apical; nervación obscura,
reticulada, con 4 a 5 hileras de areolas entre la costa y el
margen; soros redondos; indusios presentes o ausentes,
cuando presentes circulares, gruesos, peltatdos; esporas
pardas; x=?.

SIMILAR GENERA: Tectaria usually differs by erect GÉNEROS PARECIDOS: Tectaria difiere usualmente
rhizomes, rachises and costa puberulent adaxially, and two por rizomas erectos, ráquises y costas puberulentos
layers of perispore separated by pillars. adaxialmente y dos capas de periesporas separadas por
columnas.

COMMENTS: Dracoglossum occurs in Jamaica, the
Lesser Antilles, from southern Mexico to Bolivia, and the
Guianas, Venezuela, and northern Brazil. It is terrestrial or
on boulders and talus slopes, 0–900 m. The sole species
is D. plantagineum (Jacq.) Christenh. Sometimes a second
species, D. sinuatum (Fée) Christenh., is distinguished by
the presence of indusia. Indusiate plants are found in the
Guyanan Shield region and northwestern Ecuador. Other
than the indusia, the two species are indistinguishable.The
vegetative characters given by Christenhusz (2007) to
distinguish the two species do not hold up. For a long
time D. plantagineum was classified in Tectaria, which it
greatly resembles, but chloroplast DNA sequence data
place in the Lomariopisdaceae, sister to Lomariopsis. The
gametophytes of Dracoglossum are elongate and resemble
those of Lomariopsis, but they occur on rock, not the bases
of tree trunks (pers. obs.). The genus name is derived
from the Greek draco, dragon + glossum, tongue, from
a fancied resemblance of the simples leaves to a tongue.
COMENTARIOS: Dracoglossum existe en Jamaica, las
Antillas Menores, desde el sur de México hasta Bolivia, y
las Guianas,Venezuela y Brasil septentrional. Es terrestre o
sobre peñas y taludes rocosos, 0–900 m. La única especie
es D. plantagineum (Jacq.) Christenh. Algunas veces
una segunda especie, D. sinuatum (Fée) Christenh., se
distingue por la presencia de indusios. Plantas indusiadas
se encuentran en la región del Escudo Guyanés y
noroeste de Ecuador. A no ser por los indusios, las dos
especies son indistinguibles. Las características dadas por
Christenhusz (2007) para distinguir las dos especies no son
consistentes. Por mucho tiempo D. plantagineum estaba
clasificada en Tectaria, pero datos de ADN cloroplástico
la ubican en Lomariopsidaceae, hermano de Lomariopsis.
Los gametofitos de Dracoglossum son alaragados y se
asemejan los de Lomariopsis, pero existen sobre rocas, no
las bases de árboles (obs. pers.). El nombre genérico se
deriva del griego draco, dragón + glossum, lengua, de una
semejanza imaginada entre las hojas simples y una lengua.

LITERATURE: Christenhusz, M. J. M. 2007. Dracoglossum, a new Neotropical fern genus (Pteridophyta). Thaiszia 17: 1–10.
Christenhusz, M. J. M., M. Jones & S. Lehtonen. 2013. Phylogenetic placement of the enigmatic fern genus Dracoglossum.
American Fern Journal 103: 131–138.

286
Figure 140. Dracoglossum plantagineum. A. Rhizome and fertile leaf. B. Transverse section of petiole. C. Apical bud of lamina.
D. Venation. E. Indusiate sorus. F. Non-indusiate sorus (Original, © Robbin C. Moran, 2014.)

287
TECTARIACEAE (EUPOLYPODS II)

Draconopteris Li Bing Zhang & Liang Zhang..

DESCRIPTION: Terrestrial; rhizomes erect to decument; DESCRIPCIÓN: Terrestres; rizomas erectos a

leaves arranged radially around the rhizome; sterile and
fertile leaves monomorphic; laminae 1-pinnatifid; rachis
winged; pinnae 3–4 pairs, entire except for a single basal
basiscopic lobe on the basal pinnae; rachises adaxially
glabrous, ungrooved; veins reticulate; sori round; indusios
ausentes; perispores cristate; x=?
decumbentes; hojas arregladas radialmente alrededor
del rizoma; hojas estériles y fértiles monomorfas; laminas
profundamente pinatifid; raquises alados; pinnas 3–4 pairs,
enteras con excepcion de las pinnas basales que tienen
un solo lobulo basiscopico; raquises adaxialmente glabros
y no surcados; nervación reticulada; soros redondos;
indusios ausentes; esporas cristadas; x=?.

SIMILAR GENERA: Tectaria usually differs by rachises
not winged, rachis and costa puberulent adaxially, and lack
of toasted almond (cyanide) odor. Hypoderris differs by
crepping rhizomes, distichous leaves, lack of a basal lobe
on the basal pinnae, and spiny spores.
GÉNEROS PARECIDOS: Tectaria difiere usualmente
por los ráquises no alados, ráquis y costa puberulento
adaxialmente y falta de olor de almendras tostados
(cianuro). Hypoderris difiere por rizomas rastreros, hojas
dísticas, falta del lóbulo en las pinnas basales y esporas
espinosas.

COMMENTS: Draconopteris contains a single species,
D. draconoptera (D. C. Eaton) Li Bing Zhang & Schuettp.,
distributed from Nicaragua to Peru and western Brazil. It
grows in wet forests from 0–700 m. It is distinctive by its
large leaves (up to 1.5 m long), basal pinnae connected
to the suprabasal ones by a wing along the rachis, and
many small nonindusiate sori. When crushed, the rhizome
and petioles have the sweet smell of toasted almonds
(cyanide). Draconopteris is sister to Malaifilix grandidentata
(Ces) Li Bing Zhang & Schuettp., of southeast Asia.The
genus name is derived from the Greek draco, dragon +
pteris, fern. What this is refers to is unclear. .
COMENTARIOS: Draconopteris contiene una sola
especie, D. draconoptera (D. C. Eaton) Li Bing Zhang &
Schuettp., distribuido desde Nicaragua hasta Perú y Brasil
occidental. Habita bosques húmedos de 0–700 m. Es
distinctiva por sus hojas grandes hasta 1.5 m de largo,
pinnas basales conectadas a las suprabasales por una ala
por el ráquis y muchos soros pequeños no indusiados.
Cuando estrujado, el rizoma y pecíolos tienen el olor
dulce de almendras tostados (cianuro). Draconopteris es
género hermano a Malaifilix grandidentata (Ces) Li Bing
Zhang & Schuettp., de sureste Asia. El nombre genérico
se deriva del griego draco, dragón + pteris, helecho. Se
desconoce a lo que se refiere.

LITERATURE: Zhang, L., E. Schuettpelz, C. J. Rothfels, X-M. Zhou, X-F. Gao & Li-Bing Zhang. 2016. Circum-
scription and phylogeny of the fern family Tectariaceae based on plastid and nuclear markers, with the description of
two new genera: Draconopteris and Malaifilix (Tectariaceae). Taxon 65: 723–738.

288
Figure 141. Draconopteris draconoptera. A. Right half of leaf. B. Venation and sori. (From Shimek, 1897.)

289
DRYOPTERIDACEAE (EUPOLYPODS I)
Dryopteris Adans.
DESCRIPTION: Terrestrial or more rarely epiphytic or
rupestral; rhizomes compact, erect or ascending, scaly;
leaves monomorphic; petioles with more than three vas-
cular bundles, these arranged in a U with the two adaxial
ones the largest; laminae 2- to 3-pinnate-pinnatifid, less
commonly 1-pinnate-pinnatifid, the apices gradually ta-
pered and pinnatifid (not imparipinnate); pinnules of the
lower pinnae arranged isodromically or catadromically;
rachises and costae grooved adaxially, the grooves more
or less confluent at their junctures; veins free; sori round;
indusia present, attached at the base of a narrow
sinus, not peltate, reniform or orbicular; spores mono-
lete, winged; x=41.
DESCRIPCIÓN: Terrestres o menos comúnamente
epífitas o rupícolas; rizomas compactos, erectos o as-
cendentes, escamosos; hojas monomorfas; pecíolos con
más tres haces vasculares, estas arregladas en forma de U
con los dos adaxiales lo más grandes; láminas 2- o 3-pin-
nadas, menos común 1-pinnada-pinnatífidas, los ápices
gradualmente reducidas y pinatífidas (no imparipinadas);
pínnulas de las pinnas inferiores arregladas isodrómica-
mente o catadrómicamente; ráquises y costas sulcados
adaxialmente, los surcos más o menos confluentes en
las uniones; venas libres; soros redondos; indusios pre-
sentes, unidos en la base de un sinus estrecho, no
peltados, reniformes o orbiculares; esporas monoletes,
aladas; x=41.

SIMILAR GENERA: Polystichum usually differs by 1- or
2-pinnate laminae, the base of the pinnae or pinnules
slightly elongate on the acroscopic side, and the spinulose
lamina margins. Arachniodes differs by anadromic blades
and creeping rhizomes. Lastreopsis differs by the upper
surfaces of the rachises and costae rounded or flat (not
grooved). Polybotrya is (mostly) a climbing epiphytic fern
and has dimorphic sterile and fertile leaves. Saccoloma dif-
fers by an omega-shaped vascular bundle in the base of
the petiole, and marginal or submarginal sori.
COMMENTS: Dryopteris consists of about 225 species
worldwide and is well represented in the northern tem-
perate zones, especially in Asia. In the Neotropics it grows
in middle to high elevation forests and contains about
12 neotropical species, most all of which occur in Me-
soamerica. A subgeneric classification of the genus was
proposed by Frasier-Jenkins (1989). The genus name is
derived from the Greek drys, tree + pteris, fern.
GÉNEROS PARECIDOS: Polystichum usualmente difi-
ere por láminas 1- a 2-pinnadas, la base de las pinnas y
pínnulas alargada en el lado acroscópico, y los márgenes
espinulosos de las láminas. Arachniodes difiere por láminas
anadrómicas y rizomas reptantes. Lastreopsis difiere por
el haz de las ráquises y costas redondeado o achatado
(no surcado). Polybotrya es (principalmente) un helecho
trepador epífito y tiene hojas estériles y fértiles dimorfas.
Saccoloma difiere por una haz vascular en forma de una
omega en la base del pecíolo y soros marginales o sub-
marginales.
COMENTARIOS: Dryopteris consta de casi 225 espe-
cies mundial y es bien representado en las zonas temp-
ladas del norte, especialmente en Asia. En el Neotrópico
crece en bosques de elevaciones medias y altas y con-
tiene alrededor de 12 especies neotropicales, la mayoría
de las cuales existen en Mesoamérica. Una clasificación
subgenérica del género fue propuesta por Frasier-Jenkins
(1989). El nombre genérico se deriva del griego drys, ar-
bol + pteris, helecho.

LITERATURE: Christensen, C. 1913. A monograph of the genus Dryopteris. Part I. The tropical American pinnatifid-bipinnati-
fid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. ser. 7, 10: 55–282. Frasier-Jenkins, C. R. 1986. A classification of
the genus Dryopteris (Pteridophyta : Dryopteridaceae). Bulletin of the British Museum (Natural History), Botany 14(3): 183–218.
Frasier-Jenkins, C. R. 1989. A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. Bulletin
of the British Museum (Natural History), Botany 18: 323–477.

290
Figure 142. A, B. Dryopteris muenchii. C, D. D. arguta. E, F. D. futura. G, H. D. filix-mas. (From Mickel & Smith, 2004.)

291
DRYOPTERIDACEAE
Elaphoglossum sect. Amygdalifolium (H. Christ) Mickel & Atehortúa
DESCRIPTION: Plants epiphytic; rhizomes long-
creeping, the scales linear to lanceolate; aerophores
absent(?); phyllopodia present; blades glabrous or
nearly so, entire, thin, sometimes reddish when young,
the scales (when present) minute, non-setose, laciniate-
pectinate to stellate, minute glandular hairs absent; veins
free; hydathodes present; spores monolete, brown or
blackish; x=41.
DESCRIPCIÓN: Plantas epífiticas; rizomas largamente
rastreros, las escamas lineares a lanceoladas; aeróforos
ausentes(?); filopódios presentes; láminas glabras a
subglabras, enteras, delgadas, a veces rojizas cuando
jóvenes, las escamas diminutas, no setosas, lacinadas-
pectinadas a estrelladas, pelos diminutos glandulares
ausentes; venas libres; hidatodos presentes; esporas
monoletes, pardas o negruzcas; x=41.

SIMILAR GENERA: The simple-leaved species of
Polypodiaceae (e.g., Campyloneurum, Microgramma,
Pleopeltis) differ by reticulate veins and round sori. The
other sections of Elaphoglossum differ by having at least
one of the following characters: hydathodes absent,
conspicuously scaly laminae, or phyllopidia absent.
GÉNEROS PARECIDOS: Las especies de las
Polypodiaceae con hojas enteras (e.g., Campyloneurum,
Microgramma, Pleopeltis) difieren por nervadura reticulada
y soros redondos. Las otras secciones de Elaphoglossum
difieren por tener a lo menos una de las características
siguientes: hidatodos ausentes, láminas conspicuosamente
escamosas, o filopodios ausentes.

COMMENTS: Section Amygdalifolium is neotropical and COMENTARIOS: Sección Amygdalifolium es neotropical

consists of a single species:E.amygdalifolium (Mett.) H.Christ.
It occurs from 0–1500 m in Cuba and Central America
to Colombia and Ecuador (restricted to the western side
of the Andes). No other species of Elaphoglossum has
the combination of characters given in boldface in the
description above. The species is sister to the rest of the
genus (Rouhan et al., 2004). Plants in Central America
have reddish young leaves—a characteristic unique in
the genus—but plants from Ecuador apparently lack this
color (pers. obs.). Elaphoglossum amygdalifolium might
eventually prove to be hemiepiphytic, a pleisiomorphic
condition. Plants appear to start near the bases of trunks
and then climb. Such plants might produce at their bases
long, straight roots that grow into the soil where they
ramify. This growth form, which occurs in the outgroup
genera (Anapausia, Lomagramma, Teratophyllum), needs
to be searched for. The sectional name is derived from
the Latin amygdalus, almond + folium, leaf. The leaves
resemble that of an almond.
y consta de una sola especie: E. amygdalifolium (Mett.) H.
Christ. Existe desde 0–1500 m en Cuba y América Central
hasta Colombia y Ecuador (restingada al lado occidental
de los Andes). Ninguna otra especie de Elaphoglossum
tiene la combinación de caracteres dados en negritos en
la descripción arriba. La especie es hermana al resto del
género (Rouhan et al., 2004). Plantas de América Central
tienen hojas jóvenes rojizas—una característica única
en el género—pero plantas de Ecuador aparentemente
faltan este color (pers. obs.). Elaphoglossum amygdalifolium
puede ser hemiepifítica, una condición pleisiomórfica.
Plantas aparecen empezar cerca las bases de los troncos
y luego suben. Es posible que tales plantas produzcan en
sus bases raíces largas rectas que van directamente al
suelo donde ramifican. Esta forma de crecimiento, que
existe en los grupos externos (Anapausia, Lomagramma,
Teratophyllum), se necesita buscar. El nombre genérico se
deriva del latín amygdalus, almendra + folium, hoja. Las
hojas se asemejan las de una almendra.

LITERATURE: Mickel, J. T. & L. Atehortúa G. 1980. Subdivision of the genus Elaphoglossum. American Fern Journal 70:
47–68. Rouhan, G., J-Y. Dubuisson, F. Rakotondrainible, T. J. Motley, J. T. Mickel, J-N. Labat & R. C. Moran. 2004.
Molecular phylogeny of the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences:
contributions of species from the Indian Ocean area. Molecular Phylogenetics and Evolution 33: 745–763.

292
Figure 143. Elaphoglossum amygdalifolium. A. Habit. B. Long-creeping rhizome and dark phyllopodium. C. Venation, showing
hydathodes. (Original, ©Robbin Moran, 2016.)

293
DRYOPTERIDACEAE
Elaphoglossum Schott ex J. Sm., sect. Elaphoglossum
DESCRIPTION: Plants epiphytic; rhizomes long-
creeping, the scales linear to lanceolate; aerophores
absent(?); phyllopodia present; blades glabrous or
nearly so, entire, thin, sometimes reddish when young,
the scales (when present) minute, non-setose, laciniate-
pectinate to stellate, minute glandular hairs absent; veins
free; hydathodes present; spores monolete, brown or
blackish; x=41.
SIMILAR GENERA: The simple-leaved species of
Polypodiaceae (e.g., Campyloneurum, Microgramma,
Pleopeltis) differ by reticulate veins and round sori. The
other sections of Elaphoglossum differ by having at least
one of the following characters: hydathodes absent,
conspicuously scaly laminae, or phyllopidia absent.
COMMENTS: Section Amygdalifolium is neotropical and
consists of a single species:E.amygdalifolium (Mett.) H.Christ.
It occurs from 0–1500 m in Cuba and Central America
to Colombia and Ecuador (restricted to the western side
of the Andes). No other species of Elaphoglossum has
the combination of characters given in boldface in the
description above. The species is sister to the rest of the
genus (Rouhan et al., 2004). Plants in Central America
have reddish young leaves—a characteristic unique in
the genus—but plants from Ecuador apparently lack this
color (pers. obs.). Elaphoglossum amygdalifolium might
eventually prove to be hemiepiphytic, a pleisiomorphic
condition. Plants appear to start near the bases of trunks
and then climb. Such plants might produce at their bases
long, straight roots that grow into the soil where they
ramify. This growth form, which occurs in the outgroup
genera (Anapausia, Lomagramma, Teratophyllum), needs
to be searched for. The sectional name is derived from
the Latin amygdalus, almond + folium, leaf. The leaves
resemble that of an almond.
LITERATURE: Mickel, J. T. & L. Atehortúa G. 1980. Subdivision of the genus Elaphoglossum. American Fern Journal 70: 47–68. Rouhan,
G., J-Y. Dubuisson, F. Rakotondrainible, T. J. Motley, J. T. Mickel, J-N. Labat & R. C. Moran. 2004. Molecular phylogeny of the fern
genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences: contributions of species from the Indian Ocean
area. Molecular Phylogenetics and Evolution 33: 745–763.
294
DESCRIPCIÓN: Plantas epífiticas; rizomas largamente
rastreros, las escamas lineares a lanceoladas; aeróforos
ausentes(?); filopódios presentes; láminas glabras a
subglabras, enteras, delgadas, a veces rojizas cuando
jóvenes, las escamas diminutas, no setosas, lacinadas-
pectinadas a estrelladas, pelos diminutos glandulares
ausentes; venas libres; hidatodos presentes; esporas
monoletes, pardas o negruzcas; x=41.
GÉNEROS PARECIDOS: Las especies de las
Polypodiaceae con hojas enteras (e.g., Campyloneurum,
Microgramma, Pleopeltis) difieren por nervadura reticulada
y soros redondos. Las otras secciones de Elaphoglossum
difieren por tener a lo menos una de las características
siguientes: hidatodos ausentes, láminas conspicuosamente
escamosas, o filopodios ausentes.
COMENTARIOS: Sección Amygdalifolium es neotropical
y consta de una sola especie: E. amygdalifolium (Mett.) H.
Christ. Existe desde 0–1500 m en Cuba y América Central
hasta Colombia y Ecuador (restingada al lado occidental
de los Andes). Ninguna otra especie de Elaphoglossum
tiene la combinación de caracteres dados en negritos en
la descripción arriba. La especie es hermana al resto del
género (Rouhan et al., 2004). Plantas de América Central
tienen hojas jóvenes rojizas—una característica única
en el género—pero plantas de Ecuador aparentemente
faltan este color (pers. obs.). Elaphoglossum amygdalifolium
puede ser hemiepifítica, una condición pleisiomórfica.
Plantas aparecen empezar cerca las bases de los troncos
y luego suben. Es posible que tales plantas produzcan en
sus bases raíces largas rectas que van directamente al
suelo donde ramifican. Esta forma de crecimiento, que
existe en los grupos externos (Anapausia, Lomagramma,
Teratophyllum), se necesita buscar. El nombre genérico se
deriva del latín amygdalus, almendra + folium, hoja. Las
hojas se asemejan las de una almendra.
Figure 144. A, B. Elaphoglossum seminudum. C-E. E. lonchophyllum. F-J. E. sartorii. K, L. E. glabellum.  M. E. parduei. (From
Mickel & Beitel, 1988.)

295
DRYOPTERIDACEAE
Elaphoglossum J. Sm. sect. Lepidoglossa
DESCRIPTION: Primarily epiphytic, occasionally
terrestrial or rupestral; stem short- to long-creeping, rarely
erect, scaly, in transverse section with an elongated ventral
vascular bundle; leaves simple, entire, dimorphic; petioles
articulate to dark phyllopodia; laminae subglabrous to
densely scaly, scales with acicular, one-celled, non-
glandular teeth; rachis grooved adaxially; veins free; sori
acrostichoid; spores bilateral, typically nongreen; x=41.
DESCRIPCIÓN: Principalmente epífitas, a veces
terrestres o rupestres; tallos cortamente a largamente
rastreros, raras veces erectos, escamosos, en sección
transversal con un haz ventral vascular alargado; hojas
simples, enteras, dimorfas; pecíolos articulados a filopódios
oscuros; láminas subglabras a densamente escamosas, las
escamas con dientes aciculares, unicelulares, no
glandulares; raquis surcado en el haz; venas libres; soros
acrosticoides; esporas bilaterales, típicamente no verdes;
x=41.

SIMILAR GENERA: Other sections of Elaphoglossum GÉNEROS PARECIDOS: Otras secciónes de

have leaf scales with marginal teeth composed of several Elaphoglossum tienen escamas foliares con dientes
cells, the terminal one swollen and gland-like. marginales compuestos de varias células, el terminal
hinchada y como una glándula.

COMMENTS: Elaphoglossum sect. Lepidoglossa is COMENTARIOS: Elaphoglossum sect. Lepidoglossa es

pantropical and contains ca. 350 species worldwide.
Most of the species are neotropical and epiphytic. They
occur in a wide variety of habitats, from rainforests to
páramos. The species are poorly known and in need of
taxonomic study. They can be identified without fertile
leaves because they are defined primarily on the basis of
leaf shape, venation, presence or absence of hydathodes
and phyllopodia, and characteristics of the laminar scales.
The laminae of sect. Lepidoglossa are often conspicuously
scaly, but in some species the scales are reduced to
reddish dots and the laminae therefore appear glabrous.
The genus name is derived from the Greek elaphos, stag
+ glossa, tongue. The leaves resemble a deer’s tongue.
Lepido- means scaly.
pantropical e consta de ca. 350 especies mundialmente.
La mayoría de las especies son neotropicales y epifíticas.
Ocurren un una amplia variedad de hábitats, desde
bosques lluviosos hasta páramos. Las especies se
conocen pobremente y necesitan estudio taxonómico.
Se puede identificarlas sin hojas fértiles porque son
definidas principalmente en base de la forma de las hojas,
venación, presencia o ausencia de hidatodos y filopódios,
y características de las escamas laminares. Las láminas
en secc. Lepidoglossa son típicamente escamosas, pero
en algunas especies las escamas son reducidas a puntos
rojizos y pore so las láminas aprecen glabras. El nombre
genérico se deriva del griego elaphos, venado + glossa,
lengua. Las hojas se asemejan a una lengua de venado.
Lepido- significa escamoso.

LITERATURE: Alston, A. H. G. 1958. The Brazilian species of Elaphoglossum. Bol. Soc. Brot. 32: 1–32. Mickel, J. T. 1980.
Relationships of the dissected elaphoglossoid ferns. Brittonia 32: 109–117. Mickel, J. T. & L. Atehortúa G. 1980. Subdivision of
the genus Elaphoglossum. American Fern Journal 70: 47–69. Rouhan, G., J-Y. Dubuisson, F. Rakotondrainible, T. J. Motley,
J. T. Mickel, J-N. Labat & R. C. Moran. 2004. Molecular phylogeny of the fern genus Elaphoglossum (Elaphoglossaceae) based
on chloroplast non-coding DNA sequences: contributions of species from the Indian Ocean area. Molecular Phylogenetics and
Evolution 33: 745–763. Vasco, A., G. Rouhan & R. C. Moran. 2009. Circumscription and phylogeny of the Elaphoglossum ciliatum
group (E. sect. Lepidoglossa, Dryopteridaceae) based on cpDNA sequences. Taxon 58: 825–834. Vasco, A., G. Rouhan & R. C. Moran.
2009. Monograph of the Elaphoglossum ciliatum group (Dryopteridaceae). Brittonia 61: 241–272..

296
Figure 145. A–F. Elaphoglossum muscosum. G–P. E. paleaceum. Q. E. vestitum. R–W. E. pringlei. (From Mickel & Beitel,
1988.)

297
DRYOPTERIDACEAE
Elaphoglossum sect. Polytricha H. Christ
DESCRIPTION: Plants generally epiphytic; rhizomes
short- to (rarely) long-creeping, the scales linear to
lanceolate; phyllopodia present; petioles glabrous or with
minute capitate-glandular hairs; blades simple, entire,
with or without glandular hairs, scaly particularly along
the costae, the scales subulate with a slightly widened
enrolled base, patent; veins free; hydathodes absent; sori
acrostichoid; spores monolete; x=41.
DESCRIPCIÓN: Plantas generalmente epifítas, rizomas
cortamente a (raras veces) largamente rastreros, las
escamas lineares a lanceoladas; filopódios presentes;
pecíolos glabros o con pelitos gladulosos-capitados;
láminas simples, enteras, con o sin pelos glandulares,
escamosas particularmente por las costas, las escamas
subuladas con una base enrollada, patentes; venas
libres; hidatodos ausentes; soros acrosticoides; esporas
monoletes; x=41.

SIMILAR GENERA: Elaphoglossum sect. Setosa also GÉNEROS PARECIDOS: Elaphoglossum sect.
has subulate leaf scales but differs by the presence of Setosa tiene también escamas foliares pero difiere
hydathodes. The other sections of Elaphoglossum lack por la presencia de hidatodos. Las otras secciones de
subulate scales. Elaphoglossum carecen de escamas subuladas.

COMMENTS: Like all groups of Elaphoglossum, sect.
Polytrichia is primarily neotropical. The exact number
of species belonging to the group is unknown. Rouhan
et al. (2004) found the section to be monophyletic. In
Elaphoglossum, subulate scales are highly distinctive and
found only in sects. Polytrichia and Setosa. These scales
are enrolled longitudinally and en masse have a bristly
appearance. They are found only on the leaves, not the
rhizomes. In some species, flat scales or scales enrolled
basally and flat distally, occur along the lamina margins.
In such species, typical subulate scales are found on the
petioles and costae. Some widespread species of sect.
Polytrichia are E. apodum (Kaulf.) Schott ex J. Sm., E. crinitum
(L.) Christ, E. decoratum (Kunze) T. Moore, E. erinaceum
(Fée) T. Moore, E. hybridum (Bory) Brack., and E. pallidum
(Baker ex Jenm.) C. Chr. The section name comes from
the Greek poly, many, and trichia, hair, referring to the
many hair-like subulate scales on the leaves.
COMENTARIOS: Como todos los grupos de
Elaphoglossum, la secc. Polytrichia es principalmente
neotropical. El número exacto que pertenece al grupo
se desconoce. Rouhan et al. (2004) encontraron que
el grupo es monofilético. En Elaphoglossum las escamas
subuladas son altamente distintivas y se encuentran solo
en las seccs. Polytrichia y Setosa. Estas escamas se enrollan
longitudinalmente y conjunto tienen una apariencia
cerdosa. En algunas especies, escamas planadas o
solamente enrolladas basalmente y planadas distalmente,
ocurren por los márgenes de las láminas. En tales
especies, escamas subuladas típicas se encuentran por los
pecíolos y las costas. Algunas especies muy difundidas de
sect. Polytrichia son E. apodum (Kaulf.) Schott ex J. Sm.,
E. crinitum (L.) Christ, E. decoratum (Kunze) T. Moore, E.
erinaceum (Fée) T. Moore, E. hybridum (Bory) Brack. y E.
pallidum (Baker ex Jenm.) C. Chr. El nombre seccional se
deriva del griego poly, mucho, y trichia, pelo, refiriéndose a
las numerosas escamas piliformes en las hojas.

LITERATURE: Mickel, J. T. & L. Atehortúa G. 1980. Subdivision of the genus Elaphoglossum. American Fern Journal 70:
47–68. Rojas-Alvarado, A. F. 2003. Notes on Elaphoglossum (Lomariopsidaceae) section Polytrichia subsection Hybrida in
Mexico and Central America. Rev. Biol.Trop. 51: 33–48. Rojas-Alvarado, A. F. 2009.Two new species and a new combination in
Elaphoglossum sect. Polytrichia subsect. Apoda (Dryopteridaceae) from Costa Rica and Panama. Brittonia 61: 293–300. Rojas-
Alvarado, A. F. 2010. New species and new records in Elaphoglossum sect. Polytrichia subsect. Hybrida (Dryopteridaceae)
from the Neotropics. American Fern Journal 100: 172–179. Matos, F. B. In press. Typification of some species names in
Elaphoglossum section Polytrichia (Dryopteridaceae) from Brazil. Acta Botanica Brasilica 00: 00–00. Matos, F. B. & J. T. Mickel.
2014. The Brazilian species of Elaphoglossum section Polytrichia (Dryopteridaceae). Brittonia 66: 371–395. Matos, F. B. & R.
C. Moran. 2013. Elaphoglossum clathratum sp. nov. (Dryopteridaceae) from the eastern side of the Andes in Ecuador. Nordic
Journal of Botany 31: 442–445.

298
Figure 146. Elaphoglossum sect. Polytrichia. A–F. E. scolopendrifolium. (From Matos & Mickel, 2014.)

299
DRYOPTERIDACEAE
Elaphoglossum sect. Setosa (H. Christ) Mickel & Atehortúa
DESCRIPTION: Plants generally epiphytic; rhizomes
short- to long-creeping, the scales linear; phyllopodia
absent; petioles with minute capitate-glandular hairs;
blades sparsely puberulent and conspicuously scaly,
the hairs minute and glandular (handlens needed),
the scales subulate with a slightly widened enrolled
base, spreading to erect, orange to brown; veins free,
spaced well apart; hydathodes present; sori acrostichoid;
spores monolete; x=41.
SIMILAR GENERA: Similar species of Elaphoglossum
can usually be distinguished by non-subulate scales, lack
of minute glandular hairs, the presence of phyllopodia, or
the absence of hydathodes.
COMMENTS: Like all groups of Elaphoglossum, sect.
Setosa is primarily neotropical. The exact number of
species belonging to the group is unknown. Rouhan
et al. (2004) found the group to be monophyletic. In
Elaphoglossum, subulate scales are highly distinctive. They
are slender and en masse often have a bristly appearance.
With magnification it can be seen that the scales are
rolled at the base to form a short cylinder-like part on
which supports the rest of the scale. Some widespread
species of sect. Elaphoglossum Setosa are E. lindenii (Bory
ex Fée) T. Moore, E. piloselloides (C. Presl) T. Moore, y E.
villosum (Sw.) J. Sm. The section name comes from the
Latin seta, bristle, and refers to the stiff, spreading subulate
scales.
LITERATURE: Mickel, J. T. & L. Atehortúa G. 1980. Subdivision of the genus Elaphoglossum. American Fern Journal 70:
47–68. Matos, F. B. & J. T. Mickel. 2018. The Brazilian species of Elaphoglossum section Setosa (Dryopteridaceae). Brittonia
70: 173–205. [DOI 10.1007/s12228-018-9521-9]
300
DESCRIPCIÓN: Plantas generalmente epifíticas, rizomas
cortamente a largamente rastreros, las escamas lineares;
filopódios ausentes; pecíolos con pelitos gladulosos-
capitados; láminas esparcidamente puberulentas y
conpicuosamente escamosas, los pelos minutos
y glandulares (se necesita una lupa), las escamas
subuladas con la base poco ensanchada y enrollada,
patentes a erectas, narangas a pardas; venas libres, bien
espaciadas; hidatodos presentes; soros acrosticoides;
esporas monoletes; x=41.
GÉNEROS PARECIDOS: Especies similares de
Elaphoglossum se pueden distinguir usualmente por
escamas no subuladas, el carecer de pelitos glandulosos,
la presencia de filopodios, o la ausencia hidatodos.
COMENTARIOS: Como todos los grupos de
Elaphoglossum, la secc. Setosa es principalmente
neotropical. El número exacto que pertenece al grupo
se desconoce. Rouhan et al. (2004) encontraron que
el grupo es monofilético. En Elaphoglossum las escamas
subuladas son altamente distintivas. Son delgadas y a
menudo en mases parecen cerdosas. Con magnificación,
se puede ver que las escamas son enrolladas en la base y
forman un breve cilíndrico que apoya el resto de la escama.
Algunas especies muy difundidas de Elaphoglossum sect.
Setosa son E. lindenii (Bory ex Fée) T. Moore, E. piloselloides
(C. Presl) T. Moore, y E. villosum (Sw.) J. Sm. El nombre
seccional se deriva del latín seta, cerdo, y se refiere a las
escamas subuladas rígidas y patentes.
Figure 147. Elaphoglossum sect. Setosa. A–E. E. pilosius. F–G. E. setosum. J–L. E. jaliscanum. M, N. E. pusillum. Q. E.
monicae. (From Mickel & Smith, 2004.)

301
DRYOPTERIDACEAE
Elaphoglossum sect. Squamipedia Mickel & Atehortúa
DESCRIPTION: Plants epiphytic, generally less than
25 cm tall; rhizomes long-creeping, the scales usually
pale brown, concolorous; aerophores 1-3 mm long,
peg-like, found slightly behind the base of the petiole;
phyllopodia absent; blades entire or either flabellately
or pinnately dissected, subglabrous to moderately scaly,
the scales non-setose, often hastate or slightly so at base;
veins free; hydathodes absent; sori acrostichoid; spores
minutely spiny; x=40.
DESCRIPCIÓN: Plantas epífiticas, generalmente
menos de 25 cm de alto; rizomas largamente
rastreros, las escamas usualmente pardo pálido,
concoloras; aeróforos 1-3 mm de largo, cilíndricos,
atrás de la base del pecíolo; filopódios ausentes; láminas
enteras or disectadas dicotómamente o pinnatamente,
subglabras a escamosas, las escamas sin setas, a menudo
hastatas or casi así en la base; venas libres; hidatodos
ausentes; soros acrostichoides; esporas minuciosamente
espinosas; x=40.

SIMILAR GENERA: Other sections of Elaphoglossum GÉNEROS PARECIDOS: Otras secciones de

may be distinguished by one or a combination of the Elaphoglossum pueden ser distinguidas por una sola
following characters: short-creeping or erect rhizomes, o una combinación de las características siguientes:
the presence of phyllopodia, leaves longer than 25 cm. rizomas cortamente reptantes o erectos, la presencia de
filopodios, hojas más de 25 cm de largo.

COMMENTS: Section Squamipedia contains about COMENTARIOS: Sección Squamipedia consta de

17 species, all of which are neotropical except for one
in Madagascar. The section is entirely epiphytic and
generally occurs above 1000 m. The peglike aerophores
on the rhizomes are often hidden by the scales and
must be carefully searched for. They are situated about
1 mm behind the petiole. Three species with dissected
laminae occur in the section: Elaphoglossum peltatum
(laminae orbicular and flabellately dissected), E. tripartitum
(laminae tripartite), and E. moorei (laminae lanceolate
pinnately dissected, often with the pinnae forked at
the apex). Because of their dissected laminae, they are
sometimes placed in Peltapteris Link, but molecular
phylogenetic results indicate that lamina division evolved
three times separately within the section. Thus Peltapteris,
defined as such, is polyphyletic. Wagner (1980) found
n=40 in E. peltatum, which differs from n=41 in other
Dryopteridaceae. This might be a synapomorphy for the
section. The name Squamipedia refers to E. squamipes
which belongs to the section.
casi 17 especies, todos de las cuales son neotropicales
con excepción de una en Madagascar. La sección es
completamente epifítica y generalmente existe arriba
arriba de 1000 m. Los aeróforos como dedos en las
rizomeas a menudo están ocultos por las escamas y hay
que buscarlas con cuidado. Están ubicados casi 1 mm atrás
del pecíolo. Dos especies con láminas disectadas existe en
el sección: Elaphoglossum peltatum (láminas redondeadas
y discectadas dicotómamente), E. tripartitum (láminae
tripartitas) y E. moorei (láminas disectadas pinatamente,
a menudo con las pinnas furcadas en el ápice). A causa
de sus láminas disectadas, están ubicadas en Peltapteris
Link., pero resultados moleculares filogenéticos indican
que división de la lámina se evolucionó tres veces dentro
dentro de la sección. Por eso Peltapteris, definido tal, es
polifilético. Wagner (1980) encontró n=40 en E. peltatum,
lo cual difiere de n=41 en otros Dryopteridaceae. Es
posible esto es una sinapomofia para la sección. El nombre
Squamipedia refiere a E. squamipes que pertenece a la
sección.

LITERATURE: Gómez, L. D. 1975. Contribuciones a la pteridología costarricense. VI. El género Peltapteris Link en Costa Rica.
Brenesia 6: 25–31. Mickel, J. T. 1980. Relationships of the dissected elaphoglossoid ferns. Brittonia 32: 109–117. Rouhan, G.,
J-Y. Dubuisson, F. Rakotondrainible, T. J. Motley, J. T. Mickel, J-N. Labat, R. C. Moran. 2004. Molecular phylogeny of
the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences: contributions of species
from the Indian Ocean area. Molecular Phylogenetics and Evolution 33: 745–763. Vasco, A., J. T. Mickel & R. C. Moran. 2013.
Monograph of the neotropical species of Elaphoglossum sect. Squamipedia (Dryopteridaceae). Annals of the Missouri Botanical
Garden 99(2): 244–286.

302
Figure 148. A–E. Elaphoglossum peltatum fo. peltatum, note peg-like aerophotes in B. F. Elaphoglossum peltatum
fo. standleyi. G, H., Elaphoglossum peltatum fo. flabellatum. J. Elaphoglossum peltatum fo. foeniculaceum.. K, N.
Elaphoglossum moorei × squamipes? L. E. moorei.M, N. Elaphoglossum colombianum O–P. Elaphoglossum tripartitum
(Vasco et al., 2013.)

303
DRYOPTERIDACEAE
Elaphoglossum sect. Wrightiana Lóriga et al.
DESCRIPTION: Plants climbering, with long-creeping
rhizomes that begin growth on the ground and eventually
climb trunks to heights of 1–2 m, maintaining the
connection to the ground by the rhizome and roots
emitted from the lower portions of the climbing rhizome;
aerophores present, finger-like; phyllopodia present;
hydathodes absent; laminar scales flat (not subulate), with
marginal processes or teeth ending in a slightly swollen
cell (i.e., scales never with acicular marginal cells as in E.
sect. Lepidoglossa); x=?.
DESCRIPCIÓN: Plantas trepadoras, con rizomas
largamente rasteros que empiezan crecimiento en el
suelo y eventualmente suben troncos hasta alutras de
1–2 m, mantaniéndose la coneccion con el suelo por
el rizoma y raíces emitidas de las porciones proximales
del rizoma trepador; aeróforos presentes, con forma de
dedo; filopodios presentes; hidatodos ausentes; escamas
foliares planas (no subuladas), con dientes marginales
terminándose en una célula levamente hinchada (i.e.,
escamas nuncas con células aciculares marginales comon
en E. secc. Lepidoglossa); x=?.

SIMILAR GENERA: Other sections of Elaphoglossum GÉNEROS PARECIDOS: Otras secciones de

may be distinguished by one or a combination of the Elaphoglossum pueden ser distinguidas por una sola o
following characters: short-creeping or erect rhizomes, una combinación de las características siguientes: rizomas
the absence of phyllopodia, leaves longer than 25 cm. cortamente reptantes o erectos, la ausencia de filopodios,
hojas más de 25 cm de largo.

COMMENTS: Section Wrightiana consists of a single
species endemic to Cuba: E. wrightii (Mett. ex D. C. Eaton)
T. Moore. It grows in wet montane forests and is the sister
species to all others in the genus except E. amygdalifolium.
Its growth form is unique in the genus: young sporophytes
begin on the forest floor and creep around until they
encounter a trunk, which is then climbed by means of
adhesive roots. Fertile leaves are produced on the
climbing portion of the rhizome. This growth form is
present in the bolbitioid outgroups to Elaphoglossum,
such as Mickelia, Lomagramma, and Teratophyllum. The
sectional name Wrightiana refers to E. wrightii, the type
species of the genus, which is named for Charles Wright
(1811–1885), a plant collector.
COMENTARIOS: Sección Wrightiana consta de una
sola especie endémica a Cuba: E. wrightii (Mett. ex D.
C. Eaton) T. Moore. Existe en bosques húmedos y es la
especie hermano a las demas especies del género con
excepción a E. amygdalifolium. Su forma de crecimiento
es único en el género: esporofitos jóvenes empiezan en
el sotobosque y trepan hasta que encuentran un tronco,
lo cual es subido por medios de raíces adhesivos. Hojas
fértiles están producidas de la porción reptante del rizoma.
Esta forma de crecimiento es presente en los grupos
externos bolbitidoides, tales como Mickelia, Lomagramma
y Teratophyllum. El nombre seccional Wrightiana refiere a
E. wrightii, la especie tipo de la sección, que es nombrado
para Charles Wright (1811–1885), un colector de plants.

LITERATURE: Lóriga, J., A. Vasco, L. Regalado, J. Heinrichs & R. C. Moran. 2013. Phylogeny and classification of the
Cuban species of Elaphoglossum (Dryopteridaceae), with description of Elaphoglossum sect. Wrightiana sect. nov. Plant
Systematics and Evolution [doi 10.1007/s00606-013-0933-4; published online 14 Nov 2013]

304
Figure 149. A–F. Elaphoglossum (sect. Wrightiana) wrightii. D. Heteroblastic series, youngest leaf at right. A. Habit. B. Petiole
scale from fertile leaf. C. Rhizome scale. (Modified from Vasco et al., 2013.)

305
POLYPODIACEAE
Enterosora Baker in Thurn
DESCRIPTION: Epiphytes, occasionally saxicolous;
rhizome scales not clathrate, yellowish-brown to golden
brown, sometimes ciliate or glandular; laminae simple
to lobed 4/5 of the way to the midrib, not pectinate,
generally thick and spongy, mesophyll completely
composed of stellate parenchyma (pallisade
parenchyma lacking); veins simple when free, forked
when fertile, but often variously areolate, especially near
the midrib; hydathodes absent; setae present, dark
reddish, often surrounding the sorus; hairs often
branched and glandular at the apex; sori slightly to
deeply sunken in the lamina tissue, round to elongate;
spores green, trilete; x=37.
DESCRIPCIÓN: Epífitas, ocasionalmente saxícolas;
escamas del rizoma no clatradas, pardo-amarillento a
pardo-doradas, a veces ciliadas o glandulares; láminas
simples a lobadas 4/5 de la longitud a la costilla
media, no pectinada, generalmente gruesa y esponjosa,
mesofilio consta completamente de parénquima
estrellado (parénquima palisade ausente); nervaduras
simples cuando libres, furcadas cuando fértiles, pero a
menudo variadamente areoladas, especialmente cerca de
la costilla media; hidatodos ausentes; setas presentes,
rojizas oscuras, a menudo rodeando el soro; pelitos
típicamente ramificados, a menudo glandulares en el ápice;
soros ligeramente a profundamente embebidos en
el tejido laminar, redondeados a alongados, esporas
verdes, triletes; x=37.

SIMILAR GENERA: The absence of hydathodes will
separate Enterosora from all the other neotropical genera
except Ceradenia and Zygophlebia. Ceradenia differs
by the waxy glands in the sori, and Zygophlebia by the
petioles articulate to the rhizome and pinnatisect to
pinnate laminae.
GÉNEROS PARECIDOS: La ausencia de hidátodos
separa Enterosora de todos los otros géneros
neotropicales con excepción de Ceradenia y Zygophlebia.
Ceradenia difiere por las glándulas cerosas en los soros,
y Zygophlebia por los pecíolos articulados al rizoma y
láminas pinnatisectas a pinnadas.

COMMENTS: Enterosora contains 8 neotropical and COMENTARIOS: Enterosora consta de 8 especies

2 African species. In the Neotropics, it ranges from
Mexico to Bolivia and Suriname and in the Antilles. The
species grow mainly in cloud forests. The thick, spongy
mesophyll can be seen where the tissue has been torn
or deteriorated. It cells bear long “arms” and between
them are large intercellular air spaces. A palisade layer is
lacking. Enterosora is monophyletic and probably sister to
Ceradenia (Ranker et al., 2004), a genus that can also have
spongy mesophyll, although less developed. The genus
name is derived from the Greek enteron, intestine + soros.
The sori are not superficial, but embedded in pockets
within the leaf tissue.
neotropicales y 2 Africanas. En el Neotrópico, se existe
desde México hasta Bolivia y Surinam y en las Antillas. Las
especies crecen principalmente en bosques nublados. El
mesofilio grueso, esponjoso se puede ver donde el tejido
ha estado roto o deteriorado. Sus células llevan “brazos”
largos e entre si están espacios intercelulares grandes.
Un estrato palisade se carece. Enterosora es monofilético
y probablemente hermana a Ceradenia (Ranker et al.,
2004), un género que puede tener mesofilio esponjoso,
aunque menos desarrollado. El nombre genérico se
deriva del griego enteron, intestino + soros. Los soros no
son superficiales, pero hundidos en bolsillos en el tejido
de la hoja.

LITERATURE: Bishop, L. E. & A. R. Smith. 1992. Revision of the fern genus Enterosora (Grammitidaceae) in the New World.
Systematic Botany 17: 345–362. Ranker, T. A., A. R. Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Staub & H.
Schneider. 2004. Phylogeny and evolution and grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy.
Taxon 53: 415–428. Rojas, A. F. 2006. Notas sobre Enterosora Baker (Filicales: Grammitidaceae) en Costa Rica. Lankesteriana
6: 9–13.

306
Figure 150. Enterosora. A–C. E. trifurcata. D–F. E. insidiosa. G–I. E. trichosora. J–L. E. parietina. (From Bishop & Smith, 1992.)

307
ATHYRIACEAE
Ephemeropteris R. C. Moran & Sundue, gen. nov. in sched..
DESCRIPTION: Terrestrial; rhizomes creeping,
without persistant petiole bases, apices protruding
2–4 cm beyond the leaf bases; leaves distichous;
petiole bases fleshy, compressed and broadly
grooved to slightly flattened, without enlarged
pneumatophores on the sides, with 2 vascular bundles,
scaly, the scales brown, narrowly lanceolate, entire; laminae
deltate to lanceolate, 1- to 3-pinnate-pinnatifid, glabrous
except for scattered scales, membranous; veins free,
ending before the lamina margin; sori linear to circular
or hamate; indusia laciniate or dentate, membranous; x=?
DESCRIPCIÓN: Terrestres; rizomas reptantes,
sin bases persistentes de los pecíolos, ápices
proyectando 2–4 cm más afuera de las bases de las
hojas; hojas dísticas; bases de los pecíolos carnosas,
comprimidas y surcadas anchamente hasta, con
2 haces vasculares, en pneumatóforros en los lados,
escamosas, las escamas pardas, angustamente lanceoladas,
enteras; láminas deltadas a lanceoladas, 1- a 3-pinnado-
pinnatífidas, glabras excepto para escamas difundidas,
membranosas; venas libres, terminándose antes del
márgen laminar; soros lineales a circulares o hamados;
indusios laciniados o dentados, membranosos; x=?

SIMILAR GENERA: Athyrium differs by thicker rhizomes
bearing persistant petiole bases, leaves polystichously
arranged around the rhizome, and indusia laciniate or
dentate. Diplazium differs by at least some of the sori
paired back-to-back along the same vein.
GÉNEROS PARECIDOS: Athyrium difiere por rizomas
más gruesos llevando bases persistentes de los pecíolos,
hojas arregladas polísticamente alrededor del rizoma y
los indusios laciniados o dentados. Diplazium difiere por
al lo menos algunos soros pareados espalda a espalda
por la misma vena.

COMMENTS: Ephemeropteris consists of 3 species, all COMENTARIOS: Ephemeropteris consta de 3especies,

Neotropical: E.. palmensis (Christ) R. C. Moran & Sundue,
E. skinneri (Baker) R. C. Moran & Sundue, and E. tejeroi
(Mickel & Tejero) R. C. Moran & Sundue. All three were
treated taxonomiclly by Moran et al. (in review). They
occur in forests on the Pacific side of Central America (E.
skinneri is also known from two collections in Colombia
y Ecuador). The distinctive rhizomes and petiole bases of
the group need study using living material, especially in
comparison with Athryium sect. Athyrium. The sori tend
to be circular in A. palmensis and A. tejeroi. Differing from
Athryium are the lanicinate or dentate indusia. The plants
appear to be seasonal, with leaves dying at the begining of
the dry season. The name Ephemeropteris is derived from
the Latin ephemerus-, ephemeral, short-lived + pteris, fern.
It refers to the leaves that day back at the onset of the
dry season.
todas Neotrópicales: E.. palmensis (Christ) R. C. Moran
& Sundue, E. skinneri (Baker) R. C. Moran & Sundue y
E. tejeroi (Mickel & Tejero) R. C. Moran & Sundue. Las
tres ueran tratadas taxonómicamente por Moran et al.
(en revisión). Existan en bosques en el lado Pacífico de
Mesoamérica (E. skinneri se conoce también desde dos
colecciones en Colombia y Ecuador). Los distinctivos
rizomas y bases de los pecíolos del grupo necesitan estudio
usando material vivo, especialmente en comparación con
Athryium sect. Athyrium. Los soros tienden ser circulares
en E. palmensis y E. tejeroi. Difieriendo de Athyrium or
los indusios laciniados o dentados. Las plantas aparecen
ser estacionales, con las hojas muriéndose en la empieza
de la estación seca. El nombre Ephemeropteris se derive
desde el Latin emphemerus, efímeral, de vida corta, más
pteris, helecho. Se refiere a las hojas que mueren en el
comienzo de la estación seca.

LITERATURE: Liu, Y-C., W-L. Chiou & M. Kato. 2011. Molecular phylogeny and taxonomy of the fern genus Anisocampium
(Athyriaceae). Taxon 60: 824–830. Mickel, J. T. & A. R. Smith. 2004. The pteridophytes of Mexico. Memoirs of the New York
Botanical Garden 88: 1–1054. Moran, R. C., J. Garrison Hanks & M. Sundue. In review. Phylogenetic relationships of Neotrop-
ical Lady Ferns (Athyriaceae), with a description of Ephemeropteris, Gen. Nov.Taxon 00: 000–000. Stolze, R. G. 1994. Athyrium.
In: Harling, G. & L. Andersson (eds.), Flora of Ecuador 14 (5B). Polypodiaceae-Dryopteridoideae-Physematieae 49: 1–108. Wei,
R., A. Ebiharab, Y-M. Zhua, C-F. Zhao, S.Hennequin, X-C. Zhang. 2017. A total-evidence phylogeny of the lady fern genus
Athyrium Roth (Athyriaceae) with a new infrageneric classification. Molecular Phylogenetics and Evolution 00: 000–000.

308
Figure 151. Ephemeropteris. A–D. Ephemeropteris skinneri. E–G. Ephemeropteris palmensis. H, J, K. Ephemeropteris
tejeroi. (From Mickel & Smith, 2004.)

309
EQUISETACEAE
Equisetum L.
DESCRIPTION: Stems jointed, green, longitudinally
grooved, usually hollow in the center. Branches (if
present) whorled at the nodes. Leaves whorled, fused
into a sheath, the tips free, tooth-like, often deciduous.
Sporophylls aggregated in terminal strobili, peltate, tan
or yellowish, bearing 5–10 sporangia on the inner surface;
spores green, with 4 arm-like appendages (elaters); x=108.
DESCRIPCIÓN: Tallos articulados, verdes, surcados
longitudinalmente, usualmente huecos en el centro.
Ramas (cuando presentes) verticilados en los nudos.
Hojas verticiladas, fusionadas en una vaina, los ápices
libres, como dientes, a menudo deciduos. Sporofilas
agregados en estróbilos terminales, peltados, leonados
o amarillos, llevando 5–10 esporángios en la superficie
interior; esporas verdes, con 4 brazos (elateres); x=108.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Equisetum has 15 species worldwide
and occurs on all continents exceopt Australia and
New Zealand. It exhibits a reverse latitudinal gradient
in diversity, being most species-rich in north-temperate
regions between 40° and 60°. It occurs in many habitats,
but most typically in sunny places and wet soils. The
stems are covered by a layer of silica modified into
spines, tubercles, or teeth. These structures are often
distinctive for the species. In Equisetum giganteum they are
quadrangular in profile, whereas in E. myriochaetum they
are angled. The nodes remain meristematic and capable
of generating new stems if placed in soil. Fragmentation of
stems enables abortive-spored hybrids to persist at their
site of origin and even spread beyond the range of their
parents.The spores bear hygroscopic appendages called
“elaters.” When moist, the elaters coil tightly around the
spore, and when dry they uncoil and spread, catching the
wind and creating drag which help disperse the spore.
The spores are green and have short viability (about one
week). They are unique by lacking haptotypic markings
(scars from their position in the tetrad). Thus, they are
neither monolete nor trilete. The genus name is derived
from the Greek equis, horse + seta, bristle. The shoots of
some species resemble a horse’s tail.
COMENTARIOS: Equisetum tiene 15 especies mundial
y existe en todos los continentes con excepción de
Australia y Nueva Zelanda. Exhibe un gradiente latitudinal
de diversidad en revés, siendo más rico en especies en
las zonas templadas norteñas entre 40° y 60°. Existe en
muchos hábitats, pero típicamente en lugares soleados
y suelos húmedos. Los tallos están cubiertos por un
estrato de sílice modificado en espinas, tubérculos o
dientes. Estas estructuras son a menudo distintivas para
la especie. En Equisetum giganteum son cuadrangulares en
perfil, mientras que en E. myriochaetum son angulados. Los
nudos se quedan meristemáticos y capaces de generar
tallos nuevos si planteados en el suelo. Fragmentación
de los tallos facilita los híbridos con esporas abortivas
persistir en su sitio de origen y también difundirse más
allá el rango de sus padres. Las esporas llevan brazos
higroscópicos que se llaman “elateres.” Cuando mojados,
los elateres se enrollan alrededor de la espora, y cuando
se secan, se extienden, recogiendo los vientos. Las
esporas son verdes y tienen una corta viabilidad (casi una
semana). Son únicas por carece de marcas haptotípicas
(cicatrices de su posición en la tetrada). Por eso no son
monoletes ni triletes. El nombre genérico se deriva del
griego equis, caballo + seta, cerda. Los vástagos de algunas
especies se asemejan como una cola de caballo.

LITERATURE: Des Marais, D. L., A. R. Smith, D. M. Britton & K. M. Pryer. 2003. Phylogenetic relationships and evolution
of extant horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F). International Journal of Plant Science
164: 737–751. Hauke, R. L. 1963. A taxonomic monograph of Equisetum subgenus Hippochaete. Beihefte Nova Hedwigia 8:
1–123. Hauke, R. L. 1979. A taxonomic monograph of Equisetum subgenus Equisetum. Nova Hedwigia 30: 385–455.

310
Figure 152. A–D. Equisetum hyemale. E–J. E. myriochaetum (From Mickel & Smith, 2004.)

311
MARATTIACEAE
Eupodium J. Sm.
DESCRIPTION: Terrestrial; rhizomes erect or nearly so,
each bearing only 1 leaf (rarely 2) and paired rounded
fleshy stipules at the base of each leaf; fleshy; sterile and
fertile leaves monomorphic; petioles with polycyclic steles;
laminae 2-4-pinnate, triangular, the apex pinnatifid, axes
of the distal segments usually winged; pinnae opposite or
nearly so; pulvini present at the junctures of the rachises,
costae, and costules; veins free, adaxial surfaces with
greenish awns; receptacles without scales; synangia
stalked, bilateral, splitting longitudinally, with 2 rows
of opposing sporangia, each sporangium dehiscing by
a vertical slit on the inner surface; spores monolete,
spinulose; x=?
DESCRIPCIÓN: Terrestres; rizomas erectos o casi así,
cada uno llevando una sola hoja (raras veces 2) y
estípulas pareadas, redondeadas y carnosas en la base
de cada hoja; hojas estériles y fértiles monomorfas;
pecíolos con estelas policíclicos; láminas 2-4-pinnadas,
triangulares, los ápices pinatífidas, los ejes de los segmentos
distales usualmente aladas; pinnas opuestas o casi así;
pulvini presentes en las junciones de las ráquises, costas y
cóstulas; nervaduras libres, en superficies adaxiales con
arístas verdosas; receptáculos con escamas; sinángios
peciolados, bilaterales, rendiéndose longitudinalmente,
con 2 hileras de esporangios opuestos, cada esporangio
abriéndose por una hendidura vertical en la superficie
interior; esporas monoletes; x= ?

SIMILAR GENERA: Marattia differs by more than one
leaf per plant, lacking scattered awns along the veins
adaxially, and sessile synangia. Danaea differs by 1-pinnate
laminae, dimorphic sterile and fertile leaves, and synangia
that open by circular pores.
GÉNEROS PARECIDOS: Marattia difiere por más
de una hoja por planta, la falta de aristas por las venas
adaxialmente y sinángios sésiles. Danaea difiere por
láminas 1-pinadas, hojas estériles y fértiles dimorfas y
sinángios que abren por poros circulares.

COMMENTS: Eupodium is entirely Neotropical and COMENTARIOS: Eupodium es completamente

occurs in wet forests. It contains two species. The first, E.
laeve (Sm.) Murdock (syn: E. pittieri (Maxon) Christenh.)
occurs in the Greater Antilles, Central and South America.
The second, E. kaulfussii (J. Sm.) J. Sm., occurs in southern
Brazil and northern Argentina. It differs from the first
species by the presence of awns on the abaxial surfaces
of the veins, pulvini not tuberculate, and synangia varying
from elongate to circular. Both species are unusual in that
they typically produce only one leaf at a time. The genus
name is derived from the Greek eu-, good, true + pous,
foot. It refers to the stalk of the synangium.
Neotropical y existe en bosques húmedos. Contiene
dos espeies. La primera, E. laeve (Sm.) Murdock (sin: E.
pittieri (Maxon) Christenh.), existe en las Antilles Mayores
y América Central y Sur. La segunda, E. kaulfussii (J. Sm.) J.
Sm., existe en el sur de Brasil y Argentinal al norte. Esta
difiere de la primera por la presencia de arístas en las
superficies abaxiales de las venas, pulvini no tuberculados
y sinángios que varian desde elongados hasta circulares.
Ambas especies son inusuales en que producen
típicamente una sola hoja a la vez. El nombre genérico se
deriva del griego eu-, bueno, verdad + pous, pie. Refiere al
pecíolo del sinángio.

LITERATURE: Christenhusz, M. J. M. 2010. Revision of the neotropical fern genus Eupodium. Kew Bulletin 65: 115–121. del
Carmen Lavalle, M. 2003.Taxonomía de las especies neotropicales de Marattia (Marattiaceae). Darwiniana 4: 61–86. Murdock,
A. G. 2008. A taxonomic revision of the eusporangiate fern family Marattiaceae, with description of a new genus Ptisana. Taxon
57: 737–755. Underwood, L. M. 1909. Marattiaceae. North American Flora 16: 15–23.

312
Figure 153. Eupodium. Note in B the characteristic stalked synangia. A-C. E. laevis. D-E. E. kaufussii. (From Tryon & Stolze,
1989.)

313
PTERIDACEAE
Gaga Pryer, Fay-Wei Li & Windham
DESCRIPTION:Terrestrial; rhizomes short and compact;
petioles castaneous to black, typically without hairs;
rachises grooved adaxially; laminae 2- to 4-pinnate,
the basal basiscopic pinnules usually exaggerated; costae
and costules grooved adaxially; veins endings forming
prominent hydathodes; sori protected by strongly
differentiated inframarginal (i.e., behind the leaf
edge) pseudoindusia, these decurent along costae
and costules; sporangia clustered at vein tips; sporangia
32- or 64-spored; spores globose, trilete, tan, brown or
black. x=30
SIMILAR GENERA: Aspidotis differs by lamina segments
more acute or mucronate and adaxial lamina surfaces
striate and shiny. Cheilanthes differs by less differentiated
pseudoindusia, production of 32 small or 16 large spores
per sporangium, and often scaly laminae.
COMMENTS: Gaga consists of 19 species, all neotropical.
It occurs from southwestern United States (Arizona
and Texas) to southern Bolivia. Mexico is the center of
diversity, with 17 species. Generally in open or semi-open
habitats, (1500–)2000–4000 m. Gaga is sister to Aspidotis,
from which it differs only slightly. Both genera are sister
to Cheilanthes, from which Gaga is nearly inseparable
morphologically. Gaga is named for the American pop
singer, songwriter, and performer, “Lady Gaga,” stage name
of Stefani Joanne Angelina Germanotta (1986– ).
Li, F.-W., K. M. Pryer & M. D. Windham. 2012. Gaga, a new genus segregated from Cheilanthes (Pteridaceae). Systematic
Botany 37: 845–860.
314
DESCRIPCIÓN: Terrestres; rizomas cortos y
compactos; pecíolos castaños a negros, típicamente sin
pelos; ráquises surcados adaxialmente; láminas 2- a
4-pinnadas, las pínnulas basales basiscópicas usualmente
exageradas; costas y cóstulas surcados adaxialmente;
nervaduras terminando en hidátodos prominentes;
soros protegidos por pseuodindusios inframarginales
(i.e., atrás del margen de la lámina) fuertemente
diferenciados, estos decurrentes por las costas y
cóstulas; esporangios agrupados en los puntos de las
venas; esporangios con 32 o 64 esporas; esporas globosas,
triletes, leonadas, pardas o negras. x=30
GÉNEROS PARECIDOS: Aspidotis difiere por las
láminas con segmentos más agudos o mucronados y
superficies adaxiales estriadas y estriadas. Cheilanthes
difiere por pseudoindusio menos diferenciados,
producción de 32 esporas pequeñas por esporangio o
16 esporas grandes por esporangio, y a mendo laminas
escamosas.
COMENTARIOS: Gaga consta de 19 especies, todas
neotropicales. Existe desde el suroeste de los Estados
Unidos (Arizona y Texas) hasta el sur de Bolivia. México es
el centro de diversidad, con 17 especies. Generalmente en
lugares abiertos o semi-abiertos, (1500–)2000–4000 m.
Gaga es hermana a Aspidotis, de lo cual difiere solamente
un pocito. Ambos géneros son hermana a Cheilanthes,
de la cual Gaga es casi morfológicamente inseparable
inseparable.. Gaga se nombre para la Americana cantante
de “pop”, escritora de cancines y artista “Lady Gaga,”
nombre de teatro de Stefani Joanne Angelina Germanotta
(1986– ).
Figure 154. Gaga. A–C. G. marginata. D–G. G. hirsuta. H,J,K. G. marginata. In C and G, note the distinctive decurrent
pseudoindiusia. (From Mickel & Smith, 2004.)

315
POLYPODIACEAE
Galactodenia Sundue & Labiak
DESCRIPTION: Epiphytes, sometimes epipetric;
rhizomes short-creeping, the scales concolorous, non-
clathrate, orange, golden-brown or castaneous, marginally
ciliate by glandular hairs; leaves pendent, setose or not,
with glandular hairs, these clavate, hyaline, milky
white to gray or golden brown; laminae 1-pinnatisect
to 1-pinnate-pinnatisect; rachises blackish, visible on both
sides of the laminae; pinnae with blackish costae, these
often visible on both sides of the laminae; veins simple,
free, not blackish, and not visible without transmitted light;
hydathodes present, non-cretaceous, sometimes
poorly developed and difficult to see; sori round; sporangia
glabrous, or when young glandular, the glands similar to
those found on the lamina; spores green, trilete. x=?
SIMILAR GENERA: Alansmia also has pendent leaves
but differs by petioles and laminae with stellate or
clustered setae and, in most species, setulose sporangial
capsules. Lellingeria, Melpomene, and Stenogrammitis differ
by lacking glandular hairs on the laminae.
COMMENTS: Galactodenia consists of five species, all
neotropical. It occurs from southern Mexico and the
West Indies to Bolivia, (1500–)2000–4000 m. The most
distinctive characters of the genus are its pendent laminae
covered by glandular hairs. Galactodenia is sister to a clade
composed of Lellingeria, Melpomene, and Stenogrammitis.
It is not closely related to Alansmia or Terpsichore, the
two genera in which two of its species were previously
classified. Its species are G. delicatula, G. pumila, G. parriseae,
G. subscabra, and G. vareschii. A key to these species is
given in Sundue et al. (2012).Galactodenia is derived from
the Greek galacto, milk + adeno, gland, and refers to the
milky white glandular hairs characteristic of these plants.
LITERATURE: Sundue, M. A., P. H. Labiak, J. Mostacero & A. R. Smith. 2012. Galactodenia, a new genus of grammitid
ferns segregated from Terpsichore (Polypodiaceae). Systematic Botany 37: 339–346.
316
DESCRIPCIÓN: Epífitas, a veces epipéticas; rizomas
cortamente rastreros, las escamas concoloras, no
clatradas, anaranjadas a doradas-pardas o castáñas,
ciliadas en el margen por pelos glandulares; hojas setosas
o no, con pelos glandulares, estos clavados, hialinos,
lechosos a grisáceos o pardo-dorados; hojas péndulas,
láminas 1-pinnatisectas a 1-pinnada-pinnatisectas, ráquises
negruzcos, visibles en ambos lados de la lamina; pinnas
con costas negruzcas, a mendo visibles en ambos lados
de las láminas; nervaduras simples, libres, no negruzcas,
no visibles sin luz transmitida; hidatodos presentes,
sin puntos blanquecinos calizos, a veces pobremente
desarrollados y difíciles ver; soros redondos; esporangios
glabros, o, cuando jóvenes glandulares, las glándulas como
las de las láminas; esporas verdes, triletes. x=?
GÉNEROS PARECIDOS: Alansmia también tiene hojas
péndulas pero difiere por los pecíolos y láminas con setas
estelladas o agrupadas y, en la mayoría de las species,
cápsulas esporangiales setulosas. Lellingeria, Melpomene y
Stenogrammitis difieren por falta de pelos glandulares en
las laminas.
COMENTARIOS: Galactodenia consta de cinco
especies, todas neotropicales. Existe desde México del
sur y las Antillas a Bolivia, (1500–)2000–4000 m. Las
características más distinctivas del género son sus laminas
péndulas cubiertas por pelos glandulares. Galactodenia es
hermana a un clado formado por Lellingeria, Melpomene
y Stenogrammitis. No es cercanamente relacionado a
Alansmia o Terpsichore, los dos géneros donde dos de sus
species han sido clasificadas préviamente. Sus especies
son G. delicatula, G. pumila, G. parriseae, G. subscabra y G.
vareschii. Una clave a estas species se presenta en Sundue
et al. (2012). Galactodenia se deriva del Griego galacto,
leche + adeno, glándula, y se refiere a los pelos glandulares
lechoso-blancos característicos de estas plantas.
Figure 155. A–C. Galactodenia subscabra (Rivero & Rondon 1681, UC). A. Habit. B. Detail of the abaxial lamina. C. Detail of
the segments showing the sori and hairs. D–F. G. pumila (Fernández et al. 6032, MO). D. Habit. E. Detail of the adaxial and
abaxial lamina. F. Margin of a segment showing the hairs. G–I. G. parrisiae (Monro & Knapp 5272, MO). G. Habit. H. Detail
of the abaxial and adaxial side of the laminae. I. Detail of a sorus. (From Sundue et al., 2012.)

317
PTERIDACEAE
Gastoniella Li Bing Zhang & Liang Zhang
DESCRIPTION: Terrestrial or saxicolous, annual;
rhizomes short and poorly developed, sparsely scaly;
sterile and fertile leaves monomorphic, 2–20 (– 40) cm long,
clumped; petioles stramineous or brown basally, rounded
abaxially, grooved adaxially, glabrous; laminae deltate,
ovate, or lanceolate, 2- to 3-pinnate, submembranaceous;
ultimate pinnules or segments narrow, not or hardly
widened distally, entire or emarginate apically; veins free,
forked, 1 vein per segment; indusia absent; sori spreading
along veins; spore tetrahedral-glbose, trilete, whitish to
tan; x=29, 58.
DESCRIPTION: Terrestres o saxícolas, anuales; rizomas
cortos y pobremente desarollados, eparcidamente
escamosos; hoas estériles y fértiles monomorfas; 2–20(–
40) cm de larago, amacolladas; pecíolos estramíneos o
pardo basalmente, aroundeados abaxialmente, surcados
adaxialmente, glabros; láminas deltadas, ovadas o
lanceoladas, 2- a 3-pinnadas, submembranáceas; pínnulas o
segmentos últimos no o casi no ensanchados distalmente,
enteros o emarginados apicalmente; venas libres furcados,
1 vena por segmento; indusios ausentes; sorios difundidos
por las venas; esporas tetrahédricas-glbosas, triletes,
blanquecinas a leonadas; x=29, 58.

SIMILAR GENERA: Anogramma differs by scaly SIMILAR GENERA: Anogramma difiere por rizomas
rhizomes, distally widened segments or pinnae, and dark escamosos, pinnas o segmentos ensanchadas distalmente,
brown spores. y esporas pardas oscuras.

COMMENTS: Gastoniella consists of 3 species and COMMENTS: Gastoniella consta de 3 especies y es

is Neotropical except for one species endemic to
Ascencion Island in the South Atlantic Ocean. The
plants are characterized by being small and delicate
with poorly developed rhizomes, leaves less than 20 cm
long, and elongate non-indusiate sori. The three species
are G. ascensionis (Hook.) Li Bing Zhang & Liang Zhang
(Ascension Island), G. chaerophylla (Desv.) Li Bing Zhang
& Liang Zhang (widespread in the Neotropics), and G.
novogaliciana (Mickel) Li Bing Zhang & Liang Zhang
(central Mexico). They are dificult to distinguish. For a
long time, these species were classified in Anogramma, but
phylogenetic results show that they form a clade sister
to Cerosora Domin (Southeast Asia) and Pityrogramma
Link (pantropical). The sporophytes are annual, the new
ones arising every year from a perennial gametophyte.
Generally in open or semi-open habitats, (1500–)2000–
2100 m. Gastoniella is named for Gerald J. Gastony,
American pteridologist (1940– ).
Neotropical con excepcion de una especie endemica
a la Isla de Ascensión en el Sur del Oceano Atlántico.
Las plantas se caracteriza por ser pequenas y delicadas
con rizomas pobremente desarrolladas, hojas menos de
20 cm de largo y soros alargados no indusiados. Las tres
especies son G. ascensionis (Hook.) Li Bing Zhang & Liang
Zhang (Isla de Ascensión), G. chaerophylla (Desv.) Li Bing
Zhang & Liang Zhang (muy difundida en el Neotrópico),
and G. novogaliciana (Mickel) Li Bing Zhang & Liang Zhang
(Mexico central). Las tres son son difíciles distinguir. Por
mucho tiempo, estas especies fueron clasificadas en
Anogramma, pero estudiios filogenéticos demuestran que
forman un clado hermano a Cerosora Domin (Sureste de
Asia) y Pityrogramma Link (pantropical). Los esporofitos
son anuales, los nuevos surgiendo cada año de un
gametofito perenial. Generalmente en ambientes abiertos
o semiabiertos (1500–)2000–2100 m. Gastoniella honora
a Gerald J. Gastony, American pteridologist (1940– ).

LITERATURE: Nakazato, T. & G. J. Gastony. 2003. Molecular phylogenetics of Anogramma species and related genera
(Pteridacae: Taenitidoideae). Systematic Botany 28: 490–502. Pangua, E. & B. Vega. 1996. Comparative study of gametophyte
development in Cosentinia and Anogramma (Hemionitidaceae) and Cheilanthes (Sinopteridaceae). Pages 497–508. In: J. M.
Camus, M. Gibby & R. J. Johns (eds.). Pteridology in perspecitive. Royal Botanic Garden, Kew. Tryon, R. M. 1962. Taxonomic fern
notes. II. Pityrogramma (including Trismeria) and Anogramma. Contributions from the Gray Herbarium 189: 52–76. Zhang, L.-
B. & L. Zhang. 2017. Phylogeny of the fern subfamily Pteridoideae (Pteridaceae; Pteridophyta), with the description of a new
genus: Gastoniella. Molecular Phylogenetics and Evolution 109: 59–72. .

318
Figure 156. D. Gastoniella novogaliciana. F–J. G. chaerophylla (Modified from Mickel & Smith, 2004.)

319
GLEICHENIACEAE
Gleichenella Ching
DESCRIPTION: Terrestrial; rhizomes solenostelic, long-
creeping, densely pubescent, the hairs stiff, multicellular,
unbranched; leaves born in a single row along the dorsal
surface of the rhizome, up to 3 m or more long, erect
or scandent, indeterminate, of periodic growth; pinnae
opposite, unequally branched, with the larger branch
alternately to one side of the axis and then the other,
the minor branches also unequally branched; pinna axes
with a narrow lateral ridges on either side; dormant
buds in the forks pubescent, the hair dark, stiff; penultimate
divisions pectinate, glabrous and often glaucous beneath;
veins free, 2-4-forked; sori round, non-indusiate; sporangia
(6–)8–16(–25), sessile, with oblique annuli; spores
monolete, ellipsoid; x=43.
SIMILAR GENERA: Dicranopteris differs by equally
forked pinnae and (often) accessory pinnae at the forks.
Sticherus differs by scaly buds in the pinna forks, veins
usually 1-forked, and sporangia 3–6 per sorus. Gleichenella
is the only genus of Gleicheniaceae with solenostelic
rhizomes and a narrow lateral ridge on either side of the
axes of the pinnae.
COMMENTS: Gleichenella is monotypic, with its sole
species occurring from Mexico to southern Brazil and
Bolivia, and in the West Indies. Like other genera in the
family, it grows in open, sunny, often disturbed habitats. It
frequently covers steep roadsides with dense colonies.
It is the most frequent species of Gleicheniaceae at low
elevations, and its light yellowish green leaves makes
them easy to identify at a distance. Gleichenella is sister
to Dicranopteris, the genus in which it was formerly
included; therefore, its separation as a distinct genus from
Dicranopteris is only a matter of opinion. It differs from
Dicranopteris by pinna branching pattern, solenostelic
rhizomes, chromosome number, and spore morphology.
The generic name comes from Gleichenia + -ella, a
suffix denoting the diminutive.
LITERATURE: Østergaard Andersen, E. & B. Øllgaard. 1996. A note on some morphological terms of the leaf in the
Gleicheniaceae. Amer. Fern J. 86: 52–57. Østergaard Andersen, E. & B. Øllgaard. 2001. Gleicheniaceae. In: G. Harling & L.
Andersson, editors. Flora of Ecuador 66: 105–170.
320
DESCRIPCIÓN: Terrestre; rizomas solenostélicos,
largamente rastreros, densamente pubescentes, los pelos
tiesos, multicelulares, no furcados; hojas nacidas en una
sola hilera en la superficie dorsal, hasta 3 m o más de largo,
erectas o escandentes, indeterminadas, de crecimiento
periódico; pinnas opuestas, desigualmente furcadas,
con la rama más grande alternadamente a un lado y luego
en el otro, las ramas menores desigualmente ramificadas
también; ejes de las pinnas con una fila estrecha
lateral en cada lado; brotes en los furcos pubescentes,
los pelos oscuros, tiesos; divisiones penúltimos pectinadas,
glabras y a menudo glaucas en el envés; venas libres,
2-4-furcadas; soros redondos, sin indusios; esporangios
(6–)8–15(–25), séssiles, con anillo oblicuo; esporas
monoletes, elipsoides; x=43.
GÉNEROS PARECIDOS: Dicranopteris difiere por
pinnas surcadas igualmente y (a menudo) pinnas accesorias
en los furcos. Sticherus difiere por brotes escamosos en
las furcaciones de las pinnas, venas usualmente 1-furcadas
y esporangios 3–6 por soro. Gleichenella es el único
género de Gleicheniaceae con rizomas solenostélicos y
una fila estrecha en cada lado de los ejes de las pinnas.
COMENTARIOS: Gleichenella es monotípico, con su
única especie ocurriéndose desde México hasta el sur de
Brasil y Bolivia, y en las Antillas. Como los otros géneros
en la familia, crece en lugares abiertos, soleados, a menudo
en ambientes perturbados. Cubre con frecuencia taludes
de caminos con colonias densas. Es la especies de
Gleicheniaceae más frecuente en tierras bajas y sus hojas
amarillentosas verde claros lo hace fácil distinguir en la
distancia.
Gleichenella es hermana Dicranopteris, el género en
lo que fue incluido anteriormente; por eso, su separación
de Dicranopteris es solo un asunto de opinión. Difiere
de Dicranopteris por el patrón de ramificación de las
pinnas, rizomas solenostélicos, número cromosómico y
morfología de las esporas.
El nombre genérico se deriva de Gleichenia + -ella, un
sufijo denotando el diminutivo.
Figure 157. Gleichenella pectinata. Note unequal bifurcation of the pinna in A. (From Mickel & Smith, 2004.)

321
THELYPTERIDACEAE
Goniopteris C. Presl
DESCRIPTION: Terrestrial; petioles with 2 vascular
bundles; lamina simple to 1-pinnate-pinnatifid;
proximal pinnae not reduced, or only a few slightly
reduced; distal pinnae gradually shortened with a
confluent, pinnatifid apex or with a conform apex
resembling the lateral pinnae; aerophores absent; veins
free or regularly united and forming a series of areoles;
indumentum typically of minute apically furcate or
stellate hairs (in some species anchor-shaped); sori
round; indusium round-reniform or absent; sporangia
glabrous or with stellate or furcate hairs on the capsule
or stalk; x=36.
DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
vasculares; lámina simple a 1- pinnada-pinnatífida;
pinnas proximales no reducidas, o sólo unas pocas
escasamente reducidas; pinnas distales gradualmente
acortadas con un ápice pinnatífido confluente o con
una pinna apical similar en forma a las pinnas laterales;
aeróforos ausentes; nervaduras libres o regularmente
anastomosadas y formando una serie de aréolas;
indumento típicamente de pelitos furcatos o
estrellados (en algunas especies en forma de ancla);
soros redondos, con un indusio reniforme o sin indusio;
esporangios glabros, setosos o con tricomas estrellados o
furcados desde las cápsulas o el pedículo; x=36.

SIMILAR GENERA: The Ctenitis submarginalis group GÉNEROS PARECIDOS: El grupo de Ctenitis
differs by 4 or more vascular bundles in the petiole base submarginalis difiere por 4 o más haces vasculares en el
and lack of minute furcate hairs. pecíolo y falta de pelitos furcados.

COMMENTS: Goniopteris is completely neotropical and COMENTARIOS: Goniopteris es completamente

consists of about 120 species. It generally grows below
800 m in wet forests and is common in Amazonia. The
most distinctive characteristic of Goniopteris is the furcate
hairs, but these are small and a handlens is necessary
to see them. They are most easily seen on the scales at
the apex of the rhizome and in the adaxial grooves of
the lamina, but also on the lower surface of the axes.
The name of the subgenus is derived from the Greek
gonia, angle + pteris, fern. It refers to the basal veins that
converge at sharp angles.
neotropical y consta de 120 especies. Generalmente
existe abajo de 800 m en bosques húmedas y es común
en la Amazonía. La característica más distinctiva del
Goniopteris es los pelitos furcados, pero son pequeños
y una lupa es necesario verlos. Estos son más evidentes
en las escamas del ápice del rizoma y en los surcos
adaxiales, pero a menudo también en el envés en los
ejes. El nombre del subgénero se deriva del griego gonia,
ángulo + pteris, helecho. Refiere a las venas basales que
convergen en un ángulo agudo.

LITERATURE: Smith, A. R. 1973. The Mexican species of Thelypteris subgenera Amauropelta and Goniopteris. American
Fern Journal 63: 116–127. Smith, A. R. 1983. 14(4). Polypodiaceae—Thelypteridoideae. In: G. Harling & B. Sparre, Flora of
Ecuador, no. 18.

322
Figure 158. Goniopteris. A–C. G. blanda. D–F. G. tetragona. G–J. G. hatchii. L–N. G. paucipinnata. O, P. G. toganetra. Q, R.
G. imbricata. (From Mickel & Smith, 2004.)

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POLYPODIACEAE
Grammitis Sw.
DESCRIPTION: Epiphytic, rarely saxícolous or terresrial;
rhizome scales not clathrate, entire; leaves up to 20 cm
long, simple, entire, glabrous, pubescent or setulose,
with a thick, dark sclerotic margin; hydathodes present
or absent; veins simple or 1-forked, free; sori borne on
a short acroscopic veinlet, round or slightly elongate,
generally inframedial and grouped near the costa when
mature, without paraphyses; spores green. x= ?
DESCRIPCIÓN: Epiftíicas, raramente saxícolas o
terrestres; escamas del rizoma no clatradas, enteras;
hojas hasta 20 cm de longitud, simples, enteras, glabras,
pubescentes o setulosas, con un margen grueso,
esclerosado, oscuro; hidátodos presentes o ausentes;
nervaduras simples o 1-bifurcadas, libres; soros sobre una
venilla corta acroscópica, redondeados o ligeramente
alargados, generalmente inframedios y aglomerándose en
la costa cuando maduros, sin parafisos; esporas verdes.
x= ?

SIMILAR GENERA: All other genera of grammitid
ferns differ by green (not black) lamina margins. Many
others differ further by pinnatifid to 1-pinnate laminae
and the prescence of reddish setae on the laminae (only
Cochlidium, Grammitis, Lellingeria and Luisma lack setae).
GÉNEROS PARECIDOS: Todos los otros géneros de
helechos grammitidoides difieren por márgenes de las
láminas verdes (no negros). Muchos otros difieren además
por láminas pinnatífidas a 1-pinnadas y la presencia
de setas rojizas en las láminas (solamente Cochlidium,
Grammitis, Lellingeria y Luisma faltan setas).

COMMENTS: Grammitis consists of about 22 species
of both the New and Old World tropics and subtropics.
The species usually grow in cloud forests. Molecular data
indicate that Grammitis as defined here (i.e., restricted
to only those species with black laminar margins) are
monophyletic (Ranker et al. 2004). Although the genus
has been revised three times this century (Bishop, 1977;
Copeland, 1952; Maxon, 1915), relatively few specimens
were available for study. The many recent collections,
some of which have been described as new species, make
necessary a new revision of the genus. The genus name
is derived from the Greek gramme, line, referring to the
elongate to linear sori.
COMENTARIOS: Grammitis consta de casi 22 especies
de los trópicos y subtrópicos del Nuevo yViejo Mundo. Las
especies usualmente crece en bosques nublados. Datos
moleculares sugieren que Grammitis como definido aquí
(i.e., restringido a las especies con márgenes laminares
negros) son monofíleticos (Ranker et al. 2004).A pesar
de que el género ha sido revisado tres veces en este siglo
(Bishop, 1977; Copeland, 1952; Maxon, 1915), se disponía
de relativamente pocos especímenes para los estudios.
La gran cantidad de colecciones recientes, algunas de las
cuales han sido descritas como especies nuevas, hacen
necesaria una nueva revisión del género. El nombre
genérico viene del griego gramme, línea, refiriéndose a
los soros alargados o linales.

LITERATURE: Bishop, L. E. 1977. The American species of Grammitis sect. Grammitis. American Fern Journal 67: 101–106.
Copeland, E. B. 1952. Grammitis. Philippine Journal of Science 80: 93–276, t. 1–6. Maxon, W. R. 1915. Polypodium marginellum
and its immediate allies. Bulletin of the Torrey Botanical Club 42: 219–225. Labiak, P. H. & J. Prado. 2003. Grammitidaceae
(Pteridophyta) no Brasil com ênfasis nos gêneros Ceradenia, Cochlidium, e Grammitis. Hoehnea 30: 243–283. Ranker, T. A., A.
R. Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Staub & H. Schneider. 2004. Phylogeny and evolution and
grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy. Taxon 53: 415–428.

324
Figure 159. A–C. Grammitis bryophila (Maxon) F. Seym. D–F. G. marginella (Sw.) Sw. G–­J. G. leptopoda (C. H. Wright)
Copel.. K–M. Lomaphlebia graminea (Sw.) comb. ined.? (Original, © Robbin Moran, 2016.)

325
CYSTOPTERIDACEAE
Gymnocarpium Newman
DESCRIPTION:Terrestrial; rhizomes long-creeping, scaly;
sterile and fertile leaves monomorphous, thin-textured;
petioles with 2 vascular bundles; laminae usually
10–30 cm long, broadly deltate, 2-3-pinnate-pinnatifid;
pinnae with a slightly swollen point of attachment to the
rachis; axes grooved on the upper surface, the grooves
not decurrent; veins free, the tips slender (not enlarged);
indument lacking or of minute (0.1 mm long) capitate-
glandular hairs; sori round, medial or inframedial; indusia
absent; spores monolete, bilateral; x=40.
DESCRIPCIÓN: Terrestres; rhizomas largamente
rastreros, escamosos; hojas estériles y fértiles
monomorfas, de textura delgada; pecíolos con 2 haces
vasculares; láminas usualmente 10­ 30 cm de largo,
anchamente deltadas, 2-3-pinnado-pinnatífidas; pinnas
con un punto de unión con el raquis levemente hinchado;
ejes surcados en la superficie superior, los surcos no
decurrentes; indumento ausente o consta de pelitos
capitados glandulares diminutos (0.1 mm de largo);
nervaduras libres, los ápices delgados (no engrosados);
soros redondeados, mediales o inframediales; indusios
ausentes; esporas monoletes, bilaterales; x=40.

SIMILAR GENERA: Cystopteris differs by (usually) GÉNEROS PARECIDOS: Cystopteris difiere por
lanceolate laminae and indusiate sori. (usualmente) láminas lanceoladas y soros indusiados.

COMMENTS: Gymnocarpium contains 8 species and COMENTARIOS: Gymnocarpium consta de 8 especies

is of temperate distribution in North America and
Eurasia. All species are terrestrial in shaded forests or
on shaded limestone. They tend to form diffuse colonies
by the action of their long-creeping rhizomes. Molecular
phylogenetic studies indicate that Gymnocarpium is
sister to Cystopteris, and the two are sister to the rest of
Eupolypods II. Thus, the two genera deserve their own
family, Cystopteridaceae. This is a deviation from Smith
et al. (2006) generally followed here. The genus name is
derived from the Greek gymnos, naked + karpos, fruit. It
refers to the absence of an indusium.
y es principalmente de distribución templada en América
del Norte y Eurasia. Todas las especies son terrestres
en bosques sombreados o en piedra caliza sombreada.
Tienden formar colonias difusas por acción de sus rizomas
largamente rastreros. Estudios filogenéticos molecular
indican que Gymnocarpium es hermana a Cystopteris, y los
dos son hermana al resto de Eupolypods II. Por ende los
dos géneros merecen su propia familia, Cystopteridaceae
.Esto es una desviación de la clasificación de Smith et al.
(2006) generalmente seguido aquí. El nombre genérico
se deriva del griego gymnos, desnudo + karpos, fruta. Se
refiere a la ausencia del indusio.

LITERATURE: Sarvela, J. 1978. A synopsis of the fern genus Gymnocarpium. Ann. Bot. Fenn. 15: 101–106. Sarvela, J., D. M.
Britton & K. M. Pryer. 1981. Studies on Gymnocarpium robertianum complex in North America. Rhodora 83: 421–431.

326
Figure 160. A–D. Gymnocarpium dryopteris. (Original, ©R. C. Moran, 2016.)

327
CYATHEACEAE
Gymnosphaera Blume
DESCRIPTION: Stems arborescent; sterile and fertile
leaves monomorphic or slightly dimorphic; petioles with
4–10 pairs of small, skeletonized pinnae (aphlebia)
near the base and distant from the normal pinnae;
petiole scales with single black apical setae, the
marginal cells different in shape and orientation from
the central ones (i.e., marginate); laminae 1-pinnate-
pinnatifid to 4-pinnate, the apex pinnatifid or rarely
conform; veins free; axes pubescent adaxially, the hairs
thick, strigose, multiseptate; sori round, the receptacle
elevated, subglobose; indusia absent or (in A. capensis)
present and hemitelioid; spores tetrahedral-globose, 64
per sporangium, ridged, without pores; x=69.
SIMILAR GENERA: Alsophila differs by 16 spores
per sporangium and lack of aphlebia. Cyathea and
Sphaeropteris differ by lacking an aerophore pit on the
rachis at the pinna juncture. Sphaeropteris further differs
by conform petiole scales (i.e., all cells of the same shape
and orientation).
COMMENTS: Gymnosphaera contains about 16 species
and is entirely Old World except for G. capensis (SE Brazil,
Africa) and G. salvinii (Central America, Peru). It occurs
in shaded forests. In a phylogenetic analysis (Korall et al.,
2007), Gymnosphaera formed a trichotomy with Alsophila
and Cyathea. All three genera have marginate petiole
scales. As in Alsophila, the aerophore in Gymnosphaera is
a sunken pit in the rachis at the juncture of the costa,
and there is a single black apical seta on the petiole
scales.Gymnosphaera has dark rachises and costae, and
this will distinguish it from many (not all) scaly tree
ferns. Many Old World species have the fertile pinnae
contracted compared to the sterile. Only G. capensis has
an indusium; the others in the subgenus are non-indusiate.
The name is derived from the Greek gymnos, naked +
sphaera, ball. It refers to the non-indusiate sori.
LITERATURE: Conant, D. S. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. Journal of the Arnold
Arboretum 64: 333–382. Korall, P., D. S. Conant, J. S. Metzgar, H. Schneider & K. M. Pryer. 2007. A molecular phylogeny
of scaly tree ferns. American Journal of Botany 94: 873–886.
328
DESCRIPCIÓN: Tallos arborescentes; hojas estériles
y fértiles monomorfas o levemente dimorfas; pecíolos
con 4–10 pares de pinas pequeñas en esqueletos
(aflebias) cerca la base del pecíolo y distantes
de las pinnas normales; escamas peciolares
con una seta apical negra, las células marginales
difieren en forma y orientación de las centrales (i.e.,
marginadas); láminas 2-pinada-pinatífidas a 3-pinadas,
el ápice gradualmente reducido; nervaduras libres; ejes
pubescentes adaxialmente, los pelos gruesos, estrigosos,
multiseptados; soros redondos, el receptáculo elevado,
subgloboso; indusios ausentes o (en A. capensis) presentes
y hemitilioides; esporas tetrahédricas-globosas, 64 por
esporangio, con crestas bajas, sin poros; x=69.
GÉNEROS PARECIDOS: Alsophila differs by 16
spores per sporangium y la falta de aflebias. Cyathea y
Sphaeropteris difieren por la falta de un aeróforo como
hueco por el raquis en la unión de la pinna. Sphaeropteris
difiere además por escames peciolares conformes (i.e.,
todas la células del mismo forma y orientación).
COMENTARIOS: Gymnosphaera consta de casi 16
especies y es completamente del Viejo Mundo con
excepción de G. capensis (SE Brasil, África) y G. salvinii
(América Central, Peru). Existe en bosques sombreados.
En un análisis filogenético (Korall et al., 2007),
Gymnosphaera formó una tricotomía con Alsophila y
Cyathea. Los tres géneros tienen escamas peciolares
marginadas. Como en Alsophila, el aeróforo de
Gymnosphaera es un hueco hundido en el raquis cerca de
la unión de la costa, y hay una sola seta negra apical en las
escamas peciolares. Gymnosphaera tiene ráquises y costas
oscuras, y esto lo distinguirá de muchos otros (no todos)
helechos arborescentes con escamas. Muchas especies
del Viejo Mundo tienen las pinnas fértiles contraídas en
comparación de las estériles. Solo G. capensis tiene un
indusio; las otras en el subgénro son no indusiados. El
nombre se derive del griego gymnos, desnudo + sphaera,
pelota. Se refiere a los soros no indusiados.
Figure 161. A–B, H–M. Gymnosphaera salvinii, in note aphlebiae (skeletonized pinnae) on petiole base. C–G. G. capensis ssp.
polypodioides (Original, © R. C. Moran, 2016.)

329
PTERIDACEAE (VITTARIOID CLADE)
Hecistopteris J. Sm.
DESCRIPTION: Epiphytic or rupicolous; rhizomes
short-erect, proliferous roots present; rhizome scales
clathrate, not obscured by densely clumped pubescent
roots; petioles absent or very short; laminae less than 4
cm long, incised, narrowly cuneate, flabellate, glabrous,
with spicular idioblasts (false veins) adaxially; veins free;
sori elongate along the veins in the distal portion of the
lamina; paraphyses simple or branched, with an enlarged
apical cell; spores trilete, nongreen; gametophytes with
unpaired gemmae; x=?
SIMILAR GENERA: None.
COMMENTS: Hecistopteris is entirely neotropical
and consists of three species. They grows primarily
as an epiphyte in wet, shaded forests below 600 m. It
sometimes grows on rotting logs.The plants form colonies
by elongate rootlets that bear buds, and it is typical to
find several plants united by a single rootlet.The rhizomes
are often descending-erect with the leaves clustered at
the apex (pers. obs.) For over 150 years Hecistopteris
was considered monotypic, containing the sole species
H. pumila (Spreng.) J. Sm., which occurs from Mexico to
southern Brazil and Bolivia, and the Antilles. But recently
a species was described from Guyana (H. kiaeteurensis
Kelloff & G. McKee), and another from Ecuador (H.
pinnatifida R. C. Moran & Øllgaard). Hecistopteris is sister
to Radiovittaria (Crane, 1997) and differs from that genus,
and all other vittarioid ferns, by incised leaves, proliferous
roots, and free veins. The genus name is derived from the
Greek hekistos, least + pteris, fern, referring to the small
size of H. pumila.
LITERATURE: Crane, E. H. 1997. A revised circumscription of the genera of the fern family Vittariaceae. Systematic Botany
22: 509–517. Kelloff, C. L. & G. S. McKee. 1998. A new species of Hecistopteris from Guyana, South America. American Fern
Journal 88: 155–157.
330
DESCRIPCIÓN: Epífitas o rupícolas; rizomas
cortamente erectos, raíces prolíferas presentes;
escamas del rizoma clatradas, no oscurecidas por raíces
cespitosas densamente pelosas; pecíolos ausentes o muy
reducidos; láminas menos de 4 cm de largo, incisas,
estrechamente cuneada en la base, flabelada, glabra, con
idioblastas spiculares (venas falsas) en el haz; nervaduras
libres; soros alargados junto a las nervaduras en la
porción distal de la lámina; parafisos simples o ramificados
con una célula apical agrandada; esporas triletes, no
verdes; gametofitos con yemas no pareadas; x=?
GÉNEROS PARECIDOS: Ningunos.
COMENTARIOS: Hecistopteris es completamente
neotropical y consta de tres especies. Crecen
principalmente como epífito en bosques húmedos y
sombreados bajo de 600 m. Algunas veces crece sobre
troncos podridos. Las plantas forman colonias por
raicillas alargadas que llevan yemas adventicias, y es típico
encontrar varias plantas unidas a una raicilla única. Los
rizomas son a menudo descendentes-erectos con las
hojas agrupadas en el ápice (pers. obs.). Por mas de 150
años se considera a Hecistopteris monotípico, conteniendo
la única especie H. pumila (Spreng.) J. Sm., la que ocurre
desde México hasta el sur de Brasil y Bolivia, y las Antillas.
Pero recientemente una especie fue descrita de Guyana
(H. kiaeteurensis Kelloff & G. McKee) y otra de Ecuador
(H. pinnatifida R. C. Moran & Øllgaard). Hecistopteris es
hermana a Radiovittaria (Crane, 1997), y difiere de ese
género y todos los demás helechos vittarioides por sus
hojas incisas, raíces prolíferas y venas libres. El nombre
genérico se deriva del griego hekistos, menos + pteris,
helecho, refiriéndose al tamaño pequeño de H. pumila.
Figure 162. Two species of Hecistopteris. A–D. Hecistopteris pinnatifida. A. Habit showing root proliferation. B. Habit. C.
Paraphyses with enlarged apical cell. D. Clathrate rhizome scale. E. H. pumila, habit. (From Moran & Øllgaard, 1995.)

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HEMIDICTYACEAE
Hemidictyum L.
DESCRIPTION: Terrestrial; leaves ca. 3 m, erect-arching;
sterile and fertile leaves monomorphous; the base of the
petiole with 2 vascular bundles, these uniting distally in
a U-shape; laminae 1-pinnate with a terminal pinnae
similar in form to the lateral ones; pinnae 23–50 X
6–12 cm, linear-oblong, entire, opposite to subopposite,
sessile, slightly cordate and overlapping the rachis; veins
areolate toward the marings, a submaringal connecting
vein present; indusia narrow, linear membranaceous;
spores bilateral, brown; x=31.
DESCRIPCIÓN: Terrestres; hojas c. 3 m, erectas-
arqueadas; hojas estériles y fértiles monomorfas; la base
del pecíolo con 2 haces vasculares, estos uniéndose
distalmente en forma de “U”; láminas 1-pinnadas
con una pinna terminal similar en forma a las
pinnas laterales; pinnas 23–50 X 6–12 cm, linear-
oblongas, enteras, opuestas o subopuestas, la base
sésil, ligeramente cordiformes y traslapadas al raquis;
nervaduras anastomosadas hacia los márgenes, una
vena colectora submarginal presente; indusios angostos,
lineares, membranáceos; esporas bilaterales; x=31.

SIMILAR GENERA: Asplenium differs by smaller leaves
(never more than 1 m long). Diplazium differs by at least
some of the sori (usually the basal ones) back to back
(diplazioid); none of its species have entire pinnae and
anastomosing veins.
GÉNEROS PARECIDOS: Asplenium difiere por hojas
más pequeñas (nunca más de un metro de longitud).
Diplazium difiere por tener a lo menos unos soros
(usualmente los basales) espalda a espalda (diplazioide);
ninguna de sus especies tienen pinnas enteras y venas
anastomosadas.

COMMENTS: Hemidictyum consists of only one species,
H. marginatum (L.) C. Presl, which occurs throughout
most of the Neotropics. It generally grows in wet forests
or forest margins from 0–1300 m. The Hemidictyaceae
is monotypic. Although sister to Aspleniaceae, it
resembles certain species of the Asian Diplaziopsis in the
Diplaziopsidaceae, the closest outgroup family. The genus
name is derived from the Greek hemi, half + diktyon, net.
The veins anastomose in the distal half of the pinnae.
COMENTARIOS: Hemidictyum consta de una sola
especie, H. marginatum (L.) C. Presl, la cual existe por
la mayoría de los Neotropicos. Generalmente existe
en bosques húmedos o por los bordes de bosques de
0–1300 m. Hemidictyaceae es monotípico. Aunque es
hermana a Aspleniaceae, se asemeja ciertas especies
del Asiático Diplaziopsis en Diplaziopsidaceae, la familia
grupo afuera más relacionada. El nombre genérico se
deriva del griego hemi, mitad + dictyon, red. Las venas se
anastomosen en la mitad distal de las pinnas.

LITERATURE: Øllgaard, B. 1994. Hemidictyum. In: G. Harling & L. Andersson, editors. Flora of Ecuador, 14 (5B). Polypodiaceae-
Dryopteridoideae-Physematieae 49: 1–108. Schuettpelz, E. & K. M. Pryer. 2007. Phylogeny of ferns inferred from 400
leptosporangiate species and three plastid genes. Taxon 56: 1037–1050.

332
Figure 163. Hemidictyum marginatum. A. leaf apex. B. sori and venation. C. pinna margin. (From Stolze, 1981.)

333
PTERIDACEAE
Hemionanthes Mickel
DESCRIPTION: Terrestrial; rhizomes compact,
horizontal, the scales linear, bicolorous with a dark central
stripe; petioles stramineous, mostly glabrous, 3-grooved,
ca. half the length of the leaf; blades lanceolate to deltate,
pinnate-pinatifid to 2-pinnate basally; pinnae 5–7 pairs,
the lobes obtuse, basiscopically somewhat exaggerated,
the adaxial surfaces glabrous, abaxial surfaces with
sparse straight jointed hairs; rachises with sparse narrow,
bicolorous scales; veins free with a few casual anastomoses
(2–5 per pinna); sporangia extending along the distal
2–4 mm of the veins; indusial absent, blade margins not
reflexed or modified; spores tetrahedral-globose, crested;
x=30.
DESCRIPCIÓN: Terrestres; rizomas compactos,
horizontales, las escamas lineares, bicolores con una raya
central oscura; pecíolos estramineosos, principalmente
glabros, 3-surcados, ca. Una mitad del largo de la hoja;
láminas lanceoladas a deltadas, pinnadas-pinnatífidas
a 2-pinnadas basalmente; pinnas 5–7 pares, los lobos
obtusos, basiscopicamnte also exagerado, superficies
adaxiales con pelos articulados esparsos; ráquises con
escamas bicoloras angostas esparsas; venas libres con
unas pocas anastomoses casuales (2–5 por pinna);
esporangios extendiéndose por el distal 2–4 mm de
las venas; indusios ausentes, márgenes de las láminas no
recurvados o modificados; esporas tetrahédrico-globosas,
crestados; x=30.

SIMILAR GENERA: Cheilanthes skinneri differs by leaves GÉNEROS PARECIDOS: Cheilanthes skinneri difiere
2- to 4-pinnate-pinnatifid and stalks of the pinnae (6– por hojas 2- a 4-pinnado-pinnatífidas y los peciolulos
)10–25 mm long, and sori at the apices of the veins. de las pinnas (6–)10–25 mm de longitud y soros en los
apices de las venas.

COMMENTS: Hemionanthes consists of three species:
H. gryphus (Mickel) Mickel, H. lozanoi (Maxon) comb. ined.
y H. seemannii (Hook.) comb. ined. All are endemic to
Mexico. They grow in deciduous woodlands and moist
shaded gullies. Hemionanthes gryphus has the appearance
of a hybrid between Cheilanthes skinneri and Hemionitis
subcordata, which can grow together with it; however,
Hemionanthes is not a hybrid. It has normal spores and is
only distantly related to the two species it resembles.The
genus name is derived from the intermediate appearance
of the species between Cheilanthes skinneri and Hemionitis
subcordata.
COMENTARIOS: Hemionanthes consta de tres species:
H. gryphus (Mickel) Mickel, H. lozanoi (Maxon) comb. ined.
y H. seemannii (Hook.) comb. ined. Todas son endémicas
a México. Crecen en bosques decíduos en arroyos
sombreados húmedos. Hemionanthes gryphus tiene
la apariencia de un híbrido entre Cheilanthes skinneri y
Hemionitis subcordata, que pueden crecer juntos con
él; no obstante, Hemionanthes no es un híbrido. Tiene
esporas normales y es solo relacionado distantamente a
las dos especies que se asemeja. El nombre genérico se
deriva de la apariencia intermedia de la especies entre
Cheilanthes skinneri and Hemionitis subcordata.

LITERATURE: Mickel, T. T. 1987. A new fern from western Mexico and its bearing on the taxonomy of the cheilanthoid
ferns. American Fern Journal 77: 109–114.

334
Figure 164. Hemionanthes gryphus. A. Habit. B. Rhizome scale. C. Pinnule. (From Mickel & Smith, 2004.)

335
PTERIDACEAE
Hemionitis L.
DESCRIPTION: Terrestrial or rupestral; rhizomes erect
to short-creeping, scaly, the scales (at least those toward
the apex of the rhizome) concolorous, but those on the
older part of the rhizome bicolorous with a dark central
strip or concolorous (in H. palmata); leaves monomorphic
or dimorphic; petioles castaneous, dark brown or straw-
colored, with 1 vascular bundle; laminae simple, orbicular
to 1-2-pinnate, densely to sparsely hairy, occasionally
on the lower surface with two-celled, gland-tipped hairs;
veins netted or free, if netted then the areoles without
included veinlets; sori running along the veins,
sometimes (in H. subcordata ) restricted to the distal one-
quarter of near the margin; indusium absent; x=30.
DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto a
cortamente rastrero, escamoso, las escamas concoloras
hacia el ápice del rizoma, aquellas en las porciones más
viejas del rizoma bicolores, con una franja media castaño
o concolora (en H. palmata); hojas monomorfas a
dimorfas; pecíolo castaño, pardo oscuro o pajizo, con un
haz vascular; lámina simple, orbicular, pedata, pinnatífida
o 1-2-pinnada, densa a esparcidamente pilosa,
ocasionalmente con tricomas 2-celulares con glándulas
en las puntas en el envés; nervaduras o anastomosadas, las
aréolas sin nérvulos incluidos; esporangios cubriendo
las nervaduras a todo lo largo o a veces restringidos al
1/4 marginal (en H. subcordata ); indusio ausente; x=30.

SIMILAR GENERA: Bommeria differs by sori restricted GÉNEROS PARECIDOS: Bommeria difiere por los
near margin, not extending completely between the soros restringidos cerca del margen, no extendiéndose
midrib and margin. completamente entre la costa y el margen.

COMMENTS: Hemionitis has 10 species, all Neotropical.
The genus typically grows in mesic habitats such as shaded
rocks, tall forests, and along rivers. Two clades occur
within the genus: one with palmate laminae and netted
veins (Hemionitis s.s.) and another with pinnate lamina
and free veins (formerly distinguished as Gymnopteris
Bernh.).Hemionitis arifolia (Burm. f.) T. Moore, a commonly
cultivated Asian species, was resolved sister to the four
neotropical Hemionitis species sampled by Zhang et
al. (2010). The genus name is derived from the Greek
hemionis, mule. Ancient name for a fern thought to be a
hybrid.
COMENTARIOS: Hemionitis tiene 10 especies,
todas Neotropicales. El género crece típicamente en
hábitats mesofíticas como rocas sombreadas, selvas
altas perennifolias, o comunidades raparías. Dos clados
existen dentro del género: uno con láminas palmadas y
venas anastomosadas (Hemionitis s.s.) y otra con laminas
1-pinnadas y venas libres (anteriormente distinguido
como Gymnopteris Bernh.). Hemionitis arifolia (Burm. f.) T.
Moore, una especie Asiática comúnmente cultivada, fue
resuelta hermano a los cuatro especies neotropicales de
Hemionitis muestradas por Zhang et al. (2010). El nombre
genérico se deriva del griego hemionis, mula. Nombre
anciano para un helecho pensado ser un híbrido.

LITERATURE: Maxon, W. R. 1913. Studies of tropical American ferns—No. 5. Contributions from the United States National
Herbarium 17: 133–179. Mickel, J. T. 1974. A redefinition of Hemionitis. American Fern Journal 64: 3–12. Ranker, T. A. 1989.
Spore morphology and generic delimitation of New World Hemionitis, Gymnopteris, and Bommeria (Adiantaceae). American
Journal of Botany 76: 297–306. Ranker, T. A. et al. 1989. Genetic evidence for allopolyploidy in the neotropical fern Hemionitis
pinnatifida (Adiantaceae). Systematic Botany 14: 439–447. Zhang, G-m., H-m Liu, W-l Yang & X-C. Zhang. 2009. Systematic
position of Hemionitis arifolia based on the analysis of rbcL sequences. Journal of Beijing Forestry University 31: 15–18.

336
Figure 165. A. Hemionitis pinnatifida. B. H. levyi. C. H. palmata. (From Mickel & Smith, 2004.)

337
DENNSTAEDTIACEAE
Histiopteris (J. G. Agardh) J. Sm.
DESCRIPTION: Terrestrial; rhizomes long-creeping, 2-5
mm wide, scaly; leaves large, distant, erect to subscandent;
petiole base in cross-section with a vascular bundle in
the form of a corrugated “U”; blades 2- to 3-pinnate,
often with large stipule-like pinnules at the pinna
base; pinnae and pinnules opposite, light pea-green,
herbaceous; lamina glabrous or with crisped, multicellular
hairs, often glaucous; veins netted, without included
veinlets; sori marginal, continuous, supplied by more than
one vein, blade margin recurved and differentiated as an
indusium (false indusium); inner indusium absent; spores
bilateral; x=96.
SIMILAR GENERA: Blotiella differs by shorter sori,
pubescent laminae and absence of stipule-like pinnules
at the base of the pinnae. Hypolepis differs by sori
supplied by a single vein. Lonchitis, which differs by free
veins, pubescent laminae, and the presence of an inner
indusium.
COMMENTS: Histiopteris is pantropical and consists of
about 7 species worldwide. Only one, H. incsa (Thunb.) J.
Sm., is Neotropical. It occurs primarily from 1000–3000
m. Six other species occur in Southeast Asia and the
South Pacific.The leaf grows intermittently.The apex rests
while the pinnae pair beneath develop. After the pinnae
are nearly fully developed, the apex resumes growth.Thus
the leaves almost always have a fiddlehead at their apex,
this usually protected by the enlarged basal pinnules.
Histiopteris is the only genus of Dennstaedtiaceae that
has scaly rhizomes (the others are pubescent). The genus
name is derived from the Greek histion, sail + pteris, fern.
It refers to the small sail-like basal pinnules.
LITERATURE: none; ninguna
338
DESCRIPCIÓN: Terrestres; rizomas largamente
rastreros, 2-5 mm de ancho, escamosos; hojas grandes,
distantes, erectas a subescandentes; la base del pecíolo
en sección cruz con un hace vascular en forma de un
“U” corrugado; láminas 2- a 3-pinnadas, a menudo con
pínnulas que asemejan a estípulas; pinnas y pínnulas
opuestas, verde claras, herbáceas, glabras o con pelos
multicelulares crespadas, a menudo glaucas; venas
anastomosadas, sin venas incluídas; soros marginales,
contínuos; suministrados por más de una vena, margen
de la lámina recurvado y diferenciado como indusio
(indusio falso); un indusio interior ausente; esporas
bilaterales; x=96.
GÉNEROS PARECIDOS: Blotiella difiere por soros más
cortos, láminas pubescentes y la ausencia de pínnulas que
asemejan como estípulas en la base de las pinnas. Lonchitis,
lo cual difiere por venas libres, láminas pubescentes, y la
presencia de un indusio interior.
COMENTARIOS: Histiopteris incsa (Thunb.) J. Sm. es
pantropical y consta de casi 7 especies mundial. Solomente
una, H. incsa (Thunb.) J. Sm., es Neotrópical. Existe
principalmente desde 1000–3000 m. Seis otras especies
existen en el sureste de Asia y el Pacífico del sur. La hoja
crece intermitente. El ápice descansa mientras que el par
de pinnas abajo se desarrollan. Después de que las pinnas
son casi completamente desarrolladas, el ápice resume
crecimiento. Por eso, las hojas casi siempre tienen un
cayado en sus ápices, esto usualmente protegido por las
pínulas basales agrandados. Histiopteris es el único género
de Dennstaedtiaceae que tiene rizomas escamosos (los
otros son pubescentes). El nombre genérico se deriva del
griego histion, vela + pteris, helecho. Se refiere a la pínnula
pequeña basal que asemeja una vela.
Figure 166. Histiopteris incsa. A. Pinnae. B. Creeping rhizome and petiole base with epipetiolar bud. C. Fertile pinnule and net
veins. (From Mickel & Smith, 2004.)

339
DIPLAZIOPSIDACEAE
Homalosorus Small ex Pic. Serm.
DESCRIPTION: Terrestrial; rhizomes creeping to erect,
scaly, the scales not clathrate; sterile and fertile leaves
subdimorphic, ceaspitose; petiole bases with 2 vascular
bundles, these uniting distally to form a “U”; laminae
1-pinnate, hairs absent, scales sparse to absent; pinnae
sessile, equilateral, truncate or nearly so at base; rachis
sulcate adaxially, lacking hairs within the groove; veins free,
usually ending before the margin in clavate hydathodes;
fertile leaves longer petiolate with narrower pinnae; sori
elongate, approximate, filling most of the laminar
tissue between the costa and margin, occasionally
paired back to back along a single vein; indusia present;
annulus of 15–20 cells; spores bilateral, brown, cristate;
x=40.
DESCRIPCIÓN: Terrestres; rizomas rastreros a erectos,
escamosos, las escamas no clatradas; hojas estériles y
fértiles subdimorphic; bases de los pecíolos con 2 haces
vasculares, éstos uniéndose distalmente para formar
un haz vascular en forma de “U”; láminas 1-pinnadas,
pelos ausentes, escamas esparcidas al ausentes; ráquises
surcados adaxialmente, sin pelos en los surcos; pinnas
séssiles, equiláterales, truncadas o casi así en la base;
venas libres, usualmente terminándose antes del margen
en hidatodos clavados; hojas fértiles más largamente
pecioluladas con pinnas más estrechas; soros alargados,
aproximados, llenando la mayoría del tejido láminar
entre la costa y márgen, a veces pareados de par en
par por una sóla vena; indusios presentes; anillo de 15–20
células; esporas bilaterales, pardas, crestadas; x=40.

SIMILAR GENERA: Deparia differs by jointed, silvery
hairs on the leaves. Athyrium and Diplazium differs by
rachises and costae grooved adaxially with the grooves
decurrent into one another, and usually by having some
hairs on the leaves, at least in the grooves adaxially. In both
genera, the sori are rarely as crowded and filling so much
space between the costa and margin as in Homalosorus.
GÉNEROS PARECIDOS: Deparia difiere por pelos
plateados articulados en las hojas. Athyrium y Diplazium
difieren por ráquises y costas surcados adaxialmente con
los surcos decurrentes, y usualmente con algunos pelos
foliares, al menos en los surcos adaxialmente. En ambos
géneros, los soros raras veces son tan congestionados y
llenando tanto espacio entre la costa y el márgen como
en Homalosorus.

COMMENTS: Homalosorus consists of a single species:
H. pycnocarpon (Spreng.) Pic. Serm. It is native to
eastern North America where it grows in wet forests.
Morphologically, it most resembles Diplazium flavoviride
Alston (Kato & Darnaedi, 1988), a species of eastern
Malesia that should be transferred to Homalosorus. The
sori are single along the veins; rarely are they back-to-
back as in Diplazium. The young indusia are inflated and
silvery, turning brown with age. Homalosorus is sister
to Diplaziopsis, an Asian genus. Together they form
the Diplaziopsidaceae, a family closely related to the
Hemidyctiaceae and Aspleniaceae (Wei et al., 2010). The
genus name is derived from the Greek homalos, even,
smooth + soros. It refers to the evenly spaced sori.
COMENTARIOS: Homalosorus consta de una sola
especie: H. pycnocarpon (Spreng.) Pic. Serm. Es nativa a
América del Norte oriental donde existe en bosques
húmedos. Morfológicamente, se asemeja más a Diplazium
flavoviride Alston (Kato & Darnaedi, 1988), una especie de
Malesia oriental que debe ser transferida a Homalosorus.
Los soros son simples por las venas; raras veces son
espaldas a espaldas como en Diplazium. Los indusios
jóvenes son infladas y plateadas, tornándose pardos con
edad. Homalosorus es hermana a Diplaziopsis, un género
asiático. Juntos, forman la Diplaziopsidaceae, una familia
emparentada con Hemidictyiaceae y Aspleniaceae (Wei
et al., 2010). El nombre genérico se deriva del griego
homalos, uniforme, plano + soros. Se refiere a los soros
uniformamente espaciados.

LITERATURE: Kato, M. & D. Darnaedi. 1998. Taxonomic and phytogeographic relationships of Diplazium flavoviride, D. pycnocarpon,
and Diplaziopsis. American Fern Journal 78: 77–85. Sano, R., M. Takamiya, M. Ito, S. Kurita & M. Hasebe. 2000. Phylogeny of the lady
fern group, Tribe Physematieae (Dryopteridaceae), based on chloroplast rbcL gene sequences. Molecular Phylogenetics and Evolution 15:
403–413. Wei, R., X-C. Zhang & X-P. Qi. 2010. Phylogeny of Diplaziopsis and Homalosorus based on two chloroplast DNA sequences:
rbcL and rps4–trnS IGS. Acata Botanica Yunnanica (Suppl.) 17: 36–54.

340
Figure 167. Homalosorus pycnocarpon (From Small, 1935.)

341
LYCOPODIACEAE
Huperzia Bernh.
DESCRIPTION: Plants terrestrial; horizontal creeping
stems absent, erect stems isodichotomously branched,
reclining with age; trophophylls isophyllous, entire or
minutely toothed; gemmae borne on short branches
(gemmiphores) on the distal portions of stems;
strobili absent, the sporangia produced in zones along the
shoot; sporophylls similar to the trophophylls, remaining
green after the sporangia dehisce; sporangia isovalvate;
sporophylls similar to the trophophylls, attached at the
base (not peltate), remaining green after sporangial
dehiscence; spores foveolate, in polar view with
truncate angles and concave sides; gametophytes
subterranean, cylindrical; x=67, 68.
DESCRIPCIÓN: Plantas terrestres; tallos horizontales
reptantes ausentes, tallos erectos ramificados
isodicotómicamente, reclinándose con edad; trofofilos
isofílicos, enteros o diminutamente dentados; yemas
se nacen en ramas cortas (gemíforos) en la parte
distal de los tallos; estróbilos ausentes, los esporangios
se nacen en zonas por el vástago; esporofilos simulares
a los trofofilos, o los esporofilos unidos en la base
(no peltados), permaneciéndose verdes después
de la dehiscencia; esporas foveoladas, en vista polar
con ángulos truncados y lados cóncavos; gametofitos
subterráneos, cilíndricos; x=67, 68.

SIMILAR GENERA: Phlegmariurus differs by lacking
gemmae and having spores in polar view that are
suborbicular with convex sides and smooth (not foveolate)
proximal faces. The other genera of Lycopodiaceae differ
by having long-creeping horizontal branches, these giving
rise to unequally branched erect stems, and peltate
sporophylls that turn yellowish or straw-colored after
sporangial dehiscence.
GÉNEROS PARECIDOS: Phlegmariurus difiere por la
falta de yemas y tener esporas suborbiculares en vista
polar con lados convexos y caras proximales lisas (no
foveoladas). Los otros géneros de Lycopodiaceae difieren
por tener ramas horizontales principales largamente
rastreras, estos dando lugar a sistemas erectas de ramitas
desigualmente ramificadas y esporofilos peltatdos
tornándose amarillentos o pajizos después de la
dehiscencia esporangial.

COMMENTS: Huperzia consists of about 15 species COMENTARIOS: Huperzia consta de casi 15 especies
worldwide. It occurs predominantly in the Northern mundial. Existe principalmente en el Hemisferio Norte,
Hemisphere, especially in temperate, arctic, and alpine especialmente en regiones templadas, árticas y alpinas.
regions. Two species occur in the Neotropics. Huperzia Dos especies existen en el Neotrópico. Huperzia
beiteliana is endemic to southern Mexico. Huperzia beiteliana es endémica al sur de México. Huperzia serrata
serrata (Thunb.) Trevis. occurs mainly in southeastern (Thunb.) Trevis. existe principalmente en el sureste de
Asia but has Neotropical records in Mexico, Greater Asia pero tiene regístros neotropicales en México, las
Antilles, and southern Brazil. Huperzia has “cortical roots” Antillas Mayores y el sur de Brasil. Huperzia tiene “raíces
that grow through the cortex for varying distances until corticales” que crecen por la corteza varias distances
they emerge at soil level. Distinctive of Huperzia are the hasta que emergen al nivel del suelo. Distintiva de
gemmae—small, six-leaved plantlets borne on specialized Huperzia son las yemas—plántulas pequeñas con seis
branches called gemmiphores. Their structure and origin hojas que se nacen en ramas especializadas llamadas
were described by Stevenson (1976). The gemmae easily gemíforos. Su estructura y origen fueron descritos por
detach, reproducing the plants asexually. Aborted-spored Stevenson (1976). Las yemas se desprenden fácilmente,
hybrids may become locally common by this method of reproduciendo las plantas asexualmente. Híbridos
vegetative reproduction. Huperzia is the only genus in the con esporas abortivas pueden llegar a ser comunes
family in which abortive spores are regularly produced. localmente por este método de reproducción. Huperzia
The genus is named for Johann Peter Huperz (d. 1816), es el único género en la familia en cual esporas abortivas
author of a book on fern propagation. son producidas regularmente. El nombre genérico honra
Johann Peter Huperz (d. 1816), autor de un libro sobre la
propagación de helechos.
LITERATURE: Haines, A. 2003. The families Huperziaceae and Lycopodiaceae of New England, a taxonomic and ecological
reference. V. F. Thomas Co., Bowdin, Maine. Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Bo-
tanica 92: 153–178. Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the Missouri Botanical Garden 79:
687–717. Øllgaard, B. & K. Tind. 1993. Scandinavian ferns. Rhodos, Copenhagen. Stevenson, D. W. 1976. Observations on
phyllotaxis, stelar morphology, the shoot ápex, and gemmae of Lycopdium lucidulum Michaux. Botanical Journal of the Linnean
Society 72: 81–100. Testo, W., A. Haines & A. V. Gilman. 2016. Huperzia continentalis (Lycopodiaceae), a new species of
gemmiferous firmoss separated from Huperzia haleakalae. Systematic Botany 41: 894–901. Ulrike, R. 1987. Growth patterns
of gemmlings of Lycopodium lucidulum. American Fern Journal 77: 50–57.

342
Figure 168. Huperzia serrata (A–C), H. beiteliana (D–H), and H. selago (J–M). A. Habit. B. Sporophylls and sporangia. C.
Trophophyll, serrate. D. Habit. E. Sporophylls and sporangia. F. Sporophyll. G. Gemma, adaxial view (left), abaxial view (right).
H. Gemmiphore, adaxial view (left), lateral view (right). J. Spore, proximal face, note concave sides. K. Spore, distal face, not
convcave sides. L. Sporeling attached to gametophyte. M. Gemma with young shoot. (A–H from Mickel & Smith, 2004; J–M
from Øllgaard & Tind, 1993.)

343
 Hymenophyllum J.E. Sm.
DESCRIPTION: Epiphytes; rhizomes usually filiform,
long-creeping, pubescent; sterile and fertile leaves typically
1-30 cm long, monomorphic; laminae membranaceous,
usually translucent, 1-cell thick except around the veins,
simple to 3-pinnate, glabrous or pubescent (the hairs
simple, forked, or stellate); veins free, anadromous, false
veins absent; sori marginal, round to ovate; indusium
(often called an “involucre”) consisting of two
lobes; receptacle usually not exerted beyond the
indusia; spores green, tetrahaedral-globose; x=11, 13, 18,
21, 22, 28, 36.
DESCRIPCIÓN: Epífitas; rizomas usualmente filiformos,
largamente rastreros, pubescentes; hojas estériles y
fértiles típicamente 1-30 cm de largo, monomorfas;
láminas membranáceas, usualmente translucentes,
una célula de grosor con excepto alrededor de las venas,
simples a 3-pinnadas, glabras o pubescentes (los pelos
simples, furcados, o estrellados); venas libres, anádromas;
soros marginales, redondos a ovados; indusio (a
menudo llamado “involucre”) consiste de dos
lóbulos; receptáculos usualmente no exertados
fuera del indusio; esporas verdes, tetrahédrico-globosas;
x=11, 13, 18, 21, 22, 28, 36.

SIMILAR GENERA: Trichomanes differs by tubular GÉNEROS PARECIDOS: Trichomanes difiere por
indusia and exerted receptacles. When dried, its laminae indusios tubulosos y receptáculos exertos. Cuando
are green, whereas those of Hymenophyllum are often se secan, sus láminas son verdes, mientras que las de
pale reddish. Hymenophyllum a menudo son rojizas pálidas.

COMMENTS: Hymenophyllum is pantropical, with rare COMENTARIOS: Hymenophyllum es pantropical, con

extension in the temperate zones, and contains about
300 species. All the species grow as epiphytes primarily in
wet, shaded forests. All of its species are epiphytes.
Three well-defined subgenera of Hymenophyllum
occur in the Neotropics: Subgen. Hymenophyllum has
ultimate segments conspicuously serrate and eciliate
(or rarely the serrations tipped by short hairs). Subgen.
Leptocionium has ultimate segments entire and pubescent
(sometimes densely so). Subgen. Mecodium has ultimate
segments entire and glabrous.
The genus name is derived from the Greek hymen,
membrane + phyllon, leaf. The leaves are one-cell thick.
extensiones raras en las zonas templadas, y consta de
casi 300 especies.Todas las especies crecen como epífitas
principalmente en bosques húmedos y sombreados.
Todas de sus especies son epífitas.
Tres subgéneros bien definidos de existen en el
Neotrópico: Subgén. Hymenophyllum tiene segmentos
últimos conspicuamente serrados y eciliados (o raras
veces con las serraciones capadas por un pelito).
Subgén. Leptocionium tiene segmentos últimos enteros
y pubescentes (a veces densamente). Subgén. Mecodium
tiene los últimos segmentos enteros y glabros.
El nombre genérico se deriva del griego hymen,
membrana + phyllon, hoja. Las hojas son una célula de
grosor.

LITERATURE: Copeland, E. B. 1938. Genera Hymenophyllacearum. Philippine Journal of Science 67: 1–110. Diem, J. &
J. Lichtenstein. 1959. Las Hymenofiláceas del área Argentina-Chilena del sud. Darwiniana 11: 611–760. Ebihara, A., J.-Y.
Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of Hymenophyllaceae. Blumea 51: 221–280.
Larsen, C., M. M. Ponce & M. Amalia Scataglini. 2013. Revisión de las especies de Hymenophyllum (Hymenophyllaceae)
del sur de Argentina y Chile. Guayana Botany 70: 274–329. Lellinger, D. B. 1984. Hymenophyllaceae, in Botany of the
Guayana Highland--Part X. Memoirs of the New York Botanical Garden 38: 9–38. Morton, C. V. 1947. The American species
of Hymenophyllum sect. Sphaerocionium. Contributions of the United States National Herbarium 29: 139–201. Morton, C. V.
1968. The genera, subgenera and sections of the Hymenophyllaceae. Contributions of the United States National Herbarium 38:
153–214. Pryer, K., A. R. Smith, J. S. Hunt & J.-Y. Dubuisson. 2001. rbcL data reveal two monophyletic groups of filmy ferns
(Filicopsida: Hymenophyllaceae). American Journal of Botany 88: 1118–1130.

344
Figure 169. A, B. Hymenophyllum asplenioides. C, D. H. polyanthos. E–G. H. myriocarpum. H, J. H. brevistipes. (From
Mickel & Smith, 2004.)

345
TECTARIACEAE
Hypoderris R. Br. ex Hook.
DESCRIPTION: Terrestrial, occasionally on rocks;
rhizomes long-creeping with two ranks of leaves
(distichous); petiole bases diverging widely from the
rhizomes (not ascending), with Eupolypods I vasculature
pattern; laminae lobed to 2-pinnate; veins finely reticulate;
sori round, indusiate or not; spores spiny; x=40.
DESCRIPCIÓN: Terrestres, a veces sobre rocas;
rizomas largamente rastreros, con dos hileras de
hojas (dísticas), bases de los pecíolos divergiéndose
ampliamente desde el rizoma (no ascendentes), con
el patrón vascular de Eupolypods I; láminas lobuladas
a 2-pinnadas; venas densamente reticuladas; soros
redondeados, inusiados o no; esporas espinosas; x=40.

SIMILAR GENERA: Tectaria differs by polystichous
rhizomes, these almost always erect or decument, and
non-spiny spores. Triplophyllum differs by laminae deltate,
basal pinnae enlarged, and non-spiny spores. Bolbitis may
have finely reticulate veins but it differs by rhizomes
with several ranks of leaves, sterile-fertile leaf dimorphy,
laminae glabrous, and acrostichoid sori.
GÉNEROS PARECIDOS: Tectaria difiere por rizomas
polísticas, estos casi siempre erectos o decumbentes,
y esporas no espinosas. Triplophyllum difiere por las
laminas deltadas, pinnas basales agrandadas y esporas
no espinosas. Bolbitis puede tener venas reticuladas pero
difiere por rizomas con varias hileras de hojas, dimorfismo
entre las hojas estériles y fertile, laminas glabras y soros
acrostichoides.

COMMENTS: Hypoderris occurs from Nicaragua hacia
las Antillas, Venezuela y Bolivia. It consists of three species:
H. brauniana (H. Christ) F. G. Wang & Christenhusz, H.
brownii Hook., and H. nicotianifolia (Baker) R. C. Moran,
Labiak & J. Prado. It typically occurs on wet, shaded
forest floors, from 200–1100 m. The genus is sister to
Triplophyllum, which also has creeping rhizomes with
distichous leaves. The indusium of H. brownii is unusual
in that it is attached around the circumference of the
receptacle below the sporangia. In contrast, H. brauniana
has a normal reniform indusium, and H. nicotianifolia is
non-indusiate. The latter two species were previously
classified in Tectaria. The genus name is derived from the
Greek hypo-, below + derris, covering or hide. It refers to
the indusial located beneath the sporangia.
COMENTARIOS: Hypolepis existe desde Nicaragua
hacia las Antillas, Venezuela y Bolivia. Consta de tres
species: H. brauniana (H. Christ) F. G.Wang & Christenhusz,
H. brownii Hook. y H. nicotianifolia (Baker) R. C. Moran,
Labiak & J. Prado.Típicamente existe en el piso de bisques
húmedos sombreados, desde 200–1100 m. The genus is
sister to Triplophyllum, lo ccual tiene rizomas reptantes
con hojas dísticas. El indusio de H. brownii es inusual
porque es únido alrededor del perímetro del receptáculo
abajo de los esporangios. En contraste, H. brauniana tiene
un indusio normal reniforme y H. nicotianifolia es no
indusiado. Los dos species últimas fueron previamente
clasificadas en Tectaria. El nombre genérico se deriva del
griego hypo, abajo + derris, cubierta o piel. Se refiere a los
indusíos ubicado debajo de los esporangios.

LITERATURE: Fa-Guo, W., S. Barratt, W. Falcón, M. F. Fay, S. Lehtonen, H. Tuomisto, F.-W. Xing & M. J. M.
Christenhusz. 2014. On the monophyly of subfamily Tectarioideae (Polypodiaceae) and the phylogenetic placement of some
associated fern genera. Phytotaxa 164: 1–16. Moran, R. C., P. Labiak, J. Garrison-Hanks & J. Prado. 2014. The phylogenetic
relationship of Tectaria brauniana and the recognition of Hypoderris (Tectariaceae). Systematic Botany 39: 384–395.

346
Figure 170. Hypoderris brownii. (From Moran et al, 2014.)

347
DENNSTAEDTIACEAE
Hypolepis Bernh.
DESCRIPTION: Terrestrial; rhizomes long-creeping,
pubescent, the hairs dark, stiff, multicellular; petiole
bases often with horizontal branches (epipetiolar buds),
containing an omega-shaped vascular bundle,; laminae
2-pinnate to 5-pinnate, glabrous or pubescent (not scaly),
sometimes long and supporting itself on the surrounding
vegetation; veins free; sori round, one at the apex of
a vein, marginal or submarginal, indusia formed by the
reflexed margin of the lamina, sometimes absent;
x=29, 52.
SIMILAR GENERA: Dennstaedtia resembles Hypolepis
but is distinguished by having cup-shaped, marginal sori.
Macrothelypteris can be distinguished by its acicular hairs
and medial (not marginal or submarginal) sori. Other
ferns with large, finely divided laminae can usually be
separated by the form of the sori and indusia.
COMMENTS: Hypolepis is pantropical and consists
of about 50 species, with about 30 species in the
Neotropics. It generally grows in open habitats such as
pastures, roadsides, and forest margins. Some species have
indeterminate leaf growth, where the basal portion of
the lamina matures but the apex remains as a fiddlehead,
continuously extending and producing more pairs of
lateral pinnae. The genus name is derived from the Greek
hypo, below + lepis, scale. It refers to the scale-like false
indusia.
LITERATURE: Brownsey, P. J. 1983. Polyploidy and aneuploidy in Hypolepis, and the evolution of the Dennstaedtiales.
American Fern Journal 73: 98–108. Schwartzburd, P. B. & J. Prado. 2015. A taxonomic revision of the South American species
of Hypolepis (Dennsteadtiaceae), Part 1. American Fern Journal 105: 263–313; Part 2, 106: 1–53. 2016.
348
DESCRIPCIÓN: Terrestre; rizomas largamente
rastreros, pubescentes, los pelos oscuros, tiesos,
multicelulares; bases de los pecíolos a menudo con ramas
horizontales (brotes epipeciolares), contienen un haz
vascular en forma de una omega; láminas 2-pinnadas a
5-pinnadas, glabras o pubescentes (no escamosas), a veces
largas y apoyándose sobre la vegetación circundante;
venas libres; soros redondeados, cada uno en el ápice
de una vena, marginales o submarginales,; indusios
formados por el margen reflexo de la lámina, a veces
ausentes; x=29, 52.
GÉNEROS PARECIDOS: Dennstaedtia se parece a
Hypolepis pero se distingue por tener soros marginales
en forma de copa. Macrothelypteris se puede distinguir
por sus pelos aciculares y soros mediales (no marginales
o submarginales). Otros helechos con láminas grandes y
divididas se pueden separar usualmente por la forma de
los soros y indusios.
COMENTARIOS: Hypolepis es pantropical y consta
de casi 50 especies, con alrededor de 30 especies en
el Neotrópico. Generalmente crece en habitats abiertos
tales como pastizales, taludes y bordes de bosques.
Algunas especies tienen crecimiento indeterminado
de la hoja, donde la porción basal de la lámina madura
pero el ápice se queda como un cayado, continuamente
extendiéndose y produciendo más pares de pinas
laterales. El nombre genérico se deriva del griego hypo,
abajo + lepis, escama. Se refiere a los indusíos falsos
como escamas.
Figure 171. A–C. Hypolepis melanochlaena. D–F. H. pulcherrima. G–J. H. diplotricha. K, N. H. eurychlaena. L–M. H.
microchlaena. (From Mickel & Smith, 2004.)

349
ISOETACEAE
Isoëtes L.
Stylites Amstutz

DESCRIPTION: Plants generally less than 30 cm tall,
tufted, evergreen aquatics or ephemeral terrestrials.
Stems usually globose, fleshy, 2- or 3-lobed. Leaves linear,
subulate, terete or nearly so, one-veined, often winged
toward the base, containing 4 air chambers, the air
chambers septate; sporangia sunken in the leaf base,
single, usually ovoid, covered to various degrees by
a hyaline membrane (the velum); spores of two kinds:
megaspores tetrahedral-globose, trilete, often drying dull
white; microspores ellipsoid, monolete, often drying tan,
brown or dark; x=11.
DESCRIPCIÓN: Plantas generalmente menos de 30
cm de largo, amacolladas, siempreverdes, acuáticas o
terrestres efímeras. Tallos usualmente globosos, carnosos,
2- o 3-lobulados. Hojas lineares, subulatas, teretes o casi
así, con una vena, con 4 cámeras para aire; esporangios
hundidos en la base de la hoja, simples, usualmente
ovoides, cubiertos hasta varios grados por un membrano
hialino (el velum); esporeas de dos tipos: los megasporas
tetrahédricas-globosas, triletes, blanqueciñas al secarse, y
microesporas elipsoides, monoletes, leonadas, pardas o
oscuras al secarse; x=11.

SIMILAR GENERA: Pilularia differs by leaves with GÉNEROS PARECIDOS: Pilularia difiere por hojas con
circinnate vernation and without air chambers. vernación circinada y sin cámeras del aire.

COMMENTS: Isoëtes has about 200 species worldwide.
It is typically a submerged or emergent aquatic, although
some species regularly grow in seasonally dry habitats.
In the Neotropics, most species grow in the páramos.
Stylites, which grows in the high Andes of Peru and
Bolivia, is sometimes distinguished from Isoëtes. Its stems
are elongate, erect, and often dichotomously branched,
with roots arising from a single lateral furrow. Its leaves
are flat, rigid, and bear sporangia nearly superficially
a few centimeters above the base. It is necessary to
examine the megaspores with at least 30X to identify
the species. In the northern temperate zones there is
much hybridization, but this phenomenon has not been
well studied in the tropics. Many species in the genus
have CAM photosynthesis, an adaptation to oligotrophic
conditions. The genus name is derived from the Greek
isos, equal + etes, year.The green leaves persit throughout
the year in many species.
COMENTARIOS: Isoëtes tiene 200 especies mundial.
Es típicamente submergido o emergente, aunque
unas especies existen regularmente en habitats
estacionalmente secos. En el Neotrópico, la mayoría de
las especies existen en los páramos. Stylites, lo cual existe
en los Andes altos de Peru y Bolivia, a veces se distingue
de Isoëtes. Sus tallos son alargados, erectos, y a menudo
furcados dicotómicamente, con raíces originándose de
un sólo surco lateral. Sus hojas son planas, rígidas y llevan
sus esporangios casi superficialmente unos centímetros
arriba de la base. Hay que examinar los megasporas con
a lo menos 30X para identificar las especies. En las zonas
norteñas templadas hay mucha hibridación, pero este
fenómeno no ha estado bien estudiado en los trópicos.
Muchas especies del género tienen fotosíntesis CAM,
una adaptación a condiciones oligotróficas. El nombre
genérico se deriva del griego isos, igual + etes, año. Las
hojas verdes persisten por el año en muchas especies.

LITERATURE: Boom, B. 1982. Synopsis of Isoetes in the southeastern United States. Castanea 47: 38–59. Fuchs-Eckert, H. P.
1982. Zur heutigen Kenntnis von Vorkommen und Verbreitung der sudamerikanischen Isoetes-arten. Proc. Ned. Akad. Wetensch.
C85: 205–260. Hickey, R. J., C. Macluf & W. C. Taylor. 2003. A re-evaluation of Isoetes savatieri Franchet in Argentina and
Chile. American Fern Journal 93: 126–136. Pfeiffer, N. E. 1922. Monograph of the Isoëtaceae. Annals Missouri Botanical Garden
9: 79–232. Small, R. L. & R. J. Hickey. 2001. Systematics of the northern Andean Isoëtes karstenii complex. American Fern
Journal 91: 41–69. Weber, U. 1922. Zur Anatomie und Systematik der Gattung Isoetes L. Nova Hedwigia 63: 219–262.

350
Figure 172. Isoëtes. A. Habit. B. Leaf base with sporangium, the triangular structure is the ligule. C, D. Megaspores. (From
Mickel & Smith, 2004.)

351
PTERIDACEAE
Jamesonia Hook. & Grev.
DESCRIPTION: Terrestrial; rhizomes 2–4 mm wide,
short- to long-creeping, pubescent, the hairs thick,
bristle-like; leaves extremely variable in size and
cutting, from 1-pinnate and linear, erect, to 4-pinnate
and scandent, , pubescent, apices tipically indeterminate;
pinnae sessile or nearly so and imbricate and coriaceous,
to stalked spreading and herbaceous, the margins flat
or enrolled; veins free; sori elongate along the veins,
non-indusiate, protected by the reflexed margins of the
pinnae or not; spores tetrahaedral-globose, nongreen;
x=87.
DESCRIPCIÓN: Terrestres; rizomas 2–4 mm de ancho,
corto- a largamente rastreros, pubescentes, los tricomas
guesos como cerdas; hojas extremamente variables,
desde 1-pinnatas, lineares y erectas hasta 4-pinnatas,
anchamente deltadas y escandendes, pubescentes, el
ápice circinado, usualmente indeterminado; pinnas los
márgenes típicamente enrrollados; venas libres; soros
a lo largo de las venas, no-indusiados, protegidos
por los margenes reflexos de las pinnas o no; esporas
tetraédrico-globosas, no verdes; x=87.

SIMILAR GENERA: Pterozonium differs pinnae often
flabellate and many close, elongate sori. It is rare outside
the Tepui region of the Guayanas. Tryonia differs by
stramineous rachises. It is restriced to southeastern Brazil.
GÉNEROS PARECIDOS: Pterozonium difiere por
pinnas a menudo flabeladas y muchos soro aproximados
alargados. Es raro a fuera de la region de los tepuis en
la Guyanas. Tryonia difiere por ráquises estramineos. Es
restrigida a sureste de Brasil.

COMMENTS: Jamesonia is completely neotropical COMENTARIOS: Jamesonia es completamente

(except for one population on Tristan da Cunha) and
contains ca. 50 species. It is found in high-elevation
vegetation (> 2500 m), especially páramos. It includes
Eriosorus, formerly maintained as distinct. Species with
stiffly erect 1-pinnate leaves, como J. alstonii, represent
a morphotype restricted to páramos. This morphotype
evolved at least four times from ancestors with spreading,
more divided leaves. Some species have highly divided
scandent leaves, such as J. flexuosa. The species hybridize
frequently when growing together, and the hybrids have
aborted spores. In the New World, Pityrogramma is most
closely related to Pterozonium and Tryonia. The generic
name honors William Jameson (1796–1873), Scottish
botanist who collected plants in Quito, Ecuador.
neotropical (con excepción de una población en Tristan
da Cunha) y consta de 50 especies. Se encuentra en
vegetación de elevaciones altas (> 2500 m), especialmente
los páramos. Incluye Eriosorus anteriormente mantenido
como distincto. Las especies con hojas tiesamente erectas
1-pinnadas, como J. alstonii, representan un morfotipo
restingido a los páramos. Este morfotipo ha tendio
al menos cuatro orígenes independientes a partir de
ancestros con hojas divergentes y más divididas. Algunas
especies tienen hojas muy divididas y escandentes, como J.
flexuosa. Las especies se hibridizan con frequencia cuando
crecen juntos, y los híbridos tienen esporas abortivas. En
el Nuevo Mundo, Pityrogramma es más cercanamente
relacionado a Pterozonium y Tryonia. El nombre genérico
honra William Jameson (1796–1873), botánico escocés
quien colectaba plantas en Quito, Ecuador.

LITERATURE: Gómez, L. D. 1979. Contribuciones a la pteridología costarricense XIII. Novitates. Brenesia 16: 95–100.
Sánchez-Baracaldo, P. 2004. Phylogenetics and biogeography of the neotropical fern genera Jamesonia and Eriosorus
(Pteridaceae). American Journal of Botany 91: 274–284. Pabón-Mora, N. & F. González. 2016. Nephopteris out of the
clouds: molecular evidence places the enigmatic N. maxonii (Pteridaceae) within the Jamesonia clade. Brittonia 68: 83–92.
Sánchez-Baracaldo, P. & G. H. Thomas. 2004. Adaptation and convergent evolution within the Jamesonia-Eriosorus
complex in high-elevation biodiverse Andean hotspots. PLoS One 9(10): e110618. Scamman, E. 1962. The genus Eriosorus
in Costa Rica. Contributions from the Gray Herbarium of Harvard University 191: 81-89. Tryon, A. F. 1962. A monograph
of the fern genus Jamesonia. Contributions from the Gray Herbarium of Harvard University. 191: 109–197. Tryon, A. F.
1970. A monograph of the fern genus Eriosorus. Contributions from the Gray Herbarium of Harvard University 200: 54–174.

352
Figure 173. A, B. Jamesonia hirsuta. C, D. J. flexuosa. E, F. J. flexuosa var. galeana. G. J. maxonii. H. J. alstonii. (A-F, G. from
Mickel & Smith, 2004; G, original, © Robbin Moran, 2017.)

353
DRYOPTERIDACEAE
Lastreopsis Ching
DESCRIPTION: Terrestrial; rhizomes short- to long-
creeping, scaly; leaves monomorphic; petiole with four
or more vascular bundles; laminae 2–4-pinnate-pinnatifid,
lanceolate, deltate, or pentagonal, the lower pinnae
enlarged basiscopically; scaly buds absent on the
distal parts of the laminae; pinnules anadromic or
catadromic; rachis with 2 prominent ridges adaxially,
these continuous with the thickened margins of
the ultimate segments (Fig. D,E); costae raised (not
grooved), hairy adaxially with 0.1-0.3 mm long, multicellular
hairs, costae, and costules scaly abaxially, hairy adaxially;
glandular hairs often present on the lower surface of the
lamina, these cylindrical, oblong appressed, bright yellow
to orange-red; sori round. x=41.
DESCRIPCIÓN: Terrestres; rizomas breves a largamente
rastreros, escamosos; hojas monomorfas; pecíolos con 4 o
más haces vasculares; láminas 2–4-pinnadas, lanceoladas,
deltadas o pentagonales, las pinnas más inferiores
agrandadas basiscópicamente; yemas escamosas
ausentes en las porciones distales de las láminas;
pínnulas dispuestas anadrómica o catadrómicamente;
raquis con 2 prominentes crestas adaxiales, éstas
continuas con los engrosados márgenes foliares de
los segmentos terminales (Fig. D,E); costas elevadas
(no surcadas), pelosas adaxialmente con pelos 0.1-0.3
mm, multicelulares; tricomas glandulosos a menudo
presentes en el haz de la lámina, cilíndricos, oblongos,
adpresos, amarillo brillante a anaranjado-rojizo; soros
redondos. x=41.

SIMILAR GENERA: Parapolystichum is indistinguishable GÉNEROS PARECIDOS: Parapolystichum es

morphologically. Dryopteris and Arachniodes differ by axes
adaxially glabrous, not finely pubescent. Megalastrum
differs by coarse, whitish, strigose hairs on the axes
adaxially, prominent hydathodes, and the basal veinlet
of the distal pinnules springing from the costa, not the
costule, with the lobe supplied by this veinlet adnate to
the costa.
indistinguible morfológicamente. Dryopteris y Arachniodes
se difieren por sus ejes adaxialmente glabros, no finamente
pubescentes. Megalastrum difiere por pelos toscos,
blanqueciños, strigosos sobre los ejes adaxialmente,
hidatodos prominentes, y la venilla basal de las pinnulas
distales surgiendo de la costa, no de la cóstula, con el
lóbulo suministrado por esta venilla adnato a la costa.

COMMENTS: Lastreopsis contains about 15 species. COMENTARIOS: Lastreopsis consta de casi 15 y existe
and occurs in southeastern Asia, Australia, New Zealand, en sureste de Asia, Australia, Nueva Zelanda y ciertas
and certain islands of the southwestern Pacific. About 27 islas del suroeste Pacífico. Casi 27 especies previamente
species previously placed in Lastreopsis are now classified ubicadas en Lastreopsis ahora están clasificadas en
in Parapolystichum. The raquis-costa architecture of Parapolystichum. La arquitectura raquis-costa de
Lastreopsis is unique and unmistakable once learned. It Lastreopsis es único y no se puede confundir después
is characterized by the shape of the axes on the upper de haber comprendido. Se caracteriza por la forma de
surface of the lamina (Fig. D,E). The rachis has two ridges los ejes en el haz de la lámina (Fig. D,E). El raquis tiene
and is densely puberulent between them. The ridges are dos crestas y es densamente puberulento entre ellas. Las
continuous with the decurrent margins of the pinnules. crestas son continuas con los márgenes decurrentes de
The central part of the axes is raised (not grooved) as las pínnulas. La parte central de los ejes es prominulo (no
in most of the Dryopteridaceae. The two species of sulcados) como en la mayoría de las Dryopteridaceae. Las
Lastreopsis that occur in the Neotropics (L. amplissima dos especies de Lastreopsis que existen en el Neotrópico
(C. Presl) Tind. and L. killipii (C. Chr. & Maxon) Tind.) (L. amplissima (C. Presl) Tind. and L. killipii (C. Chr. &
belong to subg. Rhopalotricha Labiak & R. C. Moran. In Maxon) Tind.) pertenecen al subgén. Rhopalotricha Labiak
contrast, subg. Lastreopsis is entirely Old World, occurring & R. C. Moran. En contraste, el subgén. Lastreopsis ocurre
in southeastern Asia, Malesia, Australia, and New Zealand. solamente en el Viejo Mundo, en sureste de Asia, Malesia,
The genus name is derived from Lastrea, a fern genus + Australia y Nueva Zelandia. El nombre genérico se deriva
opsis, like. de Lastrea, un género de un helecho + opsis, parecido.
LITERATURE: Labiak, P. H., M. Sundue, G. Rouhan, J. Garrison Hanks, J. T. Mickel & R. C. Moran. 2014. Phylogeny
and historical biogeography of the lastreopsid ferns (Dryopteridaceae). American Journal of Botany 101: 1207–1228. Labiak, P.
H., M. Sundue, G. Rouhan & R. C. Moran. 2014. New combinations in Lastreopsis and Parapolystichum (Dryopteridaceae).
Brittonia 67: 79–86. Labiak, P. H., M. Sundue, G. Rouhan & R. C. Moran. 2015. Rhopalotricha, a new subgenus of the fern
genus Lastreopsis (Dryopteridaceae). American Fern Journal 105: 20–30. Tindale, M. D. 1965. A monograph of the genus
Lastreopsis Ching. Contributions to the New South Wales National Herbarium 3: 249–339, t. 1–23.

354
Figure 174. A-E. Lastreopsis amplissima. A. Basal pinna, showing a detail of the abaxial surface and the sori. B. Adaxial side of
a pinnule, showing the hairs on the axes. C. Scale from the rachis. D. Clavate hair from the adaxial side of costa. E. Glandular
hair from the abaxial side of the lamina. F-I. L. davallioides. F. Basal pinna, showing a detail of the abaxial surface and the sori.
G. Adaxial side of a pinnule, showing the hairs on the axes. H. Club-shaped hair from the adaxial side. I. Scale from the rachis.
J-N. L. hispida. J. Basal pinna, showing a detail of the abaxial surface and the sori. K. Adaxial side of a pinnule, showing the hairs
on the axes. L. Glandular hair from the abaxial side of the lamina. M. Club-shaped hair from the adaxial side. N. Scale from the
rachis. O–S. L. killipii. O. Basal pinna, showing a detail of the abaxial surface and the sori. P. Adaxial side of a pinnule, showing the
hairs on the axes. Q. Glandular hair from the abaxial side of the lamina. R. Club-shaped hair from the adaxial side. S. Scale from
the rachis. (From Labiak et al., 2013.)

355
POLYPODIACEAE
Lellingeria A. R. Smith, R. C. Moran, & L. E. Bishop
DESCRIPTION: Plants epiphytic or epipetic; rhizome
short- to long-creeping, ascending, or erect, the scales
clathrate, usually blackish, glabrous or provided with
hyaline marginal setulae, attached across the entire width
of the base; phyllopodia absent; petioles absent or shorter
than the lamina, continuous with (not articulate to) the
rhizome; laminae pinnatifid to 1-pinnate-pinnatifid, lacking
reddish setae, or pubescent (at least along the petiole
and rachis), the hairs usually with a pale basal cell and
forked unequally, the smaller cell oblique, glandular,
the larger cell acicular; hydathodes present; veins
simple, free, several per segment; sori round or elliptic,
often somewhat sunken, lacking paraphyses; sporangial
capsules glabrous or (in 3 species) setose; x=37.
DESCRIPCIÓN: Plantas epífitas o epipéticas; rizomas
cortamente reptantes, ascendentes o rectos, las escamas
clatradas, usualmente negruzcas, glabras o provistas de
sétulas marginales hialinas, unidas por el ancho de la base;
filopodios ausentes; pecíolos ausentes o más cortos que
la lámina, contínuos (no articulados) al rizoma; láminas
de pinnatífida a 1-pinnada-pinnatífida, setas rojizas
ausentes, o pubescente (al menos a lo largo del pecíolo
y raquis), los tricomas usualmente con una célula
basal pálida y furcada desigualmente, la célula
más pequeña oblicua, la célula más larga acicular;
hidatodos presentes; venas simples, libres, varias por
segmento; soros redondos o elípticos, a menudo levemente
hundidos, paráfisis ausentes; cápsulas de los esporangios
glabras o (en 3 especies) setosas; x=37.

SIMILAR GENERA: Cochlidium differs by tan,
nonclathrate scales and lack of forked hairs. Grammitis
differs by entire laminae with black borders. Stenogrammitis
differs by narrow leaves (< 5 mm) and only one sorus and
vein per segment. Other neotropical grammitids genera
differ by the presence of dark reddish setae on the leaves.
GÉNEROS PARECIDOS: Cochlidium difiere por
escamas leonadas, no clatradas y la ausencia de tricomas
furcados. Grammitis difiere por laminas enteras con
bordes negras. Stenogrammitis difiere por hojas estrechas
(< 5 mm) y solo un soro por segmento. Otros géneros
gramitidoides neotropicales difieren por la presencia de
setas oscuras rojizas en las hojas.

COMMENTS: Lellingeria contains about 50 species and
is entirely neotropical. Most of the species are epiphytes
in cloud forests, from 1000–4000 m. The genus is
most diverse in the Andes. Lellingeria is one of the few
grammitids that lack reddish setae on the leaves. Its hairs
are distinctive, being often forked unequally—a character
it shares with Stenogrammitis (see illustration of that
genus). The genus is named for David B. Lellinger (1937–
), American pteridologist who worked at the Smithsonian
Institution.
COMENTARIOS: Lellingeria consta de casi 50 especies
y es completamente neotropical. La mayoría de las
especies son epifitas en bosques nublados, desde 1000–
4000 m. El género es más diverso en los Andes. Lellingeria
es uno de los pocos helechos grammitidoides que faltan
setas rojizas en las hojas. Sus pelos son distintivos, siendo
a menudo furcados desigualmente—una característica
que comparta con Stenogrammitis (vea la ilustración de
ese género). El género se nombra para David B. Lellinger
(1937– ), pteridólogo Americano quien trabajaba en el
Instituto Smithsonio.

LITERATURE: Copeland, E. B. 1952.The American species of Xiphopteris. American Fern Journal 42: 41–52, 93–110. Labiak,
P. H. 2013. Grammitid ferns (Polypodiaceae), I. Lellingeria. Flora Neotropica Monograph 111: 1–130. Labiak, P. H. & J. Prado.
2005. As espécies de Lellingeria A. R. Sm. & R. C. Moran (Grammitidaceae-Pteridophyta) do Brasil. Revista Brasileira de Botânica
28: 1–22. Labiak, P. H. & M. A. Sundue. 2010. Molecular phylogeny, character evolution, and biogeography in the grammitid fern
genus Lellingeria (Polypodiaceae). American Journal of Botany 97: 1354–1364. Maxon, W. R. 1914. Studies of tropical American
ferns–No. 5. Notes upon Polyodium duale and its allies. Contributions from the U.S. National Herbarium 17: 398–406. Ranker, T.
A., A. R. Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Staub & H. Schneider. 2004. Phylogeny and evolution
and grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy. Taxon 53: 415–428. Smith, A. R., R. C.
Moran, & L. E. Bishop. 1991. Lellingeria, a new genus of Grammitidaceae. American Fern Journal 81: 76–88.

356
Figure 175. Lellingeria. A–E. L. apiculata (Alston 6899, MO). A. Habit. B. Detail of the segments. C. Detail of the abaxial side of the rachis.
D. Types of hairs in the rachis abaxially. E. Rhizome scale. F–J. L. flagellipinnata (Luteyn 13482, MO). F. Habit. G. Detail of the segments.
H. Detail of the rachis, showing one sorus. I. Rhizome scale. J. Detail of the rhizome scale. K–O. L. ciliolepis (Labiak 2904, UPCB). K. Habit.
L. Detail of the segments. M. Detail of the abaxial side of the rachis. N. Rhizome scale. O. Detail of the rhizome scale. P–R. L. suspensa
(Proctor 21715, US). P. Habit. Q. Detail of the segments. R. Rhizome scale. (From Labiak, 2013.)

357
POLYPODIACEAE
Leucotrichum Labiak
DESCRIPTION: Plants epiphytic, rarely epipetric; roots
not proliferous; rhizomes erect, radial, scaly, the scales
clathrate, dark brown to blackish, basifixed, margins ciliate;
petioles absent to ca. 1 cm long, pubescent, the hairs up
to 2 mm long, whitish, simple or 1–furcate; laminae erect
to slightly arcuate, determinate, linear to linear-oblong,
pinnatifid to pinnatisect, with reduced basal segments, the
lowermost decurrent and forming short and broad wings,
apices obtuse, subconforme to the lateral segments;
rachis with dark sclerenchyma, the sclerenchyma not
visible on the abaxial side, often obscured by the laminar
tissue adaxially, setose and pubescent, the setae whitish,
up to 3 mm long, the hairs simple to 1-furcate; segments
1–3 × 0.5–1.5 mm, entire, symmetric or asymmetric at
base, setose and pubescent, the setae whitish, ca. 1–3
mm long, the hairs 0.2–1.0 mm long, 1(2)-furcate; veins
1-furcate, obscured by the laminar tissue; hydathodes
present, lacking whitish deposits; sori one per segment,
inframedial, rounded to elongate, superficial, surrounded
by the hairs of the laminae; sporangial capsules glabrous or
ciliate; spores trilete, green. x=?.
DESCRIPCIÓN: Plantas epífitas, raras veces epipetricas;
raíces no prolíferas; rizomas erectos, radiales, escamosos,
las escamas clatradas, pardo oscuros a negruzcos;
basifijadas, márgenes ciliados; pecíolos ausentes hasta
1 cm de largo, pubescentes, los pelos hasta 2 mm de
longitud, blanquecinos, simples o 1-furcados; láminas
erectas a levemente arcuadas, determinadas, lineares
a linear-oblongas, pinnatífidas a pinnatisectas, con
segmentos basales reducidos, los más basales decurrentes
y formando alas cortas y anchas, ápices obtusos,
subconformes a los segmentos laterales; ráquises con
esclerenquima oscura no visible abaxialmente, a menudo
oculta por el tejido laminar adaxialmente, setosos y
pubescentes, las setas blanquecinas, hasta 3 mm de
longitud, los pelos simples o 1-furcados; segmentos 1–3
× 0.5–1.5 mm, enteros, simétricos o asimétricos en la
base, setosos y pubescentes, las setas blanquecinas, ca.
1–3 mm de longitud, los pelos 0.2–1.0 mm de longitud,
1(2)-furcados; venas 1-furcadas, ocultas por el tejido
laminar; hydatodos presentes, faltando de depósitos
blanquecinos; soros uno por segmento, inframediales,
redondos o elípticos, superficiales, rodeados por los pelos
de la lamina; cápsulas de los esporangios glabras o ciliados;
esporas triletes, verdes; x=?.

SIMILAR GENERA: Alansmia differs by non-clathrate
rhizome scales, indeterminate leaf growth, pinnate veins,
and branched setae. Lellingeria differs by the absence of
setae and whitish hairs, and the presence of pinnate veins.
Stenogrammitis differs by eciliate rhizome scales, absence
of whitish hairs and setae on the leaves, and one vein per
segment.
GÉNEROS PARECIDOS: Alansmia difiere por las
escamas de los rizomas no clatradas, crecimiento
indeterminado de las hojas, venas pinnadas y setas
ramificadas. Lellingeria difiere por la ausencia de setas
y pelos blanquecinos, y la presencia de venas pinnadas.
Stenogrammitis difiere por escamas del rizoma no ciliadas,
ausencia de pelos y setas blanquecinos en las hojas y una
vena por segmento.

COMMENTS: Leucotrichum contains 6 species, all of COMENTARIOS: Leucotrichum contiene 6 especies,

which are epiphytes in wet forests.The genus occurs in the todos de las cuales son epifitas en bosques húmedos. El
West Indies, Central America, SE Brazil, and Madagascar. género existe en las Antillas, América Central, SE de Brasil
This distribution is unusual among neotropical fern y Madagascar. Esta distribución es inusual entre los géneros
genera by being entirely absent from the Andes and the neotropicales de los helechos por ser completamente
Guayanas. The genus name is derived from the Greek ausente en los Andes y las Guayanas. El nombre genérico
leuco-, white, and thrix, hair. It refers to the distinctive se deriva del griego leuco-, blanco, y thrix, pelo. Refiere a
whitish hairs and setae on the leaves. los distintivos pelos y setas blanquecinos de las hojas.
LITERATURE: Copeland, E. B. 1952.The American species of Xiphopteris. American Fern Journal 42: 41–52, 93–110. Labiak,
P. H. & J. Prado. 2005. As espécies de Lellingeria A. R. Sm. & R. C. Moran (Grammitidaceae-Pteridophyta) do Brasil. Rev.
Brasil. Bot. 28: 1–22. Labiak, P., G. Rouhan & M. Sundue. 2010. Phylogeny and taxonomy of Leucotrichum (Polypodiaceae):
a new genus of grammitid ferns from the Neotropics. Taxon 59: 911–921. Rouhan, G., P. H. Labiak, E. Randrianjohany
& F. Rakotondrainibe. 2012. Not so neotropical after all: the grammitid fern genus Leucotrichum (Polypodiaceae is also
Paleotropical, as revealed by a new species from Madagascar. Systematic Botany 37: 331–338. Smith, A. R., R. C. Moran, & L.
E. Bishop. 1991. Lellingeria, a new genus of Grammitidaceae. American Fern Journal 81: 76–88.

358
Figure 176. Leucotrichum. A–D. L. mortonii (Liogier 12876, NY). A. Habit. B. Rhizome scale. C. Sporangium. D. Hairs from the abaxial
surface of the lamina. E–H. L. organense (Labiak 4499, UPCB). E. Habit. F. Rhizome scale. G. Branched hair from the costae. H. Sporangium.
I–L. L. schenckii (Labiak 2938, UPCB). I. Habit. J. Rhizome scale. K. Hairs from the abaxial side of the laminar tissue. L. Sporangium. M. L.
pseudomitchellae (Mickel 3033, NY). Habit. N–P. L. mitchellae (Ekman 14884, K). N. Habit. O. Rhizome scale. P. Sporangium. (From
Paulo Labiak, 2010.)

359
LINDSAEACEAE
Lindsaea Dryander ex J.E. Sm.
DESCRIPTION: Terrestrial, rarely rupestral or epiphytic;
rhizome short- to long-creeping, protoselic, scaly, the scales
lanceolate to narrow and bristle-like; laminae anadromic,
1-pinnate to 2(-4)-pinnate, when 2-pinnate the apex
(in the neotropics) a conform terminal pinna; ultimate
segments usually dimidiate, with the costule along
the basiscopic margin, the margins entire; rachises and
costae grooved or sharply angled adaxially, usually
greenish or stramineous; veins free or anastomosing; sori
following the margin of the segments, the indusia
opening toward the margin; annulus of 7-15 cells;
spores monolete or trilete, 16 or 32 per sporangium; x =
34, 41, 47, 50.
DESCRIPCIÓN: Terrestres, ráramente rupestres o
epífitas; rizoma corto o largamente rastrero, protostélico,
escamoso, las escamas lanceoladas a angostas y como
una cerda; lámina anádroma, 1-pinnada a 2(-4)-pinnada,
cuando 2-pinnada, el ápice (en el neotrópico) consiste
de una pinna terminal conforme; últimos segmentos
usualmente dimidiados, con la costilla media a lo largo
del margen basiscópico, los margenes enteros; ráquises
y costas surcados o angulados adaxialmente,
usualmente verdosos o pajizos; nervaduras libres o
anastomosadas; soros siguiendo el margen de los
segmentos, los indusios abriéndose hacia el margen;
anillo de 7-15 células; esporas monoletes o triletes, 16 o
32 por esporangio; x = 34, 41, 47, 50.

SIMILAR GENERA: Adiantum differs by its terete, GÉNEROS PARECIDOS: Adiantum difiere por
castaneous to black petioles and rachises, indusia formed pecíolos y ráquises teretes y castaños a negros, indusios
by the enrolled margin of the segment (false indusium) formados por el margen reflejo del segmento (indusio
opening toward the costule. falso) abriéndose hacia la cóstula.

COMMENTS: Lindsaea contains about 150 species and
is pantropical. About 50 species occur in the Neotropics,
mostly in the Guayanas. Lindsaea has an unusual type of
protostele because the phloem is surrounded by xylem. It
is closely related to Odontosoria. The genus name honors
John Lindsay, a British physician who lived in Jamaica during
the late 1700s and made observations on the germination
of fern spores and formation of gametophytes.
COMENTARIOS: Lindsaea consta de casi 50 especies
y es pantropical. Alrededor de 50 especies existen en
el Neotrópico, mayormente en las Guayanas. Lindsaea
tiene un protostelo raro porque el floema está rodeado
de xilema. Es cercanamente relacionado a Odontosoria.
El nombre genérico honora a John Lindsay, un médico
británico quien vivó en Jamaica en las postrimerías de los
1700s y hizo estudios sobre la germinación de esporas y
la formación de gametofitas.

LITERATURE: Kramer, K. U. 1957. A revision of the genus Lindsaea in the New World. Acta Botanica Neerlandica 6: 97–290.
Kramer, K. U. 1991. Dennstaedtiaceae. Pages 20–81. In: A. R. A. Görts-van Rijn, editor. Flora of the Guianas, fascicle 4. Koeltz
Scientific Books. Lehtonen, S., H. Tuomisto, G. Rouhan & M. J. M. Christenhusz. 2010. Phylogenetics and classification of
the pantropical fern family Lindsaeaceae. Botanical Journal of the Linnean Society 163: 305–359. Rojas-Alvarado, A. F. & J. D.
Tejero-Diez. 2017. Novelties and notes in Lindsaea (Lindsaeaceae) from Mexico and Central America. Phytotaxa 296: 147–160.

360
Figure 177. Top: L. portoricensis. G–K: L. quadrangularis subsp. antillensis. L–O: L. lancea. P–R: L. arcuata. S–U: L. stricta.
(From Mickel & Smith, 2004.)

361
PTERIDACEAE
Llavea Lag.
DESCRIPTION: Terrestrial or rupestral; rhizomes short-
creeping or erect, the apex with lanceolate, stramineous
to tan scales, sometimes with a black central portion
(especially on petiolar scales); leaves hemidimorphic,
with the apical pinnae fertile and contracted; laminae
2–3-pinnate; pinnules oblong to elliptic; veins free; indusia
formed by the revolute margin of the pinnule, with short
glandular hairs on the inner surface; sporangia along the
veins but confluent at maturity; spores trilete, tetrahedral;
x=29.
DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto
a cortamente rastrero, el ápice con escamas lanceoladas,
pajizas, leonadas, a veces con una porción central negra
(especialmente en las escamas del pecíolo); hojas
hemidimorfas, con las pinnas apicales fértiles y
contraídas; pecíolos teretes a subcuadrangulars; láminas
2–3-pinnadas; pínnulas oblongas a elípticas; nervaduras
libres; indusios formado por el margen del segmento
reflexo, con pelitos glandulares en la superficie interior;
esporangios a lo largo de las nervaduras, pero confluentes
en la madurez; esporas triletes, tetraédricas; x=29.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Llavea occurs in Texas, Mexico and
Guatemala where it grows on rocky wooded slopes from
1200–2500 m. It contains only one species: L. cordifolia
Lag., distinctive by its hemidimorphic leaves. Within the
Pteridaceae, Llavea forms a clade with Conigramme (Asian)
and Cryptogramma. The genus name honors Pablo de la
Llave (1773–1833), a traveler in Mexico.
COMENTARIOS: Llavea occurs en Texas, México y
Guatemala donde crece en taludes rocosos bosqueados
desde 1200–2500 m. Contiene una sola especie: L. cordifolia
Lag., distinctiva por sus hojas hemidimorfas.Dentro de
las Pteridaceae, Llavea forma un clado con Coniogamme
(Asiático) Cryptogramma. El nombre genérico honora a
Pablo de la Llave (1773–1833), un viajero en México.

LITERATURE: Gastony, G. J. & D. R. Rollo. 1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern
United States and adjacent Mexico—a molecular phylogenetic reassessment of generic lines. Aliso 17: 131–141. Vaganov, A.
V., A. A. Kunznetsov & A. I. Shamakov. 2011. Taxonomy and morphology of Llavea cordifolia Lag. (Cryptogrammaceae).
Turczanianowia 14: 19–22. [Russian]

362
Figure 178. A–D. Llavea cordifolia Lag. (From Mickel & Smith, 2004.)

363
BLECHNACEAE
Lomaria Willd.
DESCRIPTION: Plants terrestrial; rhizomes erect,
sometimes forming small trunks, stoloniferous or not, the
scales concolorous or bicolorous with a blackish central
stripe, entire; leaves dimorphic; laminae pinnatisect or
1-pinnate, lanceolate or oblong-elliptic, gradually reduced
proximally to auriculate pinnae, apices pinnatisect, dark
green adaxially, pallid green abaxially; rachises
deeply grooved adaxially, glabrous or sparsely scaly
abaxially; buds absent; aerophores absent; pinnae adnate
to subpetiolulate; veins free, tips ending at pinna margins,
sometimes hydathodous, or slightly enlarged; sori linear,
indusia continuous, lacerate or erose at maturity; x=28.
DESCRIPCIÓN: Plantas terrestres; rizomas erectos, a
veces formando troncos pequeños, estoloníferos o no,
las escamas concoloras o bicoloras con una raya central
negruzco, enteras; hojas dimorfas; láminas pinnatisectas a
1-pinnadas, lanceoladas o oblongo-elípticas, gradualmente
reducidas proximalmente hasta pinnas auriculadas, ápices
pinnatisectos, verde oscuras adaxialmente, pállidas
abaxialmente; ráquises profundamente surcados
adaxialmente, glabros o esparcidamente escamosos
abaxialmente; brotes ausentes; aeróforos ausentes; pinnas
adnatas a subpetioluladas; venas libres, a veces con
hidatodos; soros lineales; indusios continuos, lacerados o
erosos en la madurez; x=28

SIMILAR GENERA: Parablechnum differs by rachises
ungrooved or only shallowly grooved adaxially and
proximal pinnae sessile or very short-petiolate.
Struthiopteris differs by creeping rhizomes and shallowly
grooved rachises.
GÉNEROS PARECIDOS: Parablechnum difiere por
los ráquises no o solamente surcados levemente y las
pinnas proximales sésiles o muy cortamente pecioluladas.
Struthiopteris difiere por rizomas reptantes y ráquises
surcados levemente.

COMMENTS: Lomaria comprises about six species, all
austral, with three or four in Australasia (New Zealand,
Australia, and New Caledonia), two in southern South
America, and one in southern Africa. Lomaria comprises
six species, and occurs in South America, South Africa,
Australia, and New Caledonia. The two South American
species are L. brunea (M. Kessler & A. R. Sm.) Gasper & V.
A. O. Dittrich (from Bolivia) and L. spannagelii (Rosenst.)
Gasper & V. A. O. Dittrich. (from southeastern Brazil).
They grow in wet shaded forests from 1900–2500 m.
Smith & Kessler (2018) discussed how to distinguish the
two species. Within the Blechnaceae, Lomaria is sister
to a clade of 10 genera that includes Austroblechnum,
Blechnum, and Parablechnum. Most of these genera have
laminae with gradually reduced pinnae towrd the base.
The genus name is derived from the Greek loma, borde,
and aris, sufix. It refers to the marginal sorus that forms
a continuous line that borders the costa of the narrow
pinnae..
COMENTARIOS: Lomaria consta de casi seis especies,
todas australes, con tres o cuatro en Australasia (Nueva
Zelandia, Australia y Nueva Caledonia), dos en el sur
de América del Sur y una en el sur de África. Las dos
especies sudamericanas son L. brunea (M. Kessler &
A. R. Sm.) Gasper & V. A. O. Dittrich (de Bolivia) y L.
spannagelii (Rosenst.) Gasper & V. A. O. Dittrich (de
sureste Brasil). Habitan bosques húmedos sombreados
desde 1900–2500 m. Smith & Kessler (2018) discutieron
como distinguir las dos especies. Dentro de Blechnaceae,
Lomaria es hermano a un clado de 10 géneros que incluye
Austroblechnum, Blechnum y Parablechnum. La mayoría de
estos géneros tiene láminas con pinnas gradualmente
reducidas hacia la base. El nombre genérico se deriva
de la Griego loma, borde, y aris, sufijo. Refiere al soro
marginal que forma una línea contínua que borde la costa
de las pinnas estrechas.

LITERATURE: Broadhurst, J. 1912. The genus Struthiopteris and its representatives in North America. Bulletin of the Torrey
Botanical Club 39: 257–278, 357–385. Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for
Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations Phytotaxa 275: 191–227. Muril-
lo, M. T. 1968. Blechnum subgénero Blechnum en Sur América, con especial referencia a las especies de Colombia. Nova Hed-
wigia 16: 329–366, t. 110–147. Rolleri, C. & C. Prada. 2006. Catálogo comentado de las especies de Blechnum L. (Blechnaceae,
Pteridophyta) en Mesoamérica y Sudamérica. Anales del Jardín Botánico de Madrid 63: 67–106. Smith A. R. & M. Kessler. 2018.
Prodromus of a fern flora for Bolivia. XXXIII. Blechnaceae. Phytotaxa 334: 99–117.

364
Figure 179. Lomaria brunea (From Smith & Kessler, 2018)

365
BLECHNACEAE
Lomaridium C. Presl

DESCRIPTION: Hemiepiphytes; rhizomes long-creep- DESCRIPCIÓN: Hemiepífitas; rizomas largamente

ing, with 8–15 vascular bundles arranged in a circle (as
seen in cross-section), scaly, the scales with a dark cen-
tral stripe; sterile and fertile leaves dimorphous; petioles
with 4 or more vascular bundles; laminae pinnatisect; seg-
ments adnate to the rachis, entire; aerophores absent at
the base of the pinnae abaxially; foliar buds absent; veins
free; sterile and fertile leaves dimorphic; sori linear, par-
allel to the costae; indusia opening toward the costae (not
towards the margin); spores bilateral, nongreen; x=29, 32.
rastreros, con 8–15 haces vasculares dispuestos en un
circulo (visto en sección transversal), escamosos, las
escamas con una raya oscura central; hojas estériles
y fértiles dimorfas; pecíolos con 4 o más haces vascu-
lares; láminas pinnatisectas; segmentos adnatos al ráquis,
enteros; aeróforos ausentes en las bases de las pinnas
abaxialmente; brotes foliares ausentes; venas libres; hojas
estériles y fértiles dimorfas; soros lineares, paralelos a
las costas; indusios abriéndose hacia la costa (no hacia el
margen); esporas bilaterales, no verdes; x=29, 32.

SIMILAR GENERA: Lomariopsis differs by sessile (not GÉNEROS PARECIDOS: Lomariopsis difiere por pin-
adnate) pinnae and imparipinnate laminae. Polypodium dif- nas sésiles (no adnatas) y láminas imparipinnadas. Poly-
fers by having round sori, usually anastomosing veins, and podium difiere por tener soros redondos, usualmente
leaves articulate to the rhizomes. nervaduras anastomsadas, y hojas articuladas al rizoma.

COMMENTS: Lomaridium consists of 16 species and oc-
curs in the Neotropics, Africa, Madagascar, and Australasia.
All the species grow in wet shaded forests at middle ele-
vations. The main Neotropics ones are L. acutum (Desv.)
Gasper & V. A. O. Dittrich, L. binervatum (Poir) Gasper &
V. A. O. Dittrich, L. ensiforme (Liebm.) Gasper & V. A. O.
Dittrich, L. fragile (Liebm.) Gasper & V. A. O. Dittrich, L.
fuscosquamatum (A. Rojas) Gasper & V. A. O. Dittrich, and
L. nigrocostatum (A. Rojas) Gasper & V. A. O. Dittrich. The
sporophytes usually begin growth as epiphytes on the
base of trees and then climb. Roots are sent from the
basal portion of the rhizome into the soil, thus the plants
become hemiepiphytes. The juvenile leaves are quite dif-
ferent than the mature ones, being simple or pinnatifid
basally. Because the fertile pinnae are contracted, it can be
difficult to interpret the typical blechnoid soral condition
(i.e., sori along the costa and indusium opening toward
the costae). Lomaridium is sister to a clade of 11 genera
that includes Austroblechnum, Blechnum, and Parablechnum.
The genus name Lomaridium means resembling Lomaria,
another genus of Blechnaceae.
COMENTARIOS: Lomaridium consta de 16 especies y
ocurre en el Neotrópico, África, Madagascar y Australasia.
Todas las especies existen en bosques húmedos som-
breados en elevaciones medias. Las especies principales
Neotropicales son L. acutum (Desv.) Gasper & V. A. O.
Dittrich, L. binervatum (Poir) Gasper & V. A. O. Dittrich,
L. ensiforme (Liebm.) Gasper & V. A. O. Dittrich, L. fragile
(Liebm.) Gasper & V. A. O. Dittrich, L. fuscosquamatum (A.
Rojas) Gasper & V. A. O. Dittrich y L. nigrocostatum (A.
Rojas) Gasper & V. A. O. Dittrich. Usualmente las esporof-
itas empiecen crecer en las bases de los árboles y luego
trepan. Se envian raíces desde la parte basal del rizoma
al suelo, por eso tornando en hemiepífitos. Las hojas ju-
veniles son diferentes que las maduras, siendo simples a
pinnatífidas basalmente. A causa de las pinnas fértiles son
contraídas, puede ser difícil interpretar la condición típica
soral blechnoide (i.e., los soros por la costa y los indusios
abriéndose hacia las costas). Se requiere la observación
esmera con una lupa. Lomaridium es hermano a un cla-
do de 11 géneros que incluye Austroblechnum, Blechnum
y Parablechnum El nombre génerico Lomaridium significa
semejante a Lomaria, otro género de Blechnaceae.

LITERATURE: Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodi-
ales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227. Murillo, M. T. 1968. Blech-
num subgénero Blechnum en Sur América, con especial referencia a las especies de Colombia. Nova Hedwigia 16: 329–366,
t. 110–147. Rojas-Alvarado, A. F. 2006. Two new species of Blechnum (Blechnaceae) from the Neotropics. Brittonia 58:
388–394. Vicent,M, J. M. Gabriel y Galán, M. V. & E. B. Sessa. 2017. Phylogenetics and historical biogeography of Lomaridium
(Blechnaceae: Polypodiopsida). Taxon 66: 1304–1316.

366
Figure 180. A–E. Lomaridium fragile, with cross-section of rhizome shown in E. F. L. ensiforme. (A, F, from Mickel & Smith,
2004; B–E original ©Robbin Moran, 2016; drawn by Susanah Graedel.)

367
BLECHNACEAE
Lomariocycas (J. Sm.) Gaspar & A. R. Sm.
DESCRIPTION: Terrestrial, rarely epiphytic; trunks up
to 4 m tall, arborescent, scaly, the scales bicolor-
ous with a dark central stripe, 2–4 cm long, curved;
sterile and fertile leaves dimorphous; petioles with
several vascular bundles, the two adaxial ones enlarged;
laminae 1-pinnate basally, pinnatisect distally; pin-
nae or segments entire or serrate, never lobed, adnate
distally; costae sulcate adaxially; aerophores absent; foliar
buds absent; veins free, bifurcate; sori linear, parallel to the
costae; indusia opening toward the costae (not towards
the margin); spores bilateral, nongreen; x=33.
SIMILAR GENERA: Plagiogyria differs by absence of
scales, three vascular bundles in the petiole base, pinnae
adnate throughout, and adaxially non-sulcate costae. Par-
ablechnum differs by concolorous petiole scales and pres-
ence of aerophores at the pinna bases.
DESCRIPCIÓN: Terrestres, raras veces epífitas; tron-
cos hasta 4 m de altura, arborescentes, escamosos,
las escamas bicolores con una raya oscura central,
2–4 cm de longitud, curvadas; hojas estériles y fértiles
dimorfas; pecíolos con varios haces vasculares, los dos
adaxiales agrandodos; láminas 1-pinnadas basalmente,
pinnatisectas distalmente; pinnas o segmentos enteros
a serrados, nunca lobados; costas surcadas adaxialmente;
aeróforos ausentes; brotes foliares ausentes; venas libres,
bifurcadas; soros lineares, paralelos a las costas; indusios
abriéndose hacia la costa (no hacia el margen); esporas
bilaterales, no verdes; x=33.
GÉNEROS PARECIDOS: Plagiogyria difiere por la au-
sencia de escamas, tres haces vasculares en la base del
pecíolo y pinnas adnatas y costas no surcadas adaxial-
mente. Parablechnum difiere por escamas peciolares
concoloras y presencia de aeróforos en las bases de las
pinnas.

COMMENTS: Lomariocycas consists of about 20 spe- COMENTARIOS: Lomariocycas consta de casi 20 es-
cies, of which 5 are neotropical. The genus occurs also pecies, de las cuales 5 son neotropicales. El género existe
in Africa and Madagascar. The species generally grow at también en África y Madagascar. Generalmente las espe-
high elevations, above 2000 m, especially in páramos and cies existen en elevaciones altas, arriba de 2000 m, es-
peat marshes. The plants tolerate ground fires because pecialmente en páramos y tuberas. Las plantas soportan
their stems are insulated by a mass of surrounding peti- fuegos porque sus tallos están protegidos por una masa
ole bases and roots. The typical blechnoid soral pattern, gruesa de bases de los pecíolos y raíces. La condición
consisting of a linear sorus on either side of the costa, típica soral blechnoide, la cual consta de un soro lineal en
is often difficult to detect because of the narrow fertile los ambos lados de las costas, es a menudo difícil detectar
pinnae. The sporangia, when mature, cover the entire ab- a causa de las pinnas fértiles estrechas. Los esporangios,
axial surface of the pinnae, obscuring the costae and in- cuando maduros, cubren toda la superficie abaxial de las
dusia. This imparts the false impression of an acrostichoid pinnas fértiles, obscureciéndo las costas y los indusios.
sorus. The only solution to the correct interpretation is Da la impresión falsa de un soro acrosticoide. La sola
careful observation. Usually a vestige of the indusium can solución para la interpretación correcta es la observación
be seen near the margin of the pinna. The genus name is cuidadosa. Usualmente se puede ver un véstigio del indu-
derived from Lomaria, a dimorphic fern genus, + Cycas. sio cerca del margen de la pinna. El nombre genérico se
Refers to the sterile-fertile leaf dimorphy and cycad-like deriva Lomaria, un género dimórfico de los helechos, +
growth habit. Cycas. Se refiere al dimorfismo foliar entre las hojas es-
tériles y fértiles y el hábito de crecimiento como un cicad.
LITERATURE: Broadhurst, J. 1912. The genus Struthiopteris and its representatives in North America. Bulletin of the Torrey Botanical Club
39: 257–278, 357–385. Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodiales: Poly-
podiopsida): New genera, resurrected names, and combinations Phytotaxa 275: 191–227. Murillo, M. T. 1968. Blechnum subgénero Blechnum
en Sur América, con especial referencia a las especies de Colombia. Nova Hedwigia 16: 329–366, t. 110–147. Rolleri, C. & C. Prada. 2006.
Catálogo comentado de las especies de Blechnum L. (Blechnaceae, Pteridophyta) en Mesoamérica y Sudamérica. Anales del Jardín Botánico de
Madrid 63: 67–106. Rolleri, C. H. C. Prada, J. M. Gabriel y Galán & L. M. Passarelli. 2013. Especies arborescentes del género Blechnum
(Blechnaceae: Pteridophyta). Revista de Biología Tropical 61: 377–408. Smith A. R. & M. Kessler. 2018. Prodromus of a fern flora for Bolivia.
XXXIII. Blechnaceae. Phytotaxa 334: 99–117.

368
Figure 181. Lomariocycas aurata. At right is a bicolorous rhizome scale. (Original, ©Robbin Moran, 2016.)

369
LOMARIOPSIDACEAE
Lomariopsis Fée
DESCRIPTION: Hempipiphytes; rhizomes long-
creeping, in cross section with a broad ventral meristele,
roots borne only on the ventral surfaces; sterile and
fertile leaves strongly dimorphic; petioles not
articulate to the rhizome, with 4-8 vascular bundles;
laminae 1-pinnate (simple in juveniles of some species),
with a conform apical pinna; pinnae entire or nearly
so, articulate to the rachis; veins free, simple or with one
fork, parallel; fertile pinnae narrower than the sterile; sori
acrostichoid; spores monolete, green; x=41.
SIMILAR GENERA: Polybotrya differs by pinnatifid leaf
apices (in all but 1 species) and pinnate groups of veins.
Also, most species of Polybotrya have decompound leaves.
Lomagramma and the climbing species of Bolbitis differ
by having anastomosing veins. Juvenile plants of certain
species with simple leaves (the Lomariopsis japurensis
group) resemble Elaphoglossum but can be distinguished
by their cartilaginous-thickened margins.
COMMENTS: Lomariopsis is a genus of wet forests. It
is pantropical with about 45 species worldwide and 15
in the Neotropics. The sporophytes start growth on the
base of trunks and climb after establishing contact by long
thick roots with the soil. The climbing rhizomes adhere
to the substrate by short lateral roots produced on the
ventral surface. In transverse section the rhizome has a
characteristic cat’s paw pattern to the vascular bundles.
The ventral bundle, which bears all the roots, is notched.
The articulations of the pinnae can be seen with a
handlens as faint dark lines at juncture with the rachis.The
articulations seem non-functional because the pinnae do
not fall away with age or upon drying. Some evidence,
however, indicates they might function in response to
damage. The genus name is derived from Lomaria, a fern
genus name + opsis, a Greek suffix meaning “like.”
LITERATURE: Moran, R. C. 2000. Monograph of the neotropical species of Lomariopsis (Lomariopsidaceae). Brittonia 52:
55–111. Rouhan, G., J. G. Hanks, D. McClelland & R. C. Moran. 2007. Preliminary phylogenetic analysis of the fern genus
Lomariopsis (Lomariopsidaceae). Brittonia 59: 115–128.
370
DESCRIPCIÓN: Hemiepífitas; rizomas largamente
rastreros, en sección transversal con un haz vascular
ancho, raíces se nacen solo en las superficies ventrales;
pecíolos no articulados al rizoma, con 4-8 haces vasculares;
hojas estériles y fértiles fuertemente dimorfas; láminas
1-pinnadas (o simples en los juveniles de algunas
especies), con una pinna apical conforme; pinnas
enteras o casi enteras, articuladas al raquis; nervaduras
libres, simples o una vez furcadas, paralelas; pinnas
fértiles más angostas que las estériles; soros acrosticoides;
esporas monoletes, verdes; x=41.
GÉNEROS PARECIDOS: Polybotrya difiere por ápices
de las hojas pinnatífidos (con excepción de una especie)
y venas en grupos pinnados. También, la mayoría de
las especies de Polybotrya tienen hojas decompuestas.
Lommagramma y las especies trepadoras de Bolbitis
difieren por venas anastomosadas. Plantas juveniles
de ciertas especies con hojas simples se asemejan
a Elaphoglossum pero se pueden distinguir por sus
márgenes engrosados cartilaginosos.
COMENTARIOS: Lomariopsis es un género de
bosques húmedos. Es pantropical con casi 45 especies
mundialmente y 15 en el Neotrópico. Los esporofitos
empiezan desarrollo en la base de los troncos y suben
después de establecer contacto por raíces largas y
gruesas con el suelo. Los rizomas trepadores adhieren
al sustrato por raíces cortos laterales producidas en la
superficie ventral. En sección transversal el rizoma tiene
un patrón característico a las haces vasculares como pata
de gato. La haz ventral, que da lugar a todas las raíces,
tiene una escotadura. Las articulaciones de las pinnas se
pueden ver con una lupa como líneas oscuras en la unión
con el raquis. La articulaciones aparecen no funcionales
porque las pinnas no se desprenden con edad o al secar.
No obstante hay evidence que funcionen cuando la
pinna se daña. El nombre genérico se deriva de Lomaria,
un nombre de un género de helecho + oasis, un sufijo
griego que significa similitud.
Figure 182. Lomariopsis prieuriana. (Original, ©R. C. Moran, 2016.)

371
LONCHITIDACEAE
Lonchitis L.
DESCRIPTION: Terrestrial or occasionally epipetric;
rhizomes 5–15 mm wide, short-creeping, hairy; leaves
0.5–2.5 m, monomorphic; petiole bases with 2
vascular bundles; laminae 2- to 3-pinnate-pinnatifid,
slightly succulent, softly hairy, the hairs numerous,
septate; pinnules catadromic throughout, but sometimes
anadromic on the basal pinnae; rachises grooved adaxially,
the groove hairy; costae prominent adaxially; veins free,
ending before the margins in clavate hydathodes; sori
marginal, borne along a connecting vein parallel to the
margin and connecting (1)2 or 3 vein tips; indusia formed
by the revolute scarious margin; spores trilete; x=50.
DESCRIPCIÓN: Terrestres u ocasionalmente rupícolas;
rizoma 5–15 mm de ancho, cortamente rastrero, peloso;
hojas 0.5–2.5 m, monomorfas; bases de los pecíolos
con 2 haces vasculares; láminas 2- a 3-pinnado-
pinnatífidas, suculentas, suavemente pelosas, los
pelos numerosos, septados; pínnulas catádromas en
toda la pinna o anádromas en la base y catádromas
distalmente; raquis sulcado adaxialmente, el surco peloso;
costas prominentes adaxialmente; nervaduras libres,
terminando antes de los márgenes en ápices claviformes;
soros marginales, dispuestos en una comisura vascular
paralela al margen y conectando (1)2 o 3 ápices de
nervaduras; indusio, formado por los márgenes
escariosos revolutos; esporas triletes; x=50.

SIMILAR GENERA: Blotiella and Histiopteris differ by GÉNEROS PARECIDOS: Blotiella y Histiopteris difieren
anastomosing veins, vascular bundles in the petiole base por venas anastomosadas, hace vascular en la base del
U-shaped, and monolete spores. pecíolo en forma de U y esporas monletes.

COMMENTS: Lonchitis contains two species: L. hirsuta
L. in the Neotropics and L. occidentalis Baker in Africa-
Madagascar. Lonchitis hirsuta typically grows on wet,
shaded forest floors from 0–1200 m. The Old World
species differs by net veins along the midribs, but this
characteristic is not constant. The petioles contain
raphides, probably of calcium oxalate. These are easily
viewed in free-hand sections of the petioles. The genus
name is derived from the Greek lonche, lance + itis, a
suffix indicating a close connection with. An ancient fern
name used by Dioscorides.
COMENTARIOS: Lonchitis consta de dos especies:
L. hirsuta L. en el Neotrópo y L. occidentale Baker en
África-Madagasscar. Lonchitis hirsuta existe en típicamente
en los pisos mojados sombreados de los bosques de
1–1200 m. La especie del Viejo Mundo difiere por venas
anastomosadas por las costas, pero esta característica
no es constante. Los pecíolos contienen rafidios,
probablemente de oxalato de calcio. Estos se observen
fácilmente en secciones transversales del pecíolo. El
nombre genérico se deriva de la Griega lonche, lanza +
itis, un sufijo indicando una conexión cercana. Un nombre
anciano usado por Dioscorides para un helecho.

LITERATURE: Tryon, R. M. 1962. Taxonomic fern notes. III. The genera Lonchitis and Blotiella. Contributions from the Gray
Herbarium of Harvard University 191: 91–107.

372
Figure 183. A–C. Lonchitis hirsuta. (From Mickel & Smith, 2004.)

373
DICKSONIACEAE
Lophosoria C. Presl
DESCRIPTION: Terrestrial; stems massive, erect
or decumbent, rarely arborescent (up to 1 m tall),
stoloniferous, densely pubescent, the hairs 2-4 cm
long, golden to dark brown, wooly; leaves 2-5 m long,
monomorphous; petiole base densely pubescent like
the stem; laminae 3-pinnate to 4-pinnate, whitish
beneath, triangular or ovate; axes pubescent beneath;
veins free; sori superficial (not marginal), without
indusia, with long tangled paraphyses; the receptacle
flat (not elevated); spores tetrahedral, with prominent
equatorial ridge; x = 65.
DESCRIPCIÓN: Terrestres; tallos masivos, erectos o
decumbentes, raras veces arborescentes (hasta 1 m de
altura), estoloníferos, densamente pubescentes, pelos
2-4 cm de largo, dorados a pardo oscuros, lanosos; hojas
2-5 m de largo, monomorfas; base del pecíolo densamente
pubescente como el tallo; láminas 3-pinnadas a
4-pinnadas, blanquecinas en el envés, triangulares u
ovadas; ejes pubescentes en el envés; nervadura libre;
soros superficiales (no marginales), sin indusios, con
paráfises largos, enmarañados; receptáculo plano (no
elevado); esporas tetraédricas, con una arista prominente
ecuatorial; x = 65.

SIMILAR GENERA: The Cyatheaceae (Alsophila, GÉNEROS PARECIDOS: Las Cyatheaceae (Alsophila,
Cyathea, and Sphaeropteris) differ by scaly (not hairy) Cyathea y Sphaeropteris) difieren por tallos y bases de
stems and petiole bases. Dicksonia, Cibotium, and Culcita los pecíolos escamosos (no pubescentes). Dicksonia,
differ by marginal sori. Cibotium, y Culcita difieren por soros marginales.

COMMENTS: Lophosoria is entirely neotropical COMENTARIOS: Lophosoria es completamente

and occurs primarily from 2000-3600 m. It contains
three species: L. contracta (Hieron.) A. R. Sm. (Ecuador
and Peru), L. quadripinnata (J. F. Gmelin) C. Chr., and L.
quesadae A. Rojas (Costa Rica). Only L. quadripinnata is
widespread.The genus is often conspicuous because of
its large size and white undersurface of the leaves. All
species of the genus tend to form colonies by stolons
(Cox & Tomlinson, 1985). Molecular data show Lophosoria
is sister to Dicksonia (Korall et al. 2006, 2007). Lophosoria
is derived from the Greek lophos, mane or tuft + soros. It
refers to the hairs between the sporangia.
neotrópical y existe principalmente desde 2000-3600
m. Contiene dos especies: L. contracta (Hieron.) A. R.
Sm. (Ecuador y Perú), L. quadripinnata (J. F. Gmelin) C.
Chr. y L. quesadae A. Rojas (Costa Rica). Solamente L.
quadripinnata es muy difundida. El género a menudo
es conspicua a causa de su tamaño grande y el envés
de la lámina blanquecino. Todos las especies del género
tienden formar colonias por estolones (Cox & Tomlinson,
1985). Datos moleculares muestran Lophosoria hermana
a Dicksonia (Korall et al. 2006, 2007). Lophosoria se deriva
del griego lophos, melena o penacho + soros. Se refiere a
los pelitos entre los esporangios.

LITERATURE: Cox, P. A. & P. B. Thomlinson. 1985. Relationships between ecological pattern and branching in the tree fern
Lophosoria quadripinnata in Veracruz, Mexico. American Fern Journal 75: 105–110. Korall, P., K. M. Pryer, J. S. Metzgar, H.
Schneider & D. S. Conant. 2006. Tree ferns: monophyletic groups and their relationships as revealed by four protein-coding
platid loci. Molecular Phylogenetics and Evolution 39: 830–845. Korall, P., D. S. Conant, J. S. Metzgar, H. Schneider & K. M.
Pryer. 2007. A molecular phylogeny of scaly tree ferns (Cyatheaceae). Systematic Botany 94: 873–886. Looser, G. 1933. Sobre
las Ciateáceas chilenas y en especial sobre Lophosoria quadripinnata. Ostenia 1933: 141–151.

374
Figure 184. A–C. Lophosoria quadripinnata. (From Mickel & Smith, 2006.)

375
BLECHNACEAE
Lorinseria C. Presl

DESCRIPTION: Terrestrial or (in nortern part of range)
epipetric; rhizomes long-creeping, scaly, the scales brown,
mostly broadly lanceolate; sterile and fertile leaves
dimorphic, reddish when young; sterile leaves 40–58 cm,
fertile leaves 49–70 cm; petioles bases not swollen; sterile
leaves pinnatifid, 13–26 cm; fertile leaves pinnate, sharply
contracted, 20–27 cm long; pinnae not articulate to rachis,
arranged in 7–12 alternate pairs; veins anastomosing into
2 or more rows of areoles between costae and margin,
free only at blade margin; sori linear-oblong, deeply
sunken into blades, nearly occupying full breadth of blade.
Indusia ± membranous, lacking thickened cells, tucked
under sporangia, not recurving but mostly disintegrating
with age; x=35.
DESCRIPCIÓN: Terrestres o (en la parte norteña de
la distribución) epipétrico; rizomas largamente rastreros,
escamosos, las escamas pardas, mayormente anchamente
lanceoladas; hojas estériles y fértiles dimorfas, rojizas
cuando jóvenes; hojas estériles 40–58 cm de largo, las
fértiles 49–70 cm; las bases de los pecíolos no hinchadas;
láminas esteriles pinnatifidas, 13–25 cm de largo; hojas
fertiles pinnadas, fuertamente contraidas, 20–27 cm
de largo; pinnas no articuladas al ráquis, arreglados en
7–12 pares alternas; nervadura anastamosada en 2 o
más hileras de areolas entre la costa y el margen; soros
lineares-oblongos, hundidos profundamente en la lámina;
indusios ± membranosos, faltando de células engrosadas,
no recurvando pero principalmente desintegrando con
la edad; x=35.

SIMILAR GENERA: Onoclea differs by the presence of GÉNEROS PARECIDOS: Onoclea difiere por la
trophopods, opposite pinnae, and fertile pinnae that are presencia de trofopodios, pinnas opuestas y pinnas fértiles
brown, indurated, and enrolled to protect the sori.. que son pardos, ascendentes, endurecidas y enrolladas
para proteger los soros.

COMMENTS: Lorinseria consists of only one species: L.
areolata (L.) C. Presl. It is endemic to the eastern United
States and most common in the coastal plain, from 0–600
m. It grows in low wet woods, seeps, bogs, and rarely
(in the northern part of its range) on sandstone. It is
most closely related to Anchistea and Woodwardia, with
which it shares netted veins and chain-like sori (the other
genera of Blechnaceae have a single long sorus parallel
to the costa). The genus name honors Gustav Lorinser
(1811–1863), an Austrian physician and botanist..
COMENTARIOS: Lorinseria consta de una sola especie:
L. areolata (L.) C. Presl. Es endémica a los Estados Unidos
oriental y es más común en la planicie costal, desde
0–600 m. Existe en bosques bajos húmedos, suelos
mojados, pántanos y raras veces (en la parte norteña de
su rango) en arenisca. Es más cercanamente relacionada
a Anchistea y Woodwardia, con las cuales comparta
venas anstomosadas y soros en forma de cadena (los
otros géneros de Blechnaceae tienen un solo soro largo
paralelo a la costa). El nombre génerico honora a Gustav
Lorinser (1811–1863), un médico austriano y botánico

LITERATURE: Cranfill, R. B. 1983. The distribution of Woodwardia areolata. American Fern Journal 73: 46–52. Gasper, A.
L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodiales: Polypodiopsida): New
genera, resurrected names, and combinations. Phytotaxa 275: 191–227.

376
Figure 185. Lorinseria areolata. (From Small, 1935.)

377
LOXOGRAMMACEAE
Loxogramme (Blume) C. Presl
DESCRIPTION: Epiphytic or saxicolous; rhizomes
short- to long-creeping; roots forming a spongy mass,
with dense, opaque, dull-brown hairs; rhizome scales
clathrate, concolorous, entire, united at the base (not
peltate); plants no sclerenchymatous except for the roots;
sterile and fertile leaves monomorphous or (rarely)
dimorphous; filopodia present or absent; laminae simple,
entire, typically linear or narrowly elliptic, glabrous except
for minute glandular or clavate hairs; veins anastomosing
with many, few or no included free veinlets; hydathodes
absent; sori usually elongate, oblique or rarely
parallel to the costa; indusia absent; paraphyses filiform
or absent; spores green, trilete or monolete. x=35, 36.
SIMILAR GENERA: Polytaenium differs by non-green
spores and more than one soral line between the costa
and margin. Pleopeltis differs by peltate scales on the
blades and (usually) among the sporangia. Microgramma
differs yellow spores and (at least in some species) scales
on the laminae.
COMMENTS: Loxogramme consists of 33 species, all
of which are paleotropical except L. mexicana (Fée) C.
Chr., which occurs from Mexico to Panama. This sole
neotropical species grows in montane forests from
600–2400 m, usually as a low-trunk epiphyte or on
mossy rocks along streams. Under dry conditions the
plants are flaccid and limp, but they revive rapidly when
wetted. Schneider et al. (2004) found that Loxogramme
forms a clade with the paleotropical genera Anarthropteris
(perhaps not distinct from Loxogramme) and Dictymia.
This clade, recognized here as the Loxogrammaceae, is
sister to the Polypodiaceae. Moran and Smith (2001)
postulated on the basis of morphology that Loxogramme
mexicana and L. abyssinica (Baker) M. G. Price were sister
species. This was supported by molecular data of Janssen
and Schneider (2007). The genus name is derived from
the Greek loxos, oblique + gramme, line, referring to the
sori that are oblique to the costa.
LITERATURE: Janssen, T., H.-P. Kreier & H. Schneider. 2007. Origin and diversification of African ferns with special emphasis on Polypodiaceae. Brittonia
59: 159–181. Mickel, J. T. & A. R. Smith. 2004. Pteridophyte flora of Mexico. Memoirs of the New York Botanical Garden 88: 1–1055. Moran, R. C. & A. R.
Smith. 2001. Phytogeographic relationships between neotropical and African-Madagascan pteridophytes. Brittonia 53: 304–351.
378
DESCRIPCIÓN: Epífitas o saxícoloas; rizomas corto
a largamente rastreros; raíces formando una masa
esponjosa, con pelitos densos, opacos, pardos; escamas
del rizoma clatradas, concoloras, enteras, unidas en la
base (no peltadas); plantas no esclerenquimatosas excepto
las raíces; hojas estériles y fértiles monomorfas o (raras
veces) dimorfas; filopodios presentes o ausentes; láminas
simples, enteros, típicamente lineares o estrechamente
elípticas, glabras excepto por diminutos pelitos
claviformes a glandulares; nervaduras anastomosadas,
con muchos, pocos o ningún nérvulo libre incluido;
hidátodos ausentes; soros generalmente alargados,
oblicuos o raramente paralelos a la costa; indusios
ausentes; parafisos filiformes o ausentes; esporas verdes,
triletes o monoletes. x=35, 36.
GÉNEROS PARECIDOS: Polytaenium difiere por
esporas no-verdes y más de una línea soral entre la
costa y margen. Pleopeltis difiere por escamas peltadas
en las láminas y (usualmente) entre los esporangios.
Microgramma difiere por esporas verdes y (a lo menos
en algunas especies) escamas en las láminas.
COMENTARIOS: Loxogramme consta de 33 especies,
todas paleotropicales excepto L. mexicana (Fée) C.
Chr., que existe desde México hasta Panamá. Esta única
especie neotropical crece en bosques montanos de 600–
2400 m, usualmente como epífito en la parte inferior de
los troncos o sobre rocas musgosas por ríochuelos. Bajo
condiciones secas las plantas están flácidas y marchitadas,
pero recuperan rápidamente cuando mojado. Schneider
et al. (2004) encontraron que Loxogramme forma un
clado con los géneros paleotropicales Anarthropteirs (tal
vez no distincto de Loxogramme) y Dictymia. Este clado,
reconocido aquí como Loxogrammaceae, es hermana a
la Polypodiaceae. Moran y Smith (2001) postularon en
la basis de morfología que Loxogramme mexicana and L.
abyssinica (Baker) M. G. Price fueron especies hermanas.
Esto fue apoyado por los datos moleculares de Janssen y
Schneider (2007). El nombre del género es derivado del
griego loxos, oblicuo + gramme, línea, referiéndose a los
soros que son oblicuos a la costa.
Figure 186. Loxogramme mexicana. A. Habit. B. Rhizome. C. Rhizome scale. D. Blade detail. E. Cross section of lamina. (From
Mickel & Smith, 2004.)

379
LOXOMATACEAE
Loxsomopsis H. Christ
DESCRIPTION: Terrestrial; rhizomes creeping,
branched, covered with abundant, rigid, dark
brown, shiny bristles; leaves monomorphic, spaced
3-10 cm apart; petioles bases with 2 vascular bundles;
laminae 2-pinnate-pinnatifid to nearly 3-pinnate;
petiole dark to atropurpureious, sparsely bristly toward
the base; laminae lanceolate to oblong subdeltate,
glabrous above, glabrous to hairy below; veins free,
extending all the way to the margin; pinnae asymmentical;
sori marginal, paraphysate; indusia narrowly cup-shaped
or uceolate, entire, the receptacle exert; sporangia with
an oblique annulus not interrupted at the stalk; spores
trilete. x=46.
SIMILAR GENERA: Trichomanes differs by
membranaceous laminae and non-paraphysate sori.
Dennsteadtia usually differs by epipetiolar buds and a
shallower cup-shaped indusium. Saccoloma has a compact,
decumbent or erect rhizome.
COMMENTS: Loxomopsis is completely neotropical,
from Costa Rica to Bolivia at 1500–3000 m. It consists of
a single species: L. pearcei (Baker) Maxon. The plants are
terrestrial and occur in humid, semi-open places, especially
on slopes and roadsides in cloud forests. Loxomopsis is
a member of the tree fern clade, Cyatheales. Only one
other genus occurs in the family: Loxsoma, endemic to
North Island, New Zealand. The genus name is derived
from the Greek Loxsoma, the name of another fern genus,
+ -opsis, like.
LITERATURE: Lehnert, M., M. Monnich, T. Pleines, A. Schmidt-Lebuhn & M. Kessler. 2001. The relictual fern genus
Loxsomopsis. American Fern Journal 91: 13–24.
380
DESCRIPCIÓN: Terrestres; rizomas reptantes,
ramificados, revestidos con raíces abundantes cerdas
pardo oscuro, rígidas, lustrosas; hojas monomorfas,
espaciadas a lo largo del rizoma 3-10.3 cm; pecíolos con
2 haces vasculares en la base; láminas 2-pinnado-
pinnatífidas a casi 3-pinnadas; pecíolo ligeramente
sulcado, pardo oscuro a atropurpúreo, con cerdas hacia
la parte basal; lámina lanceolada a oblonga subdeltada,
glabra en el haz, glabra a pelosa en el envés; nervaduras
libres, bifurcadas, que se extienden a los márgenes;
pinnas asimétricas; soros marginales, parafisados;
indusio estrechamente ciatiforme a urceolado, enteros,
el receptáculo exerto; esporangios con anillo oblicuo no
interrumpido por el pedículo; esporas triletes. x=46.
GÉNEROS PARECIDOS: Trichomanes difiere por
laminas membranáceas y soros parfisados. Dennsteadtia
difiere usualmente por brotes epipeciolares e un indusio
de forma de copa menos profundo. Saccoloma tiene un
rizoma compacto o erecto.
COMENTARIOS: Loxsomopsis es completamente
neotropical, desde Costa Rica hasta Bolivia, de 1500–
3000 m. Consta de una sola especie: L. pearcei (Baker)
Maxon. Las plantas son terrestres y existen un lugares
húmedos semiabiertos, especialmente por los taludes y
lados de caminos en bosques nublados. Loxomopsis es
miembro del clado de los helechos arborescentes, las
Cyatheales. Solamente otro género existe en la familia:
Loxoma, endémico a la Isla del Norte, Nueva Zelandia.
El nombre del género se deriva del griego Loxsoma, el
nombre de otro género de helecho, + -opsis, parecido.
Figure 187. Loxomopsis pearcei. (Original, © Robbin Moran, 2007.)

381
POLYPODIACEAE
Luisma M. T. Murillo & A. R. Sm.
DESCRIPTION: Epiphytic; rhizomes radially symmetrical,
erect or suberect, scaly, the scales 2.2 X 0.3 mm, clathrate,
ciliate, lanceolate, dark gray, shiny, peltate; petioles 1.6–
2.5 cm long, terete, with 2 vascular bundles in cross
section; leaves 9–20 0.3–0.4 cm, simple, entire, narrowly
elliptic, glabrous on both surfaces, with green borders
(not blackish); hydathodes prominent; veins free, forked;
sori ca. 1 mm wide, round, adjacent to (touching) the
midrib; paraphyses absent; spores green, trilete; x=?
SIMILAR GENERA: Campyloneurum differs by
anastomosing veins. Grammitis differs by dark laminar
margins and entire rhizome scales. Cochlidium differs by
non-clathrate, entire rhizome scales.
COMMENTS: Luisma consists of one species, L.
bivascularis M. T. Murillo & A. R. Sm., endemic to the
department of Risaralda, Colombia. It grows in wet forests
at 1550 m. It was claimed that Luisma was the only genus
of grammitids with two vascular bundles in the petiole,
but this character is more common among grammitids
ferns than previously believed. Like Lellingeria, it has dark
clathrate rhizome scales, hydathodes, and lacks reddish
setae on the leaves. It probably represents a simple-
leaved species of Lellingeria, but molecular are needed to
test this idea. The genus is named in honor of Luis María
Murillo (1896–1974), pioneer Colombian entomologist.
LITERATURE: Murillo, M. T. & A. R. Smith. 2003. Luisma, a new genus of Grammitidaceae (Pteridophyta) from Colombia.
Novon 13: 313–317.
382
DESCRIPCIÓN: Epífitas; rizomas simétricos radialmente,
erectos o suberectos, escamosos, las escamas 2.2 X 0.3
mm, clatradas, ciliatas, lanceolatas, grisiacas, lustrosas,
peltatas; pecíolos 1.6–2.5 cm de largo, teretes, con dos
haces vasculares en sección transversal; hojas 9–20 X
0.3–0.4 cm, simples, enteras, angostamente elípticas,
glabras en ambas superficies, con bordes verdes (no
negruzcos); hidátodos prominentes; nervaduras libres,
furcadas; soros ca. 1 mm de ancho, redondos, adyacente
(tocando) a la costa; paráfisis ausentes; esporas verdes,
triletes; x=?
GÉNEROS PARECIDOS: Campyloneurum difiere por
venas anastomosadas. Grammitis difiere por un margen
laminar oscuro y escamas del rizoma enteras. Cochlidium
difiere por escamas del rizoma no clatradas, enteras.
COMENTARIOS: Luisma consta de una sola especie,
L. bivascularis M. T. Murillo & A. R. Sm., endémica al
departmento de Risaralda, Colombia. Crece e bosques
húmedos alrededor de 1550 m. Se alejó que Luisma fue
el único género grammitidoide con dos haces vasculares
en el pecíolo, pero esta condición es más común entre
estos helechos que creido previamente (M. Sundue, pers.
com.). Como Lellingeria, tiene escamas del rizoma oscuras
clatradas, hidatodos y falta de setas rojizos en las hojas.
Probablemente representa una especie de Lellingeria
con hojas simples, pero se necesitan datos moleculares
para probar esta idea. El género se nombra en honor
de Luis María Murillo (1896–1974), pionero entomólogo
colombiano.
Figure 188. Luisma bivascularis. (From Murillo & Smith, 2003.)

383
LYCOPODIACEAE
Lycopodiella Holub
DESCRIPTION: Terrestrial; horizontal stems creeping,
superficial, indeterminate, branching dichotomously in the
horizontal plane, also branching in the vertical plane to
give rise to single dorsal row of erect unbranched
strobilus-bearing shoots; leaves of the erect stems
arranged in 12–20 rows, either similar to those of the
creeping stems or (in P. caroliniana) reduced, distant,
polystichous, ; gemmiferous branchlets and gemmae
absent; strobili sessile, erect, solitary; sporophylls
peltate, with a decurrent medial wing, ca. 1–1.5 times as
wide as the trophophylls, stramineous at time of sporangial
dehiscence, entire or nearly so; sporangia globose or
nearly so, isovalvate, the walls with incomplete semi-
annular thickenings; spores rugulate, the sides convex at
the equator; gametophytes green, tuberous, part below
and above soil surface, with lamellae on dorsal surface;
n=78.
DESCRIPCIÓN: Terrestres; tallos horizantales
reptantes, superficiales, indeterminados, ramificándose
dicotomicamente en el plano horizontal, también
ramificándose en el plano vertical para dar lugar a una
hilera dorsal de vástagos erectas no ramificadas
llevando estróbilos; hojas de los tallos erectos arregladas
en 12–20 hileras, simulares a las de los tallos reptantes o
(en P. caroliniana) reducidas, distantes, dimórfas; ramitas
gemíferas y yemas ausentes; estróbilos sésiles, erectos,
solitarios; esporofilos peltatos, con una ala medial
decurrente, ca. 1–1.5 veces más anchas que las trofofilos,
estramíneos en el tiempo de dehiscencia esporangial,
enteros o casi así; esporángios globosos o casi asi,
isovalvados, células epidérmicas con engrosamientos
nodulares; esporas rugulosas, los lados convexos en el
ecuador; gametofitos verdes, tuberosos, parcialmente
abajo y arriba del suelo; n=78.

SIMILAR GENUS: Pseudolycopodiella differs by GÉNERO PARECIDO: Pseudolycopodiella difiere por

sporophylls ca. 2 times wider than the trophophylls on
the peduncles and in 4–10 rows. Palhinhaea differs by
erect stems branched and tree-like, nodding or pendulous
strobili, and sporangia sunken the cortex of the strobili.
esporofilos ca. 2 veces más anchos que los trofofílos en
los pedúnculos y arreglados en ca. 4–10 hileras. Palhinhaea
difiere por los tallos erectos ramificados y semejantes
a pequeños árboles, estróbilos cernuos o péndulos y
esporangios hundidos en la corteza do los estróbilos.

COMMENTS: Lycopodiella consists of ca. 12 species
worldwide. It occurs in the northern temperate regions
of North America and Eurasia, to tropical America south
to northern Argentina. It typically grows on open wet
soils, especially on sand. In the temperate zones, many
species pass the winter by drying back to the tips of
the creeping shoots, and persisting through the winter
as short, green, buds just below the soil. These resume
growth in the spring. The sporophylls and trophophylls
are superficially similar, but the sporophylls differ greatly
by their modified bases. Hybrids between species of the
same ploidy level are apparently fertile, exhibiting normal
meiosis and well-formed spores, but hybrids between
different ploidy levels are sterile. The genus name is
formed from the Lycopodium, a related genus, and the
Greek suffix -ella, a diminutive.
COMENTARIOS: Lycopodiella consta de ca. 12 especies
mundial. Existe en las regiones templadas de América
del Norte y Eurasia, a América tropical hasta Argentina
norteña. Ocurre típicamente en suelos abiertos mojados,
especialmente sobre arena. En las zonas templadas,
muchas especies pasan el invierno por moriéndose hasta
el ápice del vástago reptante, lo cual persiste como brotes
verdes justo abajo del suelo. Renueven su crecimiento
en el verano. Los esporofilos y trofofilos son simulares
superficialmente, pero los esporofilos difieren mucho
por sus bases modificadas. Híbridos entre las especies
del mismo nivel de poliplodía son aparentemente fértiles,
exhibiéndose meiosis normal y esporas bien formadas,
pero los híbridos entre especies de niveles diferentes de
poliploidia son estériles. El nombre del género se deriva
de Lycopodium y el sufijo griego diminutivo –ella.

LITERATURE: Haines, A. 2003. The families Huperziaceae and Lycopodiaceae of New England, a taxonomic and ecological ref-
erence. V. F. Thomas Co., Southwest Harbor, Maine. Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the
Missouri Botanical Garden 79: 687–717. Øllgaard, B. & P. G. Windisch. 2016. Lycopodiaceae in Brazil. Conspectus of the family
II. The genera Lycopodiella, Palhinhaea, and Pseudolycopodiella. Rodriguésia 67: 691–719. Wikström, N. & P. Kenrick. 2000.
Relationships of Lycopodium and Lycopodiella based on combined plastid rbcL gene and trnL intron sequence data. Systematic
Botany 25: 495–510.

384
Figure 189. Lycopodiella. A. L. andicola. B. L matthewsii. (From Øllgaard, 2004.)

385
LYCOPODIACEAE
Lycopodium L.
DESCRIPTION: Terrestrial; main horizontal stems
creeping, superficial, indeterminate, giving rise to two
dorsolateral ranks of erect determinate branching systems;
ultimate branchlets terete (not flattened), isophyllous,
with leaves isophyllous, polystichous (12–20 rows), the
apices hair-like, hyaline; strobili erect, pedunculate or
sessile; sporophylls subpeltate with a decurrent medial
wing, with a basal cavity that contains mucilage; sporangia
isovalvate, reniform, short-stipitate, the epidermal cells
with sinuate anticlinal walls; spores reticulate on all faces;
gametophytes subterranean, discoid, convoluted; x=34.
DESCRIPCIÓN: Terrestres; tallos principales
horizantales reptantes, superficiales, indeterminados,
dando lugar a dos hileras dorsiventrales de erectas
determinadas sistemas de ramitas; ramitas últimas teretes
(no achatados), isofílicas, con hojas polísticas (12–20
hileras), los ápices filiformes, hialinos; estróbilos
erectos, pedunculados o sésiles; esporofilos subpeltatos
con una ala decurrente medial, con una cavidad basal que
contiene mucílago; esporangios isovalvados, reniformes,
brevamente estípitatas, con células epidérmicas sinuosas;
esporas reticuladas en ambas caras; gametofitos
subterráneos, discoides, convólutos; x=34.

SIMILAR GENERA: Diphasium and Diphasiastrum differ
by flattened ultimate branchlets. Dendrolycopodium and
Spinulum, which are boreal and north-temperate genera,
differ by sessile strobili and green acute (not hairlike) leaf
tips. Austrolycopodium differs by lacking piliform, hyaline
leaf tips and sporophylls without a decurrent medial wing.
GÉNEROS PARECIDOS: Diphasiastrum y Diphasium
difieren por las ramitas últimas achatadas. Spinulum y
Dendrolycopodium, los cuales son boreales y templado-
septentrionales, difieren por estróbilos sésiles y apices de
las hojas verdes y agudos (no piliformes). Austrolycopodium
difiere por la falta de apices de las hojas piliformes hialinos
y los esporfilos sin una ala medial decurrente.

COMMENTS: Lycopodium consists of about 10 species
worldwide and occurs on all continents except Australia.
In the tropics it is found at high elevations, usually 2000–
3800 m. The hair-like, colorless leaf apices are absolutely
diagnostic for the genus. Lycopodium clavatum L., the type
of the genus, is remarkable by being nearly cosmopolitan.
In the Neotropics it intergrades with L. contiguum Klotzsch,
a species that differs only by sessile strobili (the plants
are sometimes recognized as L. clavatum subsp. contiguum
(Klotzsch) B. Øllg.). Similar is L. vestitum Poir. from Ecuador
and Peru. It differs by the apical half of the leaves hyaline
and scarious, imparting a grayish to silvery color to the
branchlets. The genus name is derived from the Greek
lycos, wolf + pous, foot. The branch tips resemble a wolf ’s
paw.
COMENTARIOS: Lycopodium consta de casi 10 especies
mundial y existe en todos los continentes con excepción
de Antarctica. En el trópico se encuentra en elevaciones
altas, usualmente de 2000–3800 m. Los ápices piliformes
sin color son absolutamente diagnósticos para el género.
Lycopodium clavatum L., el tipo del género, es notable por
ser casi cosmopolita. En el Neotrópico se intergrada con
L. contiguum Klotzsch, una especies que difiere solamente
por estróbilos sésiles (las plantas a veces se reconocen
como L. clavatum subsp. contiguum (Klotzsch) B. Øllg.).
Parecido es L. vestitum Poir. de Ecuador y Peru. Difiere
por la mitad apical de las hojas hyalina y escariosa, dando
un apariencia grisácea a plateada a las ramitas. El nombre
genérico se deriva del griego lycos, lobo + pous, pie. Los
ápices de las ramas se asemejan a un pato de un lobo.

LITERATURE: Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Øllgaard,
B. 1988. Lycopodiaceae. In: G. Harling & L. Andersson, editors. Flora of Ecuador, no. 33. 1992. Neotropical Lycopodiaceae—an
overview. Annals of the Missouri Botanical Garden 79: 687–717. Øllgaard, B. 1989. Index of the Lycopodiaceae. Biologiske Skrift-
er 34: 1–135. Øllgaard, B. & K. Tind. 1993. Scandinavian ferns. Rhodos, Copenhagen. Øllgaard, B. & P. G. Windisch. 2014.
Lycopodiaceae in Brazil. Conspectus of the family I. The genera Lycopodium, Austrolycopodium, Diphasium, and Diphasiastrum.
Rodriguésia 62(2): 2014 (on-line only).

386
Figure 190. Lycopodium. A. Spores. B. Lycopodium lagopus. Note the annual growth contrictions. C. Sporophylls; note
medial wing decurrent onto the axis of the strobilus. D. Lycopodium clavatum, habit; note note piliform apices of leaves.. (A–C
from Øllgaard & Tind, 1993; D from Mickel & Smith, 2004.)

387
LYGODIACEAE
Lygodium Sw.
DESCRIPTION: Terrestrials; rhizome creeping,
branched, pubescent; leaves vine-like, of indeterminate
growth, climbing by means of a twinning rachis;
pinnae alternate, short-stalked, the stalk forked with
an arrested bud in the axil; veins free or areolate;
sporangia borne singly on the vein tips (not clustered
in sori), covered by a flap of lamina tissue, arranged in
2 rows on narrow marginal projections of the lamina;
annulus apical; x=29, 30.
DESCRIPCIÓN: Terrestres; rizomas reptantes,
ramificados, pubescentes; hojas de crecimiento
indefinido, trepa por medio de un raquis que
enrosca como un bejuco; pinnas alternas, cortamente
pecioluladas, peciolulo furcado con un brote latente
en la axila; nervaduras libres o areoladas; esporangios
naciendo individualmente en los ápices de las
nervaduras (no agrupados en soros), cubiertos por una
excresencia pequeña de tejido laminar, arreglados en 2
hileras sobre proyecciones marginales estrechas de la
lámina; anillo apical; x=29, 30.

SIMILAR GENERA: In the Neotropics, the only other GÉNEROS PARECIDOS: En el Neotrópico, el único
fern genus that climbs by means of twinning rachises is otro género de los helechos que trepa por medio de
Salpichlaena. It differs by its much larger size and sori los ráquises que enroscan es Salpichlaena. Difiere por tu
parallel to the costules. tamaño más grande y los soros paralelos a las cóstulas.

COMMENTS: Lygodium is pantropical and contains COMENTARIOS: Lygodium es pantropical y contiene

about 25 species, only one of which is temperate (L.
palmatum, of the northeastern United States). Four species
occur in the Neotropics. Two of these are common and
widespread, especially in seasonally dry forests: Lygodium
venustum Sw. and L. volubile Sw.
Lygodium is distinctive by its twining rachises. The
young leaves have reduced pinna and elongated rachises
that circumnutate widely until they find a support.
After a support is found, then larger, expanded pinnae
are produced. The short bifurcated pinna stalk is also
distinctive but easy to overlook, as is the small bud in its
axil.
The genus is placed in the Schizaeales along with
Lygodiaceae and Anemiaceae. The three families share
the synapomorphy of a complete apical annulus.
The genus name is derived from the Greek lygodes,
flexible, and refers to the long, flexible rachis.
alrededor de 25 especies, de las cual solo una es templada
(L. palmatum, del noreste de los E.E.U.U.). Cuatro especies
existen en el Neotrópico. Dos de ellas son comunes y muy
difundidas, especialmente en bosques secos estacionales:
Lygodium venustum Sw. y L. volubile Sw.
Lygodium es distinctivo por sus ráquises enredadores.
Las hojas jóvenes tienen pinnas reducidas y ráquises
muy alargados que hacen circumnutaciones hasta que
encuentran un apoyo. Después, producen pinnas más
grandes, expandidas. El peciolulo corto y bifurcado es
distinctivo también pero fácil pasar por alto, tanto como el
brote pequeño en su axila. Se ubica en las Schizaeales, con
Lygodiaceae y Anemiaceae. Las tres familias comparten la
sinapomorfía un anillo apical completo.
El nombre genérico se deriva del griego lygodes, flexible,
y se refiere al raquis largo y flexible.

LITERATURE: Duek, J. J. 1978. A taxonomic revision of Lygodium (Filicinae) in America. Feddes Repertorium. Zeitschrift
für Botanische Taxonomie und Geobotanik 89: 411–423. Garrison-Hanks, J. 1998. Monograph of Lygodium Swartz
(Pteridophyta: Lygodiaceae). Unpublished doctoral dissertation, City University of New York. Lellinger, D. B. 1969.
Schizaeaceae (Filicales). In: B. Maguire and collaborators. The botany of the Guayana Highland—Part VIII. Memoirs of the New
York Botanical Garden 17(1): 2–11. Maxon, W. R. 1909. Schizaeaceae. North American Flora 16: 31–52.

388
Figure 191. Upper left: Lygodium volubile. G: L. heterodoxum. H, J: L. venustum. (From Mickel & Smith, 2004.)

389
THELYPTERIDACEAE
Macrothelypteris (H. Itô) Ching

DESCRIPTION: Terrestrial; rhizome scales pubescent
on the surfaces and margins; petioles with 2 vascular
bundles; laminae 2- to 3-pinnate-pinnatifid; rachises
and costae not grooved adaxially; pinnae with adnate
pinnules, the proximal pinnae not reduced, or only slightly
so; aerophores absent; veins free, 1-forked, ending before
the margin in a clavate apex (seen on the upper surface)
indumentum of acicular hairs; sori round; indusium
round-reniform, rarely absent; sporangia generally with
capitate glands near the annulus; x=31.
DESCRIPCIÓN: Terrestres; escamas del rizomas
pubescentes en las superfícies y márgenes; pecíolos
con 2 haces vasculares; láminas 2- a 3-pinnada-
pinnatífidas o más divididas; ráquises y costas no
sulcadas adaxialmente; pinnas con pínnulas adnatas,
las pinnas proximales no reducidas o poco reducidas;
aeróforos ausentes; nervaduras libres, 1-furcadas,
terminándose antes del margen en un ápice claviforme
(vistas en el haz); indumento de pelos aciculares; soros
redondos; indusios rotundos-reniformes, raramente
exindusiados; esporangios generalmente con glándulas
capitadas cerca del anillo; x=31.

SIMILAR GENERA: All other American species of
Thelypteridaceae differ by less divided laminae and (with
the exception of Phegopteris) veins ending at the lamina
margin. Because of its leaf size and division, Macrothelypteris
might be mistaken for dryopteroid genera such as Ctenitis,
Megalastrum, y Lastreopsis, but it differs from them by
having needle-shaped hairs and two vascular bundles in
the petiole.
GÉNEROS PARECIDOS: Todos las otras especies
Americánas de Thelypteridaceae difieren por las láminas
menos divididas y (con la excepción de Phegopteris) las
venas terminándose en el márgen de la lámina. A causa
del tamaño y división de sus hojas, Macrothelypteris puede
ser confundido con géneros driopteroides como Ctenitis,
Megalastrum y Lastreopsis, pero difiere de ellos por tener
tricomas aciculares y dos haces vasculares en el pecíolo.

COMMENTS: All the 10 species of Macrothelypteris are
native to the Paleotropics. Only M. torresiana (Gaudich.)
Ching is naturalized in the Americas. This species has
spread rapidly and widely and is now one of the most
common weedy species. It exists in the United States as
far north as southern Illinois. It generally grows in disturbed
habitats.The genus forms a clade with Phegopteris (North
Temperate Zone) and Pseudophegopteris (Paleotropical).
These three have rachises and costae ungrooved adaxially
and veins that end before the lamina margin. The surfaces
of the scales are pubescent, which is a rare character in
the family. The genus name is derived from the Greek
macros, large + Thelypteris. The species of the genus are
large compared to other thelypteroid ferns.
COMENTARIOS: Todos las 10 especies de
Macrothelypteris son nativas en los Paleotrópicos. Sólo
M. torresiana (Gaudich.) Ching está naturalizada en el
las Américas. Esta especie ha difundida rapidamente
y ampliamente y ahora es una de las especies mas
comunes y agresivas. Existe en los Estados Unidos
hasta Illinois del sur. Generalmente existe en ambientes
abiertos perturbados. El género forma un clado con
Phegopteris (de las zonas templadas) y Pseudophegopteris
(paleotropical). Estos tres tienen ráquises y costas no
surcadaos adaxialmente y venas que terminan antes el
márgen de la lámina. Las superficies de las escamas son
pubescentes, que es una característica rara en la familia. El
nombre genérico se deriva del griego makros, grande +
Thelypteris. Las especies del género son grandes cuando
comparadas a otros helechos thelypteroides.

LITERATURE: Holttum, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris and
Macrothelypteris. Blumea 17: 5–32. Leonard, S. W. 1972. The distribution of Thelypteris torresiana in the southeastern United
States. American Fern Journal 62: 97–99. Smith, A. R. 1983. 14(4). Polypodiaceae—Thelypteridoideae. In: G. Harling & B. Sparre,
Flora of Ecuador, no. 18. Smith, A. R. & R. Cranfill. 2002. Intrafamilial relationships of the thelypteroid ferns (Thelypteridaceae).
American Fern Journal 92: 131–149.

390
Figure 192. Macrothelypteris torresiana (From Tryon & Stolze, 1992.)

391
MARATTIACEAE
Marattia Sw.
DESCRIPTION: Terrestrial, fleshy; stems massive, erect;
sterile and fertile leaves monomorphic, with several
borne at a time; petiole bases with 2 rounded stipules,
without nodes, scaly, the scales narrow, often filiform;
lamina triangular, 2-5-pinnate, the apex pinnatifid;
pinnae attached to the rachis by swollen nodes,
opposite or nearly so; axes of the distal segments usually
winged; veins free; synangia sessile, splitting longitudinally
to reveal two faces each with a row of elongate pores;
paraphyses absent or subtending the synangia in a nest-
like ring; spores monolete; x=40.
SIMILAR GENERA: Eupodium differs by stalked
synangia, the presence of awns along the veins adaxially,
and leaves typically one per plant. Danaea differs by
1-pinnate leaves, dimorphy between the sterile and fertile
leaves, and sporangia opening by circular pores.
COMMENTS: Marattia contains 7 species and occurs
from southern Mexico to Panama, the Caribbean,
southern Brazil, and Hawaii. The species, all of which
inhabit wet forests, are: Marattia alata Sw. (Jamaica and
Cuba); M. cicutifolia Kaulf. (southern Brazil), M. douglasii
(Hawaii); M. excavata Underw. (from Mexico to Panama),
M. laxa Kunze (from Mexico to Panama), M. interposita H.
Christ (from Guatemala to Panama), and M. weinmannifolia
(southern Mexico and Guatemala). The genus name
honors Giovanni Francesco Maratti (1723–1777), a
Benedictine abbot and later professor and head of the
botanical garden at Rome.
LITERATURE: del Carmen Lavalle, M. 2003.Taxonomía de las especies neotropicales de Marattia (Marattiaceae). Darwiniana
4: 61–86. Murdock, A. G. 2008. A taxonomic revision of the eusporangiate fern family Marattiaceae, with description of a new
genus Ptisana. Taxon 57: 737–755. Underwood, L. M. 1909. Marattiaceae. North American Flora 16: 15–23.
392
DESCRIPCIÓN: Terrestres, carnosas; tallos masivos,
erectos; hojas estériles y fértiles monomorfas, con
varias nacidas a la vez; bases de los pecíolos con 2
estípulas redondeadas, sin nudos, escamosas, las escamas
angostas, a menudo filifomes; láminas triangulares,
2-5-pinnadas, el ápice pinnatífido; pinnas unidas al raquis
por nudos hinchados, opuestas o casi así; ejes de los
segmentos distales usualmente alados; nervaduras libres;
sinángios sésiles, partiéndose longitudialmente revelar
dos caras cada una con una hilera de poros elongados;
parafisos ausentes o subtendiendo los sinángios en un
anillo nidiforme; esporas monoletes; x=40.
GÉNEROS PARECIDOS: Eupodium difiere por los
snángios peciolados, la presencia de aristas por las venas
adaxialmente y típicamente hojas una por planta. Danaea
difiere por láminas 1-pinnadas, dimorfismo foliar entre
las hojas estériles y fértiles y esporangios abriéndose por
poros circulares.
COMENTARIOS: Marattia consta de 7 especies y
existe desde el sur de México hasta Panamá, el Caribeo,
sur de Brazil y Hawaii. Las especies, todos de las cuales
existen en bosques húmedos, son: Marattia alata Sw.
(Jamaica y Cuba), M. cicutifolia Kaulf. (sur de Brasil), M.
douglasii (Hawai), M. excavata Underw. (México a
Panamá), M. laxa Kunze (México a Panamá); M. interposita
H. Christ (Guatemala a Panamá) y M. weinmannifolia (
sur de México y Guatemala). El nombre genérico honra
Giovanni Francesco Maratti (1723–1777), un abato
Benedictino and más tarde un profesor y jefe del jardín
botánico en Roma.
Figure 193. A–B: Marattia excavata. C–F: M. weinmanniifolia. G–J: M. laxa. (From Mickel & Smith, 2004.)

393
MARSILEACEAE
Marsilea L.
DESCRIPTION: Aquatic or semiaquatic; rhizomes
long-creeping, branched, rooting at the nodes and (in
many species) the internodes; leaves 1-pinnate, with 4
wedge-shaped pinnae clustered at the apex of a
long petiole; floating leaves with petioles longer than in
terrestrial plants, lax or flexuous, the largest pinnae often
with red bands on the lower surface (hidropotonia);
fertile leaves identical in appearance to the sterile ones,
with short-stalked sporocarps along the petiole
base or towards its base; mega- and microsporangia in
the same sporocarps; x=20.
DESCRIPCIÓN: Acuáticos o semiacuáticos; rizomas
largamente rastreros, ramificados, enraizandos en los
nudos y (en muchas especies) además en los entrenudos;
hojas 1-pinnadas, con 4 pinnas cuneiformes agrupadas
en el ápice de un largo pecíolo; hojas flotantes con
pecíolos más largos que en plantas terrestres, laxos o
flexuosos, las pinnas más grandes a menudo con bandas
rojas en el envés (hidropotonia); hojas fértiles idénticas
en apariencia a las hojas estériles, con esporocarpos
cortamente pediculados a lo largo del pecíolo o
cerca de su base; mega y microsporangios en el mismo
esporocarpo; x=20.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Marsilea is cosmopolitan and contains
about 45–70 species, most of which occur in Africa and
Australia. It occurs in wet or seasonally wet habitats,
and is frequently found around the margins of ponds
and marshes. The sporocarps represent conduplicate,
indurated pinnae fused along their margins to form a
closed structure. Phylogenetically, the genus belongs
to the clade of heterosporous water ferns, which also
includes Azolla, Salivia, and Regnellidium. The leaves of
Marsilea exhibit sleep movements, or nyctinasty. During
the day the pinnae are oriented horizontally. Each is
equipped with pulvinus at the base. Toward nightfall, the
two distal pinnae rise up, twist half around and come into
contact with one another. Afterwards, they are embraced
by the basal pinnae, so that the four pinnae with their
lower surfaces turned outward form a vertical packet.
Marsilea is the only fern that exhibits nyctinasty (Darwin,
1896).Marsilea is named for Count Luigi Ferndinando
Marsigli (1656–1730), Italian botanist at Bologna.
COMENTARIOS: Marsilea es cosmopólito y cuenta con
45–70 especies, la mayoría de las cuales existen en África y
Australia. Occure en hábitats mojadas o estacionalmente
así y se encuentra con frequencia alrededor de los
márgenes de chacras y pantanos. Los esporocarpos
representan pinnas conduplicadas, endurecidas y
fusionadas por los márgines para forma una estructura
cerrada. Filogeneticamente, el género pertenece al clado
de los helechos acuáticos heterospóricos, que incluye
también Azolla, Salivia y Regnellidium. Las hojas de Marsilea
exhiben movimientos de dormir, o nictinasty. Durante el
día las pinnas están orientadas horizontalmente. Cada
una tiene un pulvínulo en la base. Al atardecer las dos
pinnas distales se levantan y se torcen media vía y llegan
a ser en contacto con cada una. Después están abrazadas
por las pinnas basales, de manera que las cuatro pinnas
con sus superficies abaxiales tornadas a fuera forman un
paquete vertical. Marsilea es el único helecho que exhibe
nictinasty (Darwin, 1896). Marsilea es nombrado para
Count Luigi Ferndinando Marsigli (1656–1730), botánico
italiano en Bologna.

LITERATURE: Darwin, C. 1896. The power of movement in plants. D. Appleton & Co., New York. Jacono, C. C. & D. M.
Johnson. 2006. Water-clover ferns, Marsilea, in the southeastern United States. Castanea 71: 1–14. Johnson, D. M. 1986.
Systematics of the New World species of Marsilea (Marsileaceae). Systematic Botany Monographs 11: 1–87. Pérez-García, B.
R. Riba & D. M. Johnson. 1999. Pteridofitas, Familia Marsileaceae. Flora de México 5: 1–17. Schneider, H. & K. M. Pryer. 2002.
Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. International Journal of Plant Science
163: 485–505.

394
Figure 194. A–D: Marsilea macropoda. A. habit. B. sporocarps borne on branched peduncle. C. sporocarps with indument
removed to show variation in shape. D. variation in attachment of sporocarps. E–F: M. ancylopoda. E. habit. F. sporocarps to
show variation in shape and in orientation of peduncle. (From Johnson, 1986.)

395
ONOCLEACEAE
Matteuccia Todaro
DESCRIPTION:Terrestrial; rhizomes erect, stoloniferous,
scaly; sterile and fertile leaves dimorphic; sterile leaves
50–130 x 15–25 cm; petioles 1/5–1/10 the length of the
lamina, with 2 vascular bundles; sterile blades 1-pinnate-
pinnatifid, oblanceolate, gradually tapered toward
the base, apices pinnatifid; sterile pinnae sessile, 20–60
pairs, truncate; veins free; fertile leaves 2-pinnate, 20–40
cm long, erect, green but turning brown and indurate
at maturity; fertile pinnae strongly ascending; pinnules
enrolled and forming a globose structure enclosing each
sorus; receptacle prominent; paraphyses absent; indusia
present as a thin hyaline scale; spores bilateral, greenish;
x=40.
SIMILAR GENERA: Onocleopsis, of southern Mexico
and Guatemala, differs by net veins and fertile leaves
2-3-pinnate. Thelypteris subg. Amauropelta differs by
aerophores present (at the pinna bases), acicular hairs,
and monomorphic sterile and fertile leaves.
COMMENTS: Matteuccia consists of 3 species native
to north temperate regions. Two are Asian, and one is
North American (M. struthiopteris). They mainly occur on
shaded forest floors and tend to form colonies by action
of the stolons. Onoclea stores starch in the thick, wide,
petiole bases that persist after the above part of the leaf
has senesced. The fertile leaves persist throughout the
winter and release their spores the following late winter
or early spring. Onoclea is sister to a clade consisting of
Onocleopsis and Matteuccia (Gastony & Ungerer, 1997).
These genera, plus the Asian Pentarhizidium (2 species)
constitute the Onocleaceae. The genus name honors
Carolo Matteucci (1800–1863), physicist at the University
of Florence, Italy.
LITERATURE: Gastony, G. J. & M. C. Ungerer. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns
(Dryopteridaceae: Onocleeae). American Journal of Botany 84: 840–849. Lloyd, R. M. 1971. Systematics of the onocleoid ferns.
University of California Publications in Botany 61: 1–86. Von Aderkas, P. 1984. Economic history of ostrich fern, Matteuccia
struthiopteris, the edible fiddlehead. Economic Botany 38: 14–23.
396
DESCRIPCIÓN: Terrestres; rizomas erectos,
estoloníferos, escamosos; hojas estériles y fértiles
dimorfas; hojas estériles 50–130 x 15–25 cm; pecíolos
1/5–1/10 la longitud de las láminas, con 2 haces vasculares;
láminas estériles 1-pinnado-pinnatífidas, oblanceoladas,
gradualmente angostadas hacia la base, ápices pinnatífidos;
pinnas estériles séssiles, 20–60 pares, truncadas; venas
libres; hojas fértiles 2-pinnadas, 20–40 cm de largo,
erectas, verdes pero tornándose pardas y induratas con
maduréz; pínulas fértiles enrolladas y formando una
estructura globosa encapsulando cada soro; receptáculo
prominente; paraphyses ausentes; indusios presentes
como una escama delgada hialina para cada soro; esporas
bilaterales, verdosas; x=40.
GÉNEROS PARECIDOS: Onocleopsis, del sur de
México y Guatemala, difiere por rizomas nervación en
forma de red y hojas fértiles 2-3-pinnadas. Thelypteris
subg. Amauropelta difiere por aeróforos presentes (en las
bases de las pinnas), pelos aciculares y hojas estériles y
fértiles monomórfas.
COMENTARIOS: Matteuccia consta de 3 especies
nativas a zonas templadas norteñas. Dos son Asiáticos,
y una es Norteamerican (M. struthiopteris). Existen
principalmente en los sotobosques sombreados y
tienden formar colonias por acción de sus estolones.
Onoclea alménese almidón en las bases gruesos y anchos
de los pecíolos que persistan después de lo demás de la
hoja ha caída. Las hojas fértiles persistan por el invierno y
libran sus esporas en el invierno siguiente o la primavera
temprano. Onoclea es hermana a un clado formado por
Onocleopsis y Matteuccia (Gastony & Ungerer, 1997).
Estos géneros, y el asiático Pentarhizidium (2 especies)
constituyen las Onocleaceae. El nombre genérico honora
Carolo Matteucci (1800–1863), físico en la Universidad
de Florencia, Italia.
Figure 195. Matteuccia struthiopteris. Note scale-like leaves on stolons in B. (original, ©Robbin Moran, 2016.)

397
DRYOPTERIDACEAE
Maxonia C. Chr.
DESCRIPTION: Lianas or hemiepiphytes; rhizomes
creeping, scaly; sterile and fertile leaves dimorphic;
petioles with 6 or more vascular bundles; laminae 2- to
3-pinnate-pinnatifid, glabrous or nearly so, anadromous,
chartaceous to coriaceous, the apex pinnatifid; longest
segments on the acroscopic side of the pinnae; rachis
and costae grooved adaxially, the grooves more or less
continuous with those of the lower order, glabrous within
the grooves; veins free; sori round; indusium round,
appearing peltate but actually attached at a narrow sinus;
spores monolete; x=41.
SIMILAR GENERA: Arachniodes differs by
monomorphic sterile and fertile leaves. Cyclodium usually
differs by 1-pinnate laminae and anastomosing veins.
Polybotrya differs by vascular bundles of the rhizome
each surrounded by a sheath of black sclerenchyma and
acrostichoid, non-indusiate sori. Lastreopsis differs by
terrestrial habit and costae puberulent within the grooves
adaxially. Rumohra differs by monomorphic sterile and
fertile leaves and peltate undusia.
COMMENTS: Maxonia is entirely neotropical, and its
sole species, M. apiifolia (Sw.) C. Chr., grows in wet forests
from 0–1200 m. It forms a clade with other dryopteroid
genera that are lianas or hemiepiphytes (Arachniodes,
Cyclodium, Olfersia, Polybotrya). Two varieties have been
recognized: var. dualis (Guatemala to western Ecuador)
and var. apiifolia (Cuba and Jamaica). They differ only by
the rhizome scales: those of var. dualis are entire, whereas
those of var. apiifolia are erose to denticulate. The genus
name honors William Ralph Maxon (1877–1948),
pteridologist at the United States National Herbarium,
Washington, D.C.
LITERATURE: Chandra, S. 1975. Revision of the neotropical fern genus Cyclodium. Brenesia 6: 1–7. Christensen, C. 1916.
Maxonia, a new genus of tropical American fens. Smithsonian Misc. Collect. 66(9): 1–4. Moran, R. C. 1987. Monograph of the
neotropical fern genus Polybotrya (Dryopteridaceae). Bulletin of the Illinois Natuarl History Survey 34: 1–138. Walker, T. G.
1972. The anatomy of Maxonia apiifolia: a climbing fern. British Fern Gazette 10: 241–250.
398
DESCRIPCIÓN: Llianas o hemiepífitas; rizomas
reptantes, escamosos; hojas estériles y fértiles
dimorfas; pecíolos con 6 o más haces vasculares; láminas
2- a 3pinnado-pinnatífidas, glabras o casi así, anádromas,
cartácea a coriáceas; ráquises y costas sulcados
adaxialmente, los surcos más o menos confluentes de un
eje al siguiente, glabros por dentro; costas abaxialmente
con tricomas uniseriados, rojizos oparduscos, adpresos;
nervaduras libres; soros redondeados; indusios
redondeados, apareciéndose peltados pero en realidad
unidos a un seno estrecho; esporas monoletes; x=41.
GÉNEROS PARECIDOS: Arachniodes difiere por
hojas estériles y fértiles monomorfas. Cyclodium difiere
usualmente por laminas 1-pinnadas y venas anastomosadas.
Polybotrya difiere por los haces vasculares del rizomas
cada uno redeado por una vaina de esclerenquima negra
y soros acrosticoides no indusiados. Lastreopsis difiere
por habito terrestre y costa puberulentas dentro de los
surcos adaxiales. Rumohra difiere por hojas estériles y
fértiles monomórfas y indusios peltados.
COMENTARIOS: Maxonia es completamente
neotropical, y su única especie, M. apiifolia (Sw.) C. Chr.,
existe en bosques húmedos desde 0–1200 m. Forma
un clado con otros géneros dryopteroides que son
lianas o hemiepífitos (Arachniodes, Cyclodium, Olfersia,
Polybotrya). Dos variedades ha sido reconocidos: var. dualis
(Guatemala a Ecuador occidental) y la var. apiifolia (Cuba
y Jamaica). Difieren solamente en las escamas del rizoma:
en var. dualis son enteras, mientras que en la var. apiifolia
son erosas a denticuladas. El nombre genérico honora
a William Ralph Maxon (1877–1948), pteridólogo en el
Herbario Nacional de los Estados Unidos., Washington,
D.C.
Figure 196. Maxonia apiifolia. (Drawn by S. Graedel; original, ©Robbin Moran, 2016.)

399
DRYOPTERIDACEAE
Megalastrum Holttum
DESCRIPTION: Terrestrial; petioles with 4 or more
vascular bundles, scaly especially toward the base; lamina
2–4 pinnate, catadromic above the basal pinnae, the
laminar tissue without cylindrical glands; basal pinnae
enlarged basiscopically; rachises, pinna rachises, and
costules adaxially with strigose, multicelular hairs; basal
vein of the distal pinnules springing from the costa,
not the costule; veins ending in hydathodes behind
the margin; sori round; indusium absent or rarely present;
spores monolete, reniforme; x=41.
DESCRIPCIÓN: Terrestres; pecíolos con 4 o más
haces vasculares, escamosos especialmente hacia la
base; lámina 2–4-pinnada, catádroma por encima de las
pinnas basales, el tejido laminar sin glándulas cilíndricas;
pinnas basales alargadas basiscópicamente; ráquises,
ráquises de las pinnas y cóstulas adaxialmente con pelos
strigosos multicelulares; nérvulo basal basiscópico de
las pínnulas distales surgiendo de la costa, no de la
cóstula; nervaduras terminando en un hidatodo detrás
de los márgenes; soros redondos; indusio ausente o raras
veces presente; esporeas monoletes, reniformes; x=41.

SIMILAR GENERA: Macrothelypteris torresiana GÉNEROS PARECIDOS: Macrothelypteris torresiana

(Thelypteridaceae) differs by 2 vascular bundles in the (Thelypteridaceae) difiere por 2 haces vasculares en el
petiole and the presence of acicular hairs on the laminae. pecíolo y la presencia de pelos aciculares en las láminas.
Lastreopsis differs by its rachis-costa architecture (which Lastreopsis difiere por su tipo de arquitectura de las
see). ráquises y costas (veáse).

COMMENTS: Megalastrum contains about 100 species COMENTARIOS: Megalastrum se presentan

in the Neotropics, 3 in Africa-Madagascan region, and alrededor de 100 especies en el Neotrópico, 3 en la
7 circumaustral region. All are terrestrial plants of wet región África-Madagascar y 7 el la región circunaustral.
forests. The genus is closely related to Lastreopsis and Todos son plantas terrestres de bosques húmedos. El
Rumohra. Megalastrum was segregated from Ctenitis on género es más cercanamente relacionado a Lastreopsis
the basis of its venation and type of hairs. The venation y Rumohra. Megalastrum fue segregado de Ctenitis con
is characteristic because the basal basiscopic veins of the base en su nervación y tipo de tricomas. La nervación
distal pinnules spring from the costae, not the costules, es distincta porque los nérvulos basiscópicos basales de
and the basiscopic lobe supplied by this vein is adnate to las pínnulas distales surgen de la costa, no de la cóstula
the costa (see G below). Another venation characteristic y el lobo basiscópico nervado por esta rama vascular
is the prominent hydathodes. Similar genera such as es ampliamente adnato a la costa (veáse G abajo). Otra
Ctenitis and Lastreopsis have small, reddish, twisted hairs característica de la nervación es los hidatodos prominentes.
on the upper surface of the axes. In contrast, the hairs Géneros simulares como Ctenitis y Lastreopsis tienen
in this position in Megalastrum are coarse, whitish, and tricomas pequeños rojizos, torcidos, sobre los ejes
antrorsely strigose hairs with a sharp apex that often adaxialmente. En contraste, Megalastrum tiene tricomas
does not twist upon drying. Although the genus is well toscos, blanquecinos, antrorsamente estrigosos, con un
defined, many of its species are not, and the group badly ápice agudo, que a menudo no se tuercen al secarse.
needs monographic study. The genus name is derived Aunque el género está bien definido, muchas especies no
from the Greek mega–, large + Lastrea, referring to the lo están, y es muy necesaria una monografía. El nombre
genus Lastrea which honors Charles Jean Louis Delastre genérico se deriva del griego mega-, grande + Lastrea,
(1792–1859), French botanist. refiere al género Lastrea lo cual fue honora Charles Jean
Louis Desastre (1792–1859), botánico francés.
LITERATURE: Moran, R. C., J. Prado & P. Labiak. 2009. Megalastrum (Dryopteridaceae) in southeastern Brazil, Paraguay,
and Uruguay. American Fern Journal 94: 1–44. Moran, R. C., J. Prado & P. Labiak. 2009. Megalastrum (Dryopteridaceae)
in the West Indies. Brittonia 61: 273–292. Rouhan, G. & R. C. Moran. 2011. Monograph of the paleotropical species of
Megalastrum (Dryopteridaceae). Annals of the Missouri Botanical Garden 98: 90–100. Smith, A. R. & R. C. Moran. 1986.
New combinations in Megalastrum (Dryopteridaceae). American Fern Journal 77: 124–130. Sundue, M., G. Rouhan & R. C.
Moran. 2010. Megalastrum (Dryopteridaceae) of the circumaustral region: Chile, Argentina, and southern islands of the Atlantic,
Pacific, and Indian Oceans. Systematic Botany 35: 461–475.

400
Figure 197. A, B. Megalastrum lunense. C–E. M. atrogriseum. F, G. M. subincisum. H–J. M. pulverulentum. (From Mickel &
Smith, 2004.)

401
POLYPODIACEAE
Melpomene A. R. Sm. & R. C. Moran
DESCRIPTION: Epiphytic, saxicolous or terrestrial;
rhizome scales cordate at base, clathrate, generally
blackish or reddish, entire (never setulose), often with one
to several papillose cells at the apex, these often falling
away and not present; laminae sweet-smelling when
dried, pinnatifid, pinnatisect or rarely 1-pinnate basally,
linear to elliptic, pubescent and setose, the hairs 0.1–0.2
mm, branched, 2–8 celled, lax, pale reddish, with most
of the color concentrated in the transverse walls, the
setae 0.3–3 mm, reddish, multicellular, erect to spreading;
hydathodes present; veins free, often not visible; sori
round, discrete, superficial or rarely lightly sunken; spores
green, trilete; x=37.
DESCRIPCIÓN: Epífitas, saxícolas o terrestres;
escamas del rizoma cordiformes basalmente,
clatradas, generalmente negruzcas o rojizas, enteras
(nunca setulosas), a menudo con una a varias células
papilosas en el ápice, éstas a menudo desprendidas y
no presentes; láminas huelen dulce cuando secas,
pinnatífidas, pinnatisectas o raramente 1-pinnadas
basalmente, lineares a elípticas, pubescentes y setosas,
los tricomas 0.1–0.2 mm, ramificados, 2–8 células, laxos,
rojizo pálido, con la mayor parte de color concentrado
en las paredes transversales, las setas 0.3–3 mm, rojizas,
multicelulares, erectas a patentes; hidátodos presentes;
nervaduras libres, no visibles; soros redondeados,
discretos, superficiales o rara vez ligeramente hundidos;
esporas verdes, triletes; x=37.

SIMILAR GENERA: Lellingeria differs by the tiny,
unequally branched hairs on the petiole and rachis, the
absence of dark setae, and sunken sori. Also, many species
of Lellingeria have setulose rhizome scales, whereas those
of Melpomene are entire.
GÉNEROS PARECIDOS: Lellingeria difiere por los
pelitos del pecíolo y raquis desigualmente furcados, la
ausencia de setas oscuras y soros hundidos. También,
muchas especies de Lellingeria tienen escamas del rizoma
setulosas, mientras que las de Melpomene nunca son
enteras.

COMMENTS: Melpomene consists of about 30 species, COMENTARIOS: Melpomene consta de casi 30

all neotropical except for M. flabelliformis, which extends especies, todos neotropicales con excepción de M.
into Africa, Madagascar, and Réunion. The species grow flabelliformis que se extiende hasta África, Madagascar y
primarily as epiphytes in wet forests.When dry, Melpomene La Réunion. Las especies crecen principalmente como
emits a sweet, spicy odor that is unique among grammitid epífitos en bosques húmedos. Cuando seca Melpomene
ferns. Also nearly unique are the papillae at the apex of tiene un olor dulce que es único entre helechos
the rhizome scales, but these fall off easily and are often grammitiodes. También casi único son las papilas en el
not seen. Most of the species have castaneous setae on ápice de las escamas del rizoma, pero éstas se desprenden
the petiole and/or rachis, and several have long-creeping fácilmente y a menudo no están vistas. La mayoría de
rhizomes—a characteristic rare among grammitid ferns. las especies tienen setas castañas sobre el pecíolo y/o
The genus is monophyletic and sister to Lellingeria (Ranker el raquis, y varias tienen rizomas largamente rastreros—
et al., 2004). Both genera have clathrate rhizome scales; una característica rara entre helechos grammitoides. El
however, those of Melpomene tend to be reddish and género es monofilético y hermano a Lellingeria (Ranker
entire (those of Lellingeria are often ciliate). Melpomene is et al., 2004). Los dos géneros tienen escamas del rizoma
the name of the Muse of Tragedy and refers indirectly to clatradas, pero las de Melpomene tienden ser rojizas
L. Earl Bishop, American student of grammitids ferns, who y enteras (las de Lellingeria a menudo son ciliadas).
died of AIDS before completing his studies of these ferns. Melpomene es el nombre de la Musa de la Tragedia y
refiere indirectamente a L. Earl Bishiop, Estado Unidense
de helechos gramitidoides, quién murio de SIDA antes de
que completar sus estudios de estos helechos.
LITERATURE: Kessler, M., E. Connor & M. Lehnert. 2014. Volatile organic compounds in the strongly fragrant fern genus
Melpomene (Polypodiaceae). Plant Biology ISSN 1435-8603. Lehnert, M. 2008. Eleven new species in the grammitid fern genus
Melpomene (Polypodiaceae). American Fern Journal 98: 214–250. 2013. Lehnert, M. 2013. Grammitid ferns (Polypodiaceae),
II. Melpomene. Flora Neotropica Monograph 112: 1–121. Lehnert, M. et al. 2009. Phylogeny of the fern genus Melpomene
(Polypodiaceae) inferred from morphology and chloroplast DNA analysis. Systematic Botany 34: 17–27. Ranker, T. A., A. R.
Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Staub & H. Schneider. 2004. Phylogeny and evolution and
grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy. Taxon 53: 415–428. Smith, A. R. & R. C.
Moran. 1992. Melpomene, a new genus of Grammitidaceae (Pteridophyta). Novon 2: 426–432.

402
Figure 198. A, B. Melpomene leptostoma. C, D. M. deltata. E, F. M. moniliformis. G, H. M. katasophistes. J, K. M.
pilosissima. L, M. M. anfractuosa [=Ascogrammitis] (From Mickel & Smith, 2004.)

403
THELYPTERIDACEAE
Meniscium Schreb.
DESCRIPTION: Terrestrial; petioles with 2 vascular
bundles; laminae 1-pinnate, rarely entire; pinnae
entire or serrate, the basal ones the largest or nearly
so; aerophores (found at the pinna base) absent; veins
netted regularly in pairs, with the transverse veins
producing excurrent veins that run toward the
margin, the areoles 4–25(–35) between the costa and
margin; hairs needle-shaped; sori on the transverse veins,
elongate (not round); indusia absent; sporangia glabrous
or setose; x=36.
SIMILAR GENERA: Some species of Serpocaulon (e.g.,
S. fraxinifolium and S. triseriale) also have 1-pinnate blades
and netted veins with excurrent veinlets in the areoles.
They differ from Meniscium by having four or more
vascular bundles in the petiole, lack of acicular hairs, and
round sori.
COMMENTS: Meniscium is completely neotropical and
consists of 27 species. It occurs mostly below 1000 m
in wet forests. It is easy to recognize by its characteristic
pattern of venation with straight and regular rows of
veins between the costa and margin. The genus name
is derived from the Greek meniskos, diminutive of mene,
moon. It refers to the shape of the sori, which spread
across the transverse veins and form a short crescent.
LITERATURE: Fernandes, R. S., J. C. Yesilyurt & A. Salino. 2014. New species and combinations in Meniscium
(Thelypteridaceae). Phytotaxa 184: 1–11. Smith, A. R. 1983. 14(4). Polypodiaceae—Thelypteridoideae. In: G. Harling & B.
Sparre, Flora of Ecuador, no. 18.
404
DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
vasculares; láminas 1-pinnatas, raras veces enteras;
pinnas enteras or serradas, los basales lo más
largas o casi así; aeróforos ausentes; nervaduras
anastomosadas regularmente en pares, con las
nervaduras transversas produciendo nervaduras
excurrentes, las aréolas en 4–25(–35) hileras entre
la costa y el margen; pelos en forma de aguja; soros en
las nervaduras transversas, alargados (no redondeados);
indusios ausentes; esporangios glabros o setosos; x=36.
GÉNEROS PARECIDOS: Algunas especies de
Serpocaulon (e.g., S. fraxinifolium and S. triseriale) también
tienen láminas 1-pinnadas y nervaduras anastomosadas
con venillas excurrentes en los aréolas. Difieren de
Meniscium por tener 4 o mas haces vasculares en el
pecíolo, pelos aciculares ausentes, y soros redondeados.
COMENTARIOS: Meniscium es completamente
neotropical y consta de 27 especies. Existe principalmente
bajo de 1000 m en bosques húmedos. Es fácil reconocer
por el patrón característico de la nervadura con hileras
rectas y regulares entre la costa y el margen. El nombre
genérico se deriva del griego meniscos, diminutivo de
mene, luna. Refiere a la forma de los soros, los cuales se
difunden por la venilla transversa y forman un creciente
corto.
Figure 199. Meniscium. A, B. M. falcata. C, D. M. angustifolia. E–G. M. reticulata. H, J. M. serrata. K–N. M. standleyi (From
Mickel & Smith, 2004.)

405
METAXYACEAE
Metaxya C. Presl
DESCRIPTION: Terrestrial; stems densely pubescent
,not arborescent, short-creeping, solenostelic; leaves up
to 2.5(–3.6) m long, monomorphous; lamina 1-pinnate
with a terminal pinna; pinnae alternate, stalked, entire,
serrate distally; veins free, parallel to each other, simple or
bifurcate at the base; rachises and costae sulcate adaxially;
sori superficial (not marginal), without indusia, scattered
close to the costa, 1–3 on each vein; paraphyses filiform;
receptacles flat (not elevated). x=ca. 94–96.
DESCRIPCIÓN: Terrestres; tallos densamente
pubescentes, no arborescentes, cortamente rasteros,
solenoestélicos; hojas de hasta 2.5(–3.6) m de largo,
monomorfas; lámina 1-pinnada con una pinna
terminal; pinnas alternas, pecioluladas, enteras, serradas
apicalmente; venas libres, paralelas entre sí, simples
o bifurcadas en la bases; ráquises y costas sulcadas
adaxialmente; soros superficiales (no marginales), sin
indusios, difundidos cerca de la costa, 1–3 sobre cada
vena; parafises filiformes; receptáculos planos (no
elevados). x=ca. 94–96.

SIMILAR GENERA: Many species of Danaea are GÉNEROS PARECIDOS: Muchas especies de Danaea
1-pinnate with a terminal pinna but, unlike Metaxya, they son 1-pinnadas con una pinna terminal, pero diferente
have scaly stems, opposite pinnae, and swollen nodes at a Metaxya, tienen tallos escamosos, pinnas opuestas y
the attachment of the pinnae. nudos hinchados en la unión de las pinnas.

COMMENTS: Metaxya is completely Neotropical and COMENTARIOS: Metaxya es completamente

the only genus in it own family. It contains six species:
M. contamanensis Tuomisto & G. G. Cárdenas, M. elongata
Tuomisto & G. G. Cárdenas, M. lanosa A. R. Sm. & Tuomisto,
M. parkeri (Hook. & Grev.) J. Sm., M. rostrata (Kunth) C.
Presl, and M. scalaris Tuomisto & G. G. Cárdenas. All grow
in wet forests, primarily below 800 m. Its juvenile leaves
are highly lacerate-serrate, quite unlike the large leaves.
Metaxya is unique among ferns because it may have more
than one sorus per vein. It belongs to the Cyatheales,
forming a clade with Cibotiaceae, Cyatheaceae y
Dicksoniaceae. A key to the species is given in Cárdenas
et al. (2016). The genus name is derived from the Greek
metaxy, between or in the middle of. It refers to the sori
borne along the veins between the costa and margin of
the pinnae.
Neotropical y el único género en su propia familia.,
Contiene sies especies: M. contamanensis Tuomisto & G.
G. Cárdenas, M. elongata Tuomisto & G. G. Cárdenas, M.
lanosa A. R. Sm. & Tuomisto, M. parkeri (Hook. & Grev.) J.
Sm., M. rostrata (Kunth) C. Presl y M. scalaris Tuomisto &
G. G. Cárdenas. Todas existen de en bosques húmedos,
principalmente bajo de 800 m. Sus hojas pequeñas son
profundamente lacerada-serradas, muy diferente que las
hojas grandes. Metaxya es único entre helechos porque
puede tener más de un soro por vena. Pertenece a
los Cyatheales, formando un clado con Cibotiaceae,
Cyatheaceae y Dicksoniaceae. Una clave a las especies se
encuentra en Cárdenas et al. (2016). El nombre genérico se
deriva del griego metaxy, entre en medio de. Se refiere a
los soros nacidos por las venas entre la costa y el margen
de las pinnas.

LITERATURE: Cárdenas, G. G., H. Tuomisto & S. Lehtonen. 2016. Newly discovered diversity in the tropical fern genus
Metaxya based on morphology and molecular phylogenetic analyses. Kew Bulletin 71: 5 (27 pages) [DOI 10.1007/S12225-
016-9618-9]. Smith, A. R., H. Tuomisto, K. M. Pryer, J. S. Hunt, P. G. Wolf. 2001. Metaxya lanosa, a second species in
the genus and fern family Metaxyaceae. Systematic Botany 26:480–486. Qiu, Y-J., R. A. White & M. D. Turner. 1995. The
developmental anatomy of Metaxya rostrata (Filicales: Metaxyaceae). American Journal of Botany 82: 969–981.

406
Figure 200. Metaxya rostrata. (From Tryon & Stolze, 1989.)

407
DRYOPTERIDACEAE
Mickelia R. C. Moran, Labiak & Sundue
DESCRIPTION: Plants terrestrial (2 species) or
terrestrial and scandent; rhizomes short- to long-creeping,
dorsiventral, the ventral surface root-bearing, the
dorsal surface leaf-bearing; sterile and fertile leaves
dimorphic; leaves 1-pinnate or (in A. furcata) with a
single free basal segment on the basal pinnae and thus
2-pinnate; laminae usually with a free apical segment
similar to the lateral pinnae; lamina buds absent or (in
the two terrestrial species) present at the pinna bases;
pinnae articulate or continuous with the rachis, entire or
serrate distally; veins variously anastomsing but never
free, either areolate throughout without main lateral veins
and without included veinlets, or with main lateral veins
connected by slightly arched cross veins, and typically with
included veinlets in the areoles; sori acrostichoid; x=41.
DESCRIPCIÓN: Plantas terrestres (2 especies)
o terretres y escandentes; rizomas cortamente a
largamente rastreros, la superficie ventral llevando raíces;
hojas estériles y fértiles dimorfas; hojas 1-pinnadas
o (en A. furcata) con un segmento basal libre en las pinnas
basales y por eso 2-pinnadas; láminas usualmente con
un segmento apical libre y parecido a las pinnas
laterales; brotes foliares ausentes o (en dos especies
terrestres) presentes en las bases de las pinnas;; pinnas
articuladas o continuas con el caquis, enteras o distalmente
serradas; venas anastomosadas valiosamente pero
nunca libre, areoladas por todas partes sin venas
laterales principales y sin venas incluidas, o con venas
conectadas por venas cruces levamente arcuadas con
venas incluidas en las areolas; soros acrosticoides; x=41.

SIMILAR GENERA: Bolbitis usually differs by terrestrial
habit, pinnatifid lamina apices, and buds (if present)
located toward the leaf or segment apices. Lomariopsis
differs by free veins. Tectaria differs by non-acrostichoid
(usually round) sori.
GÉNEROS PARECIDOS: Bolbitis usualmente difiere
por hábito terrestre, ápices de las láminas pinnatifidos, y
brotes (si presentes) ubicados hacia el ápice de la hoja o
segmentos. Lomariopsis difiere por venas libres. Tectaria
difiere por soros no acrosticoides (usualmente redondos).

COMMENTS: Mickelia consists of 10 species, all COMENTARIOS: Mickelia consta de 10 especies,

neotropical. The genus grows in wet forests, generally
below 1200 m, and all but two species (M. hemiotis and M.
oligarchica) are scandent on trunks.The species have been
classified in Bolbitis, but Moran et al. (in press) showed
that Mickelia is distinct and sister to Elaphoglossum. A
synopsis of the genus, including a key and illustrations
of all species, is given by Moran et al. (in review). Only
two species of Mickelia (both terrestrial) have leaf buds,
and those buds are located in a different position than
in Bolbitis: In Mickelia the buds occur at the pinna bases
on the acroscopic side of the costae where the costa is
joined by the lamina tissue (i.e., the buds are produced
just outside of the pinna “axil”). In contrast, the buds
in Bolbitis are adaxial and associated with the apical
portion or terminal segment of the lamina. The rhizome
architecture of M. guianensis was studied by Gay (1993,
as Lomagramma). The genus name honors John T. Mickel,
American pteridologist (1934– ).
todas neotropicales. El género se encuentra en bosques
húmedos, generalmente abajo de 1200 m, y todas las
especies con excepción de dos (M. hemiotis y M. oligarchica)
son escandentes sobre troncos. Las especies han sido
clasificado principalmente en Bolbitis, pero Moran et al.
(en prensa) mostraron que Mickelia es distinta y hermana
a Elaphoglossum. Un sinopsis de las especies, incluyendo
una clave y ilustraciones, es dado por Moran et al. (en
revisión). Solamente dos especies de Mickelia (ambas
terrestres) tienen brotes foliares, y estos están ubicados
en una posición diferente que en Bolbitis. En Mickelia se
ubican en las bases de las pinnas en el lado acroscópico,
mientras que en Bolbitis están asociados con la porción
apical o segmento terminal de las lámina. La arquitectura
del rizoma de M. guianensis fue estudiado por Gay (1993,
como Lomagramma). El nombre genérico honora John T.
Mickel, pteridólogo estadounidense (1934– ).

LITERATURE: Gay, H. 1993. The architecture of a dimorphic clonal fern, Lomagramma guianensis (Aublet) Ching
(Dryopteridaceae). Botanical Journal of the Linnean Society 111: 343–358. Hebant-Mauri, R. & H. Gay. 1993. Morphogenesis
and its relation to architecture in the dimorphic clonal fern Lomagramma guianensis (Aublet) Ching (Dryopteridaceae). Botanical
Journal of the Linnean Society 112: 257–276. Moran, R. C., Labiak & M. Sundue. 2010. Phylogeny and character evolution
of the bolbitidoid ferns (Dryopteridaceae). International Journal of Plant Science 171: 547–559. Moran, R. C., P. Labiak & M.
Sundue. 2011. Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae). Brittonia 62: 337–356.

408
Figure 201. Mickelia. A. M. bernoullii. B. M. oligarchica. C. M. pergamentacea. D. M. lindigii (original, ©R. C Moran, 2010.)

409
POLYPODIACEAE
Microgramma C. Presl
DESCRIPTION: Epiphytic or rarely saxicolous;
rhizomes long-creeping, the scales peltate, not
clathrate, lanceolate to acicular; sterile and fertile leaves
monomorphous or (more commonly) dimorphous with
the fertile leaves longer and narrower than the sterile;
laminae simple, entire, ovate to elliptic or oblong; veins
anastomosing, the areoles with included veinlets;
sori round or rarely oblong, in a single row between
the rachis and margin; paraphyses present, consisting of
narrow scales. Spores monolete, yellow; x=37.
DESCRIPCIÓN: Epífitas o raramente saxícolas;
rizomas largamente rastreros, las escamas peltadas,
no clatradas, lanceoladas a aciculares; hojas estériles y
fértiles monomorfas a (más comúnmente) dimorfas con
las fértiles más angostas y largas que las estériles; láminas
simples, enteras, ovadas a elípticas u oblongas; venas
anastomosadas, las aréolas con venillas incluídas;
soros redondos u raras veces oblongos, en una sola hilera
entre el raquis y el margen; parafisos presentes y consisten
de escamas angostas. Esporas monoletes, amarillas; x=37.

SIMILAR GENERA: Pleopeltis differs by clathrate
rhizome scales and circular peltate scales in and around
the sori, especially when young. Microgramma lacks such
scales. Campyloneurum and Niphidium differ by their larger
size (over 20 cm long) and (usually) by having more than
one row of sori between the costa and margin.
GÉNEROS PARECIDOS: Pleopeltis difiere por
las escamas del rizoma clatradas y escamas peltadas
circulares adentro y alrededor de los soros, especialmente
los jóvenes. Microgramma carece de dichas escamas.
Campyloneurum y Niphidium difieren por su tamaño más
grande (más de 20 cm) y (usualmente) más de una hilera
de soros entre la costa y el margen.

COMMENTS: Microgramma has about 30 species in
the Neotropics and 2 in Africa and islands of the Indian
Ocean. Most of the species grow in lowland forests from
0–600 m. The genus forms a clade with Niphidium and
Campyloneurum. Many species of Microgramma have
slightly flattened stems. This tendency reaches its greatest
development in M. megalophylla (Desv.) Sota, which may
have stems up to 30 mm wide and only 3 mm thick. The
genus name is derived from the Greek mikros, small +
gramme, line. The sori are slightly elongated in the type
species.
COMENTARIOS: Microgramma tiene casi 30 especies
en el Neotrópico y 2 en África e islas del Océano Indico.
La mayoría de especies crecen en bosques de tierras
bajas desde 0–600 m. El género forma un clado con
Niphidium y Campyloneurum. Muchas especies tienen tallos
levemente aplanados. Esta tendencia es más desarrollada
en M. megalophylla (Desv.) Sota, que puede tener tallos
hasta 30 mm de ancho y sólo 3 mm de grosor. El nombre
genérico se deriva del griego mikros, pequeño + gramme,
línea. Los soros son levemente alargados en la especie
tipo.

LITERATURE: de la Sota, E. R. 1963. Microgramma megalophylla (Polypodiaceae, s. str.) una interesante especie de
Amazonas. Boletín de la Sociedad Argentina de Botánica 10: 158–165. de la Sota, E. R. 1986. Sobre la posición sistemática de
Polypodium fuscopunctatum Hook. y Polypodium percussum Cav. (Polypodiaceae s. str., Pteridophyta). Physis (Buenos Aires),
Sección C, 44(106): 19–28. de la Sota, E. R. & B. Pérez-García. 1982. Nervación y dimorfismo foliar en Microgramma Presl
(Polypodiaceae s. str.) Biotica 7: 45–64; de la Sota, E. R. & B. Pérez-García. 1982. Diversidad y estructura de las escamas
rizomáticas en Microgramma Presl. (Polypodiaceae s. str., Pteridophyta). Biotica 7: 463–472. Salino, A., T. E. Almeida, A. R.
Smith, A. N. Gómez, H.-P. Kreier & H. Schneider. 2008. A new species of Microgramma (Polypodiaceae) from Brazil and
recircumscription of the genus based on phylogenetic evidence. Systematic Botany 33: 630–635.

410
Figure 202. A, B. Microgramma piloselloides. C, D. M. reptans. F, G. M. nitida. H–K. M. tecta. L–O. M. percussa. (From
Mickel & Smith, 2004.)

411
POLYPODIACEAE
Microgramma subgen. Solanopteris (Copel.) Lellinger
DESCRIPTION: Epiphytic; stems slender, long-creeping,
bearing tubers 2–4 cm wide, globose, hollow, scaly; stem
scales not clathrate, circular or rarely linear, peltate; sterile
and fertile leaves monomorphous or dimorphous, entire
or pinnatifid, glabrous or sparsely and minutely scaly;
veins anastomosing, the areolae containing free included
veinlets; sori round, elongate or linear; paraphyses
numerous, narrowly clavate, with enlarged apical cells;
spores echinate; x=37.
DESCRIPCIÓN: Epífitas; tallos delgados, largamente
rastreros, con tubérculos 2–4 cm de ancho, globosos,
huecos, escamosos, las escamas no clatradas, circulares,
peltadas; hojas estériles y fértiles monomorfas a
dimorfas, enteras o pinnatífidas, glabras o esparcida y
diminutamente escamosas; nervaduras anastomosadas
con venillas libres incluidas; soros redondos, alargados o
lineares; parafisos numerosos, estrechamente clavados,
con células agrandadas en el ápice; esporas equinadas;
x=37.

SIMILAR GENERA: Microgramma s.s. differs by the GÉNEROS PARECIDOS: Microgramma s.s. difiere
absence of tubers, narrowly lanceolate stem scales, and en la ausencia de tubérculos, escamas estrechamente
smooth spores. lanceoladas y esporas lisas.

COMMENTS: Subg. Solanopteris is completely COMENTARIOS: Subg. Solanopteris es completamente

neotropical and contains 4 species: M. bifrons (Hook.)
Lellinger (Amazon basin of Colombia, Ecuador, and
Peru), M. bismarkii (Rauh) L. León (eastern Andes of
Peru), M. brunei (Wercklé ex H. Christ) Lellinger (Costa
Rica, Panama, western Colombia and Ecuador), and M.
tuberosa (Maxon) Lellinger (Andes of S. Ecuador). All
are high canopy epiphytes, often occurring on the outer
portions of tree branches. Subg. Solanopteris is the only
myrmecophytic fern genus in the Neotropics. Its tubers
have hollow chambers inhabited by fierce biting ants. The
ants bring into the tubers organic material that eventually
decays and liberates nutrients that are absorbed by the
plant. The subgenus name is derived from Solanum, the
genus of potato + pteris, fern (Greek). It refers to the ant-
inhabited tubers which resemble potatoes.
neotropical y consta de 4 especies: M. bifrons (Hook.)
Lellinger (cuenca Amazónica de Colombia, Ecuador,
y Perú), M. bismarkii (Rauh) L. León (Andes oriental
de Perú), M. brunei (Wercklé ex H. Christ) Lellinger
(Costa Rica, Panama, lados occidentales de Colombia
and Ecuador) y M. tuberosa (Maxon) Lellinger (Andes
del sur de Ecuador). Todos son epifitos del alto dosel, a
menudo inhabitando las partes extremas de las ramas de
árboles. Subg. Solanopteris el único género mirmecofítico
de helechos en los neotropicos. Los tubérculos tienen
cámaras huecas las cuales están habitadas por hormigas
mordazes feroces. Las hormigas llevan en los tubérculos
materia orgánica que eventualmente se pudre y libera
nutrientes que están absorbidos por la planta. El nombre
del subgénero se deriva de Solanum, el género de papas
+ pteris, helecho (griego). Se refiere a los tubérculos,
inhabitados por hormigas, que asemejan a papas.

LITERATURE: Gómez, L. D. 1974. Biology of the potato fern Solanopteris brunei. Brenesia 4: 37–61. Hagemann, W. 1969.
Zur morphologie der knolle von Polypodium bifrons Hook. und P. brunei Wercklé. Société Botanique de France, Mémoires 17–27.
León, B. & H. Beltrán. 2002. A new Microgramma subgenus Solanopteris (Polypodiaceae) from Peru and a new combination
in the subgenus. Novon 12: 481–485. Rauh, W. 1973. Solanopteris bismarckii Rauh, ein neuer knollenbildender Ameisenfarn aus
Zentral-Peru. Tropische und subtropische Pflanzenwelt 5: 223–256. Wagner, W. H., Jr. 1972. Solanopteris brunei, a little-known
fern epiphyte with dimorphic stems. American Fern Journal 62: 33–43.

412
Figure 203. Microgramma subgen. Solanopteris. A. Circular stem scale of M. brunei. B. M. bifrons. C. Tubers of M. brunei
cut dorsiventrally. D. Tubers of M. brunei, ventral surface showing entrance hole for ants. E. Leaf of M. bifrons. F. Leaf of M.
bismarckii. G. M. bismarckii. H. tuber of M. brunei cut dorsiventrally; note roots appressed to chamber walls. I. Paraphyses of M.
bifrons. J. paraphyses of M. brunei. K. Tuber of M. bismarckii. (Original, ©R. C Moran, 2010.)
413
POLYPODIACEAE
Moranopteris R. Y. Hirai & J. Prado
DESCRIPTION: Epiphytic, rarely saxicolous; rhizome
scales either golden, orangish, yellowish or orangish
brown, often shiny, not clathrate, setulose or not, setulae
when present hyaline, concoloras or, often, darker than
the body of the scale; leaves short-petiolate or nearly
sessile; petioles often setose; laminae (or pinnae if lamina
is bipinnatifid) commonly 2–15(–25) × 0.2–0.8(–1.2)
cm, linear, shallowly pinnatifid to bipinnatisect; segments
generally 20–80 pares, decurrent, sometimes with an
acroscopic lobe; veins simple or with a single acroscopic
fertile branch; hydathodes present on the upper surface
of the lamina; sori one per segment, round, superficial;
spores tetrahedral, green; x=37.
DESCRIPCIÓN: Epifíticas, rara vez saxícolas; escamas
del rizoma doradas, anaranjadas o amarillas a
anaranjado-pardusco, a menudo lustrosas, no
clatradas, setulosas o no, sétulas cuando presentes
hialinas, concoloras o, a menudo, más oscuras que el
cuerpo de la escama; hojas cortamente pecioladas
o casi séssiles; pecíolo a menudo setoso; láminas
comúnmente 2–15(–25) × 0.2–0.8(–1.2) cm, lineares,
someramente pinnatífidas a bipinnatisectas; segmentos
generalmente 20–80 pares, decurrentes, a veces con
una joroba acroscópica; nervaduras simples o con sólo una
ramificación acroscópica fértil; hidatodos presentes en
el haz; soros uno por segmento, redondos, superficiales;
esporas tetrahédricas, verdes.

SIMILAR GENERA: Terpsichore has more than one GÉNEROS PARECIDOS: Terpsichore tiene más de
sorus per segment. Lellingeria and Melpomene have un soro por segmento. Lellingeria y Melpomene tienen
clathrate rhizome scales. Cochlidum and Lellingeria lack escamas del rizoma clatradas. Cochlidum y Lellingeria
reddish setae on the leaves. carecen de setas rojizas en las hojas.

COMMENTS: Moranopteris has about 30 species, COMENTARIOS: COMENTARIOS: Moranopteris

all neotropical. They were previously placed in tiene 28 especies, todas neotropicales. Ellas fueron
Micropolypodium Hayata, which is now defined as an Asian previamente ubicadas en Micropolypodium Hayata, la cual
genus of only three species (M. okuboi, M. sikkimense, ahora se define como un género Asiático de tres especies
and M. pulojense). The other genera of grammitid ferns (M. okuboi, M. sikkimense y M. pulojense). Los otros géneros
rarely have shiny, colored rhizome scales and typically de helechos gramitidoides raras veces tienen escamas del
have more than two veins per segment. In contrast, rizoma lustrosas y coloreadas y típicamente tienen más
Moranopteris has only one or two veins per segment, de dos venas por segmento. En contraste, Moranopteris
which are represented by hydathodes on the upper tiene solo una o dos venas por segmento, las cuales son
surface of the leaf). The cells of the rhizome scales representadas por hidatodos en el haz). Las células de las
appear swollen or inflated—another difference from escamas del rizoma aparecen hinchadas o túrgidas—otra
many grammitid ferns.Moranopteris achilleifolum (Kaulf.) diferencia de muchos helechos gramitoides. Moranopteris
R. Y. Hirai & J. Prado M. longisetosum (Hook.) R. Y. Hirai achilleifolum (Kaulf.) R. Y. Hirai & J. Prado y M. longisetosum
& J. Prado were formerly classified in Terpsichore. They (Hook.) R. Y. Hirai & J. Prado fueron previamente
are unusual in Moranopteris by their bipinnatisect laminae. clasificadas en Terpsichore. Son atípicas en Moranopteris
Also unusual are M. grisebachii (Underw. Ex C. Chr.) R. Y. por sus láminas bipinatisectas. También inusual son M.
Hirai & J. Prado and M. perpusillum (Maxon) R. Y. Hirai & grisebachii (Underw. Ex C. Chr.) R. Y. Hirai & J. Prado y
J. Prado because, unlike their congeners, they lack setae M. perpusillum (Maxon) R. Y. Hirai & J. Prado porque, a
on the leaves. The genus name honors Robbin Moran, diferencia de las otras especies en el género, faltan setas
American pteridologist (1956– ). en las hojas. El nombre genérico honora a Robbin Moran,
pteridólogo americano (1956– ).
LITERATURE: Copeland, E. B. 1952. The American species of Xiphopteris. American Fern Journal 42: 41–52, 93–110.
Hirai, R. Y. & J. Prado. 2011, Moranopteris: a new neotropical genus of grammitid ferns (Polypodiaceae) segregated from
Asian Micropolypodum. Taxon 60: 1123–1137. Hirai, R. Y. & J. Prado. 2012. Monograph of Moranopteris (Polypodiaceae).
Pteridologia 4: 1–113. Labiak, P. H. & F. B. Matos. 2007. A new hybrid and two new combinations in neotropical grammitid
ferns. Brittonia 59: 182–185. Maxon, W. R. 1916. Studies of tropical American ferns—no. 6. Contributions from the United States
National Herbarium 17: 541–557. Smith, A. R. 1992. A review of the fern genus Micropolypodium (Grammitidaceae). Novon
2: 419–425.

414
Figure 204. A–C. Moranopteris trichomanoides (Sw.) R. Y. Hirai & J. Prado. D, E. M. basiattenuata (Jenman) R. Y. Hirai & J.
Prado. F, G. M. blepharodes (Maxon) R. Y. Hirai & J. Prado. (From Mickel & Smith, 2004.)

415
POLYPODIACEAE
Mycopteris Sundue
DESCRIPTION: Mostly epiphytic; roots inserted
ventrally; rhizomes dorsiventral, creeping, scaly, often with
branch buds and phyllopodia; rhizome scales orange to
castaneous, setose, the cells turgid, lustrous; fronds
beset with a black clavate fungus (Acrospermum),
monomorphic, setose throughout, the setae usually
1–2 mm long, reddish,; petiole with one or two vascular
bundles, castaneous, often pubescent, the hairs ca 0.1 mm
long, 3-celled, 1-branched; rachises castaneous; laminae
proximally 1-pinnate, distally pinnatisect, generally oblong,
with numerous segments, the margins entire; pinna
costae and veins with dark sclerenchyma, the dark color
of which is easily seen on the abaxial side of the lamina;
hydathodes present, usually cretaceous; sori round;
sporangia glabrous; spores green; x=37.
SIMILAR GENERA: Ascogrammitis differs by blackish
clathrate rhizome scales and green veins abaxially.
Alansmia, Micropolypodium, and Terpsichore differ by lack
of black clavate fungi and dark veins abaxially. Ceradenia
and Zygophlebia differ by lacking hydathodes.
COMMENTS: Mycopteris is entirely neotropical,
occurring from Mexico to Bolivia, the Guianas, southeastern
Brazil, and the West Indies. It comprises ca. 30 species
and grows in humid montane forests. The genus consists
of two main clades: the first, which includes Mycopteris
taxifolia, has rhizome scales with only apical setae, dark
sclerenchyma along the costae and veins, and medial
sori. The second clade, which includes M. semihirsuta
and M. alsopteris, has phyllopodia, lack of receptacular
setae, supramedial sori, and rachis setae of two distinctly
different lengths. Mycopteris and Ascogrammitis are the
only two grammitid fern genera associated with a black,
clavate fungus (Acrospermum, a bitunicate ascomycete).
The fungus can be found on nearly every plant; however,
it does not appear to harm the plants. The genus name
is derived from the Greek myco-, fungus, + pteris, fern,
referring to the relationship between this fern genus and
the fungus Acrospermum.
LITERATURE: Sundue, M. A., M. Islam & T. A. Ranker. 2010. Systematics of grammitid ferns (Polypodiaceae): using
morphology and plastid sequence data to resolve the circumscription of Melpomene and the polyphyletic genera Lellingeria
and Terpsichore. Systematic Botany 35: 701–715. Sundue, M. A. 2014. Mycopteris, a new neotropical genus of grammitid ferns
(Polypodiaceae). Brittonia 66: 174–185.
416
DESCRIPTION: Principalmente epifítas; raíces
insertadas ventralmente; rizomas dorsiventralses,
reptantes, escamosos, a menudo con brotes y filopodios;
escamas de rizoma anaranjadas a castañas, setosas, las
células túrjidas, lustrosas; hojas con hongos negros
calvados (Acrospermum), monomorfas, setosas por
todas partes, las setas usualmente 1–2 mm de largo,
rojizas; pecíolos con uno o dos haces vasculares, castaños,
a menudo pubescentes; los pelos ca. 0.1mm de largo,
3-celulares, 1-ramificados; ráquises castaños; laminas
próximamente 1-pinnadas, distalmente pinatisectas,
generalmente oblongas con muchos segmentos, enteras;
costas y venas con esclerínquima oscura, esto más visible
abaxialmente; hidatodos presentes, usualmente
cretásicos; sori round; sporangios glabros; esporas
verdes; x=37.
GÉNEROS PARECIDOS: Ascogrammitis difiere por
escamas del rizoma negruzcas clatradas y venas verdes
abaxialmente. Alansmia, Micropolypodium y Terpsichore
difieren por falta del hongo negro clavado y venas
oscuras abaxialmente. Ceradenia y Zygophlebia difieren
por carece de hidatodos.
COMENTARIOS: Mycopteris es completamente
neotropical, desde Mexico hasta Bolivia, las Guianas,
sureste de Brasil y las Antillas. Consta de ca. 30 especies
y crece en los bosques húmedos montanos. El género
tiene dos clados principales: el primero, que incluye
Mycopteris taxifolia, lleva escamas del rizoma con solo
setas apicales, esclerinquima oscura por las costas y
venas y soros mediales. El segundo clado, que incluye M.
semigirsuta y M. alsopteris, tiene filopodios, ausencia de
setas, soros suprmediales y setas del caquis de dos largos
diferentes.Mycopteris y Ascogrammitis son los únicos dos
géneros de helechos grammitidoides asociados con
un hongo negro clavado (Acrospermum, a bitunicate
ascomycete). El hongo se encuentra en cada planta, pero
parece que no daña las plantas. El nombre genérico se
deriva del griego myco-, hongo, + pteris, helecho. Refiere
a la relación entre el género de los helechos y el hongo
Acrospermum.
Figure 205. Mycopteris. A–G Mycopteris attenuatissima (Copel.) Sundue (Sundue & Martin 1041, NY). A. Habit. B. Rhizome
scale. C. Detail of rhizome scale. D. Adaxial lamina. E. Abaxial lamina. F. Detail of setae. G. Detail of setae. H–K Mycopteris
longipilosa Sundue (Sundue & Martin 1037, NY). H. Habit. J. Abaxial lamina. K. Detail sorus. L–O Mycopteris cretata (Maxon)
Sundue (Underwood 3181, NY). L. Habit. M. Detail of rhizome scales. N. Adaxial lamina. O. Abaxial lamina. (Original, ©Robbin
Moran, 2010.)

417
PTERIDACEAE
Myriopteris Fée
DESCRIPTION: Rupestral or terrestrial; rhizomes
compact to long-creeping, scaly, the scales concolorous
or bicolorous with dark central stripe; petioles dark, scaly
and/or pubescent; rachises adaxially terete or flattened
or grooved; blades 2- to 4-pinnate (rarely 1-pinnate-
pinnatifid), adaxially glabrous or pubescent, abaxially
glabrous, pubescent, or scaly; ultimate segments round
to oblong-ovate, minute to rarely > 1 cm long; veins
obscure and not ending in prominent hydathodes;
segment margins usually recurved, with a poorly to
strongly differentiated false indusium; sori usually partly
to completely covered by the recurved segment margins;
sporangia clustered at the vein tips, 64-spored in
sexual species, 32-spored in apomictic ones; spores
globose-tetrahedral, tan to brown; x=29, 30.
DESCRIPCIÓN: Rupestres o terrestres; rizomas
compactos a largamente rastreros, escamosos, las
escamas concoloras a bicoloras con una raya central;
pecíolos oscuros, escamosos y/o pubescentes; ráquises
adaxialmente teretes o planos o surcados; láminas
2- a 4-pinnadas (raras veces pinnadas-pinnatífidas),
adaxialmente glabras o pubescentes, abaxialmente
glabras, pubescentes o escamosas; segmentos últimos
circulares a oblongos-ovados, diminuos a raras veces > 1
cm long; venas oscuras, no terminándose en hidatodos
prominentes; los márgines de los segmentos usualmente
recurvados, con un indusio falso poco a fuertemente
diferenciado; soros usualmente parcialmente a
completamente cubiertos por los márgenes recurvados
de los segmentos; esporangios agrupados en los
ápices de las venas, con 64 esporas en las especies
sexuales y 32 esporas en las apomícticas; esporas
globosas-tetrahédricas, lenonadas a pardas; x=29, 30.

SIMILAR GENERA: Cheilanthes differs by prominent GÉNEROS PARECIDOS: Cheilanthes difiere por
hydathodes and sporangia spread along the veins. hidatodos prominentes y esporangios difundiéndose por
las venas.

COMMENTS: Myriopteris contains 47 species. It occurs COMENTARIOS: Myriopteris consta de 47 especies.

from southern Canada through the Caribbean and
Central America to southern Chile, with one species
endemic to Namibia and South Africa. Mexico is the
center of diversity of the genus; 34 of the 47 species can
be found in Mexico, 7 of which are endemic. Until recently,
most species of Myriopters were classified in Cheilanthes.
The genus, however, is more closely related to Pellaea and
Argyrochosma than to Cheilanthes. A “microphyllous” leaf
morphology is common in the genus and presumed to
be an adaptation to arid habitats. It consists of numerous
rounded or rarely acute ultimate segments. This
morphology has evolved in other cheilanthoid genera.
The genus name is derived from the Greek myrio-, many,
+ pteris, fern, referring to the many (myriad) ultimate
segments of the type species.
Existe desde el sur de Canadá por el Caribe y América
Central hasta el sur de Chile, con una especie (M.
rawsonii) endémica a Namibia y África del Sur. México es
el centro de diversidad del género; 34 de las 47 especies
se pueden encontrar en México, 7 de las cuales son
endémicas. Hasta recién, la mayoría de las especies de
Myriopteris fueron clasificadas en Cheilanthes. El género,
no obstante, es más relacionado a Pellaea y Argyrochosma
que Cheilanthes. Una morfología foliar “microfilia” es
común en el género y presumida ser una adaptación a
ambientes áridos. Consiste de numerosos segmentos
últimos. Esta morfología se han evolucionado en otros
géneros cheilantoides. El nombre genérico se deriva del
griego myrio-, mucho, + pteris, helecho, referienéndose a
los mucho segmentos últimos de la especie tipo.

LITERATURE: Grusz, A. L. & M. D. Windham. 2013. Toward a monophyletic Cheilanthes: the resurrection and
recircumscription of Myriopteris (Pteridaceae). PhytoKeys 32: 49–64. Grusz, A. L., M. D. Windham, G. Yatskievych, L.
Huiet, G. J. Gastony & K. M. Pryer. 2014. Patterns of diversification in the xeric-adapted fern genus Myriopteris (Pteridaceae).
Systematic Botany 39: 698–714. Mickel, J. T. 1979. The fern genus Cheilanthes in the continental United States. Phytologia 41:
431–437.

418
Figure 206. Myriopteris. A M. myriophylla. B–H. M. windhamii. J–O. M. clevelandii. P–V. M. lindheimeri. W–Bb. M.
yavapensis. (From Mickel & Smith, 2004.)

419
BLECHNACEAE
Neoblechnum Gasper & V. A. O. Dittrich
DESCRIPTION: Plants terrestrial; rhizomes erect,
trunk-like, up to 40 cm long, non-stoloniferous, clothed
with black, glossy, acicular, curved, entire scales; fronds
monomorphic; petioles short (<5 cm long), brown to
blackish; laminae oblanceolate, pinnatisect, gradually
reduced basally, with small auricles at the bases,
the distal half pinnatifid with ascending pinnae; buds
absent; aerophores absent; pinnae adnate to rachises,
decurrent, serrulate throughout; veins free, ending at
pinna margins, without hydathodes; sori linear, indusia
entire to subentire; spores smooth; x = 66.
DESCRIPCIÓN: Plantas terrestres; rizomas erectos,
como troncos, hasta 40 cm de largo, no estoloníferos,
vestidos con escamas lúcidas, aciculares, curvadas
enteras; hojas monomorfas; pecíolos cortos (< 5 cm
de largo), pardos a negruzcos; láminas oblanceoladas,
pinnatisectas, gradualmente reducidas hacia la
base, con aurículas pequeñas en las bases, la mitad distal
pinnatífida con pinnas ascendentes; brotes ausentes;
aeróforos ausentes; pinnas adnatas a los ráquises,
decurrentes, serruladas por todas partes; venas
libres, terminándose en los márgines de las pinnas, sin
hidatodos; soros lineales; indusios enteros a subenteros;
esporas lisas; x=66.

SIMILAR GENERA: Blechnum differs by stolons,
and generally smaller leaves (<0.5 m). Austroblechnum
differs by hydathodes and sterile-fertile leaf dimorphy.
Parablechnum differs by 1-pinnate, dimorphic sterile and
fertile leaves, pinnae sessile or short-stalked (at least in
the proximal part of the lamina), and hydathodes.
GÉNEROS PARECIDOS: Blechnum difiere por
estolones y hojas generalmente más pequeñas.
Austroblechnum difiere por todas las pinnas adnatas al ráquis
y hojas estériles y fértiles dimorfas. Parablechnum difiere por
hojas estériles y fértiles dimorfas y 1-pinnadas, pinnas
sésiles o cortamente pecioluladas (al menos en la parte
basal de la lámina) y hidatodos.

COMMENTS: Neoblechnum consists of a single species,
N. brasiliense (Desv.) Gasper & V.A.O. Dittrich, which
is entirely Neotropical, occuring from Guatemala to
southern Brazil. It grows in wet, shaded forest floors.
Often cultivated for the attractive arranged in the forrm
of a basket. The proximal pinnae are often reduced to
small (< 1cm long) obtuse lobes. The genus is sister to two
Old World genera, Doodia and Oceanopteris. The genus
name refers to the exclusively Neotropical occurrence
of the genus.
COMENTARIOS: Neoblechnum consta de una sola
especie, N. brasiliense (Desv.) Gasper & V.A.O. Dittrich,
que es completamente Neotroical, exitiéndose desde
Guatemala hasta Brasil del sur. Existe en sotobosques
sombreados húmedos. Se cultiva con frecuencia por
sus atractivas hojas arregladas en forma de cesta. Las
pinnas proximales son a menudo reducidas hasta
lóbulos pequeñas (< 1 cm). El género es hermano a
dos géneros del Viejo Mundo, Doodia y Oceanopteris. El
nombre genérico refiere a la distribución exclusivamente
Neotropical.

LITERATURE: V. A. O. Dittrich, A. L. Gasper, A. Salino, R. Monteiro & A. L. de Gasper. 2018. The family Blechnaceae
(Polypodiopsida) in Brazil: key to the genera and taxonomic treatment of Austroblechnum, Cranfillia, Lomaridium, Neoblechnum
and Telmatoblechnum for southern and southeastern Brazil. Phytotaxa 303: 1–33. Gasper, A. L. & V. A. O. Dittrich, A. R.
Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and
combinations. Phytotaxa 275: 191–227. Rolleri, C. H. C. Prada, J. M. Gabriel y Galán & L. M. Passarelli. 2013. Especies
arborescentes del género Blechnum (Blechnaceae: Pteridophyta). Revista de Biología Tropical 61: 377–408.

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Figure 207. Neoblechnum brasiliense. Habit from Robinson (1871). Leaf at lright from Santiago et al. (20014)..

421
NEPHROLEPIDIACEAE
Nephrolepis Schott
DESCRIPTION: Terrestrial or epiphytic; stems erect,
usually short, bearing long, slender stolons, scaly, the
scales peltate; sterile and fertile leaves monomorphous;
laminae 1-pinnate, linear to linear-oblong; pinnae
numerous, articulate to the rachis by a circular abscission
zone, usually cleanly deciduous; rachis scales with the
apices pointing downward; veins free, hydathodous,
the hydathodes often white; sori round, terminal on the
first acroscopic vein branch, in a single row between the
costa and margin, medial to submarginal; indusia lunate
to circular, attached by a sinus; spores bilateral, brown;
x=41.
DESCRIPCIÓN: Terrestres o epifitas; tallos erectos,
usualmente cortos, llevando estolones largos, delgados,
escamosos, las escamas peltadas; hojas estériles y fértiles
monomorfas; láminas 1-pinnadas, lineares a lineares-
oblongas; pinnas numerosas, articuladas al raquis por
una zona de abscisión circular, usualmente limpiamente
deciduas; esquamas del raquis con los ápices orientados
hacia abajo; venas libres, con hidatodos, los hidatodos
a menudo blancos; soros redondos, terminales en la
primera rama acroscópica de una vena, en una sola hilera
entre la costa y el margen, mediales a supramediales;
indusios lunatos a circulares, unidos por un seno;
esporas bilaterales, pardas; x=41.

SIMILAR GENERA: Pecluma and Polypodium differ by GÉNEROS PARECIDOS: Pecluma y Polypodium
pinnatisect laminae, non-indusiate sori, and lack stolons. difieren por láminas pinnatisectas, soros no indusiados y
Cyclopeltis semicordata differs by the basal auricle of the falta de estolones. Cyclopeltis semicordata difiere por la
pinna that overlaps the rachis. auricula basal de la pinna que traslapa el raquis.

COMMENTS: Nephrolepis is pantropical and consists of COMENTARIOS: Nephrolepis es pantropical consta de

23 species. Ten species are native to the Neotropics, and 23 especies. Diez especies son nativas al Neotrópico, y una
one (N. brownii (Desv.) Hovencamp & Miyam.) is introduced (N. brownii (Desv.) Hovencamp & Miyam.) es introducida
from tropical Asia.The leaves appear indeterminate; that is, de Asia tropical. Las hojas parecen indeterminadas; es decir,
the apex never completely unfurls. There is almost always los ápices nunca despliegan completamente. Casi siempre
a small fiddlehead. The pinnae fall and leave well-defined hay un pequeño cayado. Las pinnas se cayen y dejan
circular scars on the rachis, which are easily visible to the cicatrices circulares por el raquis, las cuales son visibles a
naked eye.The 1-pinnate laminae are characteristic of the simple vista. Las láminas 1-pinnadas son características del
genus, but horticultural forms are often highly divided género, pero formas horticulturales son a menudo muy
(Morton, 1958).Two species (N. cordifolia (L.) C. Presl and divididas (Morton, 1958). Dos especies (N. cordifolia (L.)
N. undulata (Afzel.) J. Sm.) have stolons that bear scaly C. Presl and N. undulata (Afzel.) J. Sm.) tienen estolones
tubers 1–3 cm long. The tubers contain water, not starch. que llevan tubérculos escamosos 1–3 cm de largo. Los
The rhizome system of Nephrolepis is unusual. It consists tubérculos contienen agua, no almidón. El sistema del
of two parts: a short, erect, poorly developed leaf-bearing rizoma es inusual. Consta de dos partes: una parte
part and a well developed, non-leaf-bearing stoloniferous corta, erecta, pobremente desarrollada y llevando hojas,
part. Unlike other fern rhizomes, the erect rhizomes of y una parte estolonífera bien desarrollada, no portando
Nephrolepis do not produce roots; they produce only hojas. A diferencia de que otros rizomas, los erectos de
stolons, and only the stolons produce the plants’ roots. Nephrolepis no producen raíces; solo estolónes, y estos
The stolons are further unusual by not bearing leaves producen las raíces de la planta. Además, los estolónes
as stolons usually do. The genus name comes from the no llevan hojas. El nombre genérico se deriva del griego
Greek nephros, kidney, and lepis, scale, referring to the nephros, riñón, y lepis, escama, refiriéndose al indusio con
kidney-bean shaped indusia. la forma de una faba.
LITERATURE: Hennequin, S., P. Hovencamp, M. J. M. Christenhusz & H. Schneider. 2010. Phylogenetics and biogeography of Nephrolepis – a
tale of old settlers and young tramps. Botanical Journal of the Linnean Society 164: 113–127. Hovencamp, P. H. & F. Miyamoto. 2005. A conspectus of the
native and naturalized species of Nephrolepis (Nephrolepidaceae) in the world. Blumea 50: 279–322. Morton, C. V. 1958. Observations on cultivated ferns,
V. The species and forms of Nephrolepis. Amer. Fern J. 48: 18–27. Nauman, C. E. 1981. The genus Nephrolepis in Florida. Amer. Fern J. 71: 65–70. Rojas-
Alvarado, A. F. 2008. The Nephrolepis pendula complex (Lomariopsidaceae) in the Neotropics. Métodos en Ecología y Sistemática 3(Supl. 1): 54–78.

422
Figure 208. A–C. Nephrolepis undulata. D–F. N. brownii. G–L. N. pectinata. M–O. N. rivularis. (From Mickel & Beitel, 1988.)

423
POLYPODIACEAE
Niphidium J. Sm.
DESCRIPTION: Epiphytic, terrestrial, or rupestral;
stems short- to long-creeping, often covered by a dense
mat of pubescent roots, scaly, the scales clathrate (at
least medially); sterile and fertile leaves monomorphous;
petioles articulate to the stem, the phyllopodia 0-1 cm
long, often darker than the petioles; laminae simple,
entire, coriaceous, narrowly elliptic to narrowly
oblanceolate, often glaucous beneath; rachis prominent;
venation of prominent lateral veins, these connected by
inconspicuous veinlets that anastomose to form irregular
areoles containing excurrent and recurrent veinlets; sori
in one row between the main lateral veins; spores
yellowish or hyaline, reniform; x=37.
SIMILAR GENERA: Campyloneurum differs by the
several rows of sori between the main lateral veins.
Microgramma and Pleopeltis, two other genera of
Polypodiaceae with simple, entire leaves, differ by the
single row of sori between the rachis and margin.
COMMENTS: Niphidium is completely neotropical and
contains about 10 species. It is most closely related to
Campyloneurum and Microgramma. Several species have
leaves that form a basket or nest. These leaves trap
organic material that eventually decomposes and forms
humus into which the roots grow. Niphidium nidiforme,
a high-canopy epiphyte endemic to Costa Rica, has
rhizomes that creep horizontally around trunks, and
leaves arranged in a nest-like manner to catch falling
organic matter. Many species of Niphidium have veins that
end in whitish hydathodes. The genus name is derived
from the Greek nipha, snow + eidos, like. The fronds of
the type species bear dense white indument.
LITERATURE: Lellinger, D. B. 1972. A revision of the fern genus Niphidium. American Fern Journal 62: 101–120.
424
DESCRIPCIÓN: Epífitas, terrestres o rupestres; tallos
cortamente o largamente rastreros, a menudo cubiertos
por un denso tapete de raíces pelosas, escamosos, las
escamas clatradas (al menos en la mitad); hojas estériles
y fértiles monomorfas; pecíolos articulados al tallo, los
filopódios 0-1 cm de largo, a menudo más oscuras
que el pecíolo; láminas simples, enteras, coriáceas,
angostamente elípticas a anogostamente oblanceoladas,
a menudo glauca en el envés; raquis prominente;
nervadura de venas laterales prominentes, conectadas
por venillas inconspicuas que se anastomosan formando
aréolas irregulares conteniendo venillas excurrentes
y recurrentes; soros en una hilera entre las venas
laterales principales; esporas amarillas o hialinas,
reniformes; x=37.
GÉNEROS PARECIDOS: Campyloneurum difiere
por varias series de soros entre las venas principales
laterales. Microgramma y Pleopeltis, dos otros géneros de
Polypodiaceae con hojas simples, enteras, difieren por la
única hilera de soros entre el raquis y el margen.
COMENTARIOS: Niphidium es completamente
neotropical y consta de casi 10 especies. Niphidium está
más cercanamente emparentado a Campyloneurum y
Microgramma. Varias especies tienen hojas que forman
una canasta o nido. Estas hojas atrapan materia orgánica
la cual, eventualmente, se descompone integrándose
al suelo dentro de la cual las raíces crecen. Niphidium
nidiforme, un epífito de los doseles y endémico a Costa
Rica, tiene rizomas que serpean horizontalmente
alrededor de troncos, y hojas arregladas como un nido
de manera que pueden recoger matéria orgánica caida.
Muchas especies de Niphidium tienen venas que terminan
en hidatodos blanquecinos. El nombre genérico se deriva
del griego nipha, show + eidos, parecido. Las hojas de la
especie tipo lleva un indumento denso blanquecino.
Figure 209. Niphidium crassifolium. (From Mickel & Smith, 2004.)

425
PTERIDACEAE
Notholaena R. Br.
DESCRIPTION: Terrestrial; rhizomes erect to short-
creeping, scaly; leaves monomorphic; petioles with 1
vascular bundle, light brown to blackish; laminae 1- or
2-pinnate-pinnatifid, coraiceous, with the apex pinnatifid;
pinnae white or yellow farinose on the lower
surface, the pinnules sessile or adnate, often scaly, with
or without hairs; rachises tan to light brown, glabrous or
puerulent or farinose; veins free, with two-celled, gland-
tipped hairs; veins free, ending in slightly clavate tips; sori
intramarginal, terminal on the vein tips; false indusia
absent or weakly recurved; x=30.
DESCRIPCIÓN: Terrestres; rizomas brevemente
rastreros a erectos, escamosos; hojas monomorfas;
pecíolos con un haz vascular, pardo-amarillentos a
negruzcos; láminas 1-2-pinnado-pinnatífidas, coriáceas,
con el ápice pinnatífido; pinnas blancas o amarillas
farinosas en el envés, las pínnulas sésiles o adnatas,
a menudo escamosas y/o pilosas; ráquises pardo-
amarillentos a negruzcos, glabros a puberulentos o
farinosos; nervaduras libres, terminando en puntas un
tanto claviformes; soros intramarginales, terminales en
las nervaduras; indusios falsos ausentes o débilmente
recurvados; x=30.

SIMILAR GENERA: Cheilanthes differs by nonfarinose
laminae and strongly developed false indusia. Argyrochosma
differs by stalked or sessile (not adnate) pinnules, which
are usually entire, elliptic or oblong unlike those typically
of Notholaena.
GÉNEROS PARECIDOS: Cheilanthes difiere por
láminas no farinosas y indusios falsos más fuertemente
desarollados. Argyrochosma tiene también la lámina
farinosa en el envés pero difiere por las pínnulas
pediceladas o sésiles (no adnatas). Usualmente tiene
pínnulas enteras, elipticas o oblongas, a diferencia de las
encontrado típicamente en Notholaena.

COMMENTS: As treated here in the strict sense,
Notholaena contains about 30 species and is entirely
neotropical. Its center of diversity is the Chihuahua
Desert of northern Mexico. The species occur mainly
in seasonally dry habitats. Molecular data show that
Notholaena is monophyletic and sister to Cheiloplecton
(Gastony & Rollo, 1998; Rothfels et al., 2008). Some species
previously placed in the genus have been transferred to
Cheilanthes or Argyrochosma.The genus name is derived
from the Greek nothos, false + chlaena, cloak. The blade
margin is not reflexed as in Cheilanthes.
COMENTARIOS: Como tratado aquí en un sentido
estricto, Notholaena contiene casi 30 especies e es
completamente neotropical. Su centro de diversidad
es el Desierto Chihuahuense de México norteño.
Las especies existen principalmente en ambientes
secas estacionales. Datos moleculares muestran que
Notholaena es monofilético y cercanamente relacionado
con Cheiloplecton (Gastony & Rollo, 1998; Rothfels et al.,
2008). Algunas especies ubicadas anteriormente en el
género han sido trasferidas a Cheilanthes o Argyrochosma.
El nombre del género se deriva del griego nothos, falso
+ chlaena, capa. El margen de la lámina no es recurvado
como en Cheilanthes.

LITERATURE: Gastony, G. J. & D. R. Rollo. 1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern
United States and adjacent Mexico–a molecular phylogenetic reassessment of generic lines. Aliso 17: 131–144. Rothfels, C. J., M.
D. Windham, A. L. Grusz, G. J. Gastony & K. M. Pryer. 2008. Toward a monophyletic Notholaena (Pteridaceae): resolving
patterns of evolutionary convergence in xeric-adapted ferns. Taxon 57: 712–724. Tryon, R. M. 1956. A revision of the American
species of Notholaena. Contributions of the Gray Herbarium of Harvard University 179: 1–106. Yatskievych, G. & A. R. Smith.
2003. Typification of Notholaena R. Br. (Pteridaceae). Taxon 52: 331–336.

426
Figure 210. Notholaena. A–C. N. standleyi. E–C. N. sulphurea. H–K. N. affinis. L–N. N. brachycaulis. (From Mickel & Smith,
2004.)

427
LINDSAEACEAE
Odontosoria Fée
DESCRIPTION: Terrestrial; rhizomes short-creeping,
scaly; leaves scandent, up to 6 m long, glabrous; petioles
and rachises spiny; laminae 3- to 5-pinnate, pinnae
slightly reflexed; costae and costules flexuose; veins free;
sori marginal, solitary, ovate, supplied by 1-3 veins;
indusium double, united by the base and the sides, the
adaxial green like the laminar tissue, the abaxial lighter
colored; spores trilete; x=96.
DESCRIPCIÓN: Terrestres; rizomas cortamente
rastreros, escamosos; hojas escandentes, hasta 6 m
de largo, glabras; pecíolos y ráquises espinosos; láminas
3- a 5-pinnadas; pinnas escasamente reflejas; costas y
costillas medias flexuosas; venas libres; soros marginales,
solitarios, ovados, abastecidos por 1-3 nervaduras; indusio
doble, unido por la base y los lados, el adaxial verde como
el tejido laminar, el abaxial claro; esporas triletes; x=96.

SIMILAR GENERA: Eriosorus differs by pubescent GÉNEROS PARECIDOS: Eriosorus difiere por láminas
laminae and non-indusiate sori that extend along the pubescentes y soros no indusiados que extienden por
veins. Sphenomeris differs by pubescent rhizomes and las venas. Sphenomeris difiere por rizomas pubescentes y
smaller, determinate leaves. hojas más pequenas, determinadas.

COMMENTS: Odontosoria is pantropical and consists of COMENTARIOS: Odontosoria es pantropical y consta

22 species, 12 of which occur in the Neotropics. It usually
occurs in open places such as slopes and roadsides. Its
leaves exhibit indeterminate growth (the apex always
unfurling) and use surrounded vegetation for support.This
is an adaptation of the scandent habit can also be seen
in Blotiella, Histiopteris, certain species of Hypolepis, and
two species of Eriosorus (E. flexuosus and E. glaberrimus).
The genus name is derived from the Greek odous, tooth
+ soros, sorus. In the type species, the sori are at the end
of segment teeth.
de 22 especies, 12 de las cuales existen en el Neotrópico.
Típicamente existe en lugares abiertos, como taludes y
los lados de los caminos. Sus hojas exhiben crecimiento
indeterminado (el ápice siempre desplegándose) y usan
la vegetación circundante como apoyo. Esta adaptación
del hábito escandente se puede ver también en Blotiella,
Histiopteris, ciertas especies de Hypolepis y dos especies
de Eriosorus (E. flexuosus and E. glaberrimus). El nombre
genérico se deriva del griego odous, diente + soros, soro.
En la especie tipo, los soros están en los ápices de dientes
de los segmentos.

LITERATURE: Caluff, M. G. 2006. The genus Odontosoria (Dennstaedtiaceae, Pteridophyta) in Cuba. Willdenowia 36: 469–
478. Maxon, W. R. 1913. Studies of tropical American ferns. No. 4. Contributions from the United States National Herbarium
17: 133–179. Ranker T. A., C. E. D Gemmill & P. G. Trapp. 2000. Microevolutionary patterns and processes of the native
Hawaiian colonizing fern Odontosoria chinensis (Lindsaeaceae). Evolution 54: 828–839.

428
Figure 211. A–C. Odontosoria aculeata. D–F. O. scandens. (Courtesy of Bobbi Angell.)

429
OLEANDRACEAE
Oleandra Cav.
DESCRIPTION: Epiphytic, hemiepiphytic(?) o terrestrial;
rhizomes long-creeping, densely scaly, the scales peltate,
non-clathrate; sterile and fertile leaves monomorphous;
petioles articulate to phyllopodia; laminae simple,
entire, the margins slightly cartilaginous; veins free,
long-parallel; sori round, often borne close to the rachis,
indusiate, the indusia reniform, peltate or attached at
a narrow sinus; spores monolete, bilateral, non-green,
perispores winged, cristate, or spiny; x=41.
DESCRIPCIÓN: Epífitas, hemiepífitas(¿), o terrestres;
rizomas largamente rastreros, densamente
escamosos, las escamas peltadas, no clatradas; hojas
estériles y fértiles monomorfas; pecíolos articulados a
filopodios; láminas simples y enteras, los márgenes
ligeramente cartilaginosos; venas libres, largamente
paralelas; soros redondos, a menudo nacen cerca del
raquis, indusiados, los indusios reniformes, peltados
o unidos en un seno estrecho; esporas monoletes,
bilaterales, no verdes; perisporios alados, cristados o
espinosos; x=41.

SIMILAR GENERA: Elaphoglossum differs by having non-
peltate rhizome scales, ascending veins, and acrostichoid
sori. Its phyllopodia are generally short and darker than
the petioles. Various Polypodiaceae with simple, entire
leaves (e.g., Campyloneuron, Microgramma) differ by net
veins.
GÉNEROS PARECIDOS: Elaphoglossum difiere
por tener escamas del rizoma no peltadas, venas
ascendentes y soros acrosticoides. Sus filopodios son
generalmente cortos y más oscuros que los pecíolos.
Varios Polypodiaceae con hojas simples y enteras (e.g.,
Campyloneuron, Microgramma) difieren por venas en
forma de red.

COMMENTS: Oleandra is pantropical with about 8
species in the Neotropics and 30 in the Paleotropics. It
grows in wet forests. In the Neotropics it often forms
dense colonies in the canopy. The articulation between
the petiole and phyllopodium is slightly swollen. The
articulations are functional, with the leaves falling away
cleanly. Some species with appressed rhizome scales often
have a shrubby habit.Their leaves can be clustered in loose
pseudowhorls along the rhizome, indicating rhythmic
growth. Presumably the leaf primordia are initiated
regularly around the apical meristem, but differential
internode elongation causes the pseudowhorls. The
species with spreading rhizome scales are apparently
hemiepiphytes, starting on trunk bases and climbing into
the canopy. This needs confirmation. The genus name
is derived from an allusion of the leaves to those of
Oleander (Nerium, Apocynaceae).
COMENTARIOS: Oleandra es pantropical con casi 8
especies en el Neotrópico y 30 en el Paleotrópico. Existe
en de bosques húmedos. En el Neotrópico a menudo
forma colonias densas en el dosel. La articulación
entre el pecíolo y filopodio es un poco hinchada. Las
articulaciones son funcionales, con las hojas cayéndose
limpiamente. Algunas especies con escamas del rizoma
apresas a menudo tienen un hábito arbustivo. Sus hojas
pueden ser agrupadas por el rizoma en pseudoverticelas
flojas, indicando crecimiento rítmico. Presumablemente
las primordias foliares están producidas regularmente
alrededor del meristemo apical, pero elongación
diferencial de los entrenudos causan los pseudoverticilos.
Las especies con escamas del rizoma divergentes son
aparentemente hemiepífitas, empezando en las bases
de troncos y trepando hasta el dosel. Esto se necesita
confirmación. El nombre genérico se deriva de una alusión
de las hojas a ellas de Oleander (Nerium, Apocynaceae).

LITERATURE: Maxon, W. R. 1914. The American species of Oleandra. Contributions from the United States National
Herbarium 17: 392–398. Scamman, E. 1961. The genus Oleandra of Costa Rica. Rhodora 63: 335–340. Schwartsburd, P. B.,
C. V. Miranda & J. Prado. 2016. Oleandra (Oleandraceae) in the Brazilian Atlantic Forest. American Fern Journal 106: 191–205.
Tryon, R. M. 1997. Systematic notes on Oleandra. Rhodora 99: 335–343. Tryon, R. M. 1997. Systematic notes on the Old
World fern genus Oleandra. Rhodora 102: 428–438.

430
Figure 212. Oleandra articulata. (Courtesy of Bobbi Angell.)

431
DRYOPTERIDACEAE
Olfersia Raddi
DESCRIPTION: Plants terrestrial or low-climbing;
stem short-creeping, in cross-section with the meristeles
arranged irregularly and each one not surrounded by a
dark sclerenchymatous shealth; sterile and fertile leaves
strongly dimorphic; sterile lamina 1-pinnate with a
conform terminal pinna; pinnae ovate to lanceolate,
excavate on the basiscopic side, entire; veins forking near
or at the base, parallel, connected by a submarginal
vein (this sometimes difficult to see in dried material);
rachis and costae not or only shallowly grooved, glabrous;
fertile leaves 2-pinnate, rarely 1-pinnate; sori nonindusiate;
spores monolete, echinulate; x=41.
SIMILAR GENERA: Bolbitis differs by having (usually)
reticulate veins and an elongate ventral meristele (seen
when the rhizome is cut in cross section). Polybotrya differs
by venation (various patterns but never long-parallel like
Olfersia), pinnatifid apex of the lamina, pubescent adaxial
grooves of the rachis, and dark sclerenchymatous sheaths
surrounding the meristeles.
COMMENTS: Olfersia consists of three species restricted
to the neotropics: O. alata Sánchez (eastern Cuba) and
O. cervina (L.) Kunze (S. Mexico to Panama, the Antilles,
and French Guiana to Bolivia), and O. corcovadensis Kaulf.
ex Raddi (se. Brazil). The fertile leaves are taller and
more erect than the sterile ones and soon wither after
the spores are shed. The connecting veins of the sterile
pinnae is absolutely diagnostic for the genus.
The generic name honors Ignaz Franz Werner Maria
von Olfers (1793–1871), a German naturalist, historian,
and diplomat who collected in Brazil ca. 1816.
LITERATURE: Moran, R. C. 1986. The neotropical fern genus Olfersia. American Fern Journal 76: 161–178. Sánchez, V.,
G. Caluff & Z. Pérez. 1991. Nueva especie cubana del género Olfersia (Polypodiaceae - Dryopteridoideae). Fontqueria 31:
229–233.
432
DESCRIPCIÓN: Plantas terrestres o bajo-reptantes;
tallo corto-reptante, en sección transversal con las
meristelas arregladas irregularmente y cada uno sin vaina
de esclerénquima oscuro; hojas estériles y fértiles
fuertamente dimórficas; lámina estéril con una
pinna terminal conforme; pinnas ovadas a lanceoladas,
excavadas en el lado basiscópico, enteras; venas furcadas
cerca de la base, paralelas, conectadas por una vena
submarginal (ésta a veces difícil de ver en ejemplares
secos); raquis y costas sin surcos adaxialmente, glabros;
hojas fértiles 2-pinnadas, raramente 1-pinnadas; soros sin
indusios; esporas monoletes, equinulatas; x=41.
GÉNEROS PARECIDOS: Bolbitis difiere por tener
venas reticuladas y una meristela ventral elongada (visto
cuando el rizoma se corta en sección transversal).
Polybotrya difiere por la venación (varios patrones pero
nunca es largo-paralela como Olfersia), el ápice pinnatífido
de la lámina, el surco adaxial del raquis pubescente, y el
patrón del tallo en sección transversal.
COMENTARIOS: Olfersia consiste de tres especies
restringidas al neotrópico: O. alata Sánchez (Cuba oriental)
y O. cervina (L.) Kunze (S. México a Panamá, las Antillas,
Guyana Francesa a Bolivia) y O. corcovadensis Kaulf. ex
Raddi (se. Brazil). Las hojas fértiles son más altas y rectas
que las estériles y se marchitan apenas después de que
las esporas son lanzadas. La vena colectora de las pinnas
estériles es absolutamente diagnóstica para el género.
El nombre génerico honora Ignaz Franz Werner Maria
von Olfers (1793–1871), una naturalista, historiador y
diplomático alemán quien colectó en Brasil ca. 1816.
Figure 213. Olfersia cervina (L.) Kunze. A. Sterile leaf (hoja estéril). B. Petiole cross section showing arrangement of vascular
bundles typical of Eupolypods I. C. Rachis-costa juncture, adaxial view. D, E. Precociously fertile juvenile plants. F. Sterile pinna;
note intramarginal connecting vein. (From Moran, 1986.)

433
ONOCLEACEAE
Onoclea L.
DESCRIPTION: Terrestrial; rhizomes creeping, scaly,
stolons absent; sterile and fertile leaves dimorphic;
sterile leaves 15-50 x 15-35 cm; petioles with 2 vascular
bundles; sterile blades 1-pinnate, widest at the base
or nearly so, apices pinnatifid; sterile pinnae sessile and
becoming adnate distally, 5-11 pairs, opposite toward
the base of the blade, entire to shallowly lobed, bases
cuneate, apices acuminate, margins serrulate; veins
netted, without included veinlets; fertile leaves
2-pinnate, 30-50 cm long, erect, green but turning brown
and indurate at maturity; fertile pinnae 5-11 pairs, strongly
ascending; pinnules enrolled and forming a globose
structure enclosing each sorus; receptacle prominent;
paraphyses absent; indusia absent or present as a thin
hyaline scale; spores bilateral, greenish; x=37.
SIMILAR GENERA: Onocleopsis, of southern Mexico
and Guatemala, differs by erect rhizomes, sterile blades
reduced toward the base, sterile pinnae truncate or
subcordate, and fertile leaves 2-3-pinnate. Woodwardia
aerolata, with which this species often grows in North
America, differs by serrate pinnae that are alternate
throughout.
COMMENTS: Onoclea is composed of a single species,
O. sensibilis L. It occurs in temperate regions of eastern
North America and eastern Asia, usually in wet habitats
such as swamps and marshes, in either sun or shade. It
often forms dense colonies from its creeping rhizome.
Onoclea stores starch in the thick, wide, petiole bases that
persist after the above part of the leaf has senesced. The
fertile leaves persist throughout the winter and release
their spores the following late winter or early spring.
Onoclea is sister to a clade consisting of Onocleopsis and
Matteuccia (Gastony & Ungerer, 1997). These genera,
plus the Asian Pentarhizidium (2 species) constitute the
Onocleaceae.The genus name is derived from the Greek,
onos, vessel, + kleiein, to close. The fertile pinnules roll up
into bead-like segments enclosing the sori.
LITERATURE: Lloyd, R. M. 1971. Systematics of the onocleoid ferns. University of California Publications in Botany 61: 1–86.
Gastony, G. J. & M. C. Ungerer. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae:
Onocleae). American Journal of Botany 84: 840–849.
434
DESCRIPCIÓN: Terrestres; rizomas reptantes,
escamosos estolones ausentes; hojas estériles y fértiles
dimorfas; hojas estériles 15-50 x 15-35 cm; pecíolos
con 2 haces vasculares; láminas estériles 1-pinnadas, más
anchas en hacia la base o casi así, ápices pinnatífidos;
pinnas estériles séssiles y llegando a ser adnatas
distalmente, 5-11 pares, opuestas en la base de la lámina,
bases cuneadas, ápices acuminados, márgines serrulados;
venas en forma de red, sin venillas incluidas; hojas
fértiles 2-pinnadas, 30-50 cm de largo, erectas, verdes
pero tornándose pardas y induratas con maduréz; pínulas
fértiles enrolladas y formando una estructura globosa
encapsulando cada soro; receptáculo prominente;
paraphyses ausentes; indusios ausentes o presentes
como una escama delgada hialina para cada soro; esporas
bilaterales, verdosas; x=37
GÉNEROS PARECIDOS: Onocleopsis, del sur de
México y Guatemala, difiere por rizomas erectos,
láminas estériles pinatífidas reducidas hacia la base,
pinnas estériles truncadas o subcordatas, y hojas fértiles
2-3-pinnadas. Woodwardia aerolata, que crece con esta
especie a menudo en América del Norte, difiere por
pinnas serratas que son alternas por todo la lámina.
COMENTARIOS: Onoclea consta de una sola especie,
O. sensibilis L. Ocurre en zonas templadas de América del
Norte oriental y Asia oriental, usualmente en ambientes
húmedos como pantanos y humedales, en el sol o
sombra. A menudo forma colonias densas por su rizoma
reptante. Onoclea alménese almidón en las bases gruesos
y anchos de los pecíolos que persistan después de lo
demás de la hoja ha caída. Las hojas fértiles persistan por
el invierno y libran sus esporas en el invierno siguiente
o la primavera temprano. Onoclea es hermana a un
clado formado por Onocleopsis y Matteuccia (Gastony &
Ungerer, 1997). Estos géneros, y el asiático Pentarhizidium
(2 especies) constituyen las Onocleaceae. El nombre
génerico se deriva del griego onos, vasija, + kleiein, cerrar.
Las pinulas fértiles se enrollan en segmentos redondeados
envolviendo de los soros.
Figure 214. Onoclea sensibilis L. (Original, ©Robbin Moran, 2016.)

435
ONOCLEACEAE
Onocleopsis F. Ballard
DESCRIPTION: Terrestrial; rhizomes erect, stout, often
stoloniferous from bases; rhizome scales concolorous,
brown, lustrous; sterile and fertile leaves dimorphic;
sterile leaves 50-200 x 25-35 cm; sterile blades
1-pinnate, reduced toward the base, apices pinnatifid;
sterile pinnae sessile, 17-28 pairs, bases truncate or
subcordate, apices acuminate, margins broadly serrate;
blades glabrous adaxially, short-pilose on veins and costae
abaxially; veins netted and without included veinlets;
fertile leaves 2-3-pinnate, 56-120 cm long, with laminar
tissue nearly absent; fertile pinnules rolled into 3 or 4
globular divisions, brown at maturity, ultimate segments
with 2 sori; indusia absent or present as a thin hyaline
scale for each sorus; spores bilateral; x=?
DESCRIPCIÓN: Terrestres; rizomas erectos, gruesos,
a menudo estolóniferos desde las bases; escamas del
rizoma concoloras, pardas, lustrosas; hojas estériles y
fértiles dimorfas; hojas estériles 50-200 x 25-35 cm;
láminas estériles 1-pinnadas, reducidas hacia la
base, ápices pinnatífidos; pinnas estériles séssiles, 17-28
pares, bases truncadas o subcordatas, ápices acuminados,
márgines anchamente serrados; láminas glabras en el haz,
cortamente pilosas por las venas y costas en el envés;
venas en forma de red y sin venillas incluidas; hojas
fértiles 2-3-pinnadas, 56-120 cm de largo, con tejido
laminar casi ausente; pínulas fértiles enrolladas en 3 o
4 divisiones globulares, pardo en madurez, segmentos
últimos con 2 soros; indusios ausentes o presentes como
una escama delgada hialina para cada soro; esporas
bilaterales; x=?

SIMILAR GENERA: Onoclea differs by creeping
rhizomes and pinnatifid, triangular blades (widest at base).
Matteuccia differs by free veins and 1-pinnate-pinnatifid
sterile fronds reduced basally to vestigial pinnae. Onoclea
and Matteuccia are north-temperate genera, whereas
Onocleopsis is tropical.
GÉNEROS PARECIDOS: Onoclea difiere por rizomas
reptantes y láminas pinatífidas triangulares (más anchas
en la base). Matteuccia difiere por venas libres y hojas
estériles 1-pinado-pinatífido y reducidos hacia la base a
pinas vestigiales. Onoclea y Matteuccia son de las zonas
templadas norteñas, mientras que Onocleopsis es tropical.

COMMENTS: Consisting of only one species, Onocleopsis
hintonii F. Ballard, occurring from central Mexico to
Guatemala, usually along streams in forests from 1500–
2500 m. Onocleopsis is the sister genus to Matteuccia
(Gastony & Ungerer, 1997). Both share erect rhizome that
are stoloniferous at the base, basally reduced laminae, and
the loss of gametophytic trichomes. The genus name is
derived from Onoclea + the Greek suffix –opsis, like.
COMENTARIOS: Consistiéndose de una sola especie,
Onocleopsis hintonii F. Ballard, ocurriéndose México
central hasta Guatemala de 1500–2500 m, usualmente
por riachuelos en bosques. Onocleopsis es el género
hermana a Matteuccia (Gastony & Ungerer, 1997). Los
dos comparten rizomas erectos que son estoloníferos
en la base, láminas reducidas basalmente, y la perdida de
pelos en el gametofito. El nombre genérico se deriva de
Onoclea + el sufijo griego –opsis, parecido.

LITERATURE: Ballard, F. 1945. A new fern genus from Mexico and Guatemala. American Fern Journal 35: 1–3. Ballard, F.
1948. Further notes on Onocleopsis. American Fern Journal 38: 125–132. Lloyd, R. M. 1971. Systematics of the onocleoid ferns.
University of California Publications in Botany 61: 1–86. Gastony, G. J. & M. C. Ungerer. 1997. Molecular systematics and a
revised taxonomy of the onocleoid ferns (Dryopteridaceae: Onocleeae). American Journal Botany 84: 840–849.

436
Figure 215. Onocleopsis hintonii F. Ballard. (From Mickel & Smith, 2004.)

437
OPHIOGLOSSACEAE
Ophioglossum L.
DESCRIPTION: Terrestrial, rarely epiphytic, fleshy,
completely lacking sclerenchyma; roots in most species
proliferous and forming clones, lacking root hairs; stems
erect, subterranean; leaves erect, conduplicate in bud
(not circinate); sterile lamina lacking a costa, simple,
entire, most species cordate to oblanceolate, often
mucronate; veins areolate, usually with free included
veilets; fertile spikes borne at the base of the sterile blade,
with two lateral rows of sporangia, usually apiculate;
sporangia opening by a transverse slit; x=30.
SIMILAR GENERA: Anemia (Schizaeaceae) differs
by two fertile spikes and pinnately divided leaves.
Cheiroglossa palmata (L.) C. Presl differs by epiphytic habit
and palmately lobed leaves.
\
COMMENTS: Ophioglossum is cosmopolitan and
consists of about 30 species. It typically occurs in open
or semi-open habitats, from sea level up to páramos.
The plants often form extensive colonies by proliferous
roots, but because of the plants’ small size (usually less
than 15 cm tall), their colonies are easily overlooked. The
lamina represents a rachis expanded by intercalary (not
marginal) growth. Therefore, it lacks a costa. This is similar
to what occurs in Plantago. Ophioglossum reticulatum
has the highest chromosome number of any living thing:
2n=1262. Higher numbers have been reported in the
genus but are undocumented by photos and therefore
not accepted here). The genus name is derived from
the Greek ophis, snake + glossa, tongue. The fertile spike
resembles a snake’s tongue.
LITERATURE: Clausen, R. T. 1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19(2): 1–177.
Hauk, W. D., C. R. Parks & M. W. Chase. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL and trnL–F
plastid DNA sequences and morphology. Molecular Phylogenetics and Evolution 28: 131–151..
438
DESCRIPCIÓN: Terrestres, raras veces epífitas,
carnosas, completamente sin esclerénquima; raíces en
la mayoría de las especies prolíferas y formando clones,
sin pelos sobre las raíces; tallos rectos, subterráneos;
hojas rectas, conduplicadas en la yema (no circinadas);
láminas estériles sin costa, simples, enteras, cordadas
a oblanceoladas (raramente lineales), a menudo
mucronadas; venas areoladas, usualmente con venillas
incluídas libres; espigas fértiles nacidas en la base de la
lámina estéril, con dos hileras laterales de esporangios,
generalmente apiculadas; esporangios se abren por una
dehiscencia transversal; x=30.
GÉNEROS PARECIDOS: Anemia (Schizaeaceae)
difiere por dos espigas fértiles y hojas divididas
pinnátamente. Cheiroglossa palmata (L.) C. Presl difiere
por hábito epífito con hajas lobuladas palmadamente.
COMENTARIOS: Ophioglossum cosmopólito y consta
de casi 30 especies.Típicamente existe en hábitats abiertos
a semi-abiertos, desde el nivel del mar hasta páramos. Las
plantas a menudo forman colonias grandes por sus raíces
prolíferas, pero a causa del tamaño pequeño de las plantas
(usualmente menos de 15 cm de largo), sus colonias
son fácilmente pasado por alto. La lámina representa
un ráquis expandido por crecimiento intercalario (no
marginal). Por eso, no presenta una costa. Esto es similar a
lo que pasa en Plantago. Ophioglossum reticulatum tiene el
número cromosómico más alto de cualquier ser viviente:
2n=1262. Números más altos han sido reportados para
el género pero son indocumentados por fotos y por eso
no aceptados aquí. El nombre genérico se deriva del
griego ophis, serpiente + glossa, lengua. La espiga fértil se
asemeja a una lengua de serpiente.
Figure 216. A. Ophioglossum engelmannii Prantl B. O. reticulatum L. C. O. nudicaule L.f. (note proliferous root; nótase raíz
prolifero). D. O. crotalophoroides Walt. (Original, ©Robbin Moran, 1992.)

439
THELYPTERIDACEAE
Oreopteris Holub

DESCRIPTION: Rhizomes erect, often hidden by old
petiole bases, the scales 4–15 mm long, thin, with many
cells bearing short spherical outgrowths, acicular hairs
lacking; leaves 1-pinnate-pinnatifid, gradually narrowed
basally, the basal pinna very short; petioles densely
scaly; pinnae deeply lobed; veins free, simple or forked,
the lower ones ending before the margin, the distal ones
running to the margin, ending above the base of the sinus;
laminae abaxially glandular, the glands sessile, spherical;
sori usually near the margin; indusia often glandular;
sporangia occasionally glandular near the annulus and on
the stalk; x=34.
DESCRIPCIÓN: Rizomas erectos, a menudo oculto
por las bases peciolares viejos, las escamas 4–15
mm de largo, delgadas con muchos células llevando
protuberáncias cortas esféricas,faltando de pelos aciculares;
hojas 1-pinnado-pinnatífidas, gradualmente reducidas
basalmente, las basales muy cortas; peciolos
densamente escamosos; pinnas profúndamente
lobuladas; venas libres, simples o furcados, las basales
terminándose antes del margen, los distales alcanzando
el margen, terminando arriba de la base del sinus; láminas
abaxialmente glandulares, las glándulas séssiles; esféricas;
soros usualmente cerca del margen; indusios a menudo
glandulares; esporángios occasionalmente glandulares
cerca del anillo y en el pecíolo; x=34.

SIMILAR GENERA: Thelypteris differs by creeping
rhizomes, laminae broadest at or near the base, margins
of the fertile segments recurved (not flat), and smaller
(1–2 mm long) scales on the rhzomes and petioles.
GÉNEROS PARECIDOS: Thelypteris difiere por
rizomas rastreros, láminas mas anchas en o cerca del
base, márgenes de los segmentos fértiles recurvados (no
planos) y escamas mas pequeñas (1–2 mm long) en los
rizomas y pecíolos.

COMMENTS: Oreopteris consists of three species and
occurs in North America and Eurasia. The only North
American species is O. quelpaertensis (Christ) Holub,
which occurs primarily along the northwest Pacific coast,
but a single collection is known from the eastern side
of the continent in Newfoundland. This species grows in
open, rocky woods and subalpine medows in acid soils.
Oreopteris limbosperma (Allioni) Holub occurs in Europe,
and O. elwesii (Hook. & Baker) Holttum in Asia. The
former species is the type of the genus. The genus name
is derived from the Greek oreos, mountain + pteris, fern.
A fern of the mountains..
COMENTARIOS: Oreopteris consta de 3 especies y
existe en América del Norte e Eurasia. La única especie
norteamericana es O. quelpaertensis (Christ) Holub, que
existe principalmente en la costa del noroeste Pacífico,
pero un una sola colecta conocida del lado oriental del
continente en Terranova. Esta especie occure en bosques
abiertos rocosos y prados subalpinos en suelos ácidos.
Oreopteris limbosperma (All.) Holub existe en Europa y
O. elwesii (Hook. & Baker) Holttum in Asia. La especie
anterior es el tipo del género. El nombre genérico se
deriva del griego oreos, montaña + pteris, helecho. Un
helecho de las montañas.

LITERATURE: He, L.-J. & X.-C. Zhang. 2012. Exploring generic delimitation within the fern family Thelypteridaceae.
Molecular Phylogenetics and Evolution 63: 757–764. Holub, J. 1969. Oreopteris, a new genus of the family Thelypteridaceae.
Folia Geobotanica et Phytotaxonomica 4: 33–53. Holttum, R. E. 1971. Studies in the family Thelypteridaceae III. A new system
of genera in the Old World. Blumea19: 17–52.. Holttum, R. E. 1971. The genus Oreopteris (Thelypteridaceae). Kew Bulletin 36:
223–226.

440
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Figure 217. A. Oreopteris.

441
OSMUNDACEAE
Osmunda L.
DESCRIPTION: Terrestrial; stems erect, surrounded
by numerous overlapping leaf bases and a dense mat of
fibrous roots; sterile and fertile leaves hemidimorphic;
petioles in cross section with a single C-shaped
vascular bundle, the bases flattened and winged; laminae
1-pinnate-pinnatifid to 2-pinnate, pubescent when young,
glabrescent, scales absent; veins free; sporangia globose,
borne in clusters, the annulus consisting of a single, slightly
thickened lateral patch; spores green; x=22.
DESCRIPCIÓN: Terrestres; tallos erectos, rodeados
por numerosas bases de pecíolos traslapadas y una
densa capa de raíces fibrosas; hojas estériles y fértiles
hemidimórficas; pecíolos en sección cruz con un
haz vascular en forma de C, las bases achatadas
y aladas; láminas 1-pinnado-pinnatífidas a 2-pinnadas,
pubescentes cuando jóvenes, a glabrescentes, escamas
ausentes; nervaduras libres; esporangios globosos, nacen
en grupos, el anillo consiste de un sólo parche lateral
ligeramente engrosado; esporas verdes; x=22.

SIMILAR GENERA: Osmundastrum differs by GÉNEROS PARECIDOS: Osmundastrum difiere

holodimorphic leaves (i.e., the fertile leaves are completely por hojas holodimórficas (i.e., las hojas fértiles son
sporangia-bearing, not partially so). completamente llevando esporangios, no parcialmente
así).

COMMENTS: A genus of about 10 species, mostly COMENTARIOS: Un género de casi 10 especies,

temperate. Only one occurs in the Neotropics: O.
regalis L., the type of the genus. It typically grows in
wet soils, in sun or shade. The young leaves are often
reddish or purplish. The Osmundaceae are sister to
all other leptosporangiates ferns. It has a long fossil
record extending to the Permian. The fossils are easy to
recognize as belonging to the family because they bear
the same stem and petiole anatomy of the present-day
species (Miller, 1967, 1970). Osmunda has long included
what is now called Osmundastrum cinnamomeum (which
see). To maintain this species in Osmunda would require
lumping two Australasian genera (Leptopteris and Todea)
in Osmunda to preserve its monophyly. The derivation
of the name Osmunda is uncertain. Perhaps named for
Osmunder, Saxon god of war.
principalmente templadas. Solamente una especie se
encuentra en el Neotrópico: Osmunda regalis L., el tipo
del género. Típicamente crece en suelos mojados, en
sol o sombra. Las hojas jóvenes a menudo son rojizas
o purpuradas. Las Osmundaceae son hermana a todos
los otros helechos leptoesporangiados. Tiene un registro
fósil largo extendiéndose al Permian. Los fósiles son
fáciles reconocer como pertenecer a la familia porque
llevan la anatomía distinta de los tallos y pecíolos de
las especies actuales (Miller, 1967, 1970). Por mucho
tiempo Osmunda ha incluido lo que se llama ahora
Osmundastrum cinnamomeum (veáse). Para mantener
esta especie en Osmunda requeriría subsumiendo dos
géneros Australasiaticos (Leptopteris y Todea) en Osmunda
para preservar su monofilia. La derivación del nombre
Osmunda es incierto. Tal vez se nombra para Osmunder,
el dios Sajón de la guerra.

LITERATURE: Benedict, R. C. 1916. Osmundaceae. North American Flora 16(1): 27–28. Hewitson, W. 1962. Comparative
morphology of the Osmundaceae. Annals of the Missouri Botanical Garden 49: 57–93. Metzgar, J. S., J. E. Skog, E. A. Zimmer
& K. M. Pryer. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany
33: 31–36. Miller, C. N., Jr. 1967. Evolution of the fern genus Osmunda. Contributions from the Museum of Paleontology
University of Michigan 21: 139–203. Miller, C. N., Jr. 1971. Evolution of the fern family Osmundaceae based on anatomical
studies. Contributions from the Museum of Paleontology University of Michigan 23: 105–169.

442
Figure 218. A–C. Osmunda regalis. D, E. Osmundastrum cinnamomeum. (From Mickel & Smith, 2004.)

443
OSMUNDACEAE
Osmundastrum C. Presl
DESCRIPTION: Terrestrial; stems erect, surrounded
by numerous overlapping leaf bases and a dense mat of
fibrous roots; sterile and fertile leaves holodimorphic;
petioles in cross section with a single C-shaped
vascular bundle, the bases flattened and winged;
laminae 1-pinnate-pinnatifid, densely pubescent when
young, glabrescent, the hairs cinnamon-colored, scales
absent; veins free; sporangia globose, borne in clusters,
the annulus consisting of a single, slightly thickened lateral
patch; spores green; x=22.
DESCRIPCIÓN: Terrestres; tallos erectos, rodeados
por numerosas bases de pecíolos traslapadas y una
densa capa de raíces fibrosas; hojas estériles y fértiles
holodimórficas; pecíolos en sección cruz con un
sólo haz vascular en forma de C, las bases achatadas
y aladas; láminas 1-pinnado-pinnatífidas, pubescentes
cuando jóvenes, a glabrascentes, los pelos de color
canelo, escamas ausentes; nervaduras libres; esporangios
globosos, nacen en grupos, el anillo consiste de un sólo
parche lateral ligeramente engrosado; esporas verdes;
x=22.

SIMILAR GENERA: Osmunda differs by hemidimorphic
leaves (i.e., the dimorphic fertile pinnae are restricted
to only a portion of the leaf, which is otherwise green
and photosynthetic). Thelypteris differs by two vascsular
bundles in the petiole bases.
GÉNEROS PARECIDOS: Osmunda difiere por hojas
hemidimórficas (i.e., the pinnas fértiles dimorfitas están
restingidas a solamente una parte de la hoja, que es en
otras partes verdes y fotosintéticas). Thelypteris difiere
por dos haces vasculares en las bases de los pecíolos.

COMMENTS: Osmundastrum includes only one extant
species: O. cinnamomea (a fossil species is also known). It
occurs in eastern North America, Central America, South
America, and Asia. Besides holodimorphic sterile and
fertile leaves, Osmundastrum differs from Osmunda by two
anatomical characteristics (Hewitson, 1961; Miller, 1971).
One pertains to the number of clusters of thick-walled
cells forming the sclerenchyma ring around the vascular
strand in the petiole base, the other is the presence of
a second endodermis in the stele (one endodermis is
located between the inner cortex and the pericycle, and
the second is located between the xylem cylinder and the
pith).The genus name is derived from Osmunda + astrum,
a Greek diminutive suffix.
COMENTARIOS: Osmundastrum incluye solo una
especie actual: O. cinnamomea (una especie fósil se
conoce también). Existe en América del Norte oriental,
América Central, América del Sur y Asia.Además de
hojas estériles y fértiles holodimórficas, Osmundastrum
difiere de Osmunda por dos características anatómicas
(Hewitson, 1961; Miller, 1971). Una pertenezca al número
de grupos de células con paredes engrosadas formando
el anillo de esclerénquima alrededor de la haz vascular en
la base del pecíolo, la otra es la presencia de un epidermis
segundo en la estela (un endodermis se ubica entre la
cortex interior y el perciclo, y la segunda se ubica entre
el cilindro de xilema y la médula). El nombre genérico se
deriva de Osmunda + astrum, un sufijo griego diminutivo.

LITERATURE: Benedict, R. C. 1916. Osmundaceae. North American Flora 16(1): 27–28. Hewitson, W. 1962. Comparative
morphology of the Osmundaceae. Annals of the Missouri Botanical Garden 49: 57–93. Metzgar, J. S., J. E. Skog, E. A. Zimmer
& K. M. Pryer. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany
33: 31–36. Miller, C. N., Jr. 1967. Evolution of the fern genus Osmunda. Contributions from the Museum of Paleontology
University of Michigan. 21: 139–203. Miller, C. N., Jr. 1971. Evolution of the fern family Osmundaceae based on anatomical
studies. Contributions from the Museum of Paleontology, University of Michigan 23: 105–169.

444
Figure 219. Osmundastrum cinnamomeum. A. Holodimorphy of sterile and fertile leaves. B. Pinna base, abaxial view, with tuft
of hairs at the base. C. Fertile pinnules. D. Cross-section of the rootstock showing stem in the center surrounded overlapping
petiole bases. E. Petiole base in cross-section showing C-shaped petiolar bundle. (Original, ©Robbin Moran, 2009.)

445
DENNSTAEDTIACEAE
Paesia J. St.-Hil.
DESCRIPTION: Terrestrial or rarely rupicolous;
rhizomes 3–5 mm wide, creeping, pubescent; scales
clathrate; leaves 0.22–2.5 m, glabrous or glandular-
pubescent, scandent; petioles without epipetiolar buds;
laminae 2-pinnate to 4-pinnate, without buds in the axils
of the distal pinnae; pinnules catadromic or anadromic;
rachises and costae grooved adaxially, flexuous; veins free,
ending at the margins or just behind in clavate apices; sori
marginal, continuous along each margin of the ultimate
segments, borne along a vein connecting two to several
vein tips; indusia double, the adaxial formed of the
enrolled margin and scarious of the lamina, the abaxial
inconspicuous, opening outward, membranaceous; spores
monolete; x=26.
SIMILAR GENERA: Odontosoria differs by sori supplied
by only one vein, narrow or linear ultimate segments, and
trilete spores. Eriosorus flexuosus differs by sori lacking
indusial and spreading along the viens away from the
margin.
COMMENTS: Paesia is pantropical and comprises
about 15 species, with 3–5 in the Neotropics. Most of
the species have laminae scandent on the surrounding
vegetation. The species occur at 1200–3300 m, most
often in cloud forests, usually along roadsides, shrubby
places, and deforested areas. Paesia resembles Pteridium
by having two indusia: an adaxial “false” one formed by
the enrolled margin of the lamina, and an inner “true”
indusium that is hyaline and easily overlooked. The two
most common and widespread species in the Neotropics
are P. anfractuosa (Christ) C. Chr. and P. glandulosa (Sw.)
Kuhn. The former has pinnae and pinnules arranged
anadromically, the latter catadromically. The genus name
honors Duke Fernando Diaz Paes Leme, who visited Brazil
in 1660 in the service of the Portuguese government.
LITERATURE:
446
DESCRIPCIÓN: Terrestres o raramente rupícolas;
rizomas 3–5 mm de ancho, creeping, pelosos; hojas 0.2–
2.5 m, glabras o glanduloso-pubescentes, escandentes;
pecíolos con yemas epipeciolares; láminas 2-pinnadas a
4-pinnadas, sin yemas en las axilas de las pinnas distales;
pínulas dispuestas catadrómica o anadrómicamente;
raquises y costas flexuosos, surcados adaxialmente;
nervaduras libres, terminando en los márgenes o
escasamente por detrás en ápices claviformes; soros
marginales, continuos por cada margen de los
segmentos últimos, nacidos por una vena que conecta 2
a varios ápices de nervaduras; indusios dobles, el adaxial
formado por el margen enrollado y escarioso de la
lámina, el abaxial inconspicuo, membranáceo, abriéndose
hacia afuera; esporas monoletes; x=26.
GÉNEROS PARECIDOS: Odontosoria difiere por soros
suministrados por una sola vena, segmentos últimos más
estrechos y esporas triletes. Eriosorus flexuosus difiere por
los soros no indusiados y difundiéndose por las venas
atrás del margen.
COMENTARIOS: Paesia es pantropical y consta de casi
15 especies, con 3–5 en el Neotrópico. La mayoría de
las especies tienen láminas escandentes en la vegetación
circundante. Las especies existen en 1200–3300 m, a
menudo en bosques nublados, usualmente por los bordes
de los caminos, matorrales y áreas desmontadas. Paesia
se parece a Pteridium por tener dos indusios: uno adaxial
“falso,” formado por el margen enrollado de la lámina,
y otro más interior “verdadero” que es membranáceo
y pasado por alto fácilmente. Los dos especies más
comúnes y difundidas en el Neotrópico son P. anfractuosa
(Christ) C. Chr. y P. glandulosa (Sw.) Kuhn. El anterior tiene
pinnas y pínulas arregladas anadromicamente, el último,
catadrómicamente. El nombre genérico honor el Duque
Fernando Diaz Leme, quien visitó a Brasil en 1660 en
servicio del gobierno Portugués.
Figure 220. Paesia glandulosa. A. Lamina, medial portion. B. Rhizome habit. C. Transverse section of rhizome showing solenostele.
D. Transverse section of petiole base showing omega-shaped vascular bundle. E. Adaxial surface of rhachis, costae, and pinnules,
with (at left) glandular hairs from adaxial surface of lamina. F. Lobe, abaxial surface, showing sori. G. Sorus cut transversely to
show inner and outer indusia (Original, ©R. C. Moran 2014.)

447
LYCOPODIACEAE
Palhinhaea Vasc. & Franco
DESCRIPTION: Terrestrial; main horizontal stems
creeping, superficial, indeterminate, branching
dichotomously in the horizontal plane, also branching in
the vertical plane to give rise to single dorsal rank of
erect determinate branching systems, these tree-
like, their stem surfaces often puberulent, the hairs ca.
0.1–0.3 mm long, unicellular; ultimate branchlets terete
(not flattened), isophyllous, with leaves polystichous,
not in distinct ranks, lacking hyaline hair-like apicies;
gemmiferous branchlets and gemmae absent; strobili
sessile, nodding or pendant; sporophylls peltate, with a
decurrent medial wing, stramineous at time of sporangial
dehiscence; sporangia almost completely enclosed
in cavities formed by the cortex of the strobilus and
covered by the basal membranes of adjacent sporophylls,
anisovalvate, reniform, the epidermal cells with nodular
thickenings; spores rugate to nearly smooth, the laesurae
raised with a fine central groove; gametophytes surficial,
green, rounded; n=104, 108, 110, 136, 156, c. 165, 208.
DESCRIPCIÓN:Terrestres; tallos principales horizantales
reptantes, superficiales, indeterminados, ramificándose
dicotomicamente en el plano horizontal, también
ramificándose en el plano vertical para dar lugar a una
hilera dorsal de sistemas de ramas determinadas,
estas arborescentes, sus superficies a menudo
puberulentos, los pelos ca. 0.1–0.3 mm long, unicelulares;
ramitas últimas teretes (no achatados), isofílicas, con
hojas polísticas, no en hileras distintas, carente de ápices
filiformes; ramitas gemiferas y yemas ausentes; estróbilos
sésiles, cernuos o péndulos; esporofilos peltatos, con
una ala medial decurrente, estramineo en el tiempo de
dehiscencia esporangial; esporangios casi completamente
envueltos en cavidades formadas por la corteza del
estróbilo y cubiertos por las membranas basales de
esporofilos adjacentes, anisovalvados, reniformes, células
epidérmicas con engrosamientos nodulares; esporas
rugosas hasta casi lisas, las laesuras elevadas con un surco
central fino; gametofitos surficiales, verdes, redondeados;
n=104, 108, 110, 136, 156, c. 165, 208.

SIMILAR GENERA: Lycopodiella and Pseudolycopodiella GÉNEROS PARECIDOS: Lycopodiella y

differ by erect strobli terminating simple (rarely forked)
erect branches, and sporangia isovalvate, not enclosed by
the cortical tissue of the strobilus axis. Austrolycopodium
and Lycopodium differ by main horizontal creeping stems
with two dorsolateral rows of erect branches.
Pseudolycopodiella difieren por los estrobilos erectos
terminando ramas simples (raras veces furcados) y
esporangios isovalvados, no envueltos por tejido cortical
del eje estrobilar. Austrolycopodium y Lycopodium difieren
por los tallos principales horizontales reptantes con dos
hileras dorsolaterales de ramas erectas.

COMMENTS: Palhinhaea consists of ca. 20 species COMENTARIOS: Palhinhaea consta de ca. 20 especies
worldwide. The species occur in wet forests and on mundial. Las especies existen en bosques húmedos y
disturbed soil, often on roadsides, especially at middle suelos perturbados, a menudo en los lados de caminos,
elevations. Palhinaea cernua (L.) Vasc. & Franco, the type of especialmente en elevaciones medias. Palhinaea cernua
the genus, is pantropical and common. Las otras especies (L.) Vasc. & Franco, el tipo del género, es pantropical
generally have narrow distributions. In Florida P. cernua y común. Las otras especies generalmente tienen
persists through the winter as a buried stem tips, the rest of distribuciones estrechas. En Florida P. cernua persiste por
the plant dying. Palhinhaea is sister to Lateristachys Holub, el invierno como apices caulinares enterrados en el suelo,
a genus of Australia and New Zealand. The genus name el resto de la planta muriéndose. Palhinhaea es hermano
honors Ruy Telles Palhinha (1871–1950), a Portuguese a Lateristachys Holub, un género de Australia y Nueva
botanist and student of the flora of the Azores. Zelanda. El nombre genérico honora a Ruy Telles Palhinha
(1871–1950), un botánico Portugués y estudiante de la
flora de las Azores.
LITERATURE: Øllgaard, B. 1988. Lycopodiaceae. In: G. Harling & L. Andersson, editors. Flora of Ecuador 33: 1–155. 1992.
Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the Missouri Botanical Garden 79: 687–717. Øllgaard,
B. & P. G. Windisch. 2014. Lycopodiaceae in Brazil. Conspectus of the family I. The genera Lycopodium, Austrolycopodium,
Diphasium, and Diphasiastrum. Rodriguésia 62(2): 261–277. Øllgaard, B. & P. G. Windisch. 2016. Lycopodiaceae in Brazil.
Conspectus of the family II. The genera Lycopodiella, Palhinhaea, and Pseudolycopodiella. Rodriguésia 67(3): 691–719.
Thomas, B. A. 1981.The vegetative growth of Lycopodiella cernua (L.) Pic. Ser. and its importance in the generic determination
of the species. Annals of Botany 47: 443–449. Vasconcellos, I. C. & J. A. Franco. 1967. Breves notas sobre Licopodiáceas.
Boletín del la Sociedade Broteriana, sér. 2, 41: 23–25.

448
Figure 221. Palhinhaea. A–C. P. cernua. portion of strobilus showing sporangia largely enclosed by stem cortex. D–F. P.
camporum. G–I. P. riofrioi. J–L. P. bradei, in K note sunken sporangia. (Modified from Øllgaard & Windisch, 2016.)

449
BLECHNACEAE
Parablechnum C. Presl
DESCRIPTION: Terrestrial; rhizomes erect or suberect,
rarely subarborescent or creeping, scaly, the scalles
concolorous or bicolorous; stolons absent; sterile and
fertile leaves dimorphic, sometimes covered by mucilage;
laminae 1-pinnate, truncate basally, with a conform or
subconform apices; buds absent or (more rarely) present;
aerophores often born at the bases of pinnae abaxially;
pinnae sessile (but not adanate) or stalked; entire to
denticulate; veins free, hydathhodous; sori linear; indusia
subentire to lacerate or erose; x=28, 31, 33.
DESCRIPCIÓN:Terrestres; rizomas erectas o suberectas,
raras veces subarborescentes o reptantes, escamosos,
las escameas concoloras o bicoloras; estolons ausentes;
hojas estériles y fértiles dimófas, a veces cubiertas por
mucílago; láminas 1-pinnadas, truncatas basalmente, con
ápices conformes o subconformes; brotes ausentes o
(raras veces) presentes; aeróforos a menudo nacidos en
las bases de las pinnas abaxialmente; pinnas sésiles (pero
no adnadas) o pecioluladas; entiras a denticuladas; venas
libres, con hidatodos; soros lineares; indusios subenteros
a lacerados o erosos; x=28, 31, 33.

SIMILAR GENERA: Austroblechnum differs by pinnae GÉNEROS PARECIDOS: Austroblechnum difiere

adnate to the rachis and pinnatifid (non-conform) lamina
apices. Blechnum differs by monomorphic sterile and
fertile leaves and the presence of stolons. Lomaridium
and Struthiopteris differ pinnae adnate to the rachis.
Lomariocycas differs by arborescent habit and linear
rhizome scales with a dark central stripe. Plagiogyria
(Plagiogyriaceae) differs by the complete of scales on all
parts of the plant, absence of indusia, oblique (vs. vertical)
annuli, and costae non-grooved adaxially.
por las pinnas adnatas al ráquis y ápices de la láminas
pinnatifidas (no conformes). Blechnum difiere por
hojas estériles y fértiles monomórfas y la presencia de
estolons. Lomaridium y Struthiopteris difieren por las
pinnaes adnadas al rachis. Lomariocycas difiere por hábito
arborescente y escamas del rizoma lineares con una raya
oscura central. Plagiogyria (Plagiogyriaceae) difiere por
la ausencia de escamas en todas partes de la plantas, la
ausencia de indusios, anillos oblícuos (vs. verticales) y
costas no surcados adaxialmente.

COMMENTS: Parablechnum is pantropical and consists COMENTARIOS: Parablechnum es pantropical y consta

of ca. 65 species. It is the largest genus in the family. Nearly
half the species occur in the Neotropics. The species
occur mostly at middle to high elevations (1500–3800 m).
All are terrestrial except for P. stuebelii, which is scandent.
The genus is sister to Lomariocycas. Its is derived from the
Greek para, beside, and blechnum, the ancient name for a
fern. It refers to the similarity to Blechnum.
de ca. 65 especies. Es el género más grande en la familia.
Casi una mitad de las especies existen en el Neotrópico.
Existen mayormente en elevaciones medias a altas
(1500–3800 m). El género es hermana a Lomariocycas.
Su nombre es derivado del griego para, al lado de, y
blechnum, un nombre antiguo para un helecho. Se refiere
a la similitud a Blechnum.

LITERATURE: Dittrich, V. A. O., A. R. Smith & A. L. de Gasper. 2017. Parablechnum roraimense and P. paucipinna spp.
nov. (Blechnaceae), lectotypification of P. stuebelii, and citation corrections in the family. Phytotaxa 292: 65–73. Gasper, A. L. &
V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae (Polypodiales: Polypodiopsida): New genera,
resurrected names, and combinations Phytotaxa 275: 191–227. Smith, A. R. & M. Lehnert. 2007.Ten new species and two new
combinatioons of Blechnum (Blechnaceae, Pteridophyta) from Bolivia. American Fern Journal 97: 66–80..

450
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Illustration to be done

Figure 222. Parablechnum.

451
DRYOPTERIDACEAE
Parapolystichum (Keyserl.) Ching
DESCRIPTION: Terrestrial; rhizomes short- to long-
creeping, scaly; leaves monomorphic; petiole with four
or more vascular bundles; laminae 3–4-pinnate-pinnatifid,
lanceolate, deltate, or pentagonal, the lower pinnae
enlarged basiscopically; scaly buds present on the
distal parts of the rachis of some species; pinnules
anadromic or catadromic; rachis with 2 prominent
ridges adaxially, these continuous with the thickened
margins of the ultimate segments (Fig. D,E); costae
raised (not grooved), hairy adaxially with 0.1–0.3 mm
long, multicellular hairs, costae, and costules scaly abaxially,
hairy adaxially; glandular hairs often present on the lower
surface of the lamina, these cylindrical, oblong appressed,
bright yellow to orange-red; sori round. x=41.
SIMILAR GENERA: Lastreopsis is indistinguishable
morphologically, but its neotropical species can be
distinguished by lack of foliar buds. Dryopteris and Rumohra
differ by axes adaxially glabrous. Megalastrum differs
by coarse, whitish, strigose hairs on the axes adaxially,
prominent hydathodes, and the basal veinlet of the distal
pinnules springing from the costa, not the costule, with
the lobe supplied by this veinlet adnate to the costa.
COMMENTS: Parapolystichum contains 27 species
distributed in the Neotropics (4 spp.), Africa (6 spp.),
Madagascar (9 spp.), and Australia and New Zealand
(8 spp.). The species within each of these geographic
areas form clades. Formerly, the species were treated
in Lastreopsis, but molecular phylogenetic studies show
that Parapolystichum is distinct. No known morphological
character distinguish the two genera. Several species of
Parapolystichum, however, have buds on their laminae
distally—a character absent in Lastreopsis. The rachis-
costa architecture of Lastreopsis and Parapolystichum is
unique and unmistakable once learned. It is characterized
by the shape of the axes on the upper surface of the
lamina (Fig. D,E). The rachis has two ridges and is densely
puberulent between them. The ridges are continuous
with the decurrent margins of the pinnules. The central
part of the axes is raised (not grooved) as in most of the
Dryopteridaceae. The genus name is derived from the
Greek para, beside + Polystichum.
LITERATURE: Labiak, P. H., M. Sundue, G. Rouhan, J. Garrison Hanks, J. T. Mickel & R. C. Moran. 2014. Phylogeny
and historical biogeography of the lastreopsid ferns (Dryopteridaceae). American Journal of Botany 101: 1207–1228. Labiak, P.
H., M. Sundue, G. Rouhan & R. C. Moran. 2014. New combinations in Lastreopsis and Parapolystichum (Dryopteridaceae).
Brittonia 66: 79–86. Tindale, M. D. 1965. A monograph of the genus Lastreopsis Ching. Contributions to the New South Wales
National Herbarium 3: 249-339, t. 1–23.
452
DESCRIPCIÓN: Terrestres; rizomas breves a largamente
rastreros, escamosos; hojas monomorfas; pecíolos con 4 o
más haces vasculares; láminas 3–4-pinnadas, lanceoladas,
deltadas o pentagonales, las pinnas más inferiores
agrandadas basiscópicamente; yemas escamosas en las
porciones distales del raquis en algunas especies;
pínnulas dispuestas anadrómica o catadrómicamente;
raquis con 2 prominentes crestas adaxiales, éstas
continuas con los engrosados márgenes foliares de
los segmentos terminales (Fig. D,E); costas elevadas
(no surcadas), pelosas adaxialmente con pelos 0.1–0.3
mm, multicelulares; tricomas glandulosos a menudo
presentes en el haz de la lámina, cilíndricos, oblongos,
adpresos, amarillo brillante a anaranjado-rojizo; soros
redondos. x=41.
GÉNEROS PARECIDOS: Lastreopsis es indistinguible
morfológicamente, pero sus especies neotropicales se
pueden ser distinguidas por la falta de brotes foliares..
Dryopteris y Rumohra se difieren por sus ejes adaxialmente
glabros. Megalastrum difiere por pelos toscos,
blanqueciños, strigosos sobre los ejes adaxialmente,
hidatodos prominentes, y la venilla basal de las pinnulas
distales surgiendo de la costa, no de la cóstula, con el
lóbulo suministrado por esta venilla adnato a la costa.
COMENTARIOS: Parapolystichum consta de 27
especies distribuidas en el Neotrópico (4 spp.), África
(6 spp.), Madagascar (9 spp.), and Australia and Neuva
Zelanda (8 spp.). En cada area geográfica las especies
forman un clado. Anteriormente, las especies fueron
tratadas en Lastreopsis, pero los studios moleculares
filogenéticos demuestran que Parapolystichum es distincto.
No hay caracteres morfológicos que separan los dos
géneros. Varias especies de Parapolystichum tienen brotes
en sus láminas distalmente—un carácter ausente en
Lastreopsis.La arquitectura raquis-costa de Lastreopsis y
Parapolystichum es único y no se puede confundir después
de haber comprendido. Se caracteriza por la forma de
los ejes en el haz de la lámina (Fig. D,E). El raquis tiene
dos crestas y es densamente puberulento entre ellas. Las
crestas son continuas con los márgenes decurrentes de
las pínnulas. La parte central de los ejes es prominulo (no
sulcados) como en la mayoría de las Dryopteridaceae.
El nombre genérico se deriva del greigo para, al lado +
Polystichum.
Figure 223. A–D, H: Parapolystichum effusum. F, G: P. excultum. (From Mickel & Smith, 2004.)

453
THELYPTERIDACEAE
Parathelypteris (H. Itô) Ching
DESCRIPTION: Terrestrial; rhizomes 1–3 mm wide,
long-creeping, scaly but not pubescent; blades pinnate-
pinnatifid throughout, gradually tapered proximally and
distally, suprabasal pinnae sessile but not adnate to rachis
nor connected by wing, lowest pinnae rudimentary and
widely spaced; veins free; laminar tissue eglandular, veins
free, meeting the margins above the sinuses; sori round;
indusia present; sporangial stalks eglandular, the capsules
non-setose; x=27, 31.
DESCRIPCIÓN: Terrestres; rizomas 1–3 mm de ancho,
largamente rastreros, escamosos pero no pubescentes;
láminas 1-pinnado-pinnatífidas por todo su extensión,
gradualmente reducidas proximalmente y distalmente,
pinnas basales sésiles pero no adandas al ráquis ni
conectadas por una ala, pinnas basales rudimentarias
y anchmente espaciadas; venas libres; tejido laminar
eglandular; venas libres, alcanzando los márgenes arriba de
los sinuses; soros redondos; indusios presentes; peciolos
esporangiales eglandulares, las cápsulas no setosas; x=27,
31.

SIMILAR GENERA: Amauropelta differs by erect or GÉNEROS PARECIDOS: Amauropelta differs by erect
suberect rhizomes. Thelypteris differs by laminae widest or suberect rhizomes. Thelypteris difiere por láminas más
at the base. anchas en la base.

COMMENTS: Parathelypteris has ca. 15 species, but it is
not monophyletic as currently circumscribed. It occurs in
eastern North America and the tropics and subtropics
of Asia and Malesia. The two American species are P.
nevedensis (Baker) Holttum and noveboracensis (L.) Ching. It
is readily distinguished by its long-creeping rhizomes and
1-pinnate-pinnatifid blades that taper toward the base
and apex. The type species of the genus is P. glanduligera
(Kunze) Ching, of southeastern Asia. The genus name
is derived from the Greek para, beside, or next to +
Thelypteris.
COMENTARIOS: Parathelypteris consta de ca. 15
especies, pero no es monofilético como circunscrito
actualmente. Existe en América del Norte oriental y
los trópicos y subtrópicos de Ásia y Malesia. La única
especie Americana es P. nevedensis (Baker) Holttum y
noveboracensis (L.) Ching. Se distingue fácilmente por
sus rizomas largamente rastreros y láminas 1-pinnadas-
pinnatifidas reducidas hacia la base y el ápice. La especie
tipo del género es P. glanduligera (Kunze) Ching, de Ásia
suroriental. El nombre genérico se deriva del greigo para,
al lado, o junto a + Thelypteris.

LITERATURE: Almeida, T. E., S. Hennequin, H. Schneider, A. R. Smith, J. A. N. Batista, A. J. Ramalho, K. Proite &
A. Salino. 2016. Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of
neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688–700 He, L.-J. & X.-
C. Zhang. 2012. Exploring generic delimitation within the fern family Thelypteridaceae. Molecular Phylogenetics and Evolution
65: 767–764. Holttum, R. E. 1971. Studies in the family Thelypteridaceae III. A new system of genera in the Old World.
Blumea 19: 17–52. Holttum, R. E. 1974. Thelypteridaceae of Africa and adjacent islands. Journal of South African Botany 40:
123–168. Holttum, R. E. 1981. Thelypteridaceae. In: C. G. G. J. van Steenis & R. E. Holttum (gen. eds.), Flora Malesiana, series
2 - Pteridophytes, ferns and fern allies 1(5): 331–599. Ogden, E. C. 1981. Field guide to northeastern ferns. New York State
Museum Bulletin, no. 444. Albany, New York.

454
Figure 224. Parathelypteris novaboracensis. (From Ogden, 1981.)

455
POLYPODIACEAE
Pecluma M. G. Price
DESCRIPTION: Epiphytic, rarely terrestrial or rupestral;
proliferous roots often present; rhizomes short-creeping,
not branched, scaly, the scales basifixed (not peltate),
not clathrate, concolorous, glabrous or pubescent
with dark hairs basally (these resembling rhizoids and
sometimes obscuring the scales); petioles articulate to low
(1–2 mm) phyllopodia, terete except for narrow lateral
ridge decurrent from the lamina base; laminae pectinate,
with 30 or more segments, usually reduced toward
the base, the apex pinnatifid; segments adnate, entire or
nearly so; rachises puberulent adaxially, not grooved,
abaxially glabrous or scaly and/or pubescent; veins free,
occasionally anastomosing; sori round, non-indusiate;
paraphyses present; sporangial capsules glabrous or
setulose; spores yellow when fresh, monolete; x=37.
DESCRIPCIÓN: Epífitas, raras veces terrestres o
rupestres; raíces prolíferas a menudo presentes; rizomas
cortamente rastreros, no ramificadas, escamosos, las
escamas basificadas (no peltadas), no clatradas,
concoloras, glabras o pubescentes en de la base con
pelitos oscuros (estos se parecen a rizoides y a veces
oscuren las escamas); pecíolos oscuros, no surcados
adaxialmente, articulados a filopódios cortos (1–2
mm),; láminas pectinadas, con 30 o más pares de
segmentos, usualmente reducidas en la base, el ápice
pinnatífido; segmentos adnatos, enteros o casi enteros;
ráquises puberulentos adaxialmente, no surcados,
abaxialmente glabros o escamosos y/o pubescentes;
venas libres, ocasionalmente anastomosadas; parafisos
presentes; soros redondos, no indusiados; cápsulas
esporangiales glabras o sétulas; esporas amarillas cuando
frescas, monoletes; x=37.

SIMILAR GENERA: Polypodium differs by petioles
and rachises adaxially glabrous and grooved, glabrous
rhizome scales, and non-setulose sporangial capsules. The
“Polypodium dulce group” differs by light-colored petioles,
fewer pinnae pairs, laminae widest at base, and petioles
with several vascular bundles. The Terpsichore taxifolia
group differs by long (0.5–2 mm long) hairs on the
petioles, and green trilete spores.
GÉNEROS PARECIDOS: Polypodium difiere por
pecíolos y ráquises surcados y glabros adaxialmente,
pecíolos usulmente estramíneos (no oscuros) y cápsulas
esporangiales no setulosas. El “grupo Polypodium dulce”
difiere por pecíolos más claros, menos pares de pinnas,
láminas más anchas en la base, y pecíolos con varios
haces vasculares. El grupo de Terpsichore taxifolia difiere
por pelos largos (0.5–2 mm de largo; más evidentes en el
pecíolo) y esporas verdes triletes.

COMMENTS: Pecluma contains about 28 species, all COMENTARIOS: Pecluma consta de casi 28 especies,
neotropical. A study by Ranker et al. (2004) suggested todos neotropicales. Un estudio por Ranker et al.
that the genus was sister to the “Polypodium dulce group” (2004) sugerió que el género es hermana al “Grupo de
of Moran (1995). Both share several distinctive characters: Polypodium dulce” de Moran (1995). Los dos comparten
comose rhizome scales, adaxially puberulent rachises, free varias características significativas: escamas del rizoma
veins, and setulose sporangial capsules.The only difference comosas, ráquises puberulentos adaxialmente, venas libres
is that Pecluma has rhizome scales attached across the y cápsulas esporangiales setulosas. La única diferencia
width of the scale base, whereas those of the P. dulce group es que Pecluma tiene escamas del rizoma unidas por la
are peltate, attached at a point. Many species of Pecluma anchura de la base, mientras que las del grupo P. dulce son
tolerate periods of dry weather. When dry, their laminae peltadas, unidas a un punto. Muchas especies de Pecluma
and pinnae curl inward (abaxially), and when the rains soportan periodos de tiempo seco. Cuando seco, sus
return, they uncurl rapidly and begin photosynthesis. The láminas y pinnas se enrollan a dentro (abaxialmente), y
genus name was formed by compounding the epithets of cuando las lluvias regresan, se despliegan rápidamente y
two widespread species: P. pectinata and P. plumula. empiezan a realizar fotosíntesis. El nombre del género se
formó por combinar los epítetos de dos especies muy
difundidas: P. pectinata y P. plumula.
LITERATURE: Tiana-Moreno, L. A. 2015. Católogo comentado de las especies de Pecluma (Polypodiaceae) de Colombia.
Boletín Científica del Museo de Historia Natural de la Universidad de Caldas 19(2): 17–59 [DOI: 10.17151/bccm.2015.19.2.2].
Evans, A. M. 1969. The Polypodium pectinatum-plumulum complex. Annals of the Missouri Botanical Garden 55: 193–293. Price,
M. G. 1983. Pecluma, a new tropical American fern genus. American Fern Journal 73:109–116.

456
Figure 225. A, B. Pecluma alfredii. C, D. P. cupreolepis. E, F. P. plumula. G, H. P. ptilodon. J, K. P. atra. L, M. P. consimilis. N,
O. P. ferruginea. (From Mickel & Smith, 2004.)

457
PTERIDACEAE
Pellaea Link
DESCRIPTION: Terrestrial or rupestral; rhizomes
creeping or erect, scaly, the scales usually bicolorous;
leaves 1–4 pinnate, monomorphic or almost dimorphic,
coriaceous; petiole with a single vascular bundle; pinnules
stalked, glabrous or slightly pubescent; petioles and
rachises tan to black, terete, glabrous or hairy; laminar
tissue bluish green, coriaceous, glabrous; veins free,
ending in clavate hydathodes; sori intramarginal on the
vein tips; indusium formed by the recurved margin of the
pinnule; spores trilete or monolete; x=29.
DESCRIPCIÓN: Terrestres or rupestres; rizomas
rastreros o erectos, escamosos, las escamas usualmente
bicoloras; hojas 1–4-pinnadas, monomorfas o casi
dimorfas, coriáceas; pecíolos con una haz vascular;
pínnulas pecioluladas, glabras o poco pubescentes;
pecíolos y ráquises leonados a negros, teretes, glabros
a pubescentes; tejido laminar oscuro azuloso verde,
glabro; venas libres, terminando en hidatodos claviformes;
soros intramarginales en los puntos de las venas; indusios
formados por el margen recurvado de la pínnula; esporas
triletas o monoletas; x=29.

SIMILAR GENERA: Cheilanthes, which differs by usually GÉNEROS PARECIDOS: Cheilanthes, lo cual difiere
more herbaceous laminae that are green (not blueish usualmente por láminas más herbáceas, verdes (no
green). The two are sometimes difficult to distinguish. verde-azuladas). Los dos son a veces difíciles de distinguir.

`
COMMENTS: Pellaea contains about 35 species most
commonly found in deserts or seasonally dry habitats,
usually in or among rocks. The genus has centers of
diversity in the southwestern United States, Mexico,
and Africa. Many species are apogamous (Tryon, 1968).
The neotropical species form a monophyletic group
related to Astrolepis and Argyrochosma. It are not closely
related African species in sect. Holochlaena, which itself
is biphyletic. The genus name is derived from the Greek
pellos, dusky. It refers to the bluish gray lamina tissue.
COMENTARIOS: Pellaea consta de casi 35 especies,
las que se encuentran más comúnmente en hábitats
desérticos o secos estacionalmente, usualmente en
o alrededor de las rocas. El género tiene centros de
diversidad en el suroeste de los Estados Unidos, México
y África. Muchas especies son apógamas (Tryon, 1968).
Las especies neotropicales forman un grupo monofilético
relacionado a Astrolepis y Argyrochosma. No está
cercanamente relacionado a las especies Africanas de la
sección Holochlaena, la cual es bifilética. El nombre
genérico se deriva del griego pellos, obscuro. Se refiere al
tejido laminar gris azulado.

LITERATURE: Gastony, G. J. & D. R. Rollo. 1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern
United States and adjacent Mexico—a molecular phylogenetic reassessment of generic lines. Aliso 17: 131–141. Tryon, A. F.
1957. A revision of the fern genus Pellaea section Pellaea. Annals of the Missouri Botanical Garden 44: 125–193. Tryon, A. F.
1968. Comparisons of sexual and apogamous races in the fern genus Pellaea. Rhodora 70: 1–24.

458
Figure 226. A–C. Pellaea ribae. D–F. P. ternifolia subsp. ternifolia. G, H. P. ternifolia subsp. arizonica. J, K: P. villosa. L–M. P.
ternifolia subsp. brandegeei. O, P. P. wrightiana. (From Mickel & Smith, 2004.)

459
PTERIDACEAE
Pentagramma Yatsk., Windham & E. Wollenw.
DESCRIPTION: Terrestrial or rupestral; rhizomes
short-creeping, erect, or ascending, scaly, the scales
bicolorous, with black central stripe and lighter margins,
entire; leaves 5–40 cm long, 1–4 pinnate, monomorphic;
petioles castaneous to black, terete, containing a single
vascular bundle, glabrous, farinose or viscid-glandular;
laminae deltate-pentagonal, about as long as wide, 1-
to 2-pinnate-pinnatifid proximally, glabrous or glandular,
dull, abaxially farinose, farina white or yellow; ultimate
segments sessile, adnate their full width to rachises and
costae, margins not enrolled; veins free, obscure; sori non-
indusiate; sporangia borne along veins, containing 64
spores; spores tan to brown, trilete; x=30.
DESCRIPCIÓN: Terrestres o rupestres; rizomas
cortamente rastreros, erectos, o ascendentes, escamosos,
las escamas bicoloras, con una raya central y
bordes más claras, enteras; hojas 5–40 cm de longitud,
1–4-pinnadas, monomorfas; pecíolos castaños a negro,
teretes, conteniendo una haz vascular, glabros, farinosos
o viscido-glandulares; láminas deltado-pentagonales,
casi tan largo como ancho, 1- a 2-pinnado-pinnatífido
proximalmente, glabras o glandulares, opacas, farinosas
abaxialmente, la farina blanca o amarilla; segmentos
últimos sésiles, adnatos por su anchura completa a los
ráquises y costas, márgenes no enrollados; venas libres,
obscuras; soros no indusiados; esporangios nacidos por
las venas, conteniéndose 64 esporas; esporas leonadas a
pardas, triletes; x=30.

SIMILAR GENUS: Pityrogramma differs by concolorous GÉNERO PARECIDO: Pityrogramma difiere por
rhizome scales and laminae lanceolate or oblong. escamas de los rizomas concoloras y láminas lanceoladas
Doryopteris differs by laminae without farina and sori that a oblongas. Doryopteris difiere por láminas sin farina y
run along the blade margins. soros que corren por los márgines de las láminas.

COMMENTS: Pentagramma contains two species of
western North America and Mexico: P. pallida (Weath.)
Yatsk., Windham & E. Wollenw. y P. triangularis (Kaulf.)
Yatsk., Windham & E. Wollenw. The latter is considered
having three subspecies. For a long time the genus was
included in Pityrogramma because of the farinose laminae
and sori running along the veins, but it is now known that
the two genera are not closely related within the family.
Pentagramma typically grows in dry, rocky habitats. The
genus name is derived from the Greek penta, five, and
gramma, line, alluding to the pentagonal outline of the
blade.
COMENTARIOS: Pentagramma consta de dos especies
de América del Norte occidental y México: P. pallida
(Weath.) Yatsk., Windham & E. Wollenw. y P. triangularis
(Kaulf.) Yatsk., Windham & E. Wollenw. La última se
considera tener tres subespecies. Por mucho tiempo el
género fue incluido en Pityrogramma a causa de las láminas
farinosas y soros corriéndose por las venas, pero hoy día
se conoce que los dos géneros no son cercanamente
emparentados dentro de la familia. Pentagramma crece
típicamente en hábitats rocosos áridos. El nombre
genérico se deriva del griego penta, cinco, y gramma, línea,
refiriéndose al contorno pentagonal de la lámina.

Alt, K. S. & V. Grant. 1960. Cytotaxonomic observations on the goldback fern. Brittonia 12: 153–170. Tryon, R. M. 1962.
Taxonomic fern notes. II. Pityrogramma (including Trismeria) and Anogramma. Contributions from the Gray Herbarium
of Harvard University 189: 52–76. Weatherby, C. A. 1920. Varieties of Pityrogramma triangularis. Rhodora 22: 113–120.
Yatskievych, G. M., M. D. Windham & E. Wollenweber. 1990. A reconsideration of the genus Pityrogramma (Adiantaceae)
in western North America. American Fern Journal 80: 9–17.

460
Figure 227. Pentagramma. A–C. P. triangularis subsp. triangularis. D, E. P. triangularis subsp. maxonii. F, G. P. triangularis
subsp. viscosa. (From Mickel & Smith, 2004.)

461
DRYOPTERIDACEAE
Phanerophlebia C. Presl
DESCRIPTION: Terrestrial or rupestral; rhizomes
creeping or erect, scaly; leaves monomorphic; petioles with
more than three vascular bundles; laminae 1-pinnate
(rarely simple) with a terminal pinna that resembles
the lateral ones; rachises and costae grooved adaxially,
the grooves more or less confluent at their junctures;
pinnae spinulose-serrulate, at least distally; veins
2–5-branched, free or with 1–3 series of marginal
anastomoses; sori round, in (1–)2–4 series between the
costa and margin; indusium absent or present, if present
then peltate, persistent or shrunken and falling at maturity,
the margins erose; spores monolete; x=41.
DESCRIPCIÓN: Terrestres o rupícolas; rizoma
cortamente rastrero o erecto, escamoso; hojas
monomorfas; pecíolos con más tres haces vasculares;
láminas 1-pinnadas (raramente simples) con una
pinna termnal que asemeja a las laterales; raquises
y costas sulcados adaxialmente, los surcos más o
menos confluentes en las uniones; pinnas espinuloso-
serruladas, al menos distalmente; nervaduras
2–5-ramificadas, libres o con 1–3 series de anastomosis
marginales; soros redondos, en (1–)2–4 series entre la
costa y el margen; indusio ausente o presente, si presente
entonces peltado, persistente o encogido y caedizo al
madurar, los márgenes erosos; esporas monoletes; x=41.

SIMILAR GENERA: Polystichum differs by having an
auricle on the acroscopic side of the pinna base and by a
tapered lamina apex. Nephrolepis is also one pinnate but
has a pinnatifid, tapered lamina apex.
GÉNEROS PARECIDOS: Polystichum difiere por
tener una aurícula en el lado acroscópico de la base de
la pinna y ápices de las láminas gradualmente reducidos.
Nephrolepis es también 1-pinnada pero tiene un ápice
laminar pinnatífido y gradualmente angostado.

COMMENTS: Phanerophlebia is completely neotropical COMENTARIOS: Phanerophlebia es completamente

and contains 9 species (Yatskievych, 1996). It occurs
from the southwestern United States to northern South
America and Hispaniola. All of its species occur in Mexico
except for P. haitiensis C. Chr., from Española y Honduras.
The species generally grow in wet forests above 1000
m, often in rocky habitats. Within dryopteroid ferns,
Phanerophlebia is sister to a clade consisting of Cyrtomium
(Asiatic) and Polystichum. This clade is characterized by
spinulose leaf margins and peltate indusia. The genus
name is derived from the Greek phaneros, evident +
phelps, veins. The veins are easily visible.
neotropical y consta de 9 especies (Yatskievych, 1996).
Se extiende desde el suroeste de los Estado Unidos hasta
América del Sur septentrional y Española. Todos existen
en México con excepción de P. haitiensis C. Chr., de
Española y Honduras. Las especies existen generalmente
en bosques húmedos arriba de 1000 m, a menudo en
ambientes rocosos. Entre los helechos dryopteroides,
Phanerophlebia es hermana a Cyrtomium (Asiático) y
Polystichum. Este clado se caracteriza por márgenes de
las hojas espinulosos e indusios peltados. El nombre
genérico se deriva del griego phaneros, evidente + phelps,
venas. Las venas son fácilmente evidentes.

LITERATURE: Stein, D. B., G. Yatskievych, & D. J. Gastony. 1989. Chloroplast DNA evolution and phylogeny of
some polystichoid ferns. Biochemical Systematics & Ecololgy 17: 93–101. Yatskievych, G. 1996. A revision of the fern genus
Phanerophlebia (Dryopteridaceae). Annals of the Missouri Botanical Garden 83: 168–199. Yatskievych, G., D. B. Stein & D. J.
Gastony. 1988. Chloroplast DNA evolution and systematics of Phaneroplebia (Dryopteridaceae) and related fern genera. Proc.
National Academy of Sciences USA 85: 2589–2593. Yatskievych, G. & R. Riba 1999. Phanerophlebia. Flora de México, vol. 6,
fasc. 4. Consejo Nacional de la Flora de Mexico.

462
Figure 228. A–E. Phanerophlebia juglandifolia. F–L. P. pumila. M–P. P. macrosora. Q, R. P. nobilis. S, T. P. remotispora. (From
Mickel & Beitel, 1988.)

463
THELYPTERIDACEAE
Phegopteris (C. Presl) Fée
DESCRIPTION: Terrestrial; rhizomes 1–4 mm wide,
long-creeping (ours) or erect, pubescent; rhizomes scales
glabrous on the surfaces and margins; petioles with
two vascular bundles; blades 2–3-pinnatifid at the base,
apex gradually reduced; suprabasal pinnae adnate to
rachis and connected by a wing; rachises and costae
not grooved adaxially, scaly and hairy abaxially; buds
absent; veins ending before the margin, free; indument
abaxially of unbranched, acicular unicellular (nonseptate)
hairs, also with minute (ca. 0.1 mm), yellowish, capitate-
glandular hairs; sori round to oblong, supramedial; indusia
absent; sporangial capsules often bearing a capitate gland
or an acicular hair (or both); x=30.
DESCRIPCIÓN: Terrestres; rizomas 1–4 mm de ancho,
largamente rastreros (nuestros) o erectos, pubescentes;
escamas del rizomas glabras en la superficie y
márgenes; pecíolos con 2 haces vasculares; láminas
2–3-pinatifidas en la base, ápice gradualmente reducido;
pinnas suprabasales adnatas al raquis y conectadas
por una ala; ráquises y costas no surcados
adaxialmente; brotes ausentes; venas terminándose
antes del márgen, libres; indumento abaxialmente de
pelos aciculares unicelulares (no septados), también con
pelos capitados-glandulares amarillentosos diminutos
(ca. 0.1 mm); soros redondos a oblongos, supramediales;
indusíos ausentes; cápsulas esporangiales a menudo
llevando una glándula capitada o un pelo acicular (o
ambos); x=30.

SIMILAR GENERA: Other thelypteroid genera differ
by stalked or sessile (not adnate) suprabasal pinnae and
lacking a connecting wing between the pinnae. Also, all
other thelypteroids, except Macrothelypteris, differ by
rachises grooved adaxially and veins terminating at the
lamina margin, not behind it. Macrothelypteris differs by
3-pinnate-pinnatifid laminae and rhizome and petiolar
scales pubescent on the surfaces.
GÉNEROS PARECIDOS: Los otros géneros
thelypteroides difieren por pinnas suprabasales
pecioluladas o sésiles (no adnatas) y faltando una ala
conectora entre las pinnas. Además, todos las otras
theypteroides, con excepción de Macrothelypteris
difieren por y ráquises surcados adaxialmente y las
venas terminándose en el márgen de la lámina, no atrás.
Macrothelypteris difiere por láminas 3-pinnato-pinnatifid
as y escamas de rizomas y pecíolos pubescentes en las
superfícies.

COMMENTS: Phegopteris consists of five species: two COMENTARIOS: Phegopteris consta de cinco especies:
north-temperate and boreal (P. connectilis (curcumboreal) dos en zonas templadas del norte y boreal (P. connectilis
Europe) and P. hexagonoptera (eastern North America)) (circunboreal) and P. hexagonoptera (América del Norte
and three in temperate and subtropical eastern Asia (P. oriental) y tres en Asia oriental templada y subtropical
decursive-pinnata, P. koreana y P. tibetica). All grow mainly (P. decursive-pinnata, P. koreana y P. tibetica). Todos crecen
on shaded forest floors. Phegopteris, Pseudophyegopteris, principalmente en sotobosque sombreado. Phegopteris,
and Macrothelypteris form a clade sister to the rest of Pseudophyegopteris y Macrothelypteris forman un
the Thelypteridaceae (Smith & Cranfill, 2002). The clade clado hermano al resto de Thelypteridaceae (Smith &
is defined by ungrooved rachises and costae, adnate Cranfill, 2002). El clado se define por ráquises y costas
pinnae and/or pinnules, veins ending before the margin, no surcados, pinnas y/o pínnulas anchamente adnatas,
and minute capitate-glandular hairs on the laminae and veinas terminando antes del margen y pelitos capitados-
sporangial capsules. The genus name is derived from glandulares en las láminas y cápsulas esporangiales. El
the Greek phegos, beech tree + pteris, fern. Presumably nombre genérico se deriva del griego phegos, árbol de
referring to a fern that grew beneath beech trees. haya + pteris, helecho. Presumablemente se refiere a un
helecho que crecía bajo de árboles de haya.
LITERATURE: Almeida, T. E., S. Hennequin, H. Schneider, A. R. Smith, J. A. N. Batista, A. J. Ramalho, K. Proite &
A. Salino. 2016. Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of
neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688–700. Holttum, R.
E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophyegopteris, and Macrothelypteris. Blumea 17:
5–32. Schneider, H., L-J. He, J. Marquardt, L. Wang, J. Heinrichs, S. Hennequin & X-C. Zhang. 2013. Exploring the origin
of the latitudinal diversity gradient: Contrasting the sister fern genera Phegopteris and Pseudophegopteris. Journal of Systematics
and Evolution 51: 61–70.

464
Figure 229. A–L. Phegopteris connectilis. M–R. P. hexagonoptera. (Original, ©Robbin Moran, 2016.)

465
POLYPODIACEAE
Phlebodium (R. Br.) J. Sm.
DESCRIPTION: Epiphytes, rupestral, or terrestrial;
petioles with several vascular bundles; rhizome creeping,
usually white-farinose, scaly, scales concolorous, usually
orangish, not clathrate, denticulate; leaves pinnatisect
or 1-pinnate, often glaucous beneath, monomorphic,
articulate to the rhizome; pinnae scaly or glabrous
beneath, the margins thickened and cartilaginous, entire or
near so; rachises and costae glabrous adaxially; veins
areolate, the areoles with or without included veinlets;
sori round, without paraphyses, typically supplied by
two included veinlets, in 1–7 series between the costa
and margin; spores monolete, ellipsoid, yellow; x=37.
SIMILAR GENERA: Pecluma and the Polypodium dulce
group differ by pubescent rhizome scales (at least at the
point of attachment), rachises and costae puberulent
adaxially, veins free (most species), and sporangial
capsules often setulose. Polypodium differs by free veins.
Serpocaulon differs by clathrate rhizome scales.
COMMENTS: Phlebodium is entirely neotropical and
typically occurs as an epiphyte in wet forests from 0–2500
m. It is sister to a clade consisting of Pecluma and the
Polypodium dulce group. Phlebodium consists of four species:
P. aureum, P. decumanum, P. inaequale, and P. pseudoaureum.
With the exception of P. inaequale, the sori are usually
supplied by two included veinlets—a unique character
in the Polypodiaceae (only one vein supplying the sorus
is the typical condition). A key to the species is given by
Tejero-Díez et al. (2009). Phlebodium aureum is a fertile
allotetraploid of hybrid origin between P. decumanum and
P. pseudoaureum. In Mexico, Guatemala, and Honduras, the
rhizomes are broken and sold in markets. It is commonly
called “calaguala.” It is prepared as a tea, which is believed
to cure kidney disorders and other maladies. The genus
name is derived from the Greek phlebodes, full of veins.
The veins are conspicuously netted.
LITERATURE: Tejero-Díez, J. D., J. T. Mickel & A. R. Smith. 2009. A hybrid Phlebodium (Polypodiaceae; Polypodiophyta)
and its influence on the circumscription of the genus. American Fern Journal 99: 109–116.
466
DESCRIPCIÓN: Epífitas, rupícolas o terrestres;
pecíolo con varias haces vasculares; rizoma rastrero,
generalmente blanco-farinoso, escamoso, escamas
concoloras, generalmente anaranjadas, no clatradas,
denticuladas; hojas pinnatisectas a 1-pinnatas, a menudo
glaucas en el envés, monomorfas, articuladas al rizoma;
pinnas escamosas o glabras en el envés, los márgenes
engrosados y cartilaginosos, enteros o casi enteros;
ráquises y costas glabros adaxialmente; nervaduras
areoladas, las aréolas con o sin nérvulos incluidos; soros
redondeados, sin parafisos, típicamente suministrados
por dos venillas incluidas, en 1–7 series entre la costa y
el margen; esporas monoletes, elipsoides, amarillas; x=37.
GÉNEROS PARECIDOS: Pecluma y el grupo de
Polypodium dulce difieren por escamas del rizoma
pubescentes (al menos al punto de unión), requises
y costas puberulentos adaxialmente, venas libres (la
mayoría de las especies) y cápsulas de los esporangios
a menudo setulosas. Polypodium difiere por venas libres.
Serpocaulon difiere por escamas del rizoma clatradas.
COMENTARIOS: Phlebodium es completamente
neotropical y típicamente es un epífito en bosques
húmedos de 0–2500 m. Phlebodium es grupo hermana a
un clado que consta de Pecluma y el grupo de Polypodium
dulce. Phlebodium consta de cuatro especies: P. aureum,
P. inaequale, P. decumanum y P. pseudoaureum. Con
excepción de P. inaequale, los soros están usualmente
suministrados por dos venillas incluidas—un carácter
único en las Polypodiaceae (solo una venilla sumistrando
el soro es la condición típica). Una clave a las especies
se encuentra en Tejero-Díez et al. (2009). Phlebodium
aureum es un allotetraploide fértil de origen híbrido entre
P. decumanum y P. pseudoaureum. En México, Guatemala
y Honduras los rizomas de son fragmentados y vendidos
en los mercados. Se llama comúnmente “calaguala.”
Se preparan en una infusión que, según se cree, alivia
trastornos renales y otros males. El nombre genérico se
deriva del griego phlebodes, lleno de venas. Las venas son
anastomosadas conspicuosamente.
Figure 230. A–B, F. Phlebodium psuedoaureum. D, E. P. decumanum. (From Mickel & Smith, 2004.)

467
LYCOPODIACEAE
Phlegmariurus Holub
DESCRIPTION: Plants terrestrial or epiphytic,
pendant, erect, or ascending; horizontal creeping stems
absent; stems isodichotomously branched, the
branches all similar, or in some terrestrial species lightly
differentiated into prostate versus erect aerial branches;
roots descending through the cortex to the stem base
and emerging in one basal tuft; leaves homophyllous or
heterophyllous, entire or (less rarely) minutely toothed;
gemmae and gemmiphores absent; strobili absent, the
sporangia produced in zones along the shoot; sporophylls
either similar to the trophophylls or more reduced
and appressed, remaining green after the sporangia
dehisce; sporangia isovalvate; sporophylls similar to the
trophophylls, attached at the base (not peltate), remaining
green after sporangial dehiscence; spores foveolate on
distal face, smooth on proximal, in polar view rounded
with convex sides and angles acute; gametophytes
subterranean, cylindrical, with pluricellular hairs among
the sporangia; x=68.
DESCRIPCIÓN: Plantas terrestres o epífitas, pendientes,
erectas o ascendentes; tallos horizontales reptantes
ausentes; tallos ramificados isodicotómicamente,
las ramas simulares, o en algunas especies terrestres
levemente diferenciadas en ramas aéreas prostrados
versus erectas; raíces descendientes por la corteza hasta
la base del tallo y emergiéndose en un grupo basal;
hojas homofílicas o heterofílicas, enteras o (raras veces)
dentadas dimminutamente; yemas y gemíferos ausentes;
yemas y gemíforos ausentes; estróbilos ausentes, los
esporangios se nacen en zonas por el vástago; esporofílios
simulares a los trofofílios o reducidos y apresos, únidos
en la base (no peltados), permaneciéndose verdes
después de la dehiscencia; esporangios con células
sinuosas; esporas foveoladas en la cara distal, lisas en la
cara proximal, en vista polar redondeadas, con lados
convexos y ángulos agudos; gametofitos subterráneos,
cilíndricos, con pelos pluricelulares entre los esporangios;
x=68.

SIMILAR GENERA: Huperzia differs by the presence
gemmae and spores in polar view that are rounded with
convex sides and smooth (not foveolate) proximal faces.
The other genera of Lycopodiaceae differ by having long-
creeping horizontal branches.
GÉNEROS PARECIDOS: Huperzia difiere por la
presencia de yemas y esporas en vista polar anguladas con
lados cóncavos y caras proximales lisas (no foveoladas).
Los otros géneros de Lycopodiaceae difieren por tener
ramas horizontales principales largamente rastreras.

COMMENTS: Phlegmariurus consists of about 300
species worldwide and is pantropical. Its greatest diversity
is in the Andes. Øllgaard (1987) outlined various species
groups (as “Huperzia”) based on morphology.The diversity
in growth habit is remarkable, with stiff erect terrestrial
species and lax long-pendent ones. The terrestrial habit
has evolved independently several times from epiphytic
ancestors (Wikstöm et al., 1999). The genus name is
derived from the Greek phlegma, flame, and oura, tail. It
refers to the tassel-like fertile portions of some species.
COMENTARIOS: Phlegmariurus consta de casi 300
especies mundial y es pantropical. Su diversidad mayor
está en los Andes. Øllgaard (1987) bosquejó varios
grupos de especies (como “Huperzia”) basado en la
morfología. La diversidad en forma de crecimiento es
notable, con especies rígidas, erectas terrestres y otras
laxas y largamente péndulas. El hábito terrestre se
evolucionó independiente varias veces de ancestros
epífitos (Wikstöm et al., 1999). El nombre genérico se
deriva del griego phlegma, llama, y oura, cola. Se refiere a
las porciones fértiles como una borla de algunas especies.

LITERATURE: Øllgaard, B. 1988. Lycopodiaceae. In: G. Harling & L. Andersson (eds.). Flora of Ecuador 33: 1–155. Øllgaard,
B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the Missouri Botanical Garden 79: 687–717. Øllgaard, B. 2012.
New combinations in Neotropical Lycopodiaceae. Phytotaxa 57: 10–22. Rolleri, C. H. 1980. Sinopsis de las especies de Ly-
copodium de la sección Crassistachys Herter. Revista Museo de La Plata, Sección Botánica. n.s., 13(71): 61–113. Testo, W., B.
Øllgaard, A. Field, T. Almeida, M. Kessler & D. Barrington. 2018. Phylogenetic systematics, morphological evolution, and
natural groups in Neotropical Phlegmariurus (Lycopodiaceae). Molecular Phylogenetics and Evolution https://doi.org/10.1016/j.
ympev.2018.03.016. Wagner, W. H. & J. M. Beitel. 1992. Generic classification of modern North American Lycopodiaceae.
Annals Missouri Botanical Garden 79: 676–686. Wikstöm, N., P. Kendrick & M. Chase. 1999. Epiphytism and terrestrialization
in tropical Huperzia. (Lycopodiaceae). Plant Systematics and Evolution 218: 221–243. Winther, J. L. & W. E. Friedman. 2008.
Arbuscular mycorrhizal associations in Lycopodiaceae. New Phytologist 177: 790–801.

468
Figure 231. A. Phlegmariurus hippurideus. B. P. crassus. C, D. P. pithyoides. E, F. P. reflexus. (From Mickel & Smith, 2004.)

469
MARSILEACEAE
Pilularia L.
DESCRIPTION: Rooted aquatics; rhizomes usually long-
creeping, slender, sparsely pubescent; leaves 1–10 cm
long, filiform, terete, pubescent or glabrate; sporocarps
subterranean, borne at the base of the petioles,
densely pubescent, indurate, globose, brown to black,
splitting into 4 valves (sporangial compartments); spores
of two types (plants heterosporous): large megaspores
and smaller microspores, these trilete, endosporic; x=10.
DESCRIPCIÓN: Acuáticas radicadas en lodo;
rizomas usualmente largamente rastreros, delgados,
esparcidamente pubescentes; hojas 1–10 cm de
largo, filiformes, teretes, pubescentes o glabras;
esporocarpos subterráneos, en la base de los
pecíolos, densamente pubescentes, indurados,
globosos, pardos a negros, hendiéndose en 4 valvas
(compartimentos esporangiales); esporas de dos
tipos (plantas heterospóricas): megaesporas grandes y
microesporas más pequeñas, estos triletes, endoesporicas;
x=10.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Pilularia consists of 6 species. It occurs
in. North America, Mesoamerica, South America, Europe,
Australia, New Zealand, Pacific Islands. A single species
of Pilularia occurs in the Neotropics: P. americana A. Br.
It grows in wet or seasonally wet habitats (shorelines,
ditches, ephemeral ponds) and can be fully submersed
but produces sporocarps only on emergent plants. The
plants form dense colonies but are often overlooked
because of their small, grass-like leaves. Colonies are
usually discovered around the margins of ponds and
marshes after the water level has receded.The fiddlehead
at the tip of the leaf is the best characteristic to distinguish
it from similar plants. Nagalingum et al. (2008) found that
P. americana was sister to P. novae-hollandiae and that the
two had low (compared to ferns in general) sequence
divergences. In North America, they found eastern
and western populations of P. americana differed for
the markers studied. The South American P. mandonii A.
Br. was not uncluded in their study. The genus name is
derived from the Latin pilula, little ball, and refers to the
globose sporocarps.
COMENTARIOS: Pilularia consta de 6 especies. Existe
en América del Norte, Mesoamérica, Sudamérica,
Europa, Australia, Nueva Zelanda, Islas Pacíficas. Una sola
especie de Pilularia existe en el Neotrópico: P. americana
A. Br. Existe hábitats mojadas o estacionalmente así
(orillas de lagos, zanjas, charcos efímeros), y se puede ser
completamente anegado pero produce esporocarpos
solamente en plantas emergidas. Las plantas forman
colonias densas, pero están pasado por alto a menudo
a causa de sus hojas pequeñas que se asemejan a
gramíneas. Usualmente se descubren colonias por las
orillas de chacras y pantanos cuando el nivel de agua ha
retrodicido. El cayado (prefoliación) en el ápice de las
hojas es la característica mejor para distinguirlo de otras
plantes similares. Nagalingum et al. (2008) hallaron que P.
Americana was sister to P. novae-hollandiae y que los dos
tuvieron divergencias de secuencias bajas (comparado
con otros helechos). En América del Norte, hallaron
que poblaciones orientales y occidentales difieren en
los marcadores estudiados. Pilularia mandonii A. Br., una
especie de América del Sur, no fue incluida en su estudio.
El nombre genérico se deriva del latín pilula, pequeña
pelota, y se refiere a los esporocarpos globosos.

LITERATURE: Dennis, W. M. & D. H. Webb. 1981. The distribution of Pilularia americana A. Br. (Marsileaceae) in North
America, north of Mexico. Sida 9: 19–24. Large, M. F. & J. E. Braggins. 1989. An assessment of characters of taxonomic
significance in the genus Pilularia (Marsiliaceae): with particular reference to P. americana, P. novae-hollandiae, and P. novae-
zelandiae. New Zealand Journal of Botany 27: 481–486. León, B., H. Beltrán, C. Carrasco-Badajoz, E. Portal-Quicaña
& M. Huaycha-Allcca. 2018. First record of Pilularia americana A. Braun (Polypodiidae, Salviniales, Marsileaceae) from Peru.
Checklist 14: 319–322. Nagalingum, N. S., M. D. Nowak & K. M. Pryer. 2008. Assessing phylogenetic relationships in extant
heterosporous ferns (Salviniales), with a focus on Pilularia and Salvinia. Botanical Journal of the Linnean Society 157: 673–685.

470
Figure 232. A–C. Pilularia americana A. Br. (From Mickel & Smith, 2004.)

471
PTERIDACEAE
Pityrogramma Link
DESCRIPTION: Terrestrial; rhizomes decumbent or
erect, scaly; sterile and fertile leaves monomorphous;
petioles often dark and lustrous, grooved on the upper
surface; laminae pinnatisect to 4-pinnate-pinnatifid,
farinose beneath, the farina usually white or yellow,
scales rare or absent, sometimes pubescent, the apex
pinnatifid, tapered; veins free; sporangia borne along
the veins, often confluent at maturity; indusia absent;
x=30, 58, 60.
DESCRIPCIÓN: Terrestres; rizomas decumbentes o
erectos, escamosos; hojas estériles y fértiles monomorfas;
pecíolos a menudo oscuros y lustrosos, surcados en la
superficie superior; láminas pinnatisectas a 4-pinnado-
pinnatífidas, farinosas abajo, la farina usualmente blancas
o amarillas abajo, escamas escasas o ausentes, algunas
veces pubescentes, el ápice pinnatífido, gradualmente
reducido; nervaduras libres; esporangios nacen a lo
largo de las venas, a menudo confluentes en la madurez;
indusios ausentes; x=30, 58, 60.

SIMILAR GENERA: Argyrochosma differs by elliptic GÉNEROS PARECIDOS: Argyrochosma se distingue

segments and terete (not grooved) petioles. The plants
are usually less than 30 cm long and grow in rocky habitats.
Pentagramma, a genus of the southwestern United States
and Mexico, differs by its terete petioles, triangular
laminae, and pinnae broadly adnate to the rachis.
por segmentos elípticos y pecíolos teretes (no surcados).
Las plantas miden usualmente menos que 30 cm de largo
y crecen en habitats rocosos. Pentagramma, un género
del suroeste de los Estados Unidos y México, difiere
por sus pecíolos teretes, láminas triangulares y pinnas
anchamente adnatas al ráquis.

COMMENTS: Pityrogramma has about 19 species, all
neotropical. They typically grow in open or semi-open
places where there has been recent disturbance such as
road banks, landslides, and gardens. The most common
and widespread species below 1000 m in the Neotropics
is P. calomelanos (L.) Link. It has become naturalized in the
Old World tropics is and spreading rapidly there. Above
1000 m in the Neotropics, it is replaced by P. ebenea
(L.) Proctor. Hybrids often occur where two or more
species of Pityrogramma grow together (Gómez, 1979).
Pityrogramma trifoliata (L.) R.Tryon is sometimes placed in
its own genus, Trismeria, because of its distinctive lamina
architecture (digitate pinnae and three-dimensional
leaves), bicolorous rhizome scales, and spores that lack
an equatorial ridge. The genus name is derived from
the Greek pityron, scurf, bran + gramme, line. The lower
surface of the lamina is covered with a farina, and the
sporangia are in lines along the veins.
COMENTARIOS: Pityrogramma tiene casi 19
especies, todos neotropicales. Crecen típicamente en
lugares abiertos o semi-abiertos donde ha estado una
perturbancia reciente, como en lados de los caminos,
deslaves, y jardines. La especie más común y difundida
abajo de 1000 m en el Neotrópico es P. calomelanos (L.)
Link. Ha llegado a ser naturalizada en el Viejo Mundo
y se difunde rápidamente allá. Arriba de 1000 m en el
Neotrópico, está reemplazado por P. ebenea (L.) Proctor.
A menudo se encuentran híbridos en lugares donde
dos o más especies de Pityrogramma crecen juntas
(Gómez, 1979). A veces Pityrogramma trifoliata (L.) R.
Tryon se ubica en su propio género, Trismeria, debido a
su arquitectura laminar distinta (pinnas digitadas y hojas
en tres dimensiones), escamas del rizoma bicoloras,
y esporas que faltan un reborde ecuatorial. El nombre
genérico se deriva del griego pityron, caspilla + gramme,
línea. El envés de la lámina es cubierto por una farina, y
los esporangios están en líneas.

LITERATURE: Domin, K. 1928. Generis Pityrogramma (Link) species ac sectiones in clavem analyticam dispositae. Spisy
vydávanè přirodovědeckou fakultou Karlovy university 88: 1–10. Gómez, L. D. 1979. Contribuciones a la pteridología
Costarricense XIII. Novitates. Brenesia 16: 95–100. Tryon, R. M. 1962. Taxonomic fern notes. II. Pityrogramma (including
Trismeria) and Anogramma. Contributions from the Gray Herbarium of Harvard Universtiy 189: 52–76. Nakazato, T. & G. J.
Gastony. 2003. Molecular phylogenetics of Anogramma species and related genera (Pteridaceae: Taenitidoideae). Systematic
Botany 28: 490–502.

472
Figure 233. Pityrogramma calomelanos (L.) Link (From Mickel and Smith, 2004.)

473
PLAGIOGYRIACEAE
Plagiogyria (Kunze) Mett.
DESCRIPTION:Terrestrial; rhizomes erect, subterranean
or forming a trunk to 1 meter tall, without hairs or
scales; sterile and fertile leaves dimorphic; petiole
bases with 3 vascular bundles, these uniting distally
to form a “U”; laminae lanceolate or elliptic; segments or
pinnae adnate to the rachis; mostly approximate, almost
all spreading, the proximal ones at times reflexed, the
distal ones at times ascending or falcate; sterile laminae
pinnatisect, the segments adnate, the margins serrate
or doubly serrate for most of their length; fertile laminae
pinnatisect or pinnate, the segments adnate or the pinnae
narrowly united, their margins entire or irregular, thin,
frequently involute; x=66.
DESCRIPCIÓN: Terrestres; rizomas erectos,
subterráneos o formando un tronco hasta 1 metro
de altura, sin pelos ni escamas; hojas estériles y
fértiles dimorfas; base de los pecíolos con 3 haces
vasculares, estos uniéndose distalmente en forma
de U; láminas lanceoladas o elípticas; segmentos o
pinnas adnatos al raquis, en su mayoría próximos, casi
todos patentes, los proximales a veces reflexos, los
distales a veces ascendentes o falcados; láminas estériles
pinnatisectas, los segmentos adnatos, sus márgenes
en su mayoría serrados o 2-serrados; láminas fértiles
pinnatisectas o pinnadas, los segmentos adnatos o las
pinnas angostamente unidas, sus márgenes enteros o
irregulares, delgados, a menudo involutos; x=66.

SIMILAR GENERA: Blechnum differs by scaly rhizomes,
4 or more vascular bundles in the petioles, reddish young
leaves, and sori in two rows along the costae. Other
similar genera have pinnae that are sessile but not adnate
to the rachis.
GÉNEROS PARECIDOS: Blechnum difiere por rizomas
escamosos, 4 o más haces vasculares en los pecíolos,
hojas jóvenes rojizas, y los soros en dos líneas en cada
lado de las costas. Otros géneros parecidos tienen pinnas
séssiles pero no adnatas al raquis.

COMMENTS: Plagiogyria is pantropical and contains COMENTARIOS: Plagiogyria es pantropical y consta

about 10 species (Zhang & Nooteboom, 1998). It is
usually found above 2000 m in open or semi-open
habitats. Only one species (P. pectinata (Liebm.) Lellinger)
is said to occur in the New World, but likely more exist.
Plants in Mesoamerica have short subterranean rhizomes,
whereas those from Bolivia have thick trunks up to one
meter tall and can easily be mistaken for an arborescent
species of Blechnum. The genus name is derived from
the Greek plagios, oblique + gyros, ring. It refers to the
oblique annulus.
de casi 10 especies (Zhang & Nooteboom, 1998).
Usualmente ocurre arriba de 2000 m en ambientes
abiertos o semi-abiertos. Se dicen que solo una especie
(P. pectinata (Liebm.) Lellinger) existe en el Nuevo Mundo,
pero sin duda más existe. Plantas en Mesoamérica tienen
rizomas cortos subterráneos, mientras que las de Bolivia
tienen trocos gruesos hasta un metro de altura, y se
puede confundir fácilmente con una especie arborescente
de Blechnum. El nombre genérico se deriva del griego
plagios, oblicuo + gyros, anillo. Se refiere al anillo oblicuo.

LITERATURE: Copeland, E. B. 1929. The fern genus Plagiogyria. Philippine Journal of Science 38: 377–415. Lellinger, D.
B. 1971. The American species of Plagiogyria sect. Carinatae. American Fern Journal 61: 110–118. Zhang, X.-C. & H. P.
Nooteboom. 1998. A taxonomic revision of Plagiogyriaceae (Pteridophyta). Blumea 43: 401–469.

474
Figure 234. A–E. Plagiogyria pectinata (From Mickel & Smith, 2004.)

475
POLYPODIACEAE
Platycerium Desv.
DESCRIPTION: Epiphytes; rhizomes short-creeping,
scaly, often hidden by the sterile leaves; sterile and
fertile leaves strongly dimorphic, the sterile (nest
leaves) erect, appressed, overlapping, rounded to lobed
on the distal margins, brown when mature, the fertile
leaves elongate, arching to pendant, deeply lobed, green
when mature; laminae pubescent throughout, the hairs
stellate; veins reticulate; sori acrostichoid; x=37.
DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
escamosos, a menudo ocultos por las hojas estériles;
hojas estériles y fértiles fuertemente dimorfas,
las estériles (hojas nidiformes) erectas, apresas,
traslapándose, redondeadas a lobuladas por el margen
distal, pardas cuando maduras, las hojas fértiles alargadas,
arcuadas a péndulas, divididas en lóbulos alargados,
verdes cuando maduras; láminas pubescentes en todas
partes, los pelos estrellados; nervadura reticulada;
soros acrosticoides; x=37.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Platycerium contains 18 species and COMENTARIOS: Platycerium consta de 18 especies y

occurs mostly in tropical Asia and Africa. Only one species
occurs in the Neotropics: P. andinum Baker. It grows in wet
forests in Peru and Bolivia, from 200–800 m. The nest
leaves, which are dead at functional maturity, are thick
with a water-absorbing mesophyll that helps the plant
survive droughts. The nest leaves are also said to collect
humus into which the plant grows roots to absorb water
and mineral nutrients. Many species are cultivated, and
in horticulture, the plants are popularly called “staghorn
ferns.” The genus name is derived from the Greek,
platys, flat, and keras, horn; referring the segments, which
resemble flattened horns.
existe mayormente en Asia y África tropical. Solamente
una especie existe en el Neotrópico: P. andinum Baker.
Crece en bosques húmedos en Peru y Bolivia, desde 200–
800 m. Las hojas nidiformes, las cuales están muertas a la
madurez funcional, son gruesas y presentan un mesófilo
que absorbe agua que ayuda la planta sobrevivir la sequía.
También se dicen que las hojas nidiformes colectan humus
en que la planta echa las raíces para absorber agua y
nutrientes minerales. Se cultivan muchas especies, y en
horticultura se llaman popularmente “cuerno de alce.”
El nombre genérico se deriva del griego platys, plano, y
keras, cuerno. Se refiere a los segmentos, los cuales se
asemejan a un cuerno achatado.

LITERATURE: Hennipman, E. & M. C. Roos. 1982. A monograph of the fern genus Platycerium (Polypodiaceae). Verhan-
delingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkund, Tweede Reeks, Deel 80: 1–126. Hoshi-
zaki, B. J. 1972. Morphology and phylogeny of Platycerium species. Biotropica 4: 93–117. Janssen, T. & H. Schneider. 2005.
Exploring the evolution of humus collecting leaves in drynarioid ferns (Polypodiaceae, Polypodiidae). Plant Systematics and
Evolution 252: 175–197.

476
Figure 235. Platycerium andinum. A. Habit, with erect sterile and pendent fertile leaves. B. Fertile leaf, turned to show
acrostichoid sori. C. Stellate hairs on lamina, below mixed with sporangia and thus appearing darker. D. Petiole, transverse
section. E. Sterile leaf, base with epidermis partially removed to reveal roots in the mesophyll. F. Lamina, adaxial surface with
stellate hairs. G. Lamina, abaxial surface with stellate hairs. (Original, ©Robbin Moran, 2014.)

477
POLYPODIACEAE
Pleopeltis Humb. & Bonpl. ex Willd.
DESCRIPTION: Epiphytic or rupestral; rhizome scales
clathrate (at least medially; the cell luminae sometimes
obscured), peltate, with or without rhizoides; sterile and
fertile leaves monomorphous or dimorphous; petioles
terete (not grooved adaxially), articulate to short
phyllopodia; laminae simple to 1-pinnate basally, rarely
more divided, scaly, the scales circular, peltate scales,
these clathrate at least centrally; veins anastomosing,
often hard to see because of the thick lamina; sori borne
at the juncture of several veinlets, non-indusiate, round or
rarely elongate or linear, in one row between the rachis
(or costa) and margin, young sori typically protected
by overlapping, circular, peltate scales; spores yellow,
monolete; x=34, 35, 37?
DESCRIPCIÓN: Epífiticas o rupestres; escamas del
rizomas clatradas (a lo menos medialmente; las lúminas
de las células a veces oscurridas), peltadas, sin o con
rizoides; hojas estériles y fértiles monomorfas o dimorfas;
pecíolos teretes (no surcados adaxialmente), articulados
al filopódios cortos; láminas simples a 1-pinnadas
basalmente, raras veces más divididas, escamosas, las
escamas circulares, peltadas, clatradas al menos en
el centro; venas anastomosadas, a menudo difíciles
ver a causa de la lámina gruesa; soros se nacen en las
uniones de varias venillas, sin indusios, redondos raras
veces alargados o lineares, en una sola hilera entre el
raquis (a costa) y margen, soros jóvenes típicamente
protegidos por escamas traslapadas, circulares,
peltadas; esporas bilaterales, amarillas; x=34, 35, 37?

SIMILAR GENERA: Other genera of Polypodiacae
differ by non-peltate, non-clathrate laminar scales or the
complete lack of laminar scales. Polypodium further differs
by (usually) free veins and non-clathrate rhizome scales.
Microgramma also differs by non-clathrate rhizome scales.
GÉNEROS PARECIDOS: Otros géneros de las
Polypodiaceae difieren por escamas laminares no peltadas
y no clatradas o la falta completa de escamas laminares.
Polypodium difiere además por (usualmente) venas libres
y escamas del rizoma no clatradas. Microgramma, difiere
también por escamas del rizoma no clatradas.

COMMENTS: Pleopeltis contains about 90 species COMENTARIOS: Pleopeltis consta de casi 90 especies
and is one of the largest, if not the largest, genus in its y es uno de los más grandes, si no lo más grande, géneros
family. It is primarily neotropical, with a few species from en su familia. Es principalmente neotropical, con unas
Africa to India and Sri Lanka. Most of the species occur especies desde África hasta India y Sri Lanka. La mayoría
in Central America. It grows in many habitats, from de las especies existen en América Central. Ocurre en
coffee plantations to undisturbed virgin forests. Nectaries muchos habitats, desde cafetales hasta bosques húmedos
(modified hydathodes) occur on the acroscopic base of vírgenes. Nectarios (hidatodos modificados) ocurren en
the segments and in simple-leaved species at the base la base acroscópica de los segmentos y en las especies
of the laminae. The peltate, clathrate lamina scales are con hojas enteras en la base de la lámina. Las escamas
unique in the family. The genus name is derived from the peltatadas clatradas son únicas en la familia.El nombre
Greek pleos, full, and pelte, shield. It refers to the peltate genérico se deriva del griego pleos, lleno, y pelte, escudo.
scales covering the young sori. Se refiere a las escamas peltadas que cubren los soros
jóvenes.
LITERATURE: Hooper, E. A. 1994. New combinations in the Pleopeltis macrocarpa group (Polypodiaceae: Polypidieae).
American Fern Journal 85: 75–82. de la Sota, E. R. 1966. Revisión de las especies Americanas del grupo “Polypodium
squamatum” L. “Polypodiaceae” (s. str.). Revista del Museo de La Plata, Sección Botánica 10: 69–186, t. I–VII. Maxon, W. R. 1916.
Studies of tropical American ferns–No. 6. Polypodium furfuraceum and related species. Contributions of the Gray Herbarium
of Harvard University 17: 557–579; Polypodium squamatum and its allies. Contr. U.S. Natl. Herb. 17: 579–596. Mickel, J. T. &
J. M. Beitel. 1987. Notes on XPleopodium and Pleopeltis in tropical America. American Fern Journal 77: 16–27. Otto, E. M.,
T. Janssen, H.-P. Kreier & H. Schneider. 2009. New insights into the phylogeny of Pleopeltis and related neotropical genera
(Polypodiaceae, Polypodiopsida). Molecular Phylogenetics and Evolution 53: 190–201. Smith, A. R. & D. Tejero-Díez. 2014.
Pleopeltis (Polypodiaceae), a redefinition of the genus and nomenclatural novelties. Botanical Sciences 92: 43–58. Weatherby, C.
A. 1922. The group of Polypodium lanceolatum in North America. Contributions of the Gray Herbarium of Harvard University
65: 3–14. Weatherby, C. A. 1939. The group of Polypodium polypodioides. Contributions of the Gray Herbarium of Harvard
University 124: 22–35.

478
Figure 236. Pleopeltis. Note the peltate scales, which distinguish the genus from other Polypodiaceae. Fijase las escamas peltadas
que distinguen el género de otros de las Polypodiaceae. A–D. Pleopeltis polylepis var. interjecta. E–H. P. polylepis var. erythrolepis.
J–M. P. polylepis var. polylepis. N. P. angusta var. angusta. O–R. P. angusta var. stenoloma. (From Mickel & Smith, 2004.)

479
POLYPODIACEAE
Pleopeltis (Dicranoglossum J. Sm. group)
DESCRIPTION: Epiphytes; roots dense, densely
brown pubescent, forming a spongy mass; rhizomes
short-creeping, scaly, the scales lanceolate, bicolorous;
sterile and fertile leaves monomorphous, bifurcate
subdichotomously into several segments, sparsely
scaly; veins free or (one species) areolate with included
veinlets; sori submarginal, rounded to elongate or
linear, partially covered by the revolute segment margins;
paraphyses absent; spores bilateral, yellow; x=36.
DESCRIPCIÓN: Epífitas; raíces densas, densamente
pubescentes con pelitos pardos, formando una masa
esponjiosa; rizoma cortamente rastrero, escamoso, las
escamas lanceoladas, bicoloras; hojas monomorfas,
bifurcadas subdicotómicamente en varios
segmentos, esparcidamente escamosas; nervaduras
libres o (una especie) anastomosadas con nérvulos
incluidos; soros submarginales, redondeados, alargados
a lineares, parcialmente cubiertos por los márgenes
laminares revolutos; parafisos ausentes; esporas bilaterales,
amarillas; x=36.

SIMILAR GENERA: The other species of Pleopeltis GÉNEROS PARECIDOS: Los otros especies de
differ by entire or pinnate blades. Pleopeltis típicamente tienen láminas enteras o 1-pinnadas.

COMMENTS: Dicranoglossum is entirely neotropical and COMENTARIOS: Dicranoglossum es completamente

contains 4 species, all of which are epiphytes en forests
from 0–1000(–1500) m. The group is monophyletic and
nested within Pleopeltis. All the species have a dense mass
of spongy roots that hold water. Dicranoglossum desvauxii
is the only species in the genus with anastomosing veins.
The name Dicranoglossum is derived from the Greek
dikranos, two-pronged + glossa, tongue. It refers to the
bifurcations of the laminae into tongue-like lobes.
neotropical y contiene 4 especies, todas de las cuales
son epifitas en bosques de 0–1000(–1500) El grupo es
monofilético y anidado dentro de Pleopeltis. Todas tienen
una masa densa esponjiosa de raíces que cargan agua.
Dicranoglossum desvauxii es la única especie del género
con venas anastomosadas. El nombre Dicranoglossum
se deriva del griego dikranos, dos proyecciones + glossa,
lengua. Refiere a las bifurcaciones de las láminas en
lóbulos que asemejan lenguas.

LITERATURE: Christensen, C. 1929.Taxonomic fern studies. I. Revision of the polypoidioid genera with longitudinal coenosori
(Cochlidiinae and “Drymoglossinae”); with a discussion of their phylogeny. Dansk Botanisk Arkiv 6(3): 1–93.

480
Figure 237. Dicranoglossum desvauxii (Klotzsch) Proctor (From Tryon & Stolze, 1993.)

481
DRYOPTERIDACEAE
Polybotrya Willd.
DESCRIPTION: Plants hemiepiphytic or (in 2 species)
terrestrial; stem long-creeping in the hemiepiphytic species,
short-creeping in the terrestrial ones, in cross-section
with 4–10 circularly arranged vascular bundles,
each surrounded by a dark sclerenchymatous
sheath; sterile and fertile leaves strongly dimorphic;
laminae 1-pinnate to 4-pinnate, the apex pinnatifid or
rarely (in South American populations of P. polybotryoides)
subconform; veins free or (in subgen. Soromanes)
anastomosing; fertile leaves like a skeletonalized version
of the sterile ones; indusia absent; x=41.
DESCRIPCIÓN: Plantas hemiepífitas o (en 2 especies)
terrestres; tallo largamente rastrero en las especies
hemiepífitas, cortamente rastrero en las terrestres, en
sección transversal con 4–10 haces vasculares
dispuestos circularmente, cada uno rodeado por
una vaina oscura esclerenquimatosa; hojas estériles
y fértiles marcadamente dimorfas; láminas 1-pinnadas
a 4-pinnadas, el ápice pinnatífido o raramente (en
poblaciones de América del Sur de P. polybotryoides)
subconformes; nervaduras libres o (en subgén. Soromanes)
anastomosadas; hojas fértiles semejan a un esqueleto de
las hojas estériles; indusios ausentes; x=41.

SIMILAR GENERA: Lomagramma, Lomariopsis, Maxonia GÉNEROS PARECIDOS: Lomagramma, Lomariopsis,

resemble Polybotrya by their hemiepiphytic habit and
sterile-fertile leaf dimorphy. The first two genera can be
distinguished by their conform lamina apices; the third
genus by its indusiate sori and glabrous adaxial grooves
of the leaf axes.
Maxonia se parecen a Polybotrya por sus hábitos
hemiepifíticos y dimorfismo foliar. Los dos primeros
géneros pueden distinguirse por sus ápices conformes
de la lámina; y el tercero por sus soros con indusios y los
surcos adaxiales glabros de la hoja.

COMMENTS: Polybotrya contains 35 species and is COMENTARIOS: Polybotrya consta de 35 especies y

entirely neotropical. It is most diverse in the Andes, where
23 species occur, 12 of which are endemic. Southeastern
Brazil has also played a role in the diversification of
Polybotrya because it has six species, all endemic. The
fertile leaves are ephemeral and held more erect than
the sterile leaves. Polybotrya forms a clade with Cyclodium,
Maxonia, Olfersia, and Polystichopsis. This clade is united
by creeping rhizomes. Often its species are climbing and
have dimorphic sterile and fertile leaves (or subdimorphic
in Cyclodium). Hybrids are known between Cyclodium and
Polybotrya. The genus name is derived from the Greek
poly, many + botrys, bunches, clusters. The sori appear as
many bunches or clusters on the fertile leaves.
es completamente neotropical. Es más diverso en los
Andes, donde se encuentran 23 especies, 12 de las cuales
son endémicas. Suroeste de Brasil ha hecho un papel
importante en la diversificación de Polybotrya porque
tiene 6 especies, todas endémicas. Las hojas fértiles son
efímeras y son más rectas que las estériles. Polybotrya
forma un clado con Cyclodium, Maxonia, Olfersia y
Polystichopsis. Este clado se une por rizomas reptantes.
A menudo sus especies son trepadores y tienen hojas
estériles y fértiles dimórfas (subdimorfas en Cyclodium).
Hybrids are known between Cyclodium and Polybotrya.
El nombre genérico se deriva del griego poly, muchos
+ botrys, grupitos. Los soros aparecen como muchos
grupitos en las hojas fértiles.

LITERATURE: Canestraro, B. K. & P. H. Labiak. 2015. The fern genus Polybotrya (Dryopteridaceae) in the Atlantic Forest
of Brazil, with the description of a new species. Brittonia 67: 216–220. Engels, M. E. & B. K. Canestraro. 2017. XCyclobotrya:
a new hybrid genus between Cyclodium and Polybotrya (Dryopteridaceae) from the Brazilian Amazon. Brittonia 69: 307–312
.Moran, R. C. 1987. Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Bulletin of the Illinois Natural
History Survey 34: 1–138. Moran, R. C. 1987. Sterile-fertile leaf dimorphy and the evolution of soral types in Polybotrya
(Dryopteridaceae). Systematic Botany 12: 617–628. Moran, R. C. & P. Labiak. 2015. Phylogeny of the polybotryoid fern clade
(Dryopteridaceae). International Journal of Plant Sciences 176: 880–891.

482
Figure 238. Polybotrya. Note in H and F the strong sterile-fertile leaf dimorphy, and in J the characteristic rhizome cross section.
In K is shown a typical Eupolyods I petiole cross section. AP aerophore; LT leaf trace; RT root trace; SH dark sclerenchymatous
sheath. (Modified from Moran, 1987.)

483
HYMENOPHYLLACEAE
Polyphlebium Copel.
DESCRIPTION: Epiphytes; roots absent or few;
rhizomes 0.1–0.8 mm wide, long-creeping,
frequently branched, densely pubescent, the hairs
reddish brown or light brown (not black); petioles
0.3–5.5 cm long; sterile and fertile monomorphic; sterile
laminae 1-cell thick between the veins, 1-pinnate to
4-pinnate, the apices not proliferous; veins anadromous,
free, false veins absent; sori marginal; indusia tubular, the
mouth usually dilated; receptacles exert; spores green,
tetrahaedral-globose; x=36.
DESCRIPCIÓN: Epífitas; raíces absent or pocas;
rizomas 0.1–0.8 mm de ancho, largamente
rastreros, ramificados con frecuencia, densamente
pubescentes, los pelos pardo-rojizos o claro-rojizos
(no negros); pecíolos 0.3–5.5 cm de largo; hojas estériles
y fértiles monomorfas; láminas estériles una célula de
grosor entre las venas, 1-pinnadas a 4-pinnadas, los ápices
no prolíferos; venas anádromas, libres, falsas venas
ausentes; soros marginales; indusios tubulares, la boca
usualmente dilata; receptáculos exertos; esporas verdes,
tetrahédricas-globosas; x=36.

SIMILAR GENERA: Crepidomanes differs by short
rhizome branchlets that resemble roots, and thhe
presence of blackish hairs on the rhizomes, petioles, and
petiole-wing margins. Neotropical Crepidomanes lack false
veins but have parallel folds on either side of the segment,
and these resemble false veins (Paleotropical species do
have false veins). Didymoglossum differs by the presence
of false veins in the laminae and smaller leaves (< 6 cm).
Vandenboschia differs by thicker rhizomes (1–2 mm), the
presence of robust roots, and usually longer leaves (> 15
cm)
GÉNEROS PARECIDOS: Crepidomanes difiere por
ramitas cortos que asemejan raíces, y la presencia de
pelos negruzcos en los rizomas, pecíolos y márgenes de
las alas de los pecíolos. Crepidomanes neotropical falta de
venas falsas pero tienen pleigas paralelas en cada lado del
segmmento, e éstas asemejan venas falsas (las especies
paleotropicales tienen venas falsas). Didymoglossum difiere
por la presencia de venas falsas en las láminas y hojas
más cortas (< 6 cm). Vandenboschia difiere por rizomas
más gruesos (1–2 mm), la presencia de raíces robustos y
usualmente hojas más largas (> 15 cm).

COMMENTS: Polyphlebium is pantropical and grows as COMENTARIOS: Polyphlebium es pantropical y

en eiphyte in wet forests. It consists of 13 species, 7 of
which occur in the Neotropics. Four of these are common
and widespread: P. angustatum, P. capillaceum, P. diaphanum,
and (if distinct from the latter) P. hymenophylloides. The
first two species are almost entirely restricted to tree fern
root mantles.The rhizomes of all species branch frequently
to produce short branches that resemble roots. Ebihara
et al. (2009) reported P. borbonicum from South America
and Polynesia, but I cannot distinguish this species from P.
diaphanum.The genus name is derived from the Greek
poly, many, and phleps, veins.
existe como epífito en bosques húmedos. Consta de
13 especies, de las cuales 7 existen en el Neotrópico.
Cuatro de estas son comunes y difundidas: P. angustatum,
P. capillaceum, P. diaphanum y (si distinta del anterior) P.
hymenophylloides. Los dos primeras especies son casi
completamente regtringidas a los mantas de raíces de
los helechos arborescentes. Los rizomas de todos las
especies ramifican con frecuencia y producen ramas
cortas que parecen a raíces. Ebihara et al. (2009)
reportaron P. borbonicum de América del Sur y Polinesia,
pero no puedo distinguir esta especie de P. diaphanum.
El nombre genérico se deriva del griego poly, mucho, y
phleps, venas.

LITERATURE: Dubuisson, J.-Y., S. Hennequin, F. Rakotondrainibe & H. Schneider. 2003. Ecological diversity and adaptive tendencies
in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to climbing and epiphytic habits. Botanical Journal of the Linnean
Society 142: 41–63. Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of Hymenophyllaceae.
Blumea 51: 1–60. Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global molecular phylogeny of the fern genus
Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of the Linnean Society 155: 1–27. Ebihara, A., J.
Nitta, D. Lorence & J.-Y. Dubuisson. 2009. New records of Polyphlebium borbonicum, an African filmy fern, in the New World and Polynesia.
American Fern Journal 99: 200–206. Hébant-Mauri, R. 1972. Le genre Trichomanes L. (Fougéres Leptosporangiées). Adansonia 12: 469–495.
Schneider, H. 2000. Morphology and anatomy of roots in the filmy fern tribe Trichomaneae H. Schneider (Hymenophyllaceae, Filicatae) and
the evolution of rootless taxa. Botanical Journal of the Linnean Society 132: 29–46.

484
Figure 239. A–C. Polyphlebium hymenophylloides [=P. diaphanum?]. D–E. P. capillaceum. (From Mickel & Smith, 2004.)

485
POLYPODIACEAE
Polypodium L.
DESCRIPTION: Epiphytic, rupestral, or rarely terrestrial;
rhizomes creeping, branched, bearing two rows of leaves
on the dorsal surface; rhizome scales nonclathrate,
concoloras, peltate, glabrous; sterile and fertile leaves
monomorphous; petioles sulcate adaxially, articulate
to short phyllopodia, falling off cleanly in old age; laminae
glabrous, pinnatisect to (rarely) 1-pinnate at the base;
veins free or rarely areolate with each areola containing
a single excurrent veinlet; sori terminal on the tips of
included veinlets, round or occasionally elliptic, in one row
between the costa and margin; indusia absent; paraphyses
often present among the sporangia; spores yellow (when
fresh), monolete. x=37.
DESCRIPCIÓN: Epífitas, rupestres, o raras veces
terrestres; rizomas rastreros, ramificados, llevando dos
hileras de hojas sobre la superficie dorsal; escamas del
rizoma no clatradas, concoloras, peltadas, glabras; hojas
estériles y fértiles monomorfas; pecíolos surcados
en el haz, articulados a filopódios cortos, cayéndose
limpiamente con edad,; láminas glabras, pinnatisectas
a (raras veces) 1-pinnadas en la base; venas libres
o areoladas con cada aréola conteniéndose una sóla
venilla excurrente; soros terminales en los puntos de
venillas incluidas, redondos o ocasionalmente elípticas, en
una hilera entre la costa y el margen; indusios ausentes;
paraphyses a menudo presentes entre los esporangios;
esporas amarillas (cuando frescas), monoletes. x=37.

SIMILAR GENERA: Serpocaulon differs by anastomosing
veins, typically longer-creeping and unbranched, rhizomes,
laminae 1-pinnate (at least basally), and clathrate rhizome
scales. Many species have more than one row of sori
between the costa and margin. Pleopeltis differs by scaly
and generally thicker laminae (the veins often hard to
see). Pecluma differs pectinate laminae with usually more
than 30 pinna pairs, and petioles not grooved adaxially.
GÉNEROS PARECIDOS: Serpocaulon difiere por venas
anastomosadas, rizomas típicamente más largamente
rastreros y no ramificados, láminas 1-pinnadas (a lo menos
en la base), y escamas del rizoma no clatradas. Muchas
especies tienen más de una hilera de soros entre la costa
y el margen. Pleopeltis difiere por láminas escamosas y
generalmente más gruesas (las venas difíciles ver). Pecluma
difiere por láminas pectinadas con usuamente más de 30
pares de pinnas y pecíolos no surcados adaxialmente.

COMMENTS: Polypodium comprises about 45 species COMENTARIOS: Polypodium consta de casi 45 especies
worldwide, with two subgroups informally recognized. mundial, con dos subgrupos reconocidos informalmente.
The P. vulgare group consists of about 14 species and El grupo de P. vulgare consta de casi 14 especies y es
is primarily circumboreal, extending south to Oaxaca, principalmente circumboreal, extendiéndose al sur hasta
Mexico. The P. plesiosorum group consists of about 30 Oaxaca, México. El grupo de P. plesiosorum consta de
species and occurs in Mexico, Central America, and casi 30 especies y existe en México, América Central
northern South America. Eight of its species are endemic y América del Sur septentrional. Ocho de sus especies
to Mexico and five to Central America. The species are son endémicas a México y cinco a América Central.
primarily epiphytes or saxicolous in wet forests from Las especies son principalmente epífitas o saxícolas en
(100–)1000–2000(–3000) m. The genus name comes bosques húmedos desde (100–)1000–2000(–3000)
from the Greek, poly, many, and podion, foot, alluding to m. El nombre genérico viene del griego, poly, mucho, y
the resemblance of the many phyllopodia to feet. podion, pie, aludiéndose a la semejanza de los numerosos
filopodios a pies.
LITERATURE: Luna-Vega, I. J. D. Tejero-Díez, R. Contreras-Medina, M. Heads & G. Rivas. 2012. Biogeographical
analysis of two Polypodium species complexes (Polypodiaceae) in Mexico and Central America. Biological Journal of the Linnean
Society 106: 940–955. Sigel, E. M., M. D. Windham, C. H. Haufler & K. M. Pryer. 2014. Phylogeny, divergence time
estimates, and phylogeography of the diploid species of the Polypodium vulgare complex (Polypodiaceae). Systematic Botany
39(4): 1042–1055. Tejero-Díez, J. D., S. Aguilar-Rodríguez, T. Terrazas & L. Pacheco. 2010. Arquitectura y anatomía foliar
del complejo Polypodium plesiosorum sensu Moran (Polypodiaceae). Revista de Biología Tropical 58: 955–967. Tejero-Diez,
J. D & L. Pacheco. 2004. Taxa nuevos, nomenclatura, redefinición y distribución de las especies relacionadas con Polypodium
colpodes Kunze (Polypodiaceae, Pteridophyta). Acta Botánica Mexicana 67: 75–115. Tejero-Diez, J. D & L. Pacheco. 2004.
Notas taxonómicas y de distribución en Polypodium plesiosorum y P. rhodopleuron (Polypodiaceae: Pteridophyta). Anales Instituto
de Biología, Universidad Nacional Autónoma México, ser. Bot. 75:11–37.

486
Figure 240. A–C. Polypodium rhodopleuron. E–F. P. hesperium. G–J. P. plesiosorum. K–M. P. californicum. N–P. P. arcanum.
Q–S. P. arcanum var. arcanum. (From Mickel & Smith, 2004.)

487
POLYPODIACEAE
Polypodium dulce Group (sensu Moran, 1995)
DESCRIPTION: Epiphytic, rarely terrestrial or rupestral;
proliferous roots absent; rhizomes short-creeping, not
branched(?), dorsiventral (i.e., the leaves borne on the
dorsal surface), scaly, the scales pubescent with dark
hairs (at least at the point of attachment) not clathrate,
concolorous; petioles dark, not grooved adaxially,
articulate to low (1-2 mm) phyllopodia; laminae with
10-25 segments, usually widest or nearly so at
the base, the apex pinnatifid or subconform; segments
adnate, entire or serrate; rachises puberulent adaxially,
not grooved; veins usualmente free (anastomosing
in three species); sori round or (less commonly) elliptic,
non-indusiate; sporangial capsules often setulose;
spores yellow, monolete; x=37.
DESCRIPCIÓN: Epífitas, raras veces terrestres o
rupestres; raíces prolíferas ausentes; rizomas cortamente
rastreros, no ramificadas(?), dorsiventrales (i.e., las hojas
nacen en la superficie dorsal), escamosos, las escamas
pubescentes con pelos oscuros (a lo menos al punto
de inserción), no clatradas, concoloras; pecíolos oscuros,
no surcados adaxialmente, articulados a filopódios cortos
(1-2 mm); láminas con 10-25 pares de segmentos,
usualmente más anchas o casi así en la base, el ápice
pinnatífido o subconforme; segmentos adnatos, enteros
o serrados; ráquises puberulentos adaxialmente, no
surcados; venas usualmente libres (anastomosadas
en tres especies); soros redondos o (menos común)
elípticos, no indusiados; cápsulas esporangiales a
menudo sétulas; esporas amarillas, monoletes; x=37.

SIMILAR GENERA: Pecluma differs by rhizomes scales
united across with entire width of their base. It generally
also differs by dark petioles with only one vascular bundle,
more than 30 pinna pairs, and laminae usually reduced
toward the base. Polypodium differs by adaxially glabrous
and grooved petioles and rachises, glabrous rhizome
scales, and glabrous sporangial capsules.
GÉNEROS PARECIDOS: Pecluma difiere por escamas
del rizomas unidas por la anchura de la base. Usualmente
difiere por pecíolos oscuros con un solo haz vascular, más
de 30 pares de pinnas y láminas usualmente reducidas
hacia la base. Polypodium difiere por pecíolos y ráquises
surcados y glabros adaxialmente, pecíolos usualmente
estamíneos (no oscuros) y cápsulas esporangiales glabras.

COMMENTS: The Polypodium dulce group contains
about 20 species, all neotropical. Its center of diversity is
Mexico; only two species occur in South America (P. dulce
and P. ursipes). Although its species are currently placed in
Polypodium, the group is sister to Pecluma (Ranker et al.,
2004), with which it shares the distinctive characters of
hairy rhizome scales, rachises ungrooved and puberulent
adaxially and, free typically veins, and sporangial capsules
often setulose. Eventually the P. dulce group should be
placed in a different genus—perhaps even combined
under Pecluma. Keys to most of its species can be found
in Mickel and Smith (2004) and Moran (1995). The group
is named for P. dulce, its most common and widespread
species.
COMENTARIOS: El grupo de Polypodium dulce consta
de casi 20 especies, todos neotropicales. Su centro de
diversidad es México; solamente dos especies existen
en América del Sur (P. dulce y P. ursipes). Aunque sus
especies están ubicadas actualmente en Polypodium, el
grupo es hermana a Pecluma (Ranker et al., 2004), con
la cual comparte las características distintivas de escamas
del rizoma peludas, ráquises puberulentos adaxialmente
y no surcados, venas típicamente libres y cápsulas
esporangiales típicamente setulosas. Eventualmente el
grupo de P. dulce debe ser ubicado en un género distinto,
tal vez aún combinado en Pecluma. Claves a la mayoría
de sus especies se puede encontrar en Mickel y Smith
(2004) y Moran (1995).El grupo se nombra para P. dulce,
su especie más común y muy difundida.

LITERATURE: Maxon, W. R. 1903. A study of certain Mexican and Guatemalan species of Polypodium. Contributions from
the United States National Herbarium 8: 271–280. Mickel, J. T. & A. R. Smith. 2004. The Pteridophytes of Mexico. Memoirs of
the New York Botanical Garden 88: 1–1055. Moran, R. C. 1995. Grupo de Polypodium dulce. Pages 351–355. In: Davidse, G., M.
Sousa S. & S. Knapp, editors. Flora Mesoamericana, vol. 1. Psilotaceae a Salviniaceae. Universidad Nacional Autónoma de México,
Ciudad Univeristaria, Distr. Federal.

488
Figure 241. Polypodium dulce group. Note the comose rhizome scale and setulose sporangial capsules—two of the diagnostic
characters of the group. Fijase las escamas del rizoma comosas y cápsulas esporangiales setulosas, dos caracteres diagnósticos
del grupo. Left (izquierda). Polypodium dulce. Upper right (derecha arriba). P. fissidens. Lower right (derecha abajo). P.
alavae. (Modified from Mickel & Smith, 2004.)

489
DRYOPTERIDACEAE
Polystichopsis (J. Sm.) C. Chr.
DESCRIPTION: Terrestrial; rhizomes creeping;
leaves monomorphic; petioles with more than three
vascular bundles; laminae deltate or pentagonal,
2- to 3-pinnate-pinnatifid, pilose on the axes or
throughout, the hairs whitish, septate; basal pinnae
prolonged basiscopically, especially the basal basiscopic
pinnule, anadromic; medial pinnae catadromic,
stalked; rachis and costae grooved adaxially, the grooves
pubescent within, without proliferous buds; veins free;
sori round; indusia present or absent, orbicular-reniform;
spores monolete, verrucate; x=41.
DESCRIPCIÓN: Terrestres; rizomas rastreros,
escamosos; hojas monomorfas; pecíolos con tres o más
haces vasculares; láminas deltadas o pentagonales, 2-
a 3-pinnado-pinnatífidas, pilosas por los ejes o por
todo, los pelos blanquecinos, septados; pinnas basales
prolongadas basiscópicamente, especialmente la pínnula
basal basiscópica; pinnas mediales catadrómicas;
raquis y costas surcados adaxialmente, los surcos
prolíferos adentro, sin yemas prolíferas; nervaduras libres;
soros redondos; indusios presentes o ausentes, orbicular-
reniformes; esporas monoletes, verrucatas; x=41.

SIMILAR GENERA: Arachniodes differs by anadromic
medial pinnae. Dryopteris differs by erect or decumbent
rhizomes and non-pilose laminae. Polystichum differs by
pinnae or pinnules with an enlarged acroscopic basal
lobe, spinulose teeth on the laminar margins, and peltate
indusia. Lastreopsis differs by the upper surfaces of the
rachises and costae rounded or flat (not grooved).
Cyclodium and Triplophyllum differs by puberulent rachises
and costae adaxially and the lack of pilose hairs on the
lamina.
GÉNEROS PARECIDOS: Arachniodes difiere por
pinnas mediales anadrómicas. Dryopteris difiere por
rizomas erectas o decumbentes y láminas non pilosas.
Polystichum difiere por las pinnas o pínnulas con un lóbulo
acroscópico basal agrandado, dientes espinulosos en los
márgenes de las láminas y indusios peltados. Lastreopsis
difiere por el haz de las raquises y costas redondeado o
achatado (no surcado). Cyclodium y Triplophyllum difieren
ráchises y costas puberulentos adaxialmente y la falta de
pelos pilosos en la lámina.

COMMENTS: Polystichopsis consists of about 8 species COMENTARIOS: Polystichopsis consta de casi 8

endemic to the Antilles. It grows at low to middle elevations
(200–500 m) in wet forests, limestone, and clay banks. It
is sister to Trichoneuron microlepidoides Ching, an Asian
species; in fact, it is uncertain how the two genera differ.
They form part of the polybotryoid fern clade, which has
creeping rhizomes. Polystichopsis consists of medium-sized
ferns that usually fit, without folding the leaf, on a single
herbarium sheet. The laminae are thin relative to other
dryopteroid ferns, and this plus the whitish pilose hairs
characterize the genus. A key to the species can be found
in Prado and Moran (2015). The genus name is derived
from Polystichum + the Greek suffix -opsis, likeness, or
resemblance.
especies endémicas a las Antillas. Existe en elevaciones
bajas a medias (200–500 m) en bosques húmedos, piedra
caliza y taludes de arcilla. Es hermano a Trichoneuron
microlepidoides Ching, un especie Asiática; de hecho, es
incierto como se distinguen. Forman parte del clado
polybotrioide, el cual tiene tiene rizomas reptantes.
Polystichopsis consta de helechos de medio tamaño
que suelen caber, sin doblar la hoja, en una sola hoja
del herbario. Las láminas son delgadas relativas a otros
helechos driopteroides, y esto más los pelos blanquecinos
pilosos se caracterizan el género. Una clave a las especies
se encuentra en Prado y Moran (2015). El nombre
genérico se deriva de Polystichum + el sufijo griego -opsis,
semejanza.

LITERATURE: Christensen, C. 1920. The a monograph to the genus Dryopteris. Part II. The tropical American bipinnate-
decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 8, 6: 101–120. Liu, H-M., X-C. Zhang, M-P.
Wang, H. Shang, S-L. Zhou,Y-H.Yan, X-P.Wei,W-B. Xu & H. Schneider. 2015. Phylogenetic placement of the enigmatic fern genus
Trichoneuron informs on the infra-familial relationship of Dryopteridaceae. Plant Systematics and Evolution [DOI 10.1007/s00606-
015-1265-3]. Morton, C. V. 1960. Observations on cultivated ferns VI. The ferns currently known as Rumohra. American Fern
Journal 50: 145–155. Prado, J. & R. C. Moran. 2016. Monograph of the West Indian fern genus Polystichopsis (Dryopteridaceae).
Brittonia 68: 1–24. Proctor, G. R. 1961. Notes on Polystichopsis. American Fern Journal 51: 145–148. Proctor, G. R. 1985. Ferns
of Jamaica. British Museum (Natural History), London.

490
Figure 242. Polystichopsis. A–C. P. leucochaete (Slosson) R. C. Moran, comb. ined. D–G. P. chaerophylloides (Poir) C. V.
Morton. (From Moran & Prado, 2016.)

491
DRYOPTERIDACEAE
Polystichum Roth.
DESCRIPTION: Terrestrial or epipetric; rhizomes
usually thick and covered by old petiole bases, erect to
decumbent; sterile and fertile leaves monomorphous;
petioles scaly, the scales ciliate; laminae usually 2-pinnate
to 3-pinnate; pinnae with a basal acroscopic lobe or
pinnules elongated, otherwise equilateral; pinnules
usually inequilateral, broader on the acroscopic side,
the margins usually spinulose; rachis and costae often
conspicuously scaly; veins free (except in P. dubium (H.
Karst.) Diels); sori round; indusia absent, or present and
peltate; x=41.
SIMILAR GENERA: Arachniodes and Rumohra differ
by creeping rhizomes. Dryopteris usually differs by
basiscopically elongate basal pinnae, and indusia attached
at a sinus. Dryopteris wallichiana (Spreng.) Hyl., which has
all pinnae equilateral, can be distinguished by its entire
(nonspinulose) segments and laterally attached indusia.
COMMENTS: Polystichum contains about 500 species
worldwide, about 70 of which are neotropical. There are
about 27 species in the northern and central Andes, 9
in austral South America, 9 in central and eastern Brazil,
31 in the West Indies, and ca. 23 in Mesoamerica and
Mexico. Nearly all grow above 2000 m (the genus is
absent below 600 m in the Neotropics). Many of the
West Indian species grow on limestone (Mickel, 1997).
Hybridization and polyploidy have been prominent in
the evolution of Polystichum, with 83 sterile interspecific
hybrids having been reported. The New World species
form a monophyletic group nested with the Old World
species (Little & Barrington, 2003). . This Neotropical clad
has diversified within the past 15 million years. The name
is derived from the Greek poly, many + stichos, rows. The
sori of the type species are in many regular rows on the
pinnae.
DESCRIPCIÓN: Terrestre o epipétrico; rizomas
usualmente gruesos y cubiertos por las bases de los
pecíolos viejos, erectos a decumbentes; hojas estériles y
fértiles monomorfas; pecíolos escamosos, las escamas
ciliadas; láminas usualmente 2-pinnadas a 3-pinnadas;
pinnas con una lóbulo basal acroscópico o una
pínnula basal alongada, de otra manera equiláteras;
pínnulas usualmente inequiláteras, más anchas en el lado
acroscópico, los márgenes usualmente espinulosos;
ráquis y costas a menudo conspicuamente escamosos;
nervadura libre (con excepción de P. dubium (H. Karst.)
Diels); soros redondos; indusios ausentes, o presentes y
peltados; x=41.
GÉNEROS PARECIDOS: Arachniodes y Rumohra
difieren por rizomas reptantes. Dryopteris usualmente
tiene pinnas basales elongadas basiscópicamente, y
indúsisos unidos en un sinus. Dryopteris wallichiana
(Spreng.) Hyl. tiene pinnas equiláteras, pero puede
distinguirse por sus segmentos enteros (no espinulosos)
e indusios unidos lateralmente.
COMENTARIOS: Polystichum contiene casi 500
especies mundialmente, casi 70 de las cuales son
neotropicales. Hay casi 27 especies en los Andes
septentrionales y centrales, 9 en América del Sur Austral,
9 en Brasil central y occidental, 31 en las Antillas, y casi
23 en Mesoamérica y México. Casi todas ocurren arriba
de los 2000 m (el género está ausente bajo los 600 m
en el Neotrópico). Muchas de las especies Antillanas
crecen sobre piedra caliza (Mickel, 1997). La hibridación
y poliplodia han sido prominentes en la evolución de
Polystichum, con 83 híbridos interspecíficos estériles
habían sido reportados. Las especies del Nuevo Mundo
forman un grupo monofilético anidado dentro de las
especies del Viejo Mundo (Little & Barrington, 2003). Este
clado neotropical ha diversificado dentro de los últimos
15 millones de años. El nombre genérico se deriva del
griego poly, mucho + stichos, hileras. Los soros de la
especie tipo están dispuestos en muchas hileras regularas
en las pinnas.

LITERATURE: Barrington, D. S. 2011.The fern genus Polystichum (Dryopteridaceae) in Costa Rica. Annals of the Missouri Botanical Garden
98: 431–446. Kessler, M., A. R. Smith & M. Sundue. 2005. Notes on the genus Polystichum (Dryopteridaceae) in Bolivia, with descriptions of ten
new species. Brittonia 57: 205–227. Little, D. P. & D. S. Barrington. 2003. Major evolutionary events in the origin and diversification of the
fern genus Polystichum (Dryopteridaceae). American Journal of Botany 90: 508–514. Le Péchon, T., H. Hai, L. Zhang, X-M. Zhou1, X-F. Gao, &
L-B. Zhang. 2016. Using a multilocus phylogeny to test morphology-based classifications of Polystichum (Dryopteridaceae), one of the largest
fern genera. BMC Evolutionary Biology16: 55 [DOI 10.1186/s12862-016-0626-z]. McHenry, M. A. & D. S. Barrington. 2014. Phylogeny and
biogeography of exindusiate Andean Polystichum (Dryopteridaceae). American Journal of Botany 101: 365–375. Mickel, J. T. 1997. A review of
the West Indian species of Polystichum. Pages 119–143. In: R. J. Johns (ed.), Holttum Memorial Volume. Royal Botanic Gardens, Kew. Rodríguez,
R. 1987. Notas taxonómicas sobre el género Polystichum Roth (Aspidiaceae-Filicidae) en Chile. Gayana, Botany 44: 45–53.

492
Figure 243. A–D. Polystichum rachichlaena. E. P. schizophyllum. (From Mickel & Smith, 2004.)

493
PTERIDACEAE
Polytaenium Desv.
DESCRIPTION: Epiphytes; rhizome dorsiventral; leaf
arrangement distichous; rhizome scales clathrate; leaves
simple, entire, costate; venation pluriseriate, areolate;
sori linear, along reticulate veins and, in one species,
sporangia also scattered between veins; indusia absent;
soral paraphyses absent; spores trilete; x=60.
DESCRIPCIÓN: Epífitas; rizomas dorsiventales; ; hojas
arregladas dísticamente; escamas del rizoma clatradas;
hojas simples, enteros, con costas presentes;
nervaduras pluriseriada, areoladas; soros lineares por
venas reticuladas y, un P. citrifolium, esporangios difundidos
entre las venas; alargados a lo largo de las nervaduras;
indusios ausentes; parafisos sorales ausentes; esporas
triletes; x=60.

SIMILAR GENERA: Anetium differs by scattered
sporangia between the veins, and long-creeping rhizomes.
Scoliosorus differs by the presence of paraphyses, monolete
spores, and a costa that is elevated abaxially at the base.
GÉNEROS PARECIDOS: Anetium difiere por
esporangios salpicados entre las nervaduras, y rizomas
largamente reptantes. Scoliosorus difiere por la presencia
de parafisos, esporas monoletes, y una costa elevada
abaxialmente en la base.

COMMENTS: Polytaenium is completely Neotropical and COMENTARIOS: Polytaenium es completamente

consists of about10 species, most of which are low-trunk
epiphytes in wet forests. It is the only genus of vittarioid
ferns that lacks paraphyses. As in other vittarioids, it has
false veins (cells with thick walls replaced by silica) . They
appear as minute, slender, prominulous streaks between
the veins. They are always present on the adaxial laminar
surface, but may also occur abaxially.They do not connect
with true veins and do not contain conducting cells (i.e.,
xylem and phloem). As defined here, the genus includes
Anetium Splitg. (Anetium citrifolium = P. citrifolium). At least
one species of Polytaenium (P. feei) harbors ants beneath
its mat of roots. The waste of the ants contribute much
nitrogen to the plant. The genus name is derived from
the Greek poly, many + taenia, ribbon, alluding to the sori.
Neotropical y consta de casi10 especies, la mayoria de las
cuales son epífitas en las bases de los troncos de bosques
húmedos. Es el único género de helechos vittarioides que
falta paráfises. Como en otros vittarioides, tiene las venas
falsas (células con paredes gruesas de sílice). Parecen
como rayos minutos delgados promínulos entre las venas.
Se presentan siempre en el haz de la lámina y a veces
en el envés. No se conectan con las verdaderas venas
y no tienen células conductoras (i.e., xilema y floema).
Como definido aquí el género incluye Anetium Splitg.
(Anetium citrifolium = P. citrifolium). Al menos una especie
de Polytaenium (P. feei) alberguen hormigas debajo de sus
raíces densas (Watkins et al., 2008). El desperdicio de
las hormigas contribuyen mucho nitrógeno a la planta.
El nombre genérico se deriva del griego poly, mucho +
taenia, cinta, aludiéndose a los soros.

LITERATURE: Benedict, R. C. 1907. The genus Antrophyum—I. Synopsis of subgenera, and the American species. Bulletin of
the Torrey Botanical Club 34: 445–458. Benedict, R. C. 1911. The genera of the fern tribe Vittarieae: Their external morphology,
venation, and relationships. Bulletin of the Torrey Botanical Club 38: 153–190. Crane, E. H. 1997. A revised circumscription of the
genera of the fern family Vittariaceae. Systematic Botany 22: 509–517. Schuettpelz, E., C-W. Chen, M. Kessler, J. B. Pinson,
G. Johnson,1 A. Davila, A. T. Cochran, L. Huiet & K. M. Pryer. 2016. A revised generic classification of vittarioid ferns
(Pteridaceae) based on molecular, micromorphological, and geographic data. Taxon 65: 708–722. Watkins, J. E., C. L. Cardelús
& M. C. Mack. 2008. Ants mediate nitrogen relations of an epiphytic fern. New Phytologist 180: 5–8.

494
Figure 244. Polytaenium. A–C. P. feei. D, E. P. lineatum. G–J. P. chlorosporum. K–M. P. cajenense. (From Mickel & Smith, 2004.)

495
DRYOPTERIDACEAE
Pradopteris R. C. Moran & Labiak, gen. nov. ined.
DESCRIPTION: Terrestrial; rhizomes creeping; leaves
monomorphic; petioles with more than three vascular
bundles; laminae widely deltate or pentagonal, 2-pinnate-
pinnatifid to 4-pinnate; pinnules anadromically
arranged, stalked; rachises and costae grooves
pubescent at least near the junctures, without proliferous
buds; veins free; sori round; indusia orbicular-reniform,
thick, brown, firm; spores monolete; x=41.
DESCRIPCIÓN: Terrestres; rizomas rastreros
escamosos; hojas monomorfas; pecíolo con tres o
más haces vasculares; lámina ampliamente deltada o
pentagonal, 2-5-pinnada; pinnas medias prolongadas
acroscópicamente, el lado basal basiscópico cuneado;
pínnulas dispuestas anadrómicamente, pediculadas;
surcos de los ráquises y costas pubescentes, al menos
cerca de las uniones, sin yemas prolíferas; nervaduras
libres; soros redondos; indusios orbicular-reniformes,
gruesos, pardos, duros; esporas monoletes; x=41.

SIMILAR GENERA: Cyclodium differs by leaves less
divided, usually 1- to 2-pinnate. Dryopteris differs by
rhizomes erect or ascending, pinnules catadromic, and (in
the Neotropics) leaves often capitate-glandular. Polybotya
and Maxonia differ by strong sterile-fertile leaf dimorphy,
Arachniodes differs by grooves of the rachises and costae
completely glabrous within. Lastreopsis and Rumohra differ
by a thickened marginal ridge of the costa or costules
surcurrent along the green laminar margin of the segment
of the next highest order (not adaxially along the midrib
of the costule).
GÉNEROS PARECIDOS: Cyclodium difiere por hojas
menos divididas, usualmente 1- a 2-pinnadas. Dryopteris
difiere por rizomas erectos o ascendentes, pínnulas
catadrómicas y (en el Neotrópico) hojas a menudo
glanduloso-capitadas. Arachniodes difiere por los surcos
de los ráquises y costas completamente glabros adentro.
Lastreopsis y Rumohra difieren por una fila marginal
engrosada de la costa o cóstula surcurrente por el margen
del tejido laminar verde del segmento del próximo orden
más alto (no adaxialmente por la cóstula del segmento).

COMMENTS: Pradopteris is neotropical and consists of COMENTARIOS: Pradopteris es neotropical y consta

only two species: P. macrostegia (Hook.) R. C. Moran &
Labiak, and P. ochropteroides (Baker) R. C. Moran & Labiak.
It occurs in wet forests from low to middle elevations.
The first species is terrestrial and occurs primarily on
sandy soils. The abaxial surfaces of its rachises and costa
are glabrous. In contrast, the second species is epiphytic
and has pubescent rachises and costae abaxially. The
genus name honors Jefferson Prado (1964–), Brazilian
pteridologist.
de solo dos species: P. macrostegia (Hook.) R. C. Moran
& Labiak y P. ochropteroides (Baker) R. C. Moran & Labiak.
Existe en bosques húmedos desde elevaciones bajas
hasta medias. La especie anterior es terrestre e existe
principalmente sobre suelos arenosos. Las superficies
abaxiales de los ráquises y costas son glabras. En contraste,
la secunda especies es epífito y tiene los ráquises y costas
pubescentes abaxialmente. El nombre genérico honora a
Jefferson Prado (1964–), pteridólogo brasileño.

LITERATURE: Christensen, C. 1920. The a monograph to the genus Dryopteris. Part II. The tropical American bipinnate-
decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 8, 6: 101–120. Moran, R. C. & P. Labiak.
2015. Phylogeny of the polybotryoid fern clade (Dryopteridaceae). International Journal of Plant Sciences 176(9): 880–891.

496
Figure 245. Pradopteris. A–C. P. macrostegia. D, E, G. P. ochropteroides. F. P. obtusa., ined. (Original, ©R. C. Moran, 2016.)

497
LYCOPODIACEAE
Pseudolycopodiella Holub

DESCRIPTION: Terrestrial; main horizontal stems DESCRIPCIÓN:Terrestres; tallos principales horizantales

creeping, superficial, isophyllous or strongly anisophyllous,
branching dichotomously in the horizontal plane, also
branching in the vertical plane to give rise to single
dorsal row of erect unbranched stems; leaves
of the horizontal stems anisophyllous (laterals larger
and divergent, dorsal ones smaller and overlapping)
or isophyllous; leaves of the erect stems either similar
to those of the creeping stems or (in P. caroliniana)
reduced, distant, polystichous; gemmiferous branchlets
and gemmae absent; strobili solitary, erect, sessile or
stalked; sporophylls peltate, with a decurrent medial wing,
stramineous at time of sporangial dehiscence, entire or
nearly so; sporangia reniform, isovalvate, the walls with
incomplete semi-annular thickenings; spores rugulate;
gametophytes surficial, green, rounded; x=35.
reptantes, superficiales, isofílicos o fuertamente anisofílicos,
ramificándose dicotomicamente en el plano horizontal,
también ramificándose en el plano vertical para dar lugar
a una hilera dorsal de ramas erectas no ramificadas;
hojas de los tallos reptantes anisofilas (las laterales más
grandes y divergentes, las dorsales más pequeñas y
sobrelapándose) o isofílicas; hojas de los tallos erectos
simulares a las de los tallos reptantes o (en P. caroliniana)
reducidas, distantes, dimórfas; ramitas gemíferas y yemas
ausentes; estróbilos solitários, erectos, sésiles o
pedunculados ; esporofilos peltatos, con una ala medial
decurrente, estramineo en el tiempo de dehiscencia
esporangial, enteros o casi así; esporangios reniformes,
isovalvados, células epidérmicas con engrosamientos
nodulares; esporas rugulosas; gametofitos superficiales,
verdes, redondeados; x=35.

SIMILAR GENERA: Lycopodiella differs by sporophylls
nearly as wide as the trophophylls beneath, anisovalvate
sporangia and sporophylls with few to several, prominent
and often recurved teeth. Palhinhaea differs by erect
stems branched and tree-like, nodding or pendulous
strobili, and sporangia sunken the cortex of the strobili.
GÉNERO PARECIDO: Lycopodiella difiere por los
esporofilos cas tan ancho como los trofofilos abajo,
esporangios anisovalvados y esporofilos con pocos a
varios dientes prominentes y a menudo recurvados.
Palhinhaea difiere por los tallos erectos ramificados y
semejantes a pequeños árboles, estróbilos cernuos o
péndulos y esporangios hundidos en la corteza do los
estróbilos.

COMMENTS: Pseudolycopodiella consists of ca. 10 COMENTARIOS: Pseudolycopodiella consta de ca.

species. The genus is represented in most tropical and
temperate regions, but is absent from Europe, western
temperate Asia, and western North America. The species
typically grow on wet sandy soils. Their creeping stems
have aerenchymatous cortices, presumably an adaptation
to occasionally submersion. In Africa, some species
are adapted to seasonal drought and may survive dry
periods by means of subterranean, tuberous stem apices
(Kornas, 1975). The species in the coastal plain of the
United States is P. caroliniana (L.) Holub., the type of the
genus. See Øllgaard & Windisch (2016) for a key to the
six Brazilian species. The genus name is formed from the
Greek pseudo, false, and Lycopodiella, a related genus.
10 especies mundial. El género es representado en la
mayoría de las regiones tropicales y templadas, pero es
ausente de Europa, Asia templada occidental y América
del Norte occidental. Las especies típicamente existen
en suelos húmedos y arenosos. Sus tallos reptantes son
arenquimatosos, presumablemente una adaptación a ser
submergidos ocasionalmente. En África algunas especies
son adaptadas a la sequidad ocasional y pueden sobrevivir
periodos secos por medios de tuberosos y subterráneos
ápices de los tallos. La especie en la planicie costal de
los Estados Unidos es P. caroliniana (L.) Holub, la especie
tipo del género. Véase Øllgaard & Windisch (2016) para
una clave a las seis especies brasileñas. El nombre del
género es formado del griego pseudo, falso, y Lycopodiella,
un género relacionado.

LITERATURE: Bruce, J. G. 1976. Comparative studies of Lycopodium carolinianum. American Fern Journal 66: 125–137. Bruce,
J. G. 1979. Gametophyte and young sporophyte of Lycopodium carolinianum. American Journal of Botany 66: 1156–1163. Kornas,
J. 1975. Tuber production and fire-resistance in Lycopodium carolinianum in Zambia. Acta Societatis Botanicorum Poloniae 44:
653–663. Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview. Annals of the Missouri Botanical Garden 79: 687–717.
Øllgaard, B. & P. G. Windisch. 2016. Lycopodiaceae in Brazil. Conspectus of the family II. The genera Lycopodiella, Palhinhaea,
and Pseudolycopodiella. Rodriguésia 67(3): 691–719.

498
Figure 246. A. Pseudolycopodiella paradoxa. B. Pseudolycopodiella meridionalis. C. Pseudolycopodiella carnosa (From
Øllgaard & Windisch, 2016.)

499
PSILOTACEAE
Psilotum Sw.
DESCRIPTION: Epiphytic or rupestral, rarely terrestrial;
rhizomes without roots but bearing short brownish hair-
like rhizoids; aerial stems green, apparently leafless,
unbranched basally, repeatedly dichotomously branched
distally; branches flattened or triangular in cross-section;
leaves 1-2 mm long, inconspicuous, scale-like, alternate,
widely scattered; synangia sessile, 3-lobed, subtended
by a forked appendage, yellowish at maturity; spores
hyaline, reniform; x=52.
SIMILAR GENERA: None.
COMMENTS: Only two species of Psilotum occur in the
Neotropics: P. complanatum Sw. and P. nudum (L.) Pal.They
can be distinguished by the cross-sections of their distal
branchlets. Psilotum complanatum has flattened, 2-ranked
branchlets; P. nudum has triangular, 3-ranked branchlets.
Hybrids between the two species are known from Hawaii
but have not yet been found in the Neotropics. Psilotum is
extremely variable in habit and habitat. It can be epiphytic
or terrestrial, pendent or erect. A good habitat to look
for Psilotum is in the humus-filled leaf axils of palms.
The Psilotaceae is sister to the Ophioglossaceae. Both
have sporangia on the adaxial surface of the leaves and
cylindrical subterranean gametophytes. A polyploid series
is known for P. nudum in Australia and New Zealand
(Barber, 1957), but this has not been investigated in the
Neotropics. The genus name is derived from the Greek
psilos, smooth or naked. The stems appear naked, without
large leaves.
LITERATURE: Barber, H. N. 1957. Polypoidy in the Psilotales. Proceedings of the Linnean Society of New South Wales. 82:
201–208.
500
DESCRIPCIÓN: Epífitas o rupestres, raras veces
terrestres; rizomas sin raíces pero portando rizoides
parduzcos que se parecen a pelos; tallos aéreos verdes,
aparentemente sin hojas, en la base no ramificados,
distalmente muchas veces ramificados dicotómicamente;
ramas aplanadas or triangulares en sección transversal;
hojas 1-2 mm de largo, inconspícuas, semejantes a
escamas, alternas, ampliamente difundidas; sinángios
sésiles, trilobados, subtendidos por un apéndice furcado,
amarillentos al madurar; esporas hialinas, reniformes;
x=52.
GÉNEROS PARECIDOS: Ningunos.
COMENTARIOS: Sólo dos especies de Psilotum
existen en el Neotrópico: P. complanatum Sw. and P.
nudum (L.) Pal. Son distinguibles por la sección cruz de
sus ramitas dístales. Psilotum complanatum tiene ramitas
aplanadas con dos ángulos agudos; P. nudum tiene ramitas
triangulares con tres ángulos agudos. Híbridos entre
los dos especies se conocen de Hawaii pero no han
sido encontrados todavía en el Neotrópico. Psilotum es
extremamente variable en habitu . Se puede ser epifito
o terrestre, pendiente o erecto. Un buen hábitat buscar
Psilotum es en las axilas de hojas de palmas las cuales han
acumulado mucho humus. Las Psilotaceae son hermana a
las Ophioglossaceae. Los dos tienen esporangios nacidos
en la superficie adaxial de hojas y gametofitos cilíndricos
subterráneos. Una serie de poliploides se conoce para P.
nudum en Australia y Nueva Zelanda (Barber, 1957), pero
esto no ha sido investigado en el Neotrópico. El nombre
genérico se deriva del griego psilos, desnudo o liso. Los
tallos aparecen desnudos, sin hojas grandes.
Figure 247. Left: Psilotum nudum. Right: P. complanatum. In both species, note the trilocular synangia in an adaxial position on
the sporophyll. En los dos especies, fijase en los sinángios triloculares en una posición en la esporofila. (From Mickel & Smith,
2004.)

501
DENNSTAEDTIACEAE
Pteridium Gled. ex Scop.
DESCRIPTION: Terrestrial; rhizomes deeply
subterranean, long-creeping, pubescent (not scaly);
sterile and fertile leaves monomorphous; petioles about
equaling the lamina, at base producing epipetiolar buds;
laminae 3-pinnate to 4-pinnate, coriaceous, deltate
or pentagonal, pubescent (not scaly), sterile margins
reflexed; veins free; sori marginal, continuous (not
interrupted) along the segment or lobe, supplied by many
veins; indusia 2, one formed by the reflexed margin of the
lamina, the other sometimes absent, but when present
delicate, hyaline, and hidden beneath the sporangia;
paraphyses absent; x=52.
DESCRIPCIÓN: Terrestres; rizomas profundamente
subterráneos, largamente rastreros, pubescentes (no
escamosos); hojas estériles y fértiles monomorfas;
pecíolos casi igualando la lámina, produciendo brotes
epipeciolares en la base; láminas 3-pinnadas a
4-pinnadas, coriáceas, deltadas o pentagonales,
pubescentes (no escamosas), márgenes estériles
reflexos; nervaduras libres; soros marginales, continuos
(no interrumpidos) a lo largo del segmento o lóbulo,
suministrado por muchas venas; indusios 2, uno formado
por el margen reflejo de la lámina, el otro a veces ausente,
pero cuando presente delicado, hialino y oculto abajo de
los esporangios; parafisos ausentes; x=52.

DIAGNOSIS: Laminae 3-pinnate to 4-pinnate, deltate or DIAGNOSIS: Láminas 3-pinnadas a 4-pinnadas, deltadas
pentagonal; sori marginal, covered by a reflexed indusium; o pentagonales; soros marginales, cubiertos por un
sterile margins also reflexed. indusio reflexo; márgenes estériles reflexos también.

SIMILAR GENERA: Some species of Pteris have
similarly divided laminae and marginal sori. They can be
distinguished by their scaly stems, paraphyses among the
sporangia, plane (not reflexed) sterile margins, and short
spines on the upper surface of the axes.
GÉNEROS PARECIDOS: Algunas especies de
Pteridium tienen láminas divididas y soros marginales.
Pueden distinguirse por sus tallos escamosos, parafisos
entre los esporangios, márgenes estériles planos (no
reflexos) y espinas cortas en la superficie superior de
los ejes.

COMMENTS: Pteridium is an aggressive weed and often
forms dense populations by means of its subterranean
rhizomes. It invades pastures where it is eaten by cows,
and the carcinogens know to be contained in its leaves
may be pass on in the cow’s milk. An unusual characteristic
of Pteridium is that the sterile segments have enrolled
margins like those of the fertile. In the past, bracken
has been classified as a single species with 12 varieties
worldwide. Nowadays, most pteridologists prefer to
recognize the varieties at the specific level. Four species
occur in the Neotropics.Their recognition is supported by
morphology and DNA fingerprinting studies (Thomson
et al., 2008). The genus name is derived from Greek,
pteridion, a small fern.
COMENTARIOS: Pteridium es una maleza agresiva y
a menudo forma poblaciones densas por medio de sus
rizomas subterraneos. Invade praderas donde se come
por ganado, y los carcinagos en sus hojas se pasan en
la leche de las vacas.Una característica rara es que los
segmentos estériles tienen márgenes recurvados como
los de los segmentos fértiles. En el pasado, Pteridium ha
sido clasificado como una sóla especie con 12 variedades
mundial. Hoy en día, la mayoría de pteridólogos prefieren
reconocer las variedades al nivel específico. Cuatro
especies se encuentran en el Neotrópico. Su recognición
se apoya por morfología y estudies de DNA (Thomson
et al., 2008). El nombre genérico se deriva del griego,
pteridion, un helcho pequeño.

LITERATURE: Schwartsburd, P. B., P. L. R. De Moraes & K. L. B. Lopes-Mattos. 2014. Recognition of two morpho-
types in easern South American brackens (Pteridium-Dennstaedtiaceae)-Polypodiopsida). Phytotaxa 170: 103–117. Thomson,
J. A., J. T. Mickel & K. Mehltreter. 2008. Taxonomic status and relationships of bracken ferns (Pteridium: Dennsteadtiaceae)
of Laurasian affinity in Central and North America. Botanical Journal of the Linnean Society 157: 1–17. Tryon, R. M. 1941.
Revision of the genus Pteridium. Contributions from the Gray Herbarium of Harvard University 134: 1–70. (Reprinted from
Rhodora 43: 1–31, 37–67, t. 650–653. 1941).

502
Figure 248. Pteridium. A–C. P. aquilinum. D–E. P. aquilinum var. pubescens. F. P. aquilinum var. latiusculum. G, J. P.
arachnoideum. L, M. P. caudatum. (From Mickel & Smith, 2004.)

503
PTERIDACEAE
Pteris L.
DESCRIPTION: Terrestrial; rhizome creeping to erect,
scaly; leaves 1-pinnate to 4-pinnate; petioles with
a single omega-shaped vascular bundle; laminae
glabrous or rarely pilose, usually with a conform or
subconform terminal segment; basal pinnae entire or, if
divided, usually with the basal basiscopic pinnule enlarged;
axes of the penultimate segments in many species aristate
adaxially; veins free or anastomosing and without included
veinlets; sori linear on a submarginal connecting vein,
covered by the reflexed margin of the lamina (false
indusium); paraphyses generally present, numerous;
spores tetrahedral, with an equatorial flange; x=29.
SIMILAR GENERA: None.
COMMENTS: Pteris has about 250 species worldwide,
about 60 of which are neotropical. It is heterogeneous,
probably polyphyletic, and it is difficult to give
characteristics that define the genus. Many species of
Pteris have awns on the adaxial surface of the costules of
the penultimate segments. In these species, the ridge of
the costule is interrupted at the juncture with the costule
of the next higher order and is prolonged into a short
awn. A well-marked group is that of Pteris vittata L., which
is characterized by 1-pinnate and cuneate laminae, and
concolorous rhizome scales. Another such group is that
of P. podophylla Sw., with huge pedate laminae. The genus
name is derived from Greek pteron, wing or feather. It is
an ancient Greek name for ferns in general, derived from
a resemblance of the pinnae to wings or feathers.
LITERATURE: Arbeláez A., A. L. 1996. La tribu Pterideae (Pteridaceae). Flora de Colombia 18: 1–105. Prado, J. & P. G.
Windisch. 2000. The genus Pteris L. (Pteridaceae) in Brazil. Bol. Inst. Botânica 13: 103–199. Scamman, E. 1961. The genus Pteris
of Costa Rica. Rhodora 63: 194–205.
504
DESCRIPCIÓN: Terrestres; rizoma rastrero a erecto,
escamoso; hojas 1-pinnadas a 4-pinnadas; pecíolos con
un hace vascular en forma de una omega; láminas
glabras or raras veces pilosas, usualmente con un segmento
terminal conforme o subconforme; pinnas basales enteras
o, si divididas, usualmente con la pínnula basal basiscópica
agrandada; ejes de los penúltimos segmentos en muchas
especies aristados adaxialmente; nervaduras libres o
anastomosadas y sin nérvulas incluídas; soros lineares
sobre una nervadura conectora submarginal,
cubiertos por el margen reflexo de la lámina (falso
indusio); parafisos generalmente presentes, numerosos;
esporas tetrahédricas, con un reborde ecuatorial; x=29.
GÉNEROS PARECIDOS: Ninguno.
COMENTARIOS: Pteris tiene casi 250 especies
mundial, de las cuales casi 60 son neotropicales. Es muy
heterogéneo, y probablemente polifiletico, y es difícil dar
características que definan el género. Muchas especies
de Pteris tienen aristas sobre la superficie adaxial de la
costilla media de los penúltimos segmentos. En estas
especies, la cresta de la costilla media se interumpe en la
unión con la costilla media del orden superior siguiente y
se prolonga en una arista corta. Un grupo bien marcado
es él de Pteris vittata L., que se caracteriza por láminas
1-pinnadas y cuneadas, y escamas del rizoma concoloras.
Otro grupo asi es eso de P. podophylla Sw., con láminas
grandes pedatas. El nombre genérico se deriva del griego
pteron, ala o pluma. Es un nombre antiguo griego para los
helechos en general, derivado de una semejanza de las
pinnas a las alas o plumas.
Figure 249. Pteris. A, B. P. paucinervata. C, D. P. muricella. E, F. P. pungens. (From Mickel & Smith, 2004.)

505
PTERIDACEAE
Pterozonium Fée
DESCRIPTION: Terrestrial or rupestral; rhizomes
short-creeping or ascending, either scaly, bristly, or hairy,
the indumenta usually dark brown or dark reddish brown;
leaves monomorphic; laminae simple and entire
or 1-pinnate with a terminal pinna, coriaceous,
glabrous or nearly so; fronds or pinnae reniform, ovate
or oblong; veins free, parallel for most or all of their length;
rachises and costa grooved adaxially, with narrow lateral
wings on either side; veins free, simple, in some species
ending in conspicuous hydathodes; sori elongate along
the veins, nonindusiate, rarely glandular, the glands
yellow to orange; spores tetrahedral-globose, trilete; x=?
DESCRIPCIÓN: Terrestres o rupícolas; rizomas
cortamente rastreros o ascendentes, escamosos o
cerdosos o peludos, el indumento usualmente pardo
oscuro o pardo rojizo oscuro; hojas monomorfas;
láminas simples y enteras o 1-pinnadas con una
pinna terminal, coriáceas, glabras o casi así; frondes
o pinnas reniformes, ovados o oblongos; nervaduras
libres, paralelas en la mayor parte de su longitud, en
algunas especies terminando en hidatodos conspicuos;
soros alargados por las venas, no indusiados, raras
veces glandulares, las glándulas amarillas a anaranjadas;
esporas tetraédrico-globosas, triletes; x=?.

SIMILAR GENERA: Pityrogramma has more finely GÉNEROS PARECIDOS: Pityrogramma tiene láminas
divided laminae and is often farinose beneath. Hemionitis mas finamente divididas y es a menudo farinosa en
is pubescent with thinner (not coriaceous) laminae. el envés. Hemionitis es pubescente con láminas más
delgadas, no coriáceas.

COMMENTS: Pterozonium is completely neotropical COMENTARIOS: Pterozonium es completamente

and contains 14 species. It is largely endemic to the
Guayana Highland of Venezuela but with isolated
populations in Costa Rica, Colombia, Guyana, Suriname,
Peru, and northeastern Brazil. Mainly on sandstone rocks.
Pterozonium is the sister genus to Jamesonia, and these
two genera are sister to Pityrogramma. All share non-
indusiate sori with sporangia that run along the veins, and
the indumenta of the laminae are jointed hairs in stead of
scales. The genus name is derived from the Greek pteris,
fern + zone, belt. The lamina has a broad submarginal
confluent band of sori.
neotropical y consta de 14 especies. En gran parte es
endémico a las Guayana Venezolana pero con poblaciones
aislados en Costa Rica, Colombia, Guyana, Surinam, Perú
y noreste de Brasil. Principalmente en rocas de piedra
arenisca. Pterozonium es el género hermana a Jamesonia
y los dos son hermana a Pityrogramma. Todos comparten
soros no indusiatos con esporangios que corren por las
venas, e indumento de lás láminas de pelos articulados
in lugar de escamas. El nombre genérico se deriva del
griego pteris, helecho + zone, faja. La lámina tiene una faja
ancha submarginal de soros.

LITERATURE: Lellinger, D. B. 1967. Pterozonium (Filicales: Polypodiaceae). The Botany of the Guayana Highland—Part VII.
Memoirs of the New York Botanical Garden 17: 2–23.

506
Figure 250. A. Pterozonium brevifrons (A. C. Sm.) Lellinger. B. P. reniforme (Mart.) Fée. C, D. P. spectabile Maxon & A. C.
Sm. (Original, ©Robbin Moran, 2016.)

507
PTERIDACEAE
Radiovittaria (Benedict) E. H. Crane
DESCRIPTION: Epiphytic; rhizomes suberect, radial;
rhizome scales clathrate; leaves 2–50 × 0.2–1(–1.8)
cm, polystichous, simple, entire, linear to narrowly
elliptic, sessile to stalked, chartaceous to fleshy, glabrous,
with spicular idoblasts (false veins) adaxially; petioles
dark, flattened to terete; costa present; veins obscure,
anastomosing to form one row of areoles and a
submarginal vein, included veinlets absent; sori linear,
submarginal, each one parallel to the margin, superficial
or sunken in grooves; indusia absent; paraphyses with
a funnelform (obconic) apical cell; spores monolete;
gametophytes with single gemmae (not T-shaped); x=?
DESCRIPCIÓN: Epífitas; rizomas suberectos, radiales;
escamas del rizoma clatradas; hojas 2–50 × 0.2–
1(–1.8) cm, polistichous, simples, enteros, lineales a
estrechamente elípticas, sésiles o pecioladas, cartáceas a
carnosas, glabras, con idioblastos espiculares (venas falsas)
adaxialmente; pecíolos oscuros, achatados a teretes; costa
presente; nervaduras obscuras, anastomosándose formar
una hilera de areolas y una vena submarginal, venas
incluidas ausentes; soros lineales, submarginales, cada
uno paralela al margen, superficiales o hundidos en surcos;
indusios ausentes; parafisos con una célula apical
infundibuliforme (obcónica); esporas monoletes;
gametofitos con gemas solitarias (no en forma de T); x=?

SIMILAR GENERA: Vittaria differs by distichous leaves, GÉNEROS PARECIDOS: Vittaria difiere por hojas
and dark purplish brown petioles, usually narrower blades, distichous, y pecíolos pardo oscuros, láminas usualmente
and paraphyses with a slender apical cell (not obconic). más estrechas, y parafises con una célula apical delgada
(no obcónica).

COMMENTS: Radiovittaria comprises about 8 species,
all neotropical. They occur as mostly as both low-trunk
epiphytes and in the canopies of wet forests. The species
of Radiovittaria were previously classified in Vittaria, but
according to a phylogenetic study by Crane (1997), such
a treatment leaves Vittaria polyphyletic. Radiovittaria is
sister to Hecistopteris, and those two genera are sister
to Haplopteris, an Old World genus of about 20 species.
Vittaria is more distantly related within the vittarioid ferns.
Most (all?) of the species in the genus have reddish young
leaves, a characteristic that it shares with Adiantum and
certain other vittarioids. The genus name refers to the
fact that this is the only genus of vittarioids ferns with
radially arranged (polystichous) leaves.
COMENTARIOS: Radiovittaria consta de casi 8 especies,
todas neotropicales. Ocurren como epífitos en las bases
de troncos y en los doseles de bosques húmedos. Estas
especies fueron clasificadas anteriormente en Vittaria,
pero conforme a un estudio filogenético por Crane
(1997), tal clasificación deja Vittaria polifilético. Radiovittaria
es hermana a Hecistopteris, y estos dos géneros son
hermanas a Haplopteris, un género del Viejo Mundo con
casi 20 especies. Vittaria es más distantamente relacionado
dentro de los helechos vittarioides. La mayoría (¿todas?)
las especies en el género tienen hojas jóvenes rojizas,
una característica que comparta con Adiantum y ciertos
otros vittarioides. El nombre genérico refiere al hecho
que esto es el único género de helechos vittarioides con
hojas radialmente arregladas (polísticas).

LITERATURE: Benedict, R. C. 1907. The genus Antrophyum—I. Synopsis of subgenera, and the American species. Bulletin of
the Torrey Botanical Club 34: 445–458. Benedict, R. C. 1911. The genera of the fern tribe Vittarieae: Their external morphology,
venation, and relationships. Bulletin of the Torrey Botanical Club 38: 153–190. Crane, E. H., D. R. Farrar & J. F. Wendel. 1995.
Convergent simplification leads to a polyphyletic Vittaria. American Fern Journal 85: 283–305. Crane, E. H. 1997. A revised
circumscription of the genera of the fern family Vittariaceae. Systematic Botany 22: 509–517. Tryon, R. M. 1964. Taxonomic fern
notes. IV. Some American vittarioid ferns. Rhodora 66: 110–117.

508
Figure 251. Radiovittaria. A–C. R. stipitata. E–J. R. remota. B. clathrate rhizome scale. J. Paraphysis with an obconic apical cell.
(From Mickel & Smith, 2004.)

509
DRYOPTERIDACEAE
Rumohra Raddi
DESCRIPTION: Terrestrial or rupestral; rhizomes
creeping, scaly; leaves monomorphic, borne alternately
in two dorso-lateral rows on the rhizome; petioles
with three or more vascular bundles, these arranged
in a U with the two adaxial ones largest; laminae 2- to
3-pinnate-pinnatifid, the apices gradually tapered and
pinnatifid (not imparipinnate); pinnules of the lower
pinnae arranged anadromically; rachises with two
adaxial grooves separated by a raised medial ridge;
veins free; sori round; indusia peltate, circular; spores
monolete; x=41.
DESCRIPCIÓN: Terrestres o rupícolas; rizomas
rastreros, escamosos; hojas monomorfas, se nacen
alternamente in dos hileras dorsolaterales en
el rizoma; pecíolos con tres o más haces vasculares,
estas arregladas en forma de U con los dos adaxiales
lo más grandes; láminas 2- o 3-pinnado-pinnatífidas,
los ápices gradualmente reducidas y pinatífidas (no
imparipinadas); pínnulas de las pinnas inferiores arregladas
anadrómicamente; ráquises con dos surcos adaxiales
separados por una fila medial elevada; venas libres;
soros redondos; indusios peltados, circulares; esporas
monoletes; x=41.

SIMILAR GENERA: Arachniodes differs by indusia
attached by a sinus (not peltate) and lacking the raised
center in the rachis groove. Lastreopsis differs by the
upper surfaces of the rachises and costae rounded or
flat (not grooved) and puberulent. Polybotrya differs by
dimorphic sterile and fertile leaves. Saccoloma differs
by an omega-shaped vascular bundle in the base of the
petiole, and marginal or submarginal sori.
GÉNEROS PARECIDOS: Arachniodes difiere por
inducíos unídos por un sinus (no peltadas) y falta de un
centro elevado en el surco adaxial del raquis. Lastreopsis
difiere por el haz de las ráquises y costas redondeado o
achatado (no surcado) y puberulentos. Polybotrya difiere
por hojas estériles y fértiles dimorfas. Saccoloma difiere
por una haz vascular en forma de una omega en la base
del pecíolo y soros marginales o submarginales.

COMMENTS: Rumohra consists of about 7 species,
three in South America, three in Madagascar and one (R.
adiantiformis (Forst. F.) Ching) widely distributed across
the Southern Hemisphere. Two other species occur in
South America: R. berteroana (Colla) Duek & Rodrig.
is endemic to the Juan Fernández Islands, and another
(R. turficola Senna) is endemic to the southern cone of
South America. The species grow in a variety of habitats,
including forests, shrubby places, on rocks, and on sandy
soils.The rachis grooves of Rumohra are distinctive among
dryopteroid ferns. The adaxial rachis surface has consists
of two grooves with an elevated ridge between them.
Other dryopteriod ferns lack the medial elevated ridge.
Rumohra is sister to Megalastrum, but the two genera
do not share any obvious morphological similarities to
distinguish them from other dryopteroid ferns. Because
its leaves remain green a long time after being cut,
Rumohra adiantiformis is widely used in the florist trade.
The leading producers are Florida and Costa Rica. The
genus is named for Karl F. von Rumohr (1785-1843), an
art student from Dresden, Germany.
COMENTARIOS: Rumohra consta de casi 7 especies
tres en America del Sur, tres en Madagascar, y una (R.
adiantiformis (Forst. F.) Ching) ampliamente difundida en
el Hemisferio Sur. Dos otras especies existen en América
del Sur: R. berteroana (Colla) Duek & Rodrig. es endémica
a las Islas Juan Fernández, y otra (R. turficola Senna) al
Cono Sur de América del Sur. Las especies existen en
varios ambientes, incluyendo bosques, lugares abustivos,
sobre rocas e en suelos areniscos. Los surcos de los
ráquises de Rumohra son distinctivos entre helechos
dryopteroides. La superficie adaxial del los requises
constan de dos surcos con una fila elevada entre sí. Los
otros helechos dryopteroides faltan la fila elevada medial.
Rumohra es hermano a Megalastrum, pero los dos géneros
no comparten ningunas simularidades morfológicas para
distinguirlos de otros helechos dryopteroides. A causa de
sus hojas permanezcan verde mucho tiempo después de
ser cortadas, Rumohra adiantiformis se usa ampliamente
para los ramos de flores. Los productores principales son
Florida y Costa Rica. El género se nombra para Karl F. von
Rumohr (1785-1843), un estudiante de arte de Dresden,
Alemania.

LITERATURE: Baurett, L., G. Rouhan, R. Y. Hirai, L. Perrie, J. Prado, A. Salino, B. Senterre, L. Shepherd, M. Sundue, M-A.
Selosse & M. Gaudeuli. 2017. Molecular data, based on an exhaustive species sampling of the fern genus Rumohra (Dryopteridaceae), reveal a
biogeographical history mostly shaped by dispersal and several cryptic species in the widely distributed Rumohra adiantiformis. Botanical Journal
of the Linnean Society 185: 463–481. Kato, M. 1974. A note on the systematic position of Rumohra adiantiformis. Acta Phytotaxonomica et
Geobotanica 26: 52–57. Senna, R. M. 2005. Uma nova espécie de Rumohra Raddi (Dryopteridaceae – Pteridophyta) do Rio Grande do Sul,
Brasil. Iheringia, Sér. Bot. 60: 253–258. Sundue, M., R. Hirai & J. Prado. 2013. Rumohra glandulosissima (Dryopteridaceae) a new species from
the Atlantic Rainforest, and revision of the species occurring in Brazil. Systematic Botany 38: 915–924.

510
Figure 252. Rumohra adiantiformis. (Original, ©Robbin Moran, 2016.)

511
SACCOLOMATACEAE
Saccoloma Kaulf.
DESCRIPTION: Terrestrial or saxicolous; rhizomes
decumbent to erect, scaly, surrounded by persistant
petiole bases, with two cycles of vascular bundles; sterile
and fertile leaves monomorphous; petioles without
buds at the base, with an omega-shaped (Ω) vascular
bundle; laminae 1-pinnate to 5-pinnate; rachises and
costae grooved on the upper surface, the grooves glabrous
within; veins free; sori marginal or submarginal, each
at the apex of a vein, often slightly sunken in the laminar
tissue and prominulous adaxially; indusia hyaline or
scarious, usually obconic with a truncate apex, opening
toward the lamina margin; spores trilete, with distinctive
pattern of parallel, anastomosed ridges; x = ca. 63.
SIMILAR GENERA: Dennstaedtia differs by creeping,
pubescent rhizomes, leaves with at least some hairs, and
deflexed cup-shaped sori. Unlike the Dennstaedtiaceae,
Saccoloma lacks epipetiolar buds. Other ferns with highly
divided laminae can be distinguished by the form and
position of their sori.
COMMENTS: Saccoloma is the only genus in the
Saccolomataceae. It is pantropical and contains about
15 species, 10 of which are neotropical. Within the
Neotropics, the genus occurs from Mexico to southern
Brazil. All of the neotropical species occur in wet shaded
forests, and all have decompound laminae except for S.
elegans Kaulf. (southeastern Brazil) and S. chartaceum G. B.
Nair ex Cremers & K. U. Kramer (widespread). The genus
name comes from the Latin saccus, bag + loma, border.
It refers to the marginal sori of the type species that are
located in sac-like depressions.
LITERATURE: Cremers, G. & K. U. Kramer. 1989. A new subspecies of Saccoloma elegans. Studies in the Flora of the
Guianas, no. 39. Botanica Helvetica 99: 45–48. Rojas-Alvarado, A. F. 2010. Novelties in the Saccoloma inaequale complex
(Saccolomataceae) from the Neotropics. Métodos en Ecología y Sistemática 5: 1–16. Tryon, R. M. 1962. Taxonomic fern notes,
III. 4. The genus Saccoloma Kaulf. Contributions to the Gray Herbarium of Harvard University 191: 100–106.
512
DESCRIPCIÓN: Terrestres o saxícolas; rizomas
decumbentes o erectos, escamosos, envueltos por
bases de los pecíolos persistentes, con dos ciclos de
haces vasculares; hojas estériles y fértiles monomorfas;
pecíolos sin brotes epipeciolares, con un haz vascular
en forma de una omega (�); láminas 1-pinnadas
a 5-pinnadas; ráquises y costas surcados en el haz, los
surcos glabros en su interior; nervaduras libres; soros
marginales o submarginales, cada uno en el ápice
de una vena, a menudo ligeramente hundidos en el
tejido laminar y promínulos adaxialmente; indusios
hialinos o escariosos, por lo común obcónicos con un
ápice truncado, se abren hasta el margen de la lámina;
esporas triletes, con un patrón distintivo de filas paralelas,
anastomosadas; x = ca. 63.
GÉNEROS PARECIDOS: Dennstaedtia se difiere por
rizomas reptantes pubescentes, hojas con al menos
algunos pelos y soros deflexos en forma de copa. A
diferencia de las Dennstaedtiaceae, Saccoloma carece de
brotes epipeciolares. Otros helechos con láminas muy
divididas pueden distinguirse por la forma y posición de
sus soros.
COMENTARIOS: Saccoloma es el único género en
las Saccolomataceae. Es pantropical y consta de casi 15
especies, de las cuales 10 existen en el Neotrópico. En
al Neotropico, el género existe desde México has el sur
de Brasil. Todas las especies neotropicales existen en
bosques húmedos y sombreados, y todas tienen láminas
compuetas con excepción de S. elegans Kaulf. (sureste
de Brasil) y S. chartaceum G. B. Nair ex Cremers & K. U.
Kramer (muy difundida). El nombre genérico se deriva
de la Latín saccus, bolsa + loma, margen. Se refiere a los
soros marginales de la especie tipo que están ubicados
en depresiones como bolsas.
Figure 253. Saccoloma. A, B. S. inaequale. C, D. S. chartaceum. (From Tryon & Stolze, 1989.)

513
BLECHNACEAE
Salpichlaena J. Sm.
DESCRIPTION: Terrestrial; rhizomes creeping, scaly;
climbing leaves vinelike by means of a twinning
rachis; laminae 2-pinnate; pinnae with a conform terminal
pinnule; pinnules entire, the margins cartilaginous-
thickened; veins free, connected to the cartilaginous
margin; sori liinear, parallel along both sides of the
costules; indusia cylindrical when young, entire but
becoming lacerate with age, opening toward the costule
(not the margin), deciduous along with the attached
sporangia; x=40.
SIMILAR GENERA: Lygodium is the only other
neotropical fern genus that has leaves climbing by
means of a twinning rachis. It can be distinguished from
Salpichlaena by its pinnae forked at the base, absence of
scales on the axes, and sporangia borne in sorophores
along the margin.
COMMENTS: Salpichlaena consists of three species and
is entirely neotropical. It and Lygodium are the only fern
genera that have a twining rachis—a growth characteristic
totally lacking in angiosperms. Salpichlaena volubilis (Kaulf.)
J. Sm. is common and widespread in the Neotropics. In
Amazonia, S. hookeriana Alston is also common and can
be distinguished from S. volubilis by its fully contracted
fertile pinnules and the frequent presence of buds in the
pinna axils and sometimes those of the pinnules. Three
morphological notes: The cartilaginous margins of the
pinnules appear as thickened and lighter-colored than
the laminar tissue. Unlike all other ferns, the sporangia
and indusia fall as a unit. The non-climbing leaves of
Salpichlaena are 1-pinnate and resemble a Blechnum
or Parablechnum. They are important taxonomically
because they may show differences between the species
(Tuomisto & Groot, 1995). The genus name is derived
from the Greek salpinx, pipe, trumpet + chlaena, cloak.
This refers to the tubular indusium..
LITERATURE: de Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae
(Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227. Tuomisto, H. &
A. T. Groot. 1995. Identification of the juveniles of some ferns from western Amazonia. American Fern Journal 85: 1–28.
514
DESCRIPCIÓN: Terrestre; rizomas reptantes,
escamosos; hojas trepadoras como bejucos por
medio de un raquis que enrosca; láminas pinnate;
pinnas con pínnulas conformes terminales; pínnulas
enteras, los márgenes engrosados cartilaginosamente;
nervadura libre, conectada al margen cartilaginoso; soros
lineares, paralelos por ambos lados de las cóstulas;
indusios tubulares cuando jóvenes, enteros pero llegando
a ser lacerados con edad, abriéndose hasta la cóstula (no
el margen), decíduos con los esporángios unidos; x=40.
GÉNEROS PARECIDOS: Lygodium es el único otro
género neotropical de los helechos que tiene hojas
que trepan por medio de un raquis enroscador. Puede
distinguirse de Salpichlaena por sus pinnas furcadas en la
base, ausencia de escamas sobre los ejes, y esporangios
que nacen en soróforos a lo largo del margen.
COMENTARIOS: Salpichlaena consta de tres especies
y es completamente neotropical. Él y Lygodium son los
únicos géneros de los helechos que tienen ráquises
trepadores—una característica de crecimiento que se falta
completamente en las angiospermas. Salpichlaena volubilis
(Kaulf.) J. Sm. es común y muy difundida en el Neotrópico.
En la Amazonía, S. hookeriana Alston también es común y
puede distinguirse de S. volubilis por sus pínnulas fértiles
completamente contraidas y por la presencia frecuente
de brotes en las axilas de las pinnas y algunas veces en
las de las pínnulas. Tres notas morfológicas: Los márgenes
cartilaginosos de las pínnulas aparecen engrosados y de
color más claro que el tejido laminar. Diferente de todos
helechos, los esporángios e indusios caen juntos. Las
hojas no trepadoras de Salpichlaena son 1-pinnadas y se
asemejan a un Blechnum o Parablechnum. Son importantes
taxonómicamente por que pueden mostrar diferencias
entre especies (Tuomisto & Groot, 1995). El nombre
genérico se deriva del griego salpinx, pipa, trompeta +
chaleña, capa. Esto se refiere al indusio tubular.
Figure 254. Salpichlaena volubilis. (From Tryon & Stolze 1993.)

515
SALVINIACEAE
Salvinia Micheli in Adans.
DESCRIPTION: Floating aquatics; roots absent; stems
hairy, thin, branched; leaves in verticels of 3, dimorphous,
with 2 green floating ones and single, whitish, submerged
one that is finely divided and resembles a root; floating
leaves 0.4–3 cm in diameter, circular, ovate or
oblong, entire, typically with papillae on the upper
surface, the papillae when present with 3 or 4 hairs at
their apices, these usually united and darkened at the
apex, forming a cage; veins anastomosing, the areoles
elongate, polygonal, without included veinlets; sori borne
on the submerged leaf; indusia globose, whitish; x=9.
DESCRIPCIÓN: Acuáticas flotantes; raíces ausentes;
tallos pelosos, delgados, ramificados; hojas en verticilios
de 3, dimorfas, con 2 verdes flotantes y una sin clorofila,
sumergida, finamente dividida y semejante a una raíz;
hojas flotantes 0.4–3 cm de diámetro, redondeadas,
ovadas u oblongas, enteras, típicamente con papilas
en el haz, las papilas cuando presentes con 3 o 4 pelos en
sus ápices, estos usualmente unidos y oscurecidos en los
ápices, formando una jaula; nervaduras anastomosadas, las
aréolas alargadas, poligonales, sin venillas includas; soros
sobre la hoja sumergida; indusios globosos, blanquecinos;
x=9.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Salvinia is pantropical and subtropical COMENTARIOS: Salvinia es pantropical y subtropical y

and consists of 9 species. It is found in stagnant or slow-
moving water at low elevations. The submerged leaf
resembles a root because it is whitish and finely divided;
however, it is actually a leaf because it bears sporangia
and its development and anatomy is that of a leaf, not a
root. Most neotropical species have long papilla on the
upper surface of the floating leaves, and these papillae
bear 4 hairs at their apex on top of each. In those species
belonging to the S. auriculata complex, these hairs are
united at their tips, and the entire structure, (papilla and
its hairs) resembles an egg beater. The only neotropical
species with hairs not united at the apices is Salvinia
minima Baker. A few species lack papillae on the upper
surface of the floating leaves, such as S. sprucei Kuhn. A well
known species is Salvinia molesta D. S. Mitch., which is a
aggressive weed introduced in Africa, Sri Lanka, Indonesia,
Australia, and the southern United States (Moran, 2004).
The genus name honors Antonio Maria Salvini (1633–
1729), an Italian professor of Greek who helped Micheli
with his botanical work.
consta de 9 especies. Se encuentra en agua estancada o
con flujo lento en elevaciones bajas. La hoja submergida
se parece a una raíz porque es blanquecino y finamente
dividida; no obstante, es en realidad una hoja por que lleva
esporangios y su desarrollo y anatomía es eso de una hoja,
no de una raíz. La mayoría de las especies neotropicales
tienen papilas largas en la superficie superior de las hojas
flotantes, y estas papilas llevan en sus ápices 4 pelitos. En
las especies que pertenecen al complejo de S. auriculata,
estos pelitos se unen en sus ápices, y la estructura entera
(papila más pelitos) se asemaja un batidor de huevos.
La única especies neotropical con pelos no unidos en
sus ápices es Salvinia minima Baker. Algunas especies
carecen de papilas en el haz de las hojas flotantes, tales
como S. sprucei Kuhn. Una especie bien conocida es
Salvinia molesta D. S. Mitch., la cual es una maleza agresiva
introducida en África, Sri Lanka, Indonesia, Australia, y
el sur de los Estados Unidos (Moran, 2004). El nombre
genérico honora Antonio Maria Salvini (1633–1729), un
profesor italiano de griego quien ayudo Micheli con sus
obras botánicas.

LITERATURE: de la Sota, E. R. 1962. Contribución al conocimiento de las “Salviniaceae” neotropicales, I–III. Darwiniana 12:
465–520; de la Sota, E. R. 1963. Contribución al conocimiento de las “Salviniaceae” neotropicales, IV. Darwiniana 12: 612–623;
1964. Contribución al conocimiento de las “Salviniaceae” neotropicales, V. Darwiniana 13: 529–536; de la Sota, E. R. 1976.
Sinopsis de las especies argentinas del género Salvinia Adanson (Salviniaceae, Pteridophyta). Bol. Soc. Argent. Bot. 17: 47–50.
Forno, I. W. 1983. Native distribution of the Salvinia auriculata complex and keys to specie identifications. Aquatic Botany 17:
71–83. Herzog, R. 1935. Ein Beitrag zur systematik der Gattung Salvinia. Hedwigia 74: 257–284. Moran, R. C. 2004. A Natural
History of Ferns. Timber Press, Portland, Oregon. Weatherby, C. A. 1937. A further note on Salvinia. American Fern Journal 27:
98–102.

516
Figure 255. Salvinia. A–D. S. minima. E–K. S. auriculata. (From Mickel & Smith, 2004.)

517
OPHIOGLOSSACEAE
Sceptridium Lyon
DESCRIPTION: Terrestrial; stems subterranean, fleshy,
erect, usually unbranched, without hairs or scales; roots
fleshy, without hairs, not proliferous, not or sparsely
branched; leaves one per plant, leathery or fleshy, win-
tergreen in the temperate zones, often bronzing in win-
ter; lf sheaths closed; lf primordia pilose; trophophores
often > 7 cm wide, 3–4-pinnate, with a midrib; veins free,
forked; sporophores attached below ground well below
the trophophore, often absent or aborted on juvenile
plants, ; x=41. .
DESCRIPCIÓN: Terrestres; tallos subterráneos, carno-
sos, erectos, usualmente no ramificados, sin escamas o
pelos; raíces carnosas, sin pelos, no prolíferas, no o es-
parcidamente ramificadas; hojas una por planta, coriáceas
o carnosas, verde a bronce durnate el invierno en las
zonas templadas; vainas de las hojas cerradas; primordias
de las hojas pilosas; trofoforos a menudo > 7 cm de an-
cho, 3–4-pinadas, con una costa central; venas libres, fur-
cadas; soróforos unidos abajo del suelo muy abajo del
trofoforo, a menudo ausentes o abortados en plantas
juveniles; x=41.

SIMILAR GENERA: The other genera of Ophioglossa-
ceae differ by the sporophores united to the petiole at or
just below the trophophore, not below ground. Anemia
differs by 2 erect fertile spikes (sorophores) and densely
pubescent rhizomes.
GÉNEROS PARECIDOS: Los otros géneros de Ophio-
glossaceae difieren por los esporoforos unidos al pecíolo
justo abajo del trofoforo, no abajo del suelo. Anemia di-
fiere por 2 espigas fértiles (soróforos) erectas y rizomas
densamente pubescentes.

COMMENTS: Sceptridium occurs mostly eastern North
America and eastern Asia; it is absent from Africa and
Australia. It contains 20–25 species, of which about five
5 occur in the Neotropics. Sceptridium typically grows
in open or semi-open habitats such as lawns, pastures
and second-growth woodlands. Usually, when one plant
is found, others are nearby. In the temperate zones, the
trophophores are produced late in the growing season,
usually August to October, and the sorophores persist
over the winter, remaining green or turning bronze. The
genus name is derived from the Latin sceptrum, scepter,
and –idium, a diminutive suffix; appl. to the fertile spike.
COMENTARIOS: Sceptridium existe principalmente
en America del Norte oriental y Asia oriental; es aus-
tente de África y Australia. Consta de 20–25 especies,
de las cua20–25 species, les 5 existen en el Neotrópi-
co. Sceptridium típicamente crece en hábitats abiertos o
semi-abiertos tales como céspedes, potreros y bosques
secondarios. Usualmente, cuando se encuentra una planta,
otras están cerca. En las zonas templadas, las trofoforos se
producen tarde en la estación de crecimiento, usualmente
en Agosto o Octubre, y los trofoforos persisten sobre
el invierno, como verdes o quedándose bronceado. El
nombre genérico se deriva del Latín sceptrum, scepter,
y –idium, un sufijo diminutivo; se refiere a la espíga fértil.

LITERATURE: Butters, F. K. 1917. Botrychium virginianum and its American varieties. Rhodora 19: 209. Clausen, R. T.
1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19(2): 1–177, figs. 1–33. Hauk, W. D., C. R.
Parks & M. W. Chase. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL and trnL–F plastid DNA sequences
and morphology. Molecular Phylogenetics and Evolution 28: 131–151. Hauk, W. D., L. Kennedy & H. M. Hawke. 2012. A
phylogenetic investigation of Botrychium s.s. (Ophioglossaceae): evidence from three plastid DNA sequence datasets. Systematic
Botany 37: 320–330.

518
Figure 256. A, B. Sceptridium decompositum. (From Mickel & Smith, 2004.)

519
SCHIZAEACEAE
Schizaea Sm.
DESCRIPTION: Terrestrial; stems compact, horizontal
or erect, pubescent; leaves simple to several times
dichotomous, the ultimate divisiones foliaceous to scarcely
laminar; petioles glabrous, as long or longer than the
lamina, with an adaxial groove; veins free; fertile segments
(sporangiophores) pinnatifid, curved, borne at the
tips of the laminae; sporangia erect, oblong, with
an apical annulus, in 2 rows; spores monolete, foveolate;
gametophytes epigeal, green, filamentous; x=77, 94, 96,
103.
DESCRIPCIÓN: Terrestres; rizomas compacto,
horizontal o erecto, peludos; hojas simples a varias veces
dicotómicas, las últimas divisiones foliáceas a escasamente
laminares; pecíolo glabro, tan largo como la lámina a más
largo que ella, con un surco adaxial; nervaduras libres;
segmentos fértiles (esporangióforos) pinnatos,
curvados, terminales; esporangios rectos, oblongos,
con anillo apical, 2 hileras; esporas monoletes, foveolate;
gametofitos epigeales, verdes, filamentosos; x=77, 94, 96,
103.

SIMILAR GENERA: Actinostachys differs by straight GÉNEROS PARECIDOS: Actinostachys difiere por
(not curved) fertile segments that are digitate and more segmentos fértiles rectos, digitados, y más que 2 cm de
than 2 cm long. largo.

COMMENTS: Schizaea is pantropical and consists of COMENTARIOS: Schizaea es pantropical y consta de

about 45 species, of which 8 occur in the Neotropics.
They typically grow below 600 m on shaded forest floors
or open sandy areas. The species with well-developed
laminae are sometimes separated as Lophidium, but it is
doubtful they represent a monophyletic group. Schizaea
elegans (Vahl.) Sw. has a broad, expanded lamina and
is widespread in the Neotropics. Several species with
grass-like stems (e.g., S. incurvata Schkuhr) exhibit a
clumped habit caused by repeated proliferations from
buds at the base of the petioles. This habit needs better
documentation by study of living plants in the field.
Schizaea has filamentous, gametophytes that form a
mycorrhizal relationships. Specialized, paired, globose cells
called rhiziodophores house the fungus. The sister genus
Actinostachys has subterranean, cylindrical gametophytes.
Schizaea is sometimes defined to include Actinostachys
(which see). The genus name is derived from the Greek
schizein, to split. The fan shaped fronds of some species
are split toward the base.
casi 45 especies, de las cuales 8 existen en el Neotrópico.
Típicamente crecen bajo de 600 m en el suelo de
bosques sombreados or áreas arenosas abiertas. Las
especies con láminas desarolladas están separadas a
veces como Lophidium, pero es dudoso si representan un
grupo monofilético. Schizaea elegans (Vahl.) Sw. tiene una
lámina expandida y está muy difundida en el Neotrópico.
Varias especies con hojas en forma de gramíneas stems
(e.g., S. incurvata Schkuhr) exhiben un hábito amacollado
se debe a las proliferaciones de brotes en la base de
los pecíolos. Este hábito se necesita documentación
mejor por estudio de plantas vivas en el campo. Schizaea
tiene gametofitos filamentosos que forman una relación
micotrófica. Un par de células especializadas, llamados
rizoidóforos, hospeden el hongo. El género hermano
Actinostachys tiene gametofitos subterráneos cilíndricos.
A veces se define Schizaea para incluir a Actinostachys
(véase).El nombre genérico se deriva del griego schizein,
partir o hender. Las hojas en forma de un abanico de
algunas especies son hendidas hacia la base.

LITERATURE: Britton, E. G. & A. Taylor. 1901. Life history of Schizaea pusilla. Bulletin of the Torrey Botanical Club 28:
1–19, + 6 plates. Lellinger, D. B. 1969. Schizaeaceae (Filicales). In: B. Maguire and collaborators. The botany of the Guayana
Highland—Part VIII. Memoirs of the New York Botanical Garden 18: 2–11. León, B., H. Beltrán & P. Fine. 2005. El género
Schizaea (Schizaeaceae) en el Perú. Revista Peruana de Biología 12: 97–102. Murillo-Pulido, M. T. 1986. Estudio preliminar del
género Schizaea en Colombia. Caldasia 15: 93–101. Takeuchi, M. 1960. O gênero Schizaea no Amazonia. Boletim de Museo
Paraense Emilio Goeldi 5: 1–26.

520
Figure 257. Three species of Schizaea. A. Schizaea elegans (silhouettes) showing lamina variation from entire to several times
divided. B. Schizaea pusilla, the curly-grass fern, exhibiting curly basal leaves and long, erect fertile ones. C. Schizaea sprucei. (A, B,
original, ©Robbin Moran, 2018; C from Hooker, Icones Filicum, 1867.)

521
PTERIDACEAE
Scoliosorus T. Moore
DESCRIPTION: Epiphytic; rhizomes short-creeping
solenostelic, short-creeping, dorsiventral; rhizome scales
clathrate; leaves 30–70 cm long × 1.5–20 cm wide,
simple, entire, glabrous, fleshy, distichous, clumped;
petioles short or absent; costae present or absent; veins
areolate, with several series of areoles between the
costa and margin; sori oblique to the costae; indusia
absent; paraphyses with a spherical apical cell;
spores monolete, nongreen; gametophytes with paired
gemmae; x=?
DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
solenostélicos, dorsiventrales; escamas del rizoma
clatradas; hojas 15–70 × 1.5–20 cm, simples, enteras,
glabras, carnosas, dísticas, amacolladas; pecíolos cortos
a ausentes; costas presentes o ausentes; nervaduras
areoladas, con varias series de areolas entre la costa y el
margen; soros oblícuos a la costa; indusios ausentes;
parafisos con una célula apical esférica; ssporas
monoletes, no verdes; gametofitos con yemas pareadas;
x=?

SIMILAR GENERA: Polytaenium differs by trilete spores
and lack of paraphyses. The rachis of this species wide,
flattened (not or only slightly raised) and often much
lighter in color than the lamina tissue on either side. In
Polytaenium the rachis is more prominent. Ananthacorus,
Radiovittaria, and Vittaria differ by a single, long submarginal
sorus.
GÉNEROS PARECIDOS: Polytaenium difiere por
esporas triletes y falta de parafisos. El raquis de esta
especie es ancho, achatado (no o poco levantado) y
a menudo más claro en color que el tejido laminar en
ambos lados. En Polytaenium el rachis es más prominentes.
Ananthacorus, Radiovittaria y Vittaria difieren por un sólo
soro largo submarginal.

COMMENTS: Scoliosorus comprises three species, one
of which grows from Mexico to Panama (S. ensiforme
(Hook.) T. Moore). The other two occur in Africa and
nearby islands of the Indian Ocean. The neotropical
species grows as an epiphyte in wet, shaded forests at
middle elevations. The genus name is derived from the
Greek scolios, curved, tortuous + soros. The sori are
linear and more or less sinuous.
COMENTARIOS: Scoliosorus consta de tres especies,
una de las cuales existe desde México hasta Panamá
(S. ensiforme (Hook.) T. Moore). Las dos otras existen
en África e islas cercanas del Océano Indio. La especie
neotropical crece como epífito en bosques húmedos
sombreados en elevaciones medias. El nombre genérico
se deriva del griego scolios, curvado, tortuo + soros. Los
soros son lineares y más o menos sinuosos.

LITERATURE: Benedict, R. C. 1907. The genus Antrophyum--I. Synopsis of subgenera, and the American species. Bulletin of
the Torrey Botanical Club 34: 445–458. Benedict, R. C. 1911. The genera of the fern tribe Vittarieae: Their external morphology,
venation, and relationships. Bulletin of the Torrey Botanical Club 38: 153–190. Crane, E. H. 1997. A revised circumscription of
the genera of the fern family Vittariaceae. Systematic Botany 22: 509–517. Tryon, R. M. 1964. Taxonomic fern notes. IV. Some
American vittarioid ferns. Rhodora 66: 110–117.

522
Figure 258. A–F. Scoliosorus ensiforme. (From Mickel & Smith, 2004.)

523
POLYPODIACEAE
Serpocaulon A. R. Sm.
DESCRIPTION: Epiphytic; rhizomes long‑creeping,
but short-creeping in a few species, the phyllopodia
(0.5–)5–20 times the rhizome width apart, sometimes
glaucous; black sclerenchyma strands absent; vascular
bundles sometimes with sclerenchyma sheaths; scales
peltate, clathrate (sometimes the luminae are occluded),
the side walls dark brown or black, surfaces glabrous
(lacking rhizoids). Petioles grooved adaxially (never
terete); laminae pinnatifid to pinnate, rarely simple (S.
levigatum) or shallowly pinnatifid, monomorphic; lamina
tissue glabrous or hairy, without scales or with sparse
clathrate scales along the costae and a few on the
rachises; veins regularly anastomosing, the areoles
with a single, free, included vein (goniophlebioid
veins). Sori round or nearly so, in 1–10 rows between
costae and margins, terminal on a free included vein;
indusia absent; paraphyses absent or present and short,
2--3-celled, glandular, not easily seen; sporangia glabrous;
spores yellowish, monolete; x=37.
DESCRIPCIÓN: Epífitas; rizomas largamente
rastreros, pero cortamente rastreros en algunas especies,
los filopodios (0.5–)5–20 veces distante la anchura del
rizoma; esclerínquima negra ausente; haces vasculares
a veces con vainas de esclerínquima; escamas peltadas,
clatradas (a veces la luminas son ocluidas), las paredes
laterales pardos o negros, superficies glabras (carece de
rizoides); pecíolos surcados adaxialmente (nunca teretes);
láminas pinatífidas a pinnadas, raras veces simples (S.
levigatum) o someramente pinatífidas, monomorfas; tejido
de las láminas glabro o pubescente, sin escamas o con
escamas clatradas esparcidas por las costas y algunas sobre
los requises; venas anastomosándose regularmente,
las areolas con una sóla venilla incluida (nervación
gonofleboide); soros redondos o casi así, en 1–10 hileras
entre la costa y margen, terminales en el ápice de
una vena incluida; inducíos ausentes; esporas amarillas,
monoletes; x=37.

SIMILAR GENERA: Polypodium differs by (usually) free
veins, non-clathrate rhizome scales, and single row of sori
between the costa and margin. Pleopeltis differs by scaly
laminae. Pecluma differs by free veins, or if anastomosing
(rare) then not in a goniophelbioid pattern, a single row
of sori on each side of the costae, often reduced proximal
pinnae, non-clathrate, often comose rhizome scales,
short-creeping rhizomes, and pinnae usually more than
30 pairs.
GÉNEROS PARECIDOS: Polypodium difiere por
(usualmente) venas libres, escamas del rizoma no
clatradas y una sola hilera de soros entre la costa y el
margen. Pleopeltis difiere por láminas escamosas. Pecluma
difiere por venas libres, o si raras veces anastomosadas
entonces no en un patrón goniofleboide, un sola hilera
de soros en cada lado de la costa, a menudo pinnas
proximales reducidas, escamas del rizoma no clatradas
y a menudo comosos, rizomas cortamente rastreros y
pinnas usualmente más de 30 pares.

COMMENTS: Serpocaulon contains about 40 species, all
neotropical. About 14 species occur in the Antilles and
Central America, and 26 are restricted to South America,
10 of which occur in southeastern Brazil. All grow
primarily as epiphytes in wet forests from about 0–2500
m. The generic name comes from the Latin serpens, snake
+ caulon, stem, referring to the long-creeping rhizome.
COMENTARIOS: Serpocaulon contiene casi 40 especies,
todos neotropicales. Casi 14 especies existen en las
Antillas y América Central, y 26 son restingidas al América
del Sur, 10 de las cuales existen en el sureste de Brasil.
Todos crecen principalmente como epífitos en bosques
húmedos de 0–2500 m. El nombre genérico se deriva
del latín serpens, serpiente + caulis, tallo, refiriéndose al
rizoma largamente rastrero.

LITERATURE: Kreier, H.-P., M. Rex, K. Weising, M. Kessler, A. R. Smith & H. Schneider. 2008. Inferring the diversification
of the epiphytic fern genus Serpocaulon (Polypodiaceae) in South America using chloroplast sequences and amplified fragment
length polymorphisms. Plant Systematics and Evolution 274: 1–16. Labiak, P. H. & J. Prado. 2008. New combinations in
Serpocaulon and a provisional key for the Atlantic rain forest species. American Fern Journal 98: 139–159. Sanin, D. 2018.
Serpocaulon (Polypodiaceae). Flora de Colombia 32: 1–132. Smith, A. R., H.-P. Kreier, C. H. Haufler, T. A. Ranker & H.
Schneider. 2006. Serpocaulon (Polypodiaceae), a new genus segregated from Polypodium. Taxon 55: 919–930.

524
Figure 259. Serpocaulon. A–C. S. pleurosorum. D–E. S. falcaria. F. S. fraxinifolium. G–J. S. triseriale. L–M. S. dissimile. (From
Mickel & Smith, 2004.)

525
CYATHEACEAE
Sphaeropteris Bernh.
DESCRIPTION: Terrestrial; stems arborescent, up
to 15 m tall; sterile and fertile leaves 2-3(-5) m long,
monomorphous; petioles without spines, scaly, the
scales 1–4 cm long, linear, concolorous, white or
pale brownish, the cells of the borders and center
equal in shape and orientation, margins dark-
denticulate; laminae 2-pinnate-pinnatisect, the apex
gradually tapered, pinnatisect; rachis and costae pubescent
adaxially, the hairs thick, strigose, multiseptate; veins free;
sori round, the receptacle elevated, subglosbose; indusia
cup-shaped or globose; spores tetrahedral-globose,
usually 64 per sporangium; x=69.
SIMILAR GENERA: All other genera in the family have
petiolar scales with the cells of the margin differing in
shape and orientation from those of the center. Dark-
denticulate margins occur only in a few species of Cyathea.
Many species of Cyathea have spiny petioles, whereas
Sphaeropteris always have nonspiny petioles.
COMMENTS: Sphaeropteris is pantropical and contains
about 110 species, 6 of which are Neotropical. They
grow in wet montane forests from 600–2700 m. The
Neotropical species were monographed by Tryon (1971),
who pointed out their correlation with geography: S. brunei
(Christ) R. M. Tryon (Costa Rica, Panama n. Colombia),
S. cuatrecasasii R. M. Tryon (se. Colombia), S. gardneri
(Hook.) R. M. Tryon (se. Brazil), S. horrida (Liebm.) R. M.
Tryon (Mexico to Nicaragua), S. insignis (D. C. Eaton) R. M.
Tryon (Greater Antilles), S. quindiuensis (Karst.) R. M.Tryon
(Colombia to Bolivia). Their petiole scales are distinctive,
being composed of cells with the same orientation and
narrow rectangular shape. Also, dark marginal teeth
are present, and they are more or less regular in size,
spacing, and orientation. In contrast, the petiole scales of
Alsophila, Cyathea, and Gymnosphaera have along their
borders sinouous diverging cells. Sphaeropteris subgen.
Sclephropteris P. G. Windisch is now classified in Cyathea.
Sphaeropteris is sister to the rest of the Cyatheaceae
(Korall et al., 2006). The genus name is derived from the
Greek sphaeros, sphere + pteris, fern. It refers to the
spherical indiusia.
LITERATURE: Korall, P., K. M. Pryer, J. S. Metzgar, H. Scheider & D. S. Conant. 2006. Tree ferns: monophyletic groups and their
relationships as revealed by four protein-coding plastid loci. Molecular Phylogenetics & Evolution 39: 830–845. Korall, P., D. S. Conant, J.
S. Metzgar, H. Schneider & K. M. Pryer, 2007. A molecular phylogeny of scaly tree ferns (Cyatheaceae). American Journal of Botany 94:
873–886. Tryon, R. M. 1971. The American tree ferns allied to Sphaeropteris horrida. Rhodora 73: 1–19.
526
DESCRIPCIÓN: Terrestres; tallos arborescentes,
hasta 15 m de alto; hojas estériles y fértiles 2-3(-5) m
de largo, monomorfas; pecíolos sin espinas, escamosos,
las escamas 1–4 cm de largo, lineares, concoloras,
blancas o cafés claros, las células de los bordes y del
centro iguales en forma y orientación, los márgenes
oscuro-denticulados; láminas 2-pinnado-pinnatisectas,
el ápice gradualmente reducido, pinnatisecto; raquis y
costas pubescentes adaxialmente, los pelos gruesos,
estrigosos, multiseptados; nervaduras libres; soros
redondos, el receptáculo elevado, subgloboso; indusios
en forma de copa o globosos; esporas tetrahédrico-
globosas, 64 por esporangio; x=69.
GÉNEROS PARECIDOS: Todos los otros géneros en
la familia tienen escamas del pecíolo con las células de los
bordes diferentes en forma y orientación a las del centro.
Márgenes oscuro-denticulados se encuentran en sólo
pocas especies de Cyathea. Muchas especies de Cyathea
tienen pecíolos espinosos, mientras que Sphaeropteris
siempre tiene pecíolos no espinosos.
COMENTARIOS: Sphaeropteris es pantropical y consta
de casi 110 especies, 6 de las cuales son Neotropicales.
Habitan bosques húmedos desde 600–2700 m. Las
especies Neotropicales fueron monografiadas por
Tryon (1971), quien señaló su correlación con geografía:
S. brunei (Christ) R. M. Tryon (Costa Rica, Panama n.
Colombia), S. cuatrecasasii R. M. Tryon (se. Colombia), S.
gardneri (Hook.) R. M.Tryon (se. Brasil), S. horrida (Liebm.)
R. M. Tryon (México a Nicaragua), S. insignis (D. C. Eaton)
R. M. Tryon (Antillas Mayores), S. quindiuensis (Karst.)
R. M. Tryon (Colombia a Bolivia). Las escamas de sus
pecíolos son distinctivas, siendo compuestas de células
con la misma orientación y forma rectangular estrecho.
También, dientes oscuros marginales son presentes. En
contraste, las escamas peciolares de Alsophila, Cyathea y
Gymnosphaera tienen por sus bordes células divergentes
sinuosas. Sphaeropteris subgén. Sclephropteris P. G.
Windisch se clasifica ahora en Cyathea. Sphaeropteris es
hermano al resto de Cyatheaceae (Korall et al., 2006). El
nombre genérico se deriva del griego sphaeros, esfera +
pteris, helecho. Refiere a los indusios esféricos.
Figure 260. A. Cyathea poeppigii. B, C. Sphaeropteris quindiuensis. C. Petiole scale showing conform cells with same shape
and orientation, and dark marginal teeth. (From Tryon & Stolze, 1989.)

527
LINDSAEACEAE
Sphenomeris Maxon
DESCRIPTION: Terrestrial or rupicolous; rhizomes
creeping, slender, hairy, the hairs septate, dark brown; leaves
3- to 4-pinnate, distichous, erect and determinate (not
scandent), glabrous; laminae often yellow-green; pinnae
anadromic; ultimate segments linear or narrowly
cuneate, 1–4 mm apically; veins few, bifurcate, free;
sori terminal on the ultimate segments, supplied by
1–4 veins; indusia erose or dentate, attached at base and
sides, pocket-like; spores trilete; x=38, 39.
DESCRIPCIÓN:Terrestres o rupícolas; rizomas rastreros,
delgados, pelosos, los pelos septados, pardo oscuros;
hojas 3- a 4-pinnadas, dísticas, erectas y determinadas (no
escandentes), glabras; láminas a menudo amarillo verdes;
pinnas anadrómicas; últimos segmentos lineares o
estrechamente cuneados, 1–4 mm apicalmente;
nervaduras escasas, bifurcadas, libres; soros terminales
en los segmentos últimos, suministrados por 1–4
venas; indusios erososos o dentados, unidos en la base
y por los lados, con forma de una bosilla; esporas triletes;
x=38, 39.

SIMILAR GENERA: Odontosoria differs by leaves longer GÉNEROS PARECIDOS: Odontosoria difiere por las
(1–6 m), scandent, and indeterminate. hojas más largas (1–6 m), escandentes e indeterminadas.

COMMENTS: Sphenomeris consists of a single species,
S. clavata (L.) Maxon, which grows on limestone from
0– 1000 m. It occurs in Florida, the Bahamas, Greater
Antilles, and southern Mexico. Other species previously
classified in Sphenomeris have been found to be nested
in Odontosoria (Lehtonen et al., 2010). The genus name
is derived from the Greek sphen-, wedge, + meris, part;
referring to the shape of the ultimate segments.
COMENTARIOS: Sphenomeris consta de una sola
especie, S. clavata (L.) Maxon, la cual existe sobre piedra
caliza desde 0–1000 m. Existe en Florida, las Bahamas,
las Antillas Mayores y México del sur. Otras especies
previamente clasificadas en Sphenomeris han sido
encontradas anidadas en Odontosoria (Lehtonen et al.,
2010). El nombre del género se deriva del griego sphen-,
wedge, + meris, parte, refiriéndose a la forma de los
segmentos últimos.

LITERATURE: Kramer, K. U. 1957. A revision of Lindsaea in the New World, with notes on allied genera. Acta Botanica
Neerlandica 6: 97–290. Lehtonen, S., H. Tuomisto, G. Rouhan & M. J. M. Christenhusz. 2010. Phylogenetics and classification
of the pantropical fern family Lindsaeaceae. Botanical Journal of the Linnean Society 163: 305–359.

528
Figure 261. Sphenomeris clavata. Note terminal sori in C. (From Mickel & Smith, 2004.)

529
LYCOPODIACEAE
Spinulum A. Haines
DESCRIPTION: Terrestrial; plants with a main long-
creeping surficial runner that alternately gives rise to
branched upright shoots, these branched 1–4 times, the
branchlets of nearly equal lengths and with prominent
annual growth constrictions; trophophylls terminated
by a minute, firm spine isophyllous, entire or toothed,
arranged in alternating pseudowhorls of 4 or 5, collectively
forming 8–10 ranks; strobili distinct, terminal, solitary,
sessile, 1–4 cm long; sporophylls becoming stramineous
before sporangial dehiscence, subpeltate, the stalk with a
basiscopic, thin wing adnate to the strobilus axis; sporangia
reniform, isovalvate, the epidermal cells with occasional
in- and evaginations on the sidewalls; spores trilete,
foveolate on the distal hemisphere, obscurely papillate
on the proximal faces; gametophytes subterranean,
holosaprophytic, disk-shaped; x=34.
DESCRIPCIÓN: Terrestres; plantas con rizoma s
largamente trepadores superficiales que dan lugar
alternamente a vástagos erectos ramificados, éstos
ramificados 1–4 veces, las ramitas de largas casi iguales y
con prominentes constricciones anuales de crecimiento;
trofofílios terminados por una espina minuta
firma, isofílios, enteros o dentados, arreglados en en
pseudoverticelos de 4 o 5, colectivamente formando 8–10
hileras; estróbilos distinctos, terminales, solitários,
séssiles, 1–4 cm de largo; esporofilas tornándose
estramíneous antes de dehiscencia esporangial,
subpeltadas, el pecíolo con una ala delgada decurrente
sobre el eje del estróbilo; esporángios reniformes,
isovalvados, las células del epidermis con invaginaciones
en las paredes laterales; esporas triletes, foveoladas en
el hemisfério distal, obscuramente papiladas en las caras
proximales; gametofitos suberráneos, holosaprofíticos,
disciformes; x=34.

SIMILAR GENERA: Austrolycopodium differs by GÉNEROS PARECIDOS: Austrolycopodium difiere

pedunculate strobili and peltate sporophylls (i.e., the base
lacks a thin wing ducrrent on the axis of the strobilus).
Lycopodium usually differs by pedunulate strobili and erect
lateral shoots whose branches are of obviously different
lengths.
por estrobilos pedunculados y esporofilios peltados (i.e.,
la base falta una ala delgada decurrente en el eje del
estrobilo). Lycopodium usualmente difiere por estrobilos
pedunculados y vastagos erectos laterales cuyos ramitos
son de largos obviamente diferentes.

COMMENTS: Spinulum consists of 3 species: S. COMENTARIOS: Spinulum consta de 3 especies: S.

annotinum (L.) A. Haines, S. canadense (Ness.) A. Haines,
and S. subarcticum (V. N. Vassil.) A. Haines. All three
occur in North America and Eurasia, primarily in north-
termperate and boreal regions. The genus name alludes
to the minute, firm spine that terminates the tips of the
trophophylls. In Latin spinula means spine; "spinulum" is
not a Latin word. This is fortunate because descripitve
morphological terms are not allowed to be used as
generic names according to the Code. .
annotinum (L.) A. Haines, S. canadense (Ness.) A. Haines y
S. subarcticum (V. N. Vassil.) A. Haines. Las tres existen en
América del Norte y Eurásia, principalmente en zonas
templadas y boreales. El nombre del género se refiere
a la espina minuta firma que termina los ápices de los
trofofílios. En latín, spinula significa espina; "spinulum" no
es una palabra latina. Esto es afortunado porque términos
descriptivos morfológicos no se permiten como nobres
genéricos conforme al Codigo.

LITERATURE: Callaghan, T. V., B. M. Svensson & A. Headley. 1986. The modular growth of Lycopodium annotinum. Fern
Gazette 13: 65–76. Haines, A. 2003. The families Huperziaceae and Lycopodiaceae of New England, a taxonomic and ecological
reference.V. F.Thomas Co., Bowdin, Maine. Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica
92: 153–178. Øllgaard, B. & K. Tind. 1993. Scandinavian ferns. Rhodos, Copenhagen. Primack, R. B. 1973. Growth patterns of
five species of Lycopodium. American Fern Journal 63: 3–7.

530
Figure 262. Spinulum annotinum. (Modified from Small, 1935; spores from Øllgaard & Tind, 1993.)

531
THELYPTERIDACEAE
Stegnogramma Blume
DESCRIPTION: Terrestrial or rupicolous; rhizomes
short-creeping to erect; petioles with 2 vascular bundles;
laminae 1-pinnate-pinnatifid, proximal pinnae not or only
slightly reduced; distal pinnae broadly adnate and
often decurrent on the rachis; aerophores absent; basal
veins ending at the base of the sinus (in our one species)
or anastomosing to form a complex web (several Old
World species); indument of acicular hairs; sori elongate,
exindusiate; sporangial capsules setose; spores spinulose.
x=36.
DESCRIPCIÓN: Terrestres o rupícolas; rizomas
cortamente rastreros a erectos; pecíolos con 2
haces vasculares; láminas 1-pinnada-pinnatífidas;
pinnas proximales no reducidas o poco reducidas;
pinnas distales anchamente adnatas y a menudo
decurrentes al raquis; aeróforos ausentes; nervaduras
libres, encontrándose con el margen en el seno
(Mesoamérica) o anastomosándose para formar una
red compleja (en algunas especies del Viejo Mundo);
indumento de tricomas aciculares; soros alargados,
exindusiados; cápsulas esporangiales setosas; esporas
finamente espinulosas; x=36.

SIMILAR GENERA: Amauropelta differs by having one
or more pairs of reduced proximal pinnae. Cyclosorus
differs by round, indusiate sori. Steiropteris differs by having
a cartilaginous keel between the base of the sinus and
the costa. Goniopteris differs by the presence of minute,
forked hairs (use handlens).
GÉNEROS PARECIDOS: Amauropelta difiere por
tener uno o más pares de pinnas proximales reducidas.
Cyclosorus difiere por soros redondeados, indusiados.
Steiropteris difiere por tener una quilla cartilaginosa
entre la base del sino y la costa. Goniopteris difiere por la
presencia de pelitos furcados (use lupa).

COMMENTS: Stegnogramma has 18 species worldwide, COMENTARIOS: Stegnogramma consta de 18 especies

only one of which occurs in the Neotropics: S. pilosa (M.
Martens & Galeotti) K. Iwats. It occurs from northwestern
Mexico to Honduras, from 1800–2700 m, usually on
limestone. In rock houses of the southeastern United
States (Alabama) occurs S. burksiorum (J. E. Watkins
& Farrar) Weakley. It differs subtly from the Central
American species. The name Stegnogramma is derived
from the Greek stegnos, cover + gramme, line. It refers to
the short, line-like sori that cover the veins.
mundial, solo una de las cuales existe en el Neotrópico:
S. pilosa (M. Martens & Galeotti) K. Iwats. Existe desde
el noroeste de México hasta Honduras, de 1800–2700
m, usualmente sobre piedra caliza. En lugares rocosos
en el sureste de los Estados Unidos (Alabama) existe
S. burksiorum (J. E. Watkins & Farrar) Weakley. Difiere
sutilmente de la especie mesoamericana. El nombre
Stegnogramma se deriva del griego stegnos, cubierto +
gramme, línea. Refiere a los soros como cortas líneas que
cubren las venas.

LITERATURE: Iwatsuki, K. 1963.Taxonomic studies of Pteridophyta VII. 9. A revision of the genus Stegnogramma emend. Acta
Phytotaxonomica et Geobotanica 19: 112–126. Watkins, J. E. & D. R. Farrar. 2002. A new name for an old fern from north
[sic] Alabama. American Fern Journal 92: 171–178. Watkins, J. E. & D. R. Farrar. 2005. The origin and taxonomic affinities of
Thelypteris (subgenus Stegnogramma) burksiorum (Thelypteridaceae). Brittonia 57: 183–201.

532
Figure 263. Stegnogramma. A–D. S. pilosa. (From Mickel & Smith, 2004.)

533
POLYPODIACEAE
Stenogrammitis Labiak
DESCRIPTION: Rhizomes erect, scaly, the scales
entire, clathrate, entire but usually with a single apical
cilia; petioles short, glarous to slightly hirsute, the hairs
simple or 1-furcate; laminae 2–5 mm wide, linear,
monomorphic or hemidimorphic, erect or slightly
arcuate, subcoriaceous, pinnatifid to pinnatisect, when
hemidimorphic the fertile portion less divided than the
sterile; rachises obscured by the laminar tissue or with the
dark sclerenchyma exposed, glabrescent to hirsute, the
hairs hyaline or reddish, simple to 1-furcate; segments to
5 × 1 mm, short decurrent basiscopically, apices acute to
obtuse, glabrous or hirsute, the hairs simple or 1-furcate;
veins simple, hidden in the sterile portion of the
laminae but visible and dark in the fertile portions;
hydathodes present, but lacking calcareous secretions;
sori one per segment, oblong to elliptic, glabrous;
sporangia glabrous; spores trilete, green; x=33.
DESCRIPCIÓN: Rizomas erectos, escamosos, las
escamas enteras, clatradas, enteras pero usualmente
con una cilia corta apical; pecíolos cortos, glabros a
levemente hirsutos, los pelos simples o 1-furcados;
laminas 2–5 mm de ancho, lineares, monomórficas
or hemidimórficas, erectas o levemente arcuatas,
subcoriaceas, pinnatífidas a pinatissectas, cuando
hemidimorficas, entonces las partes fértiles usualmente
menos divididas que las estériles; raquises escondidos
por el tejido laminar o visible con su esclerenchima
expuesta, glabrescentes a hirsutos, los pelos hialinos o
rojizos, simples o 1-furcados; segmentos hasta 5 × 1 mm,
cortamente decurrente basiscopicamente, los ápices
agudos a obtusos, glabros o hirsutos, los pelos simples o
1-furcados; venas simples, escondidas en las partes
estériles de las láminas pero visibles y oscuras
en las partes fértiles; hidatodos presentes, faltandos
de depósitos blanquecinos; sori one per segment,
oblongos a elípticos, glabros; sporangios glabros; esporas
triletes, verdes; x=33.

SIMILAR GENERA: Lellingeria differs by wider laminae
(usually > 1 cm), several veins per segment, usually several
sori per segment, and x=37. Leucotrichum differs by ciliate
rhizome scales, whitish hairs of the leaves, and several
veins per segment.
GÉNEROS PARECIDOS: Lellingeria difiere por
laminas más anchas (usualmente > 1 cm), venas
pinnadas, usualmente varios soros por segmento y
x=37. Leucotrichum difiere por escamas de los rizomas
ciliadas, pelos blanquecinos en las hojas y varias venas por
segmento.

COMMENTS: Stenogrammitis contains about 25 species, COMENTARIOS: Stenogrammitis consta de ca. 25

mostly neotropical but with some in Africa, Madagascar, especies, mayormente neotropical pero con algunas en
and islands of the Pacific. Within the Neotropics, the África, Madagascar e islas del Pacífico. Dentro de los
genus is widespread. Most species are epiphytes in Neotrópico, el género es muy difundido. La mayoría son
wet forests at above 1000 m, a few are epipetric. epifitas en bosques húmedos arriba de 1000 m; unas
Previously, Stenogrammitis was classified in Lellingeria as pocas son epipétricas. Previamente Stenogrammitis fue
the “myosuroides group” (Smith et al., 1991). It is sister clasificado en Lellingeria como el “grupo de myosuroides”
to Lellingeria and recognized here for convenience. (Smith et al., 1991). Es hermana a Lellingeria y se reconoce
Stenogrammitis is easily recognized by its narrow (2–5 aquí por conveniencia. Se reconoce fácilmente por sus
mm wide) linear leaves. Also distinctive are the fertile hojas estrechas (2–5 mm), lineares. Distintivo también
veins that are dark because their sclerenchyma sheath son las venas fértiles que son oscuras a causa de su vaina
is exposed, not covered by green laminar tissue as in the de esclerenquima es expuesta, no cubiertas por tejido
sterile portions of the lamina. The genus name is derived laminar verde como en las partes estériles de la lámina. El
from the Greek steno-, narrow, and Grammitis. It refers to nombre del género se deriva del griego steno-, estrecho,
the narrow (< 5mm wide) leaves. y Grammitis. Se refiere a las hojas estrechas (< 5 mm de
ancho).
LITERATURE: Labiak, P. H. & J. Prado. 2005. As espécies de Lellingeria A. R. Sm. & R. C. Moran (Grammitidaceae-Pteridophyta)
do Brasil. Revista Brasileira Botânica 28: 1–22. Labiak, P. H., M. Sundue & G. Rouhan. 2010. Molecular phylogeny, character
evolution, and biogeography of the grammitid fern genus Lellingeria (Polypodiaceae). American Journal of Botany 97: 1354–1364.
Ranker, T. A., A. R. Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Staub & H. Schneider. 2004. Phylogeny
and evolution and grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy. Taxon 53: 415–428. Ranker,
T. A., M. Sundue, P. Labiak, B. Parris & G. Rouhan. 2010. New insights into the phylogeny and historical biogeography of
the Lellingeria myosuroides clade (Polypodiaceae). PLoS Currents 18 Nov 2010; 2: RRN1197.doi: 10.1371/Currents.RRN1197.
Smith, A. R., R. C. Moran, & L. E. Bishop. 1991. Lellingeria, a new genus of Grammitidaceae. American Fern Journal 81: 76–88.

534
Figure 264. Stenogrammitis. A–D. S. paucipinnata (Hemp 1636, K - holotype). A. Habit. B. Fertile portion of the lamina. C.
Sterile portion of the lamina. D. Rhizome scale. E–G. S. limula (Tonduz 12595, US - holotype). E. Habit. F. Fertile portion of
the lamina. G. Rhizome scale. H–M. S. mysouroides (Steyermark 12429, US). H. Habit. I. Sterile portion of the lamina. J. Fertile
portion of the lamina. K. Branched hair from the midrib. L. Fertile portion of the lamina with elliptic sori. M. Rhizome scale.
(Courtesy of Paulo Labiak, 2009.)

535
GLEICHENIACEAE
Sticherus C. Presl
DESCRIPTION: Terrestrial; rhizomes long-creeping,
scaly, protostelic; leaves indeterminate, of periodic growth,
once to several times forked pseudodichotomously, rarely
(in S. simplex (Desv.) Ching) simple, bearing pairs of
opposite pinnae, the distalmost pair with a scaly resting
bud between them; pinnae forked pseudodichotomously,
with a scaly bud between the divisions; penultimate
segments pectinate; veins 1-forked, free; sori round, not
indusiate; sporangia 3–5(-6) per sorus, sessile or nearly
so, opening by a transverse slit that passes through the
apex; spores monolete; x=34.
DESCRIPCIÓN: Terrestres; rizomas largamente
rastreros, escamosos, protostélicos; hojas indeterminadas,
de crecimiento periódico, una a varias veces
pseudodicotómicamente furcadas, raras veces (en S.
simplex (Desv.) Ching) simples, llevando pares opuestas de
pinnas, el par más distal con una yema latente escamosa
entre sí; pinnas furcadas pseudodicotómicamente, con
una yema escamosa entre las divisiones; segmentos
penúltimos pectinados; venas una vez furcadas, libres;
soros redondos, no indusiados; esporangios 3–5(-6) por
soro, séssiles o casi séssiles, se abren por una abertura
transveral que pasa por el ápice; esporas monoletes;
x=34.

SIMILAR GENERA: Dicranopteris and Gleichenella differ
from Sticherus by bristly rhizomes, pubescent pinna buds
(not scaly), veins 2–4 times forked, and sporangia 8–15
per sorus. Gleichenella further differs by solenostelic
rhizomes.
GÉNEROS PARECIDOS: Dicranopteris y Gleichenella
difieren de Sticherus por rizomes cerdosos, yemas de
las pinnas pubescentes (no escamosas), venas 2–4 veces
furcadas, y esporangios 8–15 por soro. Gleichenella difiere
además por rizomas solenostélicos.

COMMENTS: Sticherus is pantropical and contains COMENTARIOS: Sticherus es pantropical y contiene

about 90 species, with about 40 in tropical America.
Like other genera in the family, it often grows in open
disturbed habitats such as roadsides. Many pteridologists
place Sticherus in Gleichenia. Sticherus is here considered
different enough in laminar division, spores shape, and
chromosome number (34 vs. 20 or 22) to merit a
distinct genus. Gleichenia is restricted to southern Africa
and Australasia. The genus name comes from the Greek
stichos, line or row. The sori are born in rows.
casi 90 especies, con alrededor de 40 en América tropical.
Como otros géneros en la familia, crece a menudo en
ambientes abiertos y pertubados tales como taludes.
Muchos pteridólogos ubican Sticherus en Gleichenia. Aquí
se considera Sticherus bastante diferente en división
laminar, forma de esporas, y número cromosómico (34
vs. 20 o 22) para merecer un género distinto. Gleichenia
está restringido al sur de África y Australasia. El nombre
genérico se deriva del griego stichos, línea o hilera. Los
soros se nacen en hileras.

LITERATURE: Ching, R. C. 1940. On the genus Gleichenia Smith. Sunyatsenia 5: 269–288. Gonzales R., J. & M. Kessler.
2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae) with descriptions of nine new species. Phytotaxa
31: 1–54. Holttum, R. E. 1957. Florae Malesianae Precursores XVI. On the taxonomic subdivision of the Gleicheniaceae with
descriptions of new Malaysian species and varieties. Reinwardtia 4: 257–280. Holttum, R. E. 1957. Morphology, growth habit, and
classification in the family Gleicheniaceae. Phytomorphology 7: 168–184. Maxon, W. R. 1909. Gleicheniaceae. North American
Flora 16: 53–63. Nakai, T. 1950. A new classification of Gleicheniales. Bull. Natl. Sci. Mus. 29: 1–71. Østergaard Andersen, E.
& B. Øllgaard. 1996. A note on some morphological terms of the leaf in the Gleicheniaceae. American Fern Journal 86: 52–57.
Østergaard Andersen, E. & B. Øllgaard. 2001. Gleicheniaceae. In: G. Harling & L. Andersson, editors. Flora of Ecuador 66:
105–170.

536
Figure 265. Sticherus bifidus. (From Mickel & Smith, 2004.)

537
THELYPTERIDACEAE
Steiropteris C. Chr.
DESCRIPTION: Terrestrial; petioles with 2 vascular DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
bundles; laminae 1-pinnate-pinnatifid or pinnatifid (with vasculares; láminas 1-pinnado-pinnatífidas o pinnatífidas
a few pinnules free in 2 spp.); proximal pinnae the (con unas cuantas pínnulas libres en 2 spp.); pinnas
longest or nearly so; distal pinnae gradually reduced or proximales las más largas o casi así; pinnas distales
sometimes with an apical pinna somewhat similar in gradualmente reducidas o a veces con una pinna apical
shape to the lateral pinnae; aerophores often strongly algo similar en forma a las pinnas laterales; aeróforos
developed at the pinnae bases; buds absent (except in a menudo fuertemente desarrollados en la base de
T. seemannii); veins free, proximal ones of the adjacent las pinnas; yemas ausentes (excepto en T. seemannii);
segments either joining the margin above the sinus base nervaduras libres, las proximales de los segmentos
(sect. Glaphyropteris), or connivent to the sinus or a adyacentes encontrándose con el margen arriba del seno
cartilaginous keel (false vein) extending from the base (sect. Glaphyropteris) o conniventes en el seno (sect.
of the sinus (sect. Steiropteris); indumentum of needle- Steiropteris) con una quilla cartilaginosa (vena falsa) que
shaped hairs, sometimes septate; sori round; indusia se extiende desde el seno hacia la costa; indumento de
present or absent; sporangia glabrous, rarely setose. x=36. tricomas aciculares, a veces septados; soros redondeados;
indusios presentes o ausentes; esporangios glabros,
raramente setulosos. x=36.

SIMILAR GENERA: Amauropelta, the most common GÉNEROS PARECIDOS: Amauropelta, el subgénero
subgenus in the Neotropics, differs by highly reduced más común en el Neotrópico, difiere por pinnas
proximal pinnae. Cyclosorus can be distinguished by lack proximales altamente reducidas. Cyclosorus se distingue
of aerophores and sinus keels. por la falta de aeróforos y quillas desde del sinus.

COMMENTS: Steiropteris consists of 21 species, all COMENTARIOS: Steiropteris consta de 21 especies,

Neotropical. It is divided into two sections: sect. Steiropteris
(5 spp.) and sect. Glaphyropteris (16 spp). The first has
the sinus keel prominent and raised, veins connivent at
the sinus or uniting below the sinus, and glands lacking
on the leaf tissue. In contrast, sect. Glaphyropteris has the
sinus keel absent, veins meeting margin above the sinus,
and glands usually present on the laminar tissue. The
aerophores at the pinna bases are usually well developed,
conical or peg-like, and whitish or yellowish. They usually
darken upon drying. It is believed that they function in
aerating the fiddlehead, which in several species (e.g., S.
decussata, S. thomsonii) is covered by a thick mucilage
through which the aerophores protrude. The genus
name is derived from the Greek steira, keel + pteris, fern.
It refers to the false vein present in sect. Steiropteris that
runs from the costa to the base of the sinus.
todas Neotropicales. Tiene dos secciones: la secc.
Steiropteris (5 spp.) y secc. Glaphyropteris (16 spp). La
primera tiene la quilla del sinus prominente y elevada,
venas conniventes en el sinus o unidas abajo del sinus, y
glándulas faltantes en el tejido laminar. En contraste, sect.
Glaphyropteris tiene la aquilla del seno ausente, venas
terminando en el márgen arriba de la base del senos y
glándulas usualmente presents en el tejido laminar. Los
aeróforos en las bases de las pinnas usualmente son bien
desarrollados, cónicos or tuberculiformes, y blanquecinos
o amarillentos. Se tornan oscuros al secar. Se cree que
funcionan en impregnar con aire la prefoliación, cual es en
varias especies (e.g., S. decussata, S. thomsonii) es cubierta
con un mucílago desde que los aeróforos proyectan. El
nombre genérico se deriva del griego steira, carina +
pteris, helecho. Refiere a la vena falsa presente en secc.
Steiropteris que se extiende desde la costa hacia la base
del sinus.

LITERATURE: Smith, A. R. 1980. Taxonomy of Thelypteris subgenus Steiropteris, including Glaphyropteris (Pteridophyta).
University of California Publications in Botany 76: 1–38, t. 1-4. Smith, A. R. 1983. Thelypteris. In: G. Harling & B. Sparre, Flora of
Ecuador, no. 18.

538
Figure 266. Steiropteris. A. S. valdepilosa. Note in B the dark aerophore at pinna base. (From Smith in Tryon & Stolze, 1992.)

539
DRYOPTERIDACEAE
Stigmatopteris C. Chr.
DESCRIPTION: Plants terrestrial; rhizome short-
creeping to erect; laminae 1-pinnate to 2-pinnate-pinnatifid,
the apex pinnatifid, the tissue usually membranaceous,
with internal punctate glands, lacking hairs (but often
with reduced, uniseriate scales that may resemble hairs);
pinnae apices long-attenuate, serrate; rachis and costae
lacking hairs abaxially or (in 2 spp.) with minute capitate-
glandular hairs, scaly, the scales thin, ciliate or papillate,
the cilia usually ending in a glandular cell, the adaxial
surface grooved, the grooves pubescent within, the hairs
ca. 0.1 mm long, 1–3-celled; veins ending before the
margin in a conspicuous hydathode, free or (in 3 spp.)
anastomoming; sori round; indusia absent; x=41.
SIMILAR GENERA: Cyclodium differs by the usually
indusiate sori, thicker laminae without internal punctate
glands, and entire scales. An important tendency is that
most species of Cyclodium have only a few scales on the
axes abaxially, in contrast to Stigmatopteris which is usually
scaly.
COMMENTS: Stigmatopteris is entirely neotropical,
occurring from Mexico to southern Brazil. It is primarily
montane, from 500–2000 m, and is absent from most
of Amazonia. The genus contains 24 species. The internal
punctate glands in the lamina when the leaf tissue is
backlit and examined with a handlens. In herbarium
specimens in oblique light, the glands often appear (under
a microscope) as minute raised bumps. The glands often
dry blackish after pressing.The genus name is derived from
the Greek stigmatos, punctate + pteris, fern. It refers to
the characteristic internal punctate glands of the laminae.
LITERATURE: Moran, R. C. 1991. Monograph of the Neotropical fern genus Stigmatopteris (Dryopteridaceae). Annals of
the Missouri Botanical Garden 78: 857–914. Moran, R. C. & P. H. Labiak. 2016. Phylogeny and character evolution of the
Neotropical fern genus Stigmatopteris (Dryopteridaceae). Brittonia 68: 476–488.
540
DESCRIPCIÓN: Plantas terrestres; rizoma corto-
reptante a recto; láminas 1-pinnadas a 2-pinnada-
pinnatífidas, el ápice pinnatífido, el tejido usualmente
membranáceo, con glándulas punctadas internas, carece
de pelos (pero a menudo con escamas uniseriadas que
simulan pelos); ápices de las pinnas largo-atenuadas,
aserradas; raquis y costas carece de pelos o (en 2 spp.)
con glandulas diminutas capitadas, escamosos, las escamas
delgadas, ciliadas o papiladas, las cilias terminando
en una célula glandular, la superficie adaxial surcada,
los surcos pubescentes adentro, los pelos ca. 0.1 mm
de largo, 1–3-celulares; venas terminando antes del
margen en un hidatodo conspicuo, libres o (en 3 spp.)
anastomosantes; soros redondos; indusios ausentes;
x=41.
GÉNEROS PARECIDOS: Cyclodium difiere por los
soros usualmente con indusios, las láminas más gruesas
y sin glándulas punctadas internas, y escamas enteras.
Una tendencia importante es que la mayoría de las
especies de Cyclodium tienen pocas escamas sobre sus
ejes abaxialmente, en contraste a Stigmatopteris que es
usualmente escamoso.
COMENTARIOS: Stigmatopteris es completamente
neotropical, extendiéndose desde México hasta el sur de
Brasil. Es principalmente de las montañas, de 500–2000
m, y es ausente de la mayoría de Amazonía. El género
consta de 24 especies. Pueden verse mejor las glándulas
punteadas internas en contraluz con una lupa. Con la
luz oblicua en ejemplares montados del herbario, las
glándulas parecen (con un microscopio) como puntitos
elevados. A menudo se vuelven negruzcas al secarse
después de ser prensadas. El nombre genérico se deriva
del griego stigmatos, punctata + pteris, helecho. Se refiere
a las glándulas características internas punteadas de las
láminas.
Figure 267. Stigmatopteris. A, B. S. heterophlebia (Baker) R. C. Moran. C. S. nephrodioides, adaxial view, note hathodes. D.
Punctate glands in lamina tissue. E. Petiole cross-section showing eupolypod I arrangement of vascular bundles. F. Costal scale,
note cilia with enlarged, darkened apical cell. G. S. longicaudata. (Original, ©Robbin Moran, 2009.)

541
BLECHNACEAE
Struthiopteris Scop..

DESCRIPTION: Plants terrestrial; rhizomes short-
creeping or elongate, non-stoloniferoous; sterile
and fertile leaves dimorphic; laminae pinnate to
pinnatifid, the bases gradually reduced to auricles,
the apices pinnatifid; rachises glabrous or sparesly scaly;
buds absent; aerophores absent; pinnae adnate, entire;
veins free, ending in submarginal hydathodes; sori
linear on both sides of the costae; indusia entire; x=31, 34.
DESCRIPCIÓN: Plantas terrestres; rizomas cortamente
rastreros a arlargados, no estoloníferos; hojas estériles
y fértiles dimorfas; láminas pinnadas a pinnatífidas,
las bases gradualmente reducidas a aurículos, los
ápices pinnatífidos; ráquises glabros o esparciadmente
escamosos; brotes foliares ausentes; aeróforos ausentes;
pinnas adnatas, enteras; venas libres, terminándose
en hidatodos conspícuosos submarginales; soros
lineares on cada lado de las costas; indusios enteros;
x=31, 34.

SIMILAR GENERA: Austroblechnum differs by crenate GÉNEROS PARECIDOS: Austroblechnum difiere por
to serrate pinnae and is generally tropical distribution. las pinnas crenadas a serradas y distribucion geografica
principalmente tropical. .

COMMENTS: Struthiopteiris is composed of 5 species, COMENTARIOS: Struthiopteiris consta de 5 especies

mostly of China and Japan, with one species, S. spicant (L.)
F. W. Weiss, being circumboreal. This species is the only
one in the New World. It occurs on the Pacific side of
North America from Alaska to California, 0–1400 m. It
grows primarily in wet confiferous woods and swamps.
The most closely related genera are Blechnidium T. Moore
and Brainea J. Sm, both Asian. The genus name is derived
from the Greek struthios, ostrich + pteris, fern. The leaves
resemble an otrich feather because they are reduced
toward the base and broadest above the middle. .
principalmente de China y Japón, con una especie, S.
spicant (L.) F. W. Weiss, siendo circunboreal. Esta especie
es la única en el Nuevo Mundo. Existie en el lado Pacífico
de América del Norte desde Alaska a California, 0–1400
m. Habita principalmetne bosques coníferos húmedos y
suampos. Los géneros más relacionades son Blechnidium
T. Moore y Brainea J. Sm, ambos Asiáticos El nombre
genérico se deriva del griego struthios, avestruz + pteris,
helecho. Las hojas se asemejan una pluma de avestruz
porque son reducidas hacia la base y más anchas arriba
de la mitad.

LITERATURE: Broadhurst, J. 1912. The genus Struthiopteris and its representatives in North America. Bulletin of the Torrey
Botanical Club 39: 257–278, 357–385. Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for
Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations Phytotaxa 275: 191–227. Rolleri,
C. H. & C. Prada. 2006. Revisión de los grupos de especies del género Blechnum (Blechnaceae-Pteridophyta): el grupo B.
penna-marina. Acta Botánica Malacitana 31: 7–50.

542
Figure 268. Struthiopteris spicant. (Original, ©Robbin Moran, 2016.)

543
POLYPODIACEAE
Synammia C. Presl
DESCRIPTION: Terrestrial, saxicolous, or epiphytic;
rhizomes pruinose, long-creeping; rhizome scales
peltate, 2-3 mm wide, thin, ovate, orangish but
blackened toward point of attachment; sterile and
fertile leaves monomorphous; laminae 1-pinnate to
2-pinnate-pinnatifid; pinnae adnate to the rachis, margins
cartilaginous-thickened and minutely notched; veins
inconspicuous, more than once forked, anastomosing to
form a single row of costal areolae; areoles with a single
included veinlet; sori round to elongate, borne on the tips
of the included veinlets; indusia absent; paraphyses dense,
filiform; spores yellow, monolete.
DESCRIPCIÓN: Terrestres, saxícoloas o epífitas;
rizomas pruinosas, largamente rastreros; escamas
del rizoma peltadas, 2-3 mm de ancho, delgadas,
ovadas; hojas estériles y fértiles monomorfas; láminas
1-pinnado a 2-pinnado-pinnatífidas, pubescentes o con
escamas densas o esparicas; pinnas adnatas al raquis,
margines cartilaginosos, engrosados, con pequeñas
hendiduras; venas inconspicuas, más que una vez surcada,
anastomosándose, formando una hilera de areolas
costales; areolas con una sola venilla incluida; soros
alargados, se nacen en los puntos de las venillas; inducíos
ausentes; parafisos densos, filiformes; esporas amarillas,
monoletes.

SIMILAR GENERA: Pleopeltis differs by peltate scales on
the blades and (usually) among the sporangia. Polypodium
differs by non-pruinose rhizomes, (often) free veins and
sporangiasters (capitate hairs) among the sporangia.
GÉNEROS PARECIDOS: Pleopeltis difiere por
escamas peltadas en las láminas y (usualmente) entre los
esporangios. Polypodium difiere por rizomas no pruinosos,
(a menudo) venas libres y esporangiastros (pelitos
capitados) entre los esporangios.

COMMENTS: Synammia consists of four species in
temperate South America, in Chile, Argentina, and the
Juan Fernandez Islands (Rodríguez, 1995). They grow
principally in wet forests. Schneider et al. (2006) found
that Synammia is strongly supported as monophyletic. It
is either sister to the rest of the Polypodiaceae or to all
other neotropical members of the family. Synammia is
derived from the Greek syn, together + ammatos, knot
or loop, referring to the veins that form a row of costal
areoles.
COMENTARIOS: Synammia consta de four especies
en las regiones templadas de América del Sur, en Chile,
Argentina, and las Islas Juan Fernández. Principalmente
existen en bosques húmedos. Schneider et al. (2006)
encontraron que Synammia es fuertemente apoyado
como monofilético. Es hermana al resto de Polypodiaceae
a todos las otras especies de la familia. =Synammia is
derivada de la Griega syn, conjunto + ammatos, knot or
loop, referiéndose a las venas que forman una hilera de
areolas costales.

LITERATURE: Looser, G. 1952. El género Polypodium L. y sus representantes chilenos. Revista Univ. (Santiago) 36: 13–82.
Rodríguez, R. 1995. Polypodium. Pages 230–236. In: C. Marticorena & R. Rodríguez, editors. Flora de Chile, vol. 1, Pteridophyta
– Gymnospermae. Univ. de Concepción, Chile. Schneider, H., H.-P. Kreier, R. Wilson & A. R. Smith. 2006. The Synammia
enigma: evidence for a temperate lineage of polygrammoid ferns (Polypodiaceae; Polypodiiae) in southern South America.
Systematic Botany 31: 31–41. Sota, E. R. 1961. Polypodiaceae y Grammididaceae argentines. Opera Lilloana 5: 1–229. Sota, E.
R. 1968. Acerca del género “Synammia” Presl. Revista Museo de La Plata, Sección Botánica 11: 129–132.

544
Figure 269. Synammia. A–D. S. sp. E–F. S. intermedia ssp. masafuerana. H–J. S. feuillei. (Original, ©Robbin Moran, 2016.)

545
TECTARIACEAE
Tectaria Cav.
DESCRIPTION: Terrestrial or saxicolous; stems
erect or long-creeping, scaly; sterile and fertile leaves
monomorphous, rarely dimorphous; laminae simple
and entire to 3-pinnate-pinnatifid; basal pinnae often
with a prolonged lobe or pinnule on the basal
basiscopic side; rachis and costae rounded or shallowly
grooved on the upper surface, the grooves not decurrent,
puberulent on the upper surface, the hairs 0.1-0.2 mm
long, reddish, pluricellular; veins reticulate, the areoles
typically with included veinlets, or free in T. brauniana
(H. Karst.) C. Chr.; sori round or (in T. athyrioides (Baker)
Mett.) irregular or J-shaped, rarely linear; indusia present,
rarely absent; spores bilateral, brown; x=40.
DESCRIPCIÓN: Terrestres o saxícolas; tallos erectos a
largamente rastreros, escamosos; hojas estériles y fértiles
monomorfas, raras veces dimorfas; láminas simples y
enteras a 3-pinnado-pinnatífidas; pinnas basales a
menudo con un prolongado lóbulo o pínnula en el
lado basal basiscópico; raquis y costas redondeados
o someramente sulcados en el haz, los surcos no
decurrentes, puberulentes en el haz, los tricomas 0.1-
0.2 mm de largo, rojizos, pluricelulares; venas reticuladas,
las aréolas típicamente con venillas incluidas, o
libres en T. brauniana (H. Karst.) C. Chr.; soros redondos
o (en T. athyrioides (Baker) Mett.) irregulares o en forma
de “J”, raras veces lineales; indusios presentes, raras veces
ausentes; esporas bilaterales, pardas; x=40.

SIMILAR GENERA: Bolbitis differs by acrostichoid sori,
axes of the lamina glabrous on the upper surface, and an
elongated ventral vascular bundle in the rhizome (seen
in cross-section). Dracoglossum differs by simple entire
leaves with a bud at the apex of the lamina.
GÉNEROS PARECIDOS: Bolbitis difiere por soros
acrosticoides, ejes de la lámina glabros en el haz, y un haz
vascular alargado ventral en el rizoma (visto en sección
transversal). Dracoglossum difiere por hojas simples
enteras con un brote en ápice de la lámina.

COMMENTS: Tectaria is pantropical and subtropical COMENTARIOS: Tectaria es pantropical y subtropical

with about 200 species. Approximately 30 species occur
in the New World and needs a monograph because
many taxonomic and nomenclatural problems require
solution. Most of the species occur in wet forests from
0–1500 m, but some are characteristic of drier habitats
and occur primarily on limestone. The genus is sister
to Hypoderris and Triplophyllum. Tectaria panamensis
(Hook.) R. M. Tryon & A. F. Tryon is sometimes put in its
own genus, Dictyoxiphium Hook., because of its simple
leaves and submarginal sori. This species hybridizes with
Tectaria incisa (see section on hybrid in this manual).
Tectaria draconopteris (D. C. Eaton) Copel. is unusual by
its cyanogenic rhizomes (pers. obs.). The genus name is
derived from the Latin tectum, roof + aria, a substantive
suffix. It alludes to the roof-like indium in many species.
y consta de casi 200 especies. Aproximadamente 30
especies existen en el Nuevo Mundo y necesitan una
monografía, ya que muchos problemas taxonómicos y
de nomenclatura requieren solución. La mayoría de las
especies existen en bosque húmedos de 0–1500 m,
pero algunas caracterizan ambientes más secas y existan
principalmente sobre piedra caliza. El género es hermano
a Hypoderris y Triplophyllum. Tectaria panamensis (Hook.)
R. M. Tryon & A. F. Tryon es algunas veces puesto en su
propio género, Dictyoxiphium Hook., a causa de sus hojas
simples y soros submarginales. Esta especie se hibridiza
con Tectaria incisa (vea la sección sobre híbridos en
este manual). Tectaria draconopteris (D. C. Eaton) Copel.
es inusual por sus rizomas cianogénicos (pers. obs.). El
nombre genérico se deriva del latín tectum, techo + aria,
un sufijo sustantivo. Alude al indusio que es como un
techo en muchas especies.

LITERATURE: Grayum, M. H. 1987. On three misunderstood neotropical species of Tectaria (Polypodiaceae: Asplenioideae).
Phytologia 64: 30–35. Morton, C. V. 1966.The Mexican species of Tectaria. American Fern Journal 56: 120–137. Wagner, W. H.
Jr., F. S. Wagner & L. D. Gómez. 1978. The singular origin of a Central American fern, Pleuroderris michleriana. Biotropica 10:
254–264. Rojas-Alvarado, A. F. & Sanín, D. 2014. Dos especies nuevas de Tectaria (Tectariaceae) para Costa Rica, Panamá y
Colombia. Anales Jard. Bot. Madrid 71(1): e004. [contains a key to all species of Tectaria in the three countries.]

546
Figure 270. Tectaria. A: T. panamensis. B–D. T. heracleifolia. E–G. T. mexicana. H–L. T. incisa. (From Mickel & Smith, 2004.)

547
BLECHNACEAE
Telmatoblechnum Perrie, D. J. Ohlsen & Brownsey
DESCRIPTION: Terrestrial; rhizomes creeping,
apices ascending-erect so that the leaves appear
clustered, the scales 0.5–3 mm long, linear-lanceolate,
black to brown, concolorous or slightly bicolorous, entire
or nearly so; sterile and fertile leaves monomorphic or
slightly dimorphic; petioles 10–40 cm long, not papillose,
glabrous or scaly, stramineous to tan; laminae 35–70 ×
10–30 cm, 1-pinnate, chartaceous to coriaceous, glabrous
abaxially, truncate at the base, without vestigial pinnae,
apices reduced gradually to a terminal pinna similar in
shape to the lateral ones; buds absent; rachises glabrous,
tan; pinnae 4–15 × 0.5–1.5 cm, 20–40 pairs, nearly
sessile, never adnate, not involute, articulate to
the rachis, eventually deciduous (leaving persistent
rachises), the margins serrulate, lighter-colored
and slightly thickened; costae tan, sparsely scaly, the
scales widely ovate; aerophores absent at the pinna bases;
veins free, the apices not or only slightly thickened;
sori linear, continuous, indusiate, typically restricted to the
apical pinnae; x=36.
SIMILAR GENERA: Nephrolepis differs by the presence
of filiform superficial stolons, pinnae with a basal basiscopic
auricle, vein apices enlarged (hydathodous) and often
whitish, and round sori.
COMMENTS: Telmatoblechnum consists of two species:
T. serrulatum (Rich.) Perrie, D. J. Ohlsen & Brownsey
(New World) y T. indicum (Burm. f.) Perrie, D. J. Ohlsen
& Brownsey (Australasia, Pacifica, and Malesia). The two
are very similar and possibly represent the same species.
In the New World, T. serrulatum is widespread, occurring
from southern Florida and the Antilles to Paraguay and
Argentina. It grows from 0–900 m en wet soils, swamps,
ditches, shrubby areas, open river borders, and grassy
fields. It is the only Blechnaceae in the New World that
has pinnae articulate to the rachis. The articulations
appear as thickened circles around the petiolules of
the pinnae with a dark line as the abscission layer. En
regions with a pronounced dry season, the pinnae fall,
leaves erect, naked rachises. The generic name is derived
from the Greek telmato-, wet soils, ponds or marshes,
and Blechnum, the genus in which the two species were
previously classified.
LITERATURE: Perrie, L. R., R. K. Wilson, L. D. Shephard, D. J. Ohlsen, E. L. Batty, P. J. Brownsey & M. J. Bayly. 2014.
Molecular phylogenetics and generic taxonomy of Blechnaceae ferns. Taxon 63: 745–758. Rolleri, C. H., C. Prada, J. M. Ga-
briel y Galán, L. M. Passarelli & M. M. Ciciarelli. 2010. Morphology of the sporophyte and gametophyte of the swamp fern,
Blechnum serrulatum (Blechnaceae, Pteridophyta). Australian Journal of Botany 58: 508–518.
548
DESCRIPCIÓN: Terrestres; rizomas rastreros, ápices
ascendientes-erectos de manera que las hojas
aparecen agrupadas, las escamas 0.5–3 mm, linear-
lanceoladas, negras a pardas, concoloras o ligeramente
bicoloras, enteras o casi enteras; hojas estériles y fértiles
monomorfas o levemente dimórfas; pecíolos 10–40
cm, no papilosos, glabros o escamosos, pajizos a pardo-
amarillentos; láminas 35–70 × 10–30 cm, 1-pinnadas,
cartáceas a coriáceas, glabras en el envés, truncadas en la
base, sin pinnas vestigiales, el ápice reducido gradualmente
a una pinna terminal algo similar en forma a las pinnas
laterales; yemas ausentes; ráquises glabros, pardo-
amarillentos; pinnas 4–15 × 0.5–1.5 cm, 20–40 pares,
casi sésiles, nunca adnatas, no involutas, articuladas
al raquis y eventualmente deciduas (dejando
ráquises persistentes), los márgenes serruladas,
más claro en color y un poco engrosados; costas
pardo-amarillas, esparcidamente escamosas, las escamas
anchamente ovadas; aeróforos ausentes en las bases de
las pinnas; venas libres, los ápices no o solamente un
poco engrosados; soros lineares, continuos, indusiados,
típicamente restringidos a los pinnas apicales; x=36.
GÉNEROS PARECIDOS: Nephrolepis difiere por la
presencia estolónes filiformes superficiales, las pinnas con
una aurícula acroscópica basal, los ápices de las venas
engrosados (hidatodos) y a menudo blanquecinos y
soros redondeados.
COMENTARIOS: Telmatoblechnum consta de dos
especies: T. serrulatum (Rich.) Perrie, D. J. Ohlsen &
Brownsey (Nuevo Mundo) y T. indicum (Burm. f.) Perrie,
D. J. Ohlsen & Brownsey (Australasia, Pacífica y Malesia).
Los dos son muy parecidos y posiblemente representan
la misma especie. En el Nuevo Mundo, T. serrulatum está
muy difundida, ocurriéndose desde el sur de Florida y las
Antillas hasta Paraguay y Argentina. Existe desde 0–900 m
en suelos húmedos, pantanos, zanjas, sabanas, matorrales,
bordos abiertos de ríos y planicies graminosas. Es el
único Blechnaceae en el Nuevo Mundo que tiene pinnas
articuladas al raquis. Las articulaciones aparecen como
círculos engrosados alrededor de los peciólulos de las
pinnas con una línea oscura en la capa de abscisión. En
regiones con una estación seca pronunciada, las pinnas
caen, dejando los ráquis erectos, desnudos. El nombre
genérico se deriva del griego telmato-, tierra mojada,
charcos o pantanos, y Blechnum, el género en que estas
dos especies fueron previamente clasificadas.
Figure 271. Telmatoblechnum serrulatum. A. Habit, note the rhizome turned upward to produce a cluster of leaves. B.
Transverse section of rhizome showing meristeles (larger circles) and leaf traces (smaller circles to right and left). C. Transverse
section of petiole showing vasculature. D. Rachis-pinnae junctures showing articulation at base of pinnae. E. Pinna showing
lighter-colored thickened margin (Original, ©Robbin C. Moran, 2016.)

549
POLYPODIACEAE
Terpsichore A. R. Sm.
DESCRIPTION: Epiphytic; rhizomes short-creeping,
compact; rhizome scales setulose on margins and
often on the surfaces, not clathrate, orange to brown,
basifixed (not cordate); petioles ca. half to equaling
the length of the laminae, setose (especially basally),
the setae 0.5–3 mm long, generally castaneous, numerous,
spreading, the hairs 0.1–0.2 mm long; laminae determinate,
pinnatisect to 1-pinnate, rarely 1-pinnate-pinnatifid,
without black clavate fungi, setose, the setae simple (not
branched); hydathodes present, without whitish chalk
deposits; veins simple, free; sori separate, round, superficial
(i.e., not sunken in the lamina); sporangial capsules
setose (except T. atroviridis); spores green, trilete. x=37.
SIMILAR GENERA: Alansmia differs by indeterminate
leaves, short petioles, laminar setae often paired, branched
or stellate, and blades tapered toward the base. The
Terpsichore taxifolia group typically has black costae, black
club-shaped fungi on the lower surface of the laminae,
white hydathodes, and glabrous sporangial capsules.
Pecluma differs by monolete, yellowish spores and lacking
setae on the petioles and blades.
COMMENTS: Terpsichore is entirely neotropical and
contains 12 species, all of which are epiphytes in wet
shaded forests from low to high elevations. Many of the
species have arching or horizontal petioles from which
the laminae are abruptly pendent. Although occurring on
various substrates, Terpsichore species are often common
on the root mantles of tree fern trunks. The genus is
recognized here in the strict sense, with the removal of
Alansmia and the T. taxifolia group.These groups, originally
included in Terpsichore (Smith, 1993), have been shown
to be distinct lineages more closely related to other
grammitid genera (Ranker et al., 2004). Terpsichore is
named for the Greek Muse of dance and choral singing.
The leaves appear to “dance” when the wind blows.
LITERATURE: Copeland, E. B. 1956. Ctenopteris in America. Philippine Journal of Science 84: 381–473, t. 1–16. Ranker, T.
A., A. R. Smith, B. B. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Staub & H. Schneider. 2004. Phylogeny and evolution
and grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy. Taxon 53: 415–428. Smith, A. R. 1993.
Terpsichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3: 478–489.
550
DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
compactos; escamas del rizoma setulosas por los
márgenes y a menudo las superficies, no clatradas,
anaranjadas a pardas, basifijas (no cordadas); pecíolos
ca. una mitad hasta igual la longitud de las láminas,
setosos (en especial basalmente), las setas 0.5–3 mm de
largo, generalmente castañas, numerosas, patentes, los
tricomas 0.1–0.2 mm de largo; láminas pinnatisectas a
1-pinnadas, raramente 1-pinnado-pinnatífidas, sin hongos
clavados negros, setosas, las setas simples (no ramificadas);
hidatodos presentes, sin puntos blanquecinos calizos;
nervaduras simples, libres; soros separados, redondos,
superficiales (i.e., no hundidos en la lámina); cápusulas
esporangiales setosos (excepto T. atroviridis) esporas
verdes, triletes. x=37.
GÉNEROS PARECIDOS: Alansmia difiere por hojas
indeterminadas, pecíolos cortos, setas de las láminas a
menudo pareadas, ramificadas o esteladas y láminas
reducidas hacia la base. El grupo de Terpsichore taxifolia
tiene costas negras y hongos negros claviformes en el
envés de la lámina, hidatodos blanquecinos y cápsulas
esporangiales glabras. Pecluma difiere por monoletes
amarillentosas y la falta de setas en los pecíolos y láminas.
COMENTARIOS: Terpsichore es completamente
neotropical y consta de 12 especies, todas epífitas en
bosques húmedos sombreados de elevaciones bajas
a altas. Muchas especies tienen pecíolos arcuados o
horizontales desde que las láminas son abruptamente
pendientes. Aun que existiendo en sustratos varios,
las especies de Terpsichore son a menudo frecuentes
sobre los manteles de raíces en los troncos helechos
arborescentes. Se reconoce el género aquí en el sentido
stricto, quitando Alansmia y el grupo de T. taxifolia. Estos
grupos, originalmente incluidos en Terpsichore (Smith,
1993), han sido mostrado ser líneas evolutivas distinctas
más relacionadas a otros géneros grammitoides (Ranker
et al., 2004). Terpsichore se nombra para la Musa griega
de la danza y cantante de coro. Las hojas parecen danzar
cuando los vientos soplan.
Figure 272. Terpsichore. A–G. T. chrysleri. H–L. T. lehmanniana. O, P. T. asplenifolia. (A–L ©R. C. Moran, 2010; O, P from
Mickel & Smith, 2004.)

551
THELYPTERIDACEAE
Thelypteris Schmidel.
DESCRIPTION: Terrestrial; rhizomes long-creeping;
rhizome scales glabrous on margins and surfaces; petiole
bases with 2 vascular bundes; laminae 1-pinnate-pinnatifid,
eglandular; basal pinnae not reduced; rachises and costae
grooved adaxially; costae abaxially sparsely scaly; veins
free, the sterile ones usually forked, all reaching the
margins above the sinus; sori round, indusiate, often
partly covered by the revolute lamina margin; sporangia
with 1 or 2 short capitate hairs; x=35.
DESCRIPCIÓN: Terrestres; rizomas largamente
rastreros, las escamas glabras por los márgines y superficies;
bases de los pecíolos con 2 haces vasculares; láminas
1-pinnado-pinnatífidas, eglandulares; pinnas basales no
reducidas; ráquises y costas surcados adaxialmente; costas
abaxialmente escamosas esparcidamente, adaxialmente
surcadas; venas libres, las estériles usualmente furcadas,
alcanzando los margines arriba del sinus; soros redondos,
indusiados, a menudo cubierto parcialmente por el
margen revoluto de la lámina; esporángios con 1 o 2
pelitos capitados; x=35.

SIMILAR GENERA: Amauropelta usually differs by
rhizomes erect and basal veins terminating at the margins
at (not above) the sinus. Cyclosorus differs by basal veins
of adjacent segments astomosing below the sinus and by
reddish spherical glands on the sporangial stalks and often
along the veins abaxially.
GÉNEROS PARECIDOS: Amauropelta difiere por
rizomas erectos y venas basales terminándos en los
márgenes en el sinus (no arriba). Cyclosorus differs by venas
basales de los segmentos adyacentes anastomosándose
abajo del sinus y por glándulas esféricas rojizas en los
pecíolos de los esporángios y a menudo por las venas
abaxialmente.

COMMENTS: Thelypteris contains two species: T.
palustris Schott and T. confluens (Thunb.) C. V. Morton.
Both are plants of wet soils, typically in marshes, swamps,
bogs, and wet river or lake margins. Thelypteris palustris,
the type species of the genus, is construed as having
two varieties: var. palustris, of Eurasia, and var. pubescens
(Lawson) Fernald, of North America., Japan, and northeast
Asia. Thelypteris confluens occurs in South America,
Madagascar, southeastern Asia, and New Zealand. The
genus name is derived from the Greek thelys, female +
pteris, fern. An old Greek name for a fern more delicate
than the male fern, Dryopteris filix-mas. .
COMENTARIOS: Thelypteris consta de dos especies: T.
palustris Schott and T. confluens (Thunb.) C. V. Morton.
Los dos son plantas de suelos mojados, típicamente en
humedales, pantanos, turberas y los márgenes de ríos
y lagos. Thelypteris palustris, la especie tipo del genero,
es interpretado tener dos variedades: la var. palustris of
Eurasia, y la var. pubescens (Lawson) Fernald, de América
del Norte, Japón y noreste Ásia. Thelypteris confluens
existe en América del Sur, Madagascar, sureste de Ásia
y Nueva Zelanda. El nombre genérico se deriva del
griego thelys, hembra + pteris, helecho. Un nombre de
los ancianos griegos para un helecho mas delicado que el
helecho macho, Dryopteris filix-mas

LITERATURE: Fernald, M. L. 1929. A study of Thelypteris palustris. Rhodora 31: 27–40. Holttum, R. E. 1970. Studies in the
family Thelypteridaceae I. A comparative study of the type-species of Thelypteris Schmidel, Cyclosorus Link, and Ampelopteris Kunze.
Blumea 18: 195–215. Holub, J. 1972. On the taxonomy and nomenclature of the European marsh fern. Taxon 21: 331–332.
Tryon, A. F. 1971. Structure and variation in spores of Thelypteris palustris. Rhodora 73: 444–460. Small, J. K. 1935. Ferns of the
Vicinity of New York. The Science Press, Lancaster, PA. Tryon, A. F., R. M. Tryon & F. Badré 1980. Classification, spores, and
nomenclature of the marsh fern. Rhodora 82: 461–473.

552
Figure 273. Thelypteris palustris (from Small, 1935, with the addition of petiolar cross-sections at left)

553
THYRSOPTERIDACEAE
Thyrsopteris Kunze
DESCRIPTION: Terrestrial; stems erect to decumbent,
to ca. 2 m tall, massive, sometimes producing stolons;
leaves 2-3.5 m long, partially dimorphic, the fertile ones
with strongly modified lower pinnae; petioles with long,
matted hairs; laminae 3–5-pinnate, broadly ovate,
tapered to the tip; pinnae and pinnules stalked, ascending;
rachises and costae grooved adaxially; veins free;
sori terminal on narrow, non-laminate segments, lacking
paraphyses, often restricted to the proximal portions of
the pinnae; receptacle columnar, clavate; indusia cup-
shaped, firm, entire; spores tetrahedral-globose, trilete.
x= ca. 76–78.
SIMILAR GENERA: Culcita has monomorphic fertile
pinnules and clam-shaped indusia. Dicksonia also differs by
clam-shaped indusia and by leaves long-tapered toward
the base.
COMMENTS: The Thyrsopteridaceae consists of a
single species, Thyrsopteris elegans Kunze, which is of
uncertain relationship to other tree ferns. It is endemic
to the Juan Fernández Islands where it grows from 500-
1000 m in woodlands and heaths, often with Dicksonia
berteriana. It is unusual by the sori restricted to only the
proximal pinnae. Many species related to Thyrsopteris
were widespread world-wide during the Mesozoic, but
nowadays only Thyrsopteris survives. Cretaceous fossils of
Thyrsopteris are known from mainland Chile. The genus
name is derived from the Greek thyrsos, bunch + pteris,
fern. The sori are arranged in grape-like clusters.
LITERATURE: Korall, P., K. M. Pryer, J. S. Metzgar, H. Schneider & D. S. Conant. 2006. Tree ferns: monophyletic groups
and their relationships as revealed by four protein-coding plastid loci. Molecular Phylogenetics and Evolution 39: 830–845.
554
DESCRIPCIÓN:Terrestres; tallos erectos a decumbentes,
hasta 2 m de altura, massivos, a veces produciendo
estolones; hojas 2-2.5 m long, parcialmente dimorfas, las
fértiles con pinnas inferiors dimorfas; pecíolos con pelos
largos y enredados; láminas 3–5-pinnadas, anchamente
ovadas, reducidas gradualmente hacia el ápice; pinnas y
pínnulas pecioluladas, ascendentes; ráquises y costas
surcados adaxialmente, los surcos decurrentes; venas
libres; soros terminales en segmentos estrechos no
laminados, carente de paráfises, a menudo restingidos a
las partes proximales de las pinnas; receptáculo columnar,
clavado; indusios en forma de una copa, firmes, enteros;
esporas tetraédrico-globosas, triletes. x= ca. 76–78.
GÉNEROS PARECIDOS: Culcita tiene pínulas fértiles
dimorfas y indusios en forma de una bivalva. También
Dicksonia difiere por indusios en forma de una bivalva y
por hojas largamente reducidas hacia la base.
COMENTARIOS: La Thyrsopteridaceae consta de una
única especie, Thyrsopteris elegans Kunze, la cual es de
una relación incierta de otros helechos arborescentes.
Es endémica a las Islas Juan Fernández donde existe de
500-1000 m en bosques y lugares abiertos abustivos,
a menudo con Dicksonia berteriana. Es inusual por
los soros retrigidos únicamente a las pinnas inferiores.
Muchas especies relaciondas a Thyrsopteris estaban muy
difundidas mundialmente durante el Mesozóico, pero hoy
día solamente Thyrsopteris sobrevive. Fósiles Cretásicos
de Thyrsopteris se conocen de Chile continental. El
nombre genérico se deriva del griego thyrsos, macolla
+ pteris, helecho. Los soros están agrupados en grupos
como uvas.
Figure 274. Thyrsopteris elegans Kunze. (Original, ©Robbin Moran, 2016.)

555
PTERIDACEAE
Trachypteris H. Christ
DESCRIPTION: Terrestrial or rupestral; rhizomes erect
or decumbent, scaly; leaves monomorphic or dimorphic,
10–25 cm long; blades densely scaly abaxially, the
scales obscuring the surface, glabrous to sparsely scaly
adaxially; veins hard to see because of the thick blades,
anastomosing without included veinlets; sori borne in
a narrow to broad band near the margin; indusia
and paraphyses absent; spores globose, trilete, strongly
cristate, without an equatorial flange; x=29 or 30.
SIMILAR GENERA: Doryopteris differs by laminae
glabrous beneath. Some species of Cheilanthes can be
densely scaly beneath, but these species usually have
finely divided leaves.
COMMENTS: Trachypteris consists of 3 species and
occurs from Ecuador (including Galapagos Islands) to
Argentina and southern Brazil, and Madagascar. It usually
grows on rocks or among rocks and in dry soils, usually
in seasonally dry habitats. The leaves have the ability to
lose most of their normal moisture content and enroll,
then after a rain rapidly absorb water and expand. The
American species grow from 700-2900 m. The three
neotropical species are: T. gilliana (Baker) Svenson, T. induta
(Maxon) R. M. Tryon & A. F. Tryon and T. pinnata (Hook. f.)
C. Chr. Trachypteris induta having a basal rosette of sterile
leaves and erect, long-petioled fertile leaves, whereas T.
pinnata lacks the rosette and has monomorphic leaves.
Trachypteris gilliana resembles T. pinnata but differs by
reticulate or verrucose spores and trifid fertile leaves.
Another species, T. drakeana C. Chr., occurs in Madagascar.
It has a basal rosette of sterile leaves, as in T. pinnata, but
each leaf bears apical bud. According to rbcL studies,
Trachypteris is the sister genus to Doryopteris (Gastony &
Rollo, 1998). The genus name is derived from the Greek,
trachys, rough + pteris, fern.The scales covering the fronds
impart a rough texture.
LITERATURE: Ballard, F. 1962. Saffordia induta. Hooker’s Icon. Plant. V, 6(4). Maxon, W. R. 1913. Saffordia, a new genus of
ferns from Peru. Smithsonian Miscellaneous Collections 61(4): 1–5. Ramos Giacosa, J. P., G. E. Giudice & M. A. Morbelli.
2008. Resurrection of the fern name Trachypteris gilliana (Baker) Svenson Pteridaceae. American Fern Journal 98: 164–169.
556
DESCRIPCIÓN: Terrestres o rupestres; rizomas erectos
o decumbentes, erectos, escamosos; hojas monomorfas o
dimorfas, 10–25 cm; láminas densamente escamosas
en el envés, las escamas oscuriendo la superficie,
adaxialmente glabras a esparcidamente escamosas;
nervadura difícil ver a causa de las láminas gruesas,
anastomosada sin venillas incluidas; soros nacen en una
faja estrecha cerca del margen; indusios y parafises
ausentes; esporas globosas, triletes, fuertemente cristadas,
sin un engrosamiento ecuatorial; x=29 o 30.
GÉNEROS PARECIDOS: Doryopteris difiere por
las láminas glabras en el envés. Algunas especies de
Cheilanthes puede ser muy escamosas en el envés, pero
estas especies usualmente tienen hojas más finamente
divididas.
COMENTARIOS: Trachypteris consta de 3 especies
y existe desde Ecuador (incluyendo los Galápagos) a
Argentina y sur de Brasil, y Madagascar. Usualmente crece
sobre o entre rocas y en suelos secos, usualmente en
ambientes estacionales secos. Las hojas tienen la capaz
de perder su contenido de agua normal y enroscarse,
y luego después de una lluvia absorber agua y expandir.
Los especies americanas existen de 700-2900 m. Las tres
especies neotropicales son: T. gilliana (Baker) Svenson,
T. induta (Maxon) R. M. Tryon & A. F. Tryon y T. pinnata
(Hook. f.) C. Chr. Trachypteris induta tiene una roseta basal
de hojas estériles y hojas fertiles erectas y largamente
pecioladas, mientras que T. pinnata falta de una roseta
basal y tiene hojas monomorfas. Trachypteris gilliana se
asemeja a T. pinnata pero difiere por esporas reticuladas
o verrucosas y hojas fértiles trífidas. Otra especie, T.
drakeana C. Chr., existe en Madagascar. Tiene una roseta
basal de hojas estériles, como en T. induta, pero cada
hoja lleva una yema apical. Conforme a estudios de rbcL,
Trachypteris es el género hermana a Doryopteris (Gastony
& Rollo, 1998). El nombre genérico se deriva del griego,
trachys, áspero + pteris, helecho. Las escamas cubriendo
las hojas imparten una textura aspera.
Figure 275. Trachypteris. A–H. T. induta. J–M. T. pinnata. (Original, ©Robbin Moran, 2016.)

557
HYMENOPHYLLACEAE
Trichomanes L. subgen. Davalliopsis
DESCRIPTION: Terrestrial; roots many, robust;
rhizomes 4-15 mm wide, erect or ascending, pubescent,
the hairs dark, multicellular; sterile and fertile leaves up
to 60 × 35 cm, monomorphic, polystichous; sterile
laminae 3 cells thick between the veins, opaque (not
translucent), 3- to 4-pinnate, usually iridescent, the
apices pinnatifid, not flagellate or proliferous; veins free,
anadromous, false veins absent; sori marginal, oriented
downward, out of the plane of the lamina; indusia
tubular, the mouth truncate or slightly dilate; receptacles
exert; spores green, tetrahaedral-globose; x=32.
DESCRIPCIÓN: Terrestres; raíces numerosas,
robustos; rizomas 4-15 mm de ancho, erectos o
ascendentes, pubescentes, los pelos oscuros, multicelulares;
hojas estériles y fértiles hasta 60 × 35 cm, monomorfas,
polistícas; láminas estériles 3 células de grosor
entre las venas, no translúcidas, 3- a 4-pinnadas,
usualmente iridiscentes, los ápices pinnatífidos, no
flagelados y prolíferos; venas libres, anadrómas, falsas
venas ausentes; soros marginales, orientados de abajo,
a fuera del plano de la lámina; indusios tubulares,
las bocas truncadas o levemente dilatas; receptáculos
exertos; esporas verdes, tetrahédricas-globosas; x=32.

SIMILAR GENERA: Abrodictyum differs by laminae non-
iridescent and only a few rows of cells on either side of
the veins. Trichomanes subgen. Feea differs by flagellate,
proliferous apices of the sterile leaves and (usually)
dimorphic sterile and fertile leaves. Trichomanes subgen.
Trichomanes differs by translucent laminae and epiphytic
habit. Vandenboschia and Trichomanes subgen. Lacostea
differ by long-creeping rhizomes that climb up trunks.
Polyphlebium differs by epiphytic habit and creeping
rhizomes.
GÉNEROS PARECIDOS: Abrodictyum difiere por
láminas no iridiscentes y solo unas hileras de células por
los lados de las venas. Trichomanes subgén. Feea difiere
por ápices de las hojas estériles flagelados y prolíferos y
(usualmente) hojas estériles y fértiles dimórfas. Trichomanes
subgén. Trichomanes difiere por láminas translucidas y
hábito epifítico. Vandenboschia y Trichomanes subgén.
Lacostea difieren por rizomas largamente rastreros
que trepan los troncos. Polyphlebium difiere por hábito
epifítico, rizomas rastreros.

COMMENTS: Trichomanes subgen. Davalliopsis consists COMENTARIOS: Trichomanes subgén. Davalliopsis

of 2 neotropical species: T. elegans (widespread) and T.
resinosum (Guayana, s. Venezuela). Both occur terrestrially
in wet forests. Their rhizomes are erect and supported
by roots on all sides. The leaves are unusual in the family
by being 3 cells thick between the veins and (if in shade)
iridescent. The sori bend downward, out of the plane of
the lamina. The chromosome number of x=32, shared
with other species of the genus, is derived within the
family (Ebihara et al., 2007) The name Davalliopsis is
derived from Davallia and the Greek suffix -opsis, like.
consta de 2 especies neotropicales: T. elegans (ampliamente
difundida) y T. resinosum (Guayana, s. Venezuela). Ambos
son terrestres en bosques húmedos. Sus rizomas son
erectos y apoyados por raíces en todos lados. Las hojas
son atípicas dentro de la familia por ser 3 células de
grosor entre las venas y (si en la sombra) iridiscentes. Los
soros doblan hacia abajo, a fuera del plano de la lámina.
El número cromosómico de x=32, compartido con las
otras especies del género, se deriva dentro de la familia
(Ebihara et al., 2007). El nombre Davalliopsis se deriva de
Davallia y el sufijo griego -opsis, semejante.

LITERATURE: Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of
Hymenophyllaceae. Blumea 51: 1–60. Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global
molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical
Journal of the Linnean Society 155: 1–27. Moran, R. C. 2000. Trichomanes resinosum (Hymenophyllaceae), a new species from
southern Venezuela and adjacent Guyana. Brittonia 52: 238–240.

558
Figure 276. Trichomanes subgen. Davalliopsis. A, B. T. elegans. C. D. T. resinosum. (From Moran, 2000.)

559
HYMENOPHYLLACEAE
Trichomanes subgen. Feea (Bory) Hook.
DESCRIPTION: Terrestrial; roots many, robust, often
stilt-like; rhizomes 2-8 mm wide, suberect or ascending,
unbranched, pubescent, the hairs dark, multicellular;
sterile and fertile leaves up to 30 × 18 cm, dimorphic
(except T. mougeotii), polystichous; sterile laminae 1-cell
thick between the veins, 1-pinnate, the apices often
becoming flagellate and proliferous; veins free,
catadromous, false veins absent; sori marginal; indusia
tubular, the mouth truncate or slightly dilate; receptacles
exert; spores green, tetrahaedral-globose; x=32.
SIMILAR GENERA: Trichomanes subgen. Trichomanes
usually differs by monomorphic sterile and fertile leaves,
and sori that tend to be clustered toward the pinna tips.
Trichomanes subgen. Davalliopsis differs by laminae opaque
(3-cells thick between the veins), usually iridescent, and
not flagellate and proliferous.
DESCRIPCIÓN: Terrestres; raíces numerosas,
robustos, a menudo como raíces zancudos; rizomas 2-8
mm de ancho, suberectos a ascendentes, no ramificados,
pubescentes, los pelos oscuros, multicelulares; hojas
estériles y fértiles hasta 30 × 18 cm, dimorfas (con
excepción de T. mougeotii), polistícas; láminas estériles
una célula de grosor entre las venas, 1-pinnadas, los
ápices a menudo se tornan flagellados y prolíferos;
venas libres, catádromas, falsas venas ausentes; soros
marginales; indusios tubulares, la boca entera, tuncada;
receptáculos exertos; esporas verdes, tetrahédricas-
globosas; x=32.
GÉNEROS PARECIDOS: Trichomanes subgén.
Trichomanes difiere usualmente por hojas estériles y
fértiles monomórfas y soros que tienden ser agrupados
hacia los ápices de los pinnas. Trichomanes subgén.
Davalliopsis difiere por láminas no translucidas (3 células
de grosor entre las venas) y no flageladas y prolíferas.

COMMENTS: Trichomanes subgen. Feea is neotropical COMENTARIOS: Trichomanes subgén. Feea es

and typically grows in wet forests on steep clay soils or
rocks. It contains 5 species: T. botryoides, T. diversifrons, T.
mougeotii, T. osmundoides, and T. trollii. These plants typically
have short erect rhizomes supported by prop roots,
imparting the aspect of a small tree.The sterile leaf apices
may become flagellate and proliferous. If a lamina is
present on the fertile leaves, it is more than one cell thick.
All but one species (T. mougeotii) have dimorphic sterile
and fertile leaves. The chromosome number of x=32,
shared with other species of the genus, is derived within
the family (Ebihara et al., 2007) The genus name honors
A.L.A. Fée (1789–1874), a pteridologist from Strasbourg.
neotropical y típicamente ocurre en bosques húmedos
sobre suelos arcillosos o rocas. Consta de 5 especies: T.
botryoides,T. diversifrons,T. mougeotii,T. osmundoides y T. trollii.
Estas plantas típicamente tienen rizomas cortas erectas
apoyadas por raíces zancudos, dando el aspecto de un
arbolito. Los ápices de las hojas estériles pueden tornarse
flagelados y prolíferos. Si una lámina está presente en
las hojas fértile, es más de una célula de grosor. Con la
excepción de una especie (T. mougeotii), todas tienen hojas
estériles y fértiles dimorfas. El número cromosómico de
x=32, compartido con las otras especies del género, se
deriva dentro de la familia (Ebihara et al., 2007). El nombre
genérico honora A. L. A. Fée (1789–1874), pteridólogo
de Strasbourg.

LITERATURE: Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of
Hymenophyllaceae. Blumea 51: 1–60. Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global
molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical
Journal of the Linnean Society 155: 1–27. Dubuisson, J.-Y., S. Hennequin, F. Rakotondrainibe & H. Schneider. 2003.
Ecological diversity and adaptive tendencies in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to
climbing and epiphytic habits. Botanical Journal of the Linnean Society 142: 41–63. Hébant-Mauri, R. 1972. Le genre Trichomanes
L. (Fougéres Leptosporangiées). Adansonia 12: 469–495.

560
Figure 277. Trichomanes subgen. Feea. A–C. Trichomanes diversifrons. D–F. T. botryoides. G–K. T. osmundoides. L, M. T.
muegotii. (Original, ©Robbin Moran, 2009.)

561
HYMENOPHYLLACEAE
Trichomanes subgen. Lacostea (Bosch) C. Chr.
DESCRIPTION: Lianas; rhizomes 0.5-1.5 mm wide,
long-creeping, climbing, often branched, pubescent, the
hairs dark, multicellular; sterile and fertile leaves up to 30 ×
9 cm, monomorphic, short-petiolate, distichous,; laminae
pressed flat against the substrate, 1-cell thick between
the veins, simple, 1-pinnate or 1-pinnate-bipinnatifid;
pinnae symetrical; veins free, anadromous, false veins
absent; sori marginal, pedicellate, projecting at right
angles to the substrate and lamina; indusia tubular, the
mouth truncate or sometimes dilate; receptacles exert;
spores green, tetrahaedral-globose; x=32.
DESCRIPCIÓN: Lianas; rizomas 0.5-1.5 mm de ancho,
largamente rastreros, a menudo ramificados, pubescentes,
los pelos oscuros, multicelulares; hojas estériles y fértiles
hasta 30 × 9 cm, monomorfas, cortamente pecioladas,
dísticas; láminas aplastadas contra el sustrato, una
célula de grosor entre las venas, simple a 1-pinnadas o
-pinnado-bipinnatífidas; venas libres, anádromas, falsas
venas ausentes; soros marginales, pedicelladas,
llevados a angulos rectos al sustrato y lámina;
indusios tubulares, las bocas tuncadas o a veces dilatas;
receptáculos exertos; esporas verdes, tetrahédricas-
globosas; x=32.

SIMILAR GENERA: Vandenboschia differs by GÉNEROS PARECIDOS: Vandenboschia difiere por

hemiepiphytic or hemiepipetric habit and leaves spreading
away from substrate (not appressed). Trichomanes subgen.
Trichomanes differs by catadromous venation and leaves
spreading away from the substrate. Hymenophyllum differs
by two-valved indusia and included receptacles.
hábito hemiepifítico o hemiepipétrico y hojas divergentes
del sustrato (no aplastadas). Trichomanes subgén.
Trichomanes difiere nervación catadrómica y hojas
divergentes desde el sustrato. Hymenophyllum difiere por
indusios bivalvados y receptáculos incluidos.

COMMENTS: Trichomanes subgen.Lacostea is neotropical COMENTARIOS: Trichomanes subgén. Lacostea es

and occurs in wet forests below 800 m. It contains five
species: T. ankersii, T. pedicellatum, T. tanaicum, T. tuerckheimii,
and T. subsessile. These species are hemiepiphytes, starting
from gametophytes around the bases of trunks. The
earliest sporeling stage sends a long root to the soil, and
after contact is made, the rhizome starts to climb. Their
leaves are unusual by being plastered against the trunk
with pedicellate sori projecting outward. The petioles are
short or apparently absent. The chromosome number of
x=32, shared with other species of the genus, is derived
within the family (Ebihara et al., 2007) The name Lacostea
honors Cornelius Marinus van der Sande Lacoste (1815-
1887), student of East Indian bryophytes.
neotropical y existe en bosques húmedos abajo de 800
m. Consta de cinco especies: T. ankersii, T. pedicellatum, T.
tanaicum, T. tuerckheimii y T. subsessile. Estas especies son
hemiepífitas, empezando en las bases de troncos. Las
etapas jóvenes del esporofitos envia una raíz larga al
suelo, y después de hacer contacto, el rizoma empieza
subir. Sus hojas son inusuales por ser aplastadas contra el
tronco con los soros pedicelados proyectándose a fuera.
Los pecíolos son cortos a aparentemente ausentes. El
número cromosómico de x=32, compartido con otras
especies del género, se deriva dentro de la familia (Ebihara
et al., 2007). El nombre Lacostea honora Cornelius
Marinus van der Sande Lacoste (1815-1887), estudiante
de briofitas de India oriental.

LITERATURE: Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of
Hymenophyllaceae. Blumea 51: 1–60. Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global
molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical
Journal of the Linnean Society 155: 1–27. Dubuisson, J.-Y., S. Hennequin, F. Rakotondrainibe & H. Schneider. 2003.
Ecological diversity and adaptive tendencies in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to
climbing and epiphytic habits. Botanical Journal of the Linnean Society 142: 41–63. Hébant-Mauri, R. 1972. Le genre Trichomanes
L. (Fougéres Leptosporangiées). Adansonia 12: 469–495.

562
Figure 278. Trichomanes subgen. Lacostea. A–D. T. ankersii. E–K. T. tuerckheimii. L–O. T. tanaicum. (Original, ©Robbin
Moran, 2010.)

563
HYMENOPHYLLACEAE
Trichomanes L. subgen. Trichomanes
DESCRIPTION: Terrestrial, rarely epiphytic; roots
many, robust; rhizomes 1-5 mm wide, erect or short- to
long-creeping, pubescent; sterile and fertile leaves up to 70
× 25 cm, monomorphic or rarely dimorphic, polystichous;
sterile laminae 1-cell thick between the veins, 1-pinnate,
the apices pinnatifid (flagellate and proliferous only in two
species), pubescent, the hairs with a distinct basal cell;
pinnae symmetric; veins free, catadromous (anadromous
in T. scandens and T. anadromum), false veins absent
(present only in T. pinnatum); sori marginal, immersed
in the laminae or (in two species) pedicellate; indusia
tubular, the mouth truncate or slightly dilate; receptacles
exert; spores green, tetrahaedral-globose; x=32.
SIMILAR GENERA: Trichomanes subgen. Feea differs
by flagellate, proliferous apices of the sterile leaves and
(usually) dimorphic sterile and fertile leaves. Trichomanes
subgen. Davalliopsis differs by laminae opaque (3-cells thick
between the veins) and usually iridescent. Vandenboschia
and Trichomanes subgen. Lacostea differ by long-creeping
rhizomes that climb the base of trunks. Polyphlebium
usually differs by epiphytic habit, creeping rhizomes, and
more divided leaves.
COMMENTS: Trichomanes subgen. Trichomanes is
neotropical except for T. crenatum of western Africa.
The subgenus contains ca. 30 species that typically are
terrestrial in wet forests. The neotropical species were
monographed by Windisch (1992).. Most have 1-pinnate
leaves with sori borne at the pinna apices, not along
the sides. Many have the laminae pubescent by long,
multicellular hairs, these with a distinct basal cell. The
chromosome number of x=32, shared with other species
of the genus, is derived within the family (Ebihara et al.,
2007) The genus name is derived from the Greek thrix,
hair + manes, cup. It alludes to the hair-like receptacle that
protrudes from the cup-like indusium.
LITERATURE: Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global molecular phylogeny of the
fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of the Linnean Society
155: 1–27. Lellinger, D. B. 1994. Trichomanes polypodioides and its allies. American Fern Journal 84: 1–4. Windisch, P. G. 1992.
Trichomanes crispum L. (Pteridophyta, Hymenophyllaceae) and allied species. Bradea 6: 78–117.
564
DESCRIPCIÓN: Terrestres, raras veces epífitas;
raíces numerosas, robustos; rizomas 1-5 mm de
ancho, erectos o cortamente a largamente reptantes,
pubescentes; hojas estériles y fértiles hasta 70 × 25 cm,
monomorfas o raras veces dimorfas, polistícas; láminas
estériles una célula de grosor entre las venas, 1-pinnadas,
los ápices pinnatífidos (flagelados y prolíferos solo en dos
especies), pubescentes, los pelos con una célula distinta
basal; venas libres, catádromas, falsas venas ausentes
(presentes solo en T. pinnatum); soros marginales,
inmersas en la lámina o (en dos especies) pediceladas;
indusios tubulares, las bocas enteras, truncadas o
levemente dilatas; receptáculos exertos; esporas verdes,
tetrahédricas-globosas; x=32.
GÉNEROS PARECIDOS: Trichomanes subgén. Feea
difiere por ápices de las hojas estériles flagelados y
prolíferos y (usualmente) hojas estériles y fértiles dimórfas.
Trichomanes subgén. Davalliopsis difiere por láminas
no translucidas (3 células de grosor entre las venas) y
usualmente iridiscentes. Vandenboschia y Trichomanes
subgén. Lacostea difieren por rizomas largamente
rastreros que trepan la base de los troncos. Polyphlebium
difiere por hábito epifítico, rizomas rastreros y hojas más
divididas.
COMENTARIOS: Trichomanes subgén. Trichomanes
es neotropical con excepción de T. crenatum de África
occidental. El subgénero consta de ca. 30 especies que
típicamente ocurre en bosques húmedos. Las especies
neotropicales fueron monografiadas por Windisch
(1992). La mayoría tienen hojas 1-pinnatas con soros
llevados en los ápices de las pinnas, no los lados. Muchas
especies tienen las láminas pubescentes por pelos
largos multicelulares, estos con una célula basal distinta.
El número cromosómico de x=32, compartido con las
otras especies del género, se deriva dentro de la familia
(Ebihara et al., 2007). El nombre genérico se deriva del
griego thrix, pelo + manes, copa. Se refiere a receptáculo
piliforme que sobresale del indusio en forma de copa.
Figure 279. Trichomanes subgen. Trichomanes. A, B. T. pinnatum. C, D. T. polypodioides. E, F. T. lucens. G, H. T. crispum.
(From Mickel & Smith, 2004.)

565
TECTARIACEAE
Triplophyllum Holttum
DESCRIPTION: Terrestrial; rhizomes creeping, scaly;
leaves monomorphous, long-petiolate; vascular bundles in
petiole 4 or more; lamina typically tripartite, deltate or
pentagonal, the basal pinnae typically conspicuously larger
than the suprabasal ones and wider on the basiscopic
side; rachis and costae not grooved, usually puberulent
below and usually above with reddish hairs, these 0.1-0.4
mm long, reddish, articulate, below provided with a few
dark scales; veins free; sori round; indusia present, circular
to reniform, attached laterally (not peltate); x=41.
SIMILAR GENERA: Tectaria typically differs by
anastomosing veins. Megalastrum and Ctenitis usually differ
by erect or decumbent rhizomes and scalier rachises and
costae. Similar dryopteroid genera, such as Arachniodes,
Dryopteris, and Polybotrya, differ by adaxially grooved
rachises and costae.
COMMENTS: Triplophyllum occurs in Africa (14 spp.) and
the Neotropics (9 spp.). A key to the neotropical species
can be found in Prado & Moran (2008).The genus is most
species-rich and abundant in the Guayanas and northern
Brazil. It usually grows on wet, shaded forest floors,
generally from 0–1200 m. The usually tripartite laminae,
with basal pinnae greatly elongated on the basiscopic
side, make this an easy genus to recognize. The species
are more challenging to identify, being distinguished by
indument characteristics that require at least a 30X
dissecting microcope and a strong light source.
The genus name is derived from the Latin triplo, three
times + the Greek phyllon, leaf. It refers to the tripartite
leaf.
LITERATURE: Holttum, R. E. 1986. Studies in the fern-genera allied to Tectaria Cav., 5. Triplophyllum, a new genus of Africa
and America. Kew Bulletin 41: 237–260. Prado, J. & R. C. Moran. 2008. Revision of the neotropical species of Triplophyllum
(Tectariaceae). Brittonia 60: 103–130.
566
DESCRIPCIÓN: Terrestre; rizomas reptantes,
escamosos; hojas monomorfas, largamente pecioladas;
haces vasculares en los pecíolos 4 o más; láminas
típicamente tripartidas, deltadas o pentagonales, las
pinnas basales típicamente conspicuosamente más grandes
que las adyacentes y más anchas en el lado basiscópico;
raquis y costas no surcados, usualmente puberulentos
en la superficie inferior e usualmente superior con
pelos 0.1-0.4 mm de largo, rojizos, articulados, en el lado
inferior provisto con pocas escamas oscuras; nervadura
libre; soros redondeados; inducíos presentes, circulares a
reniformes, unidos lateralmente (no peltados); x=41.
GÉNEROS PARECIDOS: Tectaria difiere típicamente
por venas anastomosadas. Megalastrum y Ctenitis
usualmente difieren por rizomas erectos o decumbentes y
ráquises y costas más escamosos. Géneros dryopteroides
parecidos, tales como Arachniodes, Dryopteris, y Polybotrya,
difieren por los raquises y pinnas surcados adaxialmente.
COMENTARIOS: Triplophyllum existe en África (14
spp.) y el Geotrópico (9 spp.). Una clave a las especies
neotropicales está presentada en Prado & Moran (2008).
El género es más diverso en las Guayanas y Brasil del
norte. Existe en sotobosques húmedos y sombreados,
generalmente de 0–1200 m. Las láminas usualmente
tripartitas, con pinas basales alargadas fuertemente en
el lado basiscópico, contribuyen a la recognición fácil de
este género. Las especies presentan un reto identificar,
siendo distinguidas por características del indumento que
requieren a lo menos a 30X microscopio de disección y
una fuente de luz fuerte. El nombre genérico se deriva
del latín triplo, tres veces + el griego phyllon, hoja. Se
refiere a la hoja tripartita.
Figure 280. Triplophyllum. A. T. chocoense. B. T. dicksonioides, pinnule. C. T. dicksonioides, habit. D. T. funestum, sorus.
E. T. funestum, adaxial surface of the pinna rachis (left), abaxial surface of same (right). F. T. dicksonoides, pinnule. G. T.
dicksonioides, abaxial surface of the pinna rachis (left), side view of pinna rachis, adaxial side to the left (left). (From Prado &
Moran, 2008.)

567
PTERIDACEAE
Tryonia Schuettp., J. Prado & A. T. Cochran
DESCRIPTION: Terrestrial or saxicolous; rhizomes
creeping, bristly; leaves 6–100 cm; petioles brown
basally and straminous distally, pubescent; laminae
linear to elongate-triangular, 1- to 3-pinnate-pinnatifid,
determinate; rachises stramineous, straight, terete or
sulcate adaxially, pubescent; pinnae 0.5–10 × 0.5–5 cm,
membranaceous to herbaceous, densely pubescent on
both surfaces, the hairs glandular or non-glandular; veins
free; soros no indusiados; sporangia borne along the
veins; spores brown, trilete, tetrahedral-globose; x=?.
DESCRIPCIÓN: Terrestres o saxícolas; rizomas
reptantes, cerdosos; hojas 6–100 cm; pecíolos pardos
basalmente e estramíneous distalmente, pubescentes;
láminas lineares a alargadas-triangulares, 1- 3-pinnado-
pinnatífidas, determinadas; ráquises estramineos,
rectos, teretes o surcados adaxialmente, pubescentes;
pinnas 0.5–10 × 0.5–5 cm, membranosas a herbáceas,
densamente pubescentes en ambos lados, los pelos
glandulosos o no; venas libres; soros no indusiados;
esporangios difundidos por las venas; esporas pardas,
triletes, tetrahédricas-globosas; x=?.

SIMILAR GENERA: Jamesonia differs castaneous GÉNEROS PARECIDOS: Jamesonia difiere por
rachises. ráquises castaños.

COMMENTS: Tryonia consists of four species of the
Atlantic forests of southeastern Brazil, with one species
occurring in adjacent Uruguay. Generally along streams
on damp, shaded sandstones; 600–2300 m. The species
of Tryonia were formerly-treated in Eriosorus, a genus
now subsumed under Jamesonia. Tryonia is distinguished
from Jamesonia only by stramineous rachises. Instead
being most closely related to Jamesonia, Tryonia is most
closely related to the Old World genera Austrogramme,
Pterozonium, Syngramma and Taenitis. The four species of
Tryonia are T. areniticola (Schwartsb. & Labiak) Schuettp.,
J. Prado & A. T. Cochran, T. myriophylla (Sw.) Schuettp., J.
Prado & A. T. Cochran, T. schwackeana (Christ) Schuettp.,
J. Prado & A. T. Cochran y T. sellowiana (Kuhn) Schuettp., J.
Prado & A.T. Cochran. Commemorates Alice Faber Tryon
(1920–2009), American pteridologist who monographed
Eriosorus and Jamesonia, the genera from which Tryonia
was segregated.
COMENTARIOS: Tryonia consta de cuatro especies
de los bosques Atlánticos del sureste de Brasil, con
una especie en Uruguay adyacente. Generalmente por
riachuelos en piedra arenosa sombreada e húmedo; 600–
2300 m. Las especies de Tryonia fueron anteriormente
clasificadas en Eriosorus, un género subsumido ahora
abajo de Jamesonia. Tryonia se distingue de Jamesonia solo
por ráquises estramíneos. Tryonia es más cercanamente
relacionados a los géneros del Viejo Mundo Austrogramme,
Syngramma y Taenitis. Las cuatro species de Tryonia son T.
areniticola (Schwartsb. & Labiak) Schuettp., J. Prado & A.
T. Cochran, T. myriophylla (Sw.) Schuettp., J. Prado & A. T.
Cochran, T. schwackeana (Christ) Schuettp., J. Prado & A.
T. Cochran y T. sellowiana (Kuhn) Schuettp., J. Prado & A. T.
Cochran. Conmemora a Alice Faber Tryon (1920–2009),
pteridólogo Americano quien hizo una monografía a
Eriosorus y Jamesonia, los generos de los cuales Tryonia
fue segregado.

LITERATURE: Cochran, A. T., J. Prado & E. Schuettpelz. 2014. Tryonia, a new taenitidoid fern genus segregated from
Jamesonia and Eriosorus. (Pteridaceae). PhytoKeys 35: 23–43. Tryon, A. F. 1970. A monograph of the fern genus Eriosorus.
Contributions from the Gray Herbarium of Harvard University 200: 54–174.

568
Figure 281. Tryonia. A–D. T. myriophylla (Sw.) Schuettp., J. Prado & A. T. Cochran. E–G. T. sellowiana (Kuhn) Schuettp., J.
Prado & A. T. Cochran. (Original, ©Robbin Moran, 2016.)

569
HYMENOPHYLLACEAE
Vandenboschia Copel.
DESCRIPTION: Hemiepiphytic or hemiepilithic;
roots many and robust; rhizomes 1-2 mm wide, long-
creeping, irregularly branching, densely pubescent, the hairs
dark, multicellular; sterile and fertile leaves up to 40 × 20
cm, monomorphic, distichous; laminae membranaceous,
translucent, 1-cell thick between the veins, 1-pinnate to
5-pinnate; veins free, anadromous, false veins absent; sori
marginal; indusium (often called an “involucre”)
tubular to campanulate, the mouth entire or flaring;
receptacles exert, extending beyond the mouth of the
indusium; spores green, tetrahaedral-globose; x=36.
DESCRIPCIÓN: Hemiepifitos o hemiepilíticos;
raíces numerosas y robustos; rizomas 1-2 mm de ancho,
largamente rastreros, ramificándose irregularmente,
típicamente densamente pubescentes, los pelos oscuros,
multicelulares; hojas estériles y fértiles hasta 40 × 20
cm, monomorfas, dísticas; láminas membranáceas,
translucientes, una célula de grosor entre las venas,
1-pinnadas a 5-pinnadas; venas libres, anádromas, falsas
venas ausentes; soros marginales; indusio (a menudo
llamado “involucre”) tubular, la boca entera o
ensanchada; receptáculos exertos, extendiéndose
fuera de la boca del indusio, filiforme; esporas verdes,
tetrahédricas-globosas; x=36.

SIMILAR GENERA: Hymenophyllum differs by two-
valved indusia and included receptacles. When dried, its
laminae are often pale reddish, whereas those of the
trichomanoid genera are green. Polyphlebium differs by
epiphytic habit, filiform rhizomes, and lack of conspicuous
roots. Trichomanes subgen. Trichomanes usually differs by
terrestrial habit, erect to short-creeping rhizomes, and
catadromous venation.
GÉNEROS PARECIDOS: Hymenophyllum difiere por
indusios bivalvados y receptáculos incluídos. Cuando se
secan, sus láminas son a menudo rojizas pálidas, mientras
que las de los géneros trichomanoides son verdes.
Polyphlebium difiere por su hábito epifítico, rizomas
filiformes y falta de raíces conspicuas. Trichomanes subgen.
Trichomanes difiere usualmente por hábito terrestre,
rizomas erectos a cortamente reptantes y nervación
catadrómica.

COMMENTS: Vandenboschia is pantropical and contains COMENTARIOS: Vandenboschia es pantropical y

about 15 species, of which 4 are in the New World: V. consta de 15 especies, de las cuales 4 están en el Nuevo
boschiana, V. collariata, V. radicans, and V. rupestris. Among Mundo: V. boschiana, V. collariata, V. radicans, and V. rupestris.
the trichomanoid genera, Vandenboschia is unique by its Entre los géneros trichomanoides, Vandenboschia es único
hemiepiphytic or hemiepilithic habit; that is, the sporelings por su hábito hemiepifítico o hemiepipétrico; es decir, los
begin growth on the bases of trees or rocks and then esporofitos empiezan crecer en las bases de troncos o
become secondarily terrestrial by growing roots into rocas y luego llegan a ser terrestres secundariamente
the soil (where they branch profusely). The roots are por enviando raíces al suelo. Estas raíces se producen de
produced from a short-creeping rhizome that also un rizoma cortamente reptante que también produce
produces long-creeping climbing rhizomes. The latter rizomas largamente rastreros y trepadoras. Estos
occasionally produce new branches and thick roots from produces ocasionalmente nuevas ramas y raíces gruesas
buds located in or near the axils of the leaves. The genus desde brotes ubicados en o cerca de las axilas de las
name honors Roelof van den Bosch (1810–1862), Dutch hojas. El nombre genérico honora Roelof van den Bosch
pteridologist and student of filmy ferns. (1810–1862), pteridólogo holandés y estudiante de las
himenofiláceas.
LITERATURE: Ebihara, A., J.-Y. Dubuisson, K. Iwatsuki, S. Hennequin & M. Ito. 2006. A taxonomic revision of
Hymenophyllaceae. Blumea 51: 1–60. Ebihara, A., K. Iwatsuki, M. Ito, S. Hennequin & J-Y. Dubuisson. 2007. A global
molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical
Journal of the Linnean Society 155: 1–27. Dubuisson, J.-Y., S. Hennequin, F. Rakotondrainibe & H. Schneider. 2003.
Ecological diversity and adaptive tendencies in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to
climbing and epiphytic habits. Botanical Journal of the Linnean Society 142: 41-63. Hébant-Mauri, R. 1972. Le genre Trichomanes
L. (Fougéres Leptosporangiées). Adansonia 12: 469–495. Nitta, J. & M. J. Epps. 2009. Hemi-epiphytism in Vandenboschia
collariata (Hymenophyllaceae). Brittonia 61: 392–397.

570
Figure 282. Vandenboschia. A–C. V. radicans. D–F. V. collariata. (From Mickel & Smith, 2004.)

571
PTERIDACEAE
Vittaria J. E. Sm.
DESCRIPTION: Epiphytic; rhizomes short-creeping,
dorsiventral; rhizome scales clathrate. Leaves distichous,
clumped or slightly remote; petioles absent or nearly so;
laminae 14–40 cm long, 1-4 mm wide, simple, entire,
elongate-linear, coriaceous to subcoriaceous, glabrous;
veins obscure, anastomosing, with a single row of areoles
between the rachis and margin, the areoles without
included veinlets; sori linear, in one row between the
rachis and margin, parallel to the margin, sunken in a
groove; paraphyses with a slender or slightly clavate apical
cell (not obconic); spores trilete or monolete, nongreen;
gametophytes with paired gemmae; x=60.
DESCRIPCIÓN: Epífito; rizomas cortamente rastreros,
dorsiventrales, escamas del rizoma clatradas; hojas
dísticous, amacolladas o un poco remotas; pecíolos
ausentes o casi así; láminas 15–40 cm longas, 1–4 mm
de ancho, simples, enteras, alargados-lineares, coriáceas
a subcoriáceas, glabras; venas obscuras, anastomosadas,
con una hilera de aréolas entre el raquis y el margen, las
aréolas sin venillas incluídas; soros lineares, en una sóla
hilera entre el raquis y el margen, paralelas al margen,
hundidos en un surco; parafisos con una célula apical
delgada o levemente clavada (pero no obcónica); esporas
triletes o monoletes, no verdes; gametofitos con gemas
pareadas; x=60.

SIMILAR GENERA: Radiovittaria differs by dark petioles,
usually wider blades (0.2–1.0(–1.8) cm), and paraphyses
with an obconic apical cell. Other epiphytic genera with
simple, entire leaves (e.g., Campyloneurum, Elaphoglossum)
usually differ by non-linear sori and the presence of hairs
or scales on the blades.
GÉNEROS PARECIDOS: Radiovittaria difiere por
pecíolos oscuros, usualmente láminas más anchas (0.2–
1.0(–1.8) cm) y parafisos con una célula apical obcónica.
Otros géneros epifíticos con hojas simples y enteras (e.g.,
Campyloneurum, Elaphoglossum) usualmente difieren por
soros no lineares y la presencia de pelos o escamas sobre
la lámina.

COMMENTS: Vittaria comprises 6 species, all neotropical
except for one that occurs in Africa and offshore islands
in the Indian Ocean. All the species typically grow in wet
forests as epiphytes on trunks or in the canopy. All species
have pendulous leaves. Because of the long, linear leaves
of some species, the genus has the common name of
“shoe-string fern.” The genus name is derived from the
Latin vitta, ribbon. The leaves are narrow and ribbon-like.
COMENTARIOS: Vittaria consta de 6 especies, todas
neotropicales con excepción de una que ocurre en África
e islas cercanas en el Océano Indio. Todas las especies
crecen típicamente en bosques húmedos como epífitas
sobre troncos o en el dosel. Todas tienen hojas péndulas.
Debido a que las hojas son largas y lineares en algunas
especies, el género tiene el nombre común de “helecho
de cordón de zapato”. El nombre genérico se deriva del
latín vitta, cinta. Las hojas son estrechas y como una cinta.

LITERATURE: Benedict, R. C. 1914. A revision of the genus Vittaria J.E. Smith. I. The species of subgenus Radiovittaria.
Bulletin of the Torrey Botanical Club 41: 391–410. Crane, E. H. 1977. A revised circumscription of the genera of the fern family
Vittariaceae. Systematic Botany 22: 509–517. Crane, E. H., D. R. Farrar, & J. F. Wendel. 1995. Phylogeny of the Vittariaceae:
convergent simplification leads to a polyphyletic Vittaria. American Fern Journal 85: 283–305. Tryon, R. M. 1964.Taxonomic Fern
Notes, IV. Some American vittarioid ferns. Rhodora 66:110–117.

572
Figure 283. Vittaria. A–C. V. dimorpha. E–F. V. flavicosta. G–O. V. graminifolia. L–O. V. lineata. (From Mickel & Smith, 2004.)

573
WOODSIACEAE
Woodsia R. Br.
DESCRIPTION: Terrestrial or saxicolous; stems erect or
decumbent, scaly, the scales thin, translucent; sterile and
fertile leaves monomorphous; petioles with 2 vascular
bundles; laminae usually 10–35 cm long, 1-pinnate-
pinnatifid to 2-pinnate-pinnatifid, scaly (on axes), usually
pubescent with articulate hairs and sometimes with
capitate-glandular hairs; pinnae sessile or nearly so; axes
grooved, the grooves decurrent; veins free, the tips slightly
thickened (hydathodous); sori round; indusia attached
around the base of the receptacle, splitting at
maturity into several lobes or many hairs that form
a loose cup around the sporangia; x=38, 39, 41.
SIMILAR GENERA: Cystopteris differs by the indusium
attached on the proximal side of the receptacle only.
Cheilanthes differs by one vascular bundle at the base of
the petiole and marginal sori protected by false indusia.
COMMENTS: Woodsia is the only genus of Woodsiaceae.
It contains about 40 species mostly in the temperate
zones of the Northern Hemisphere, but with extensions
into Central and South America, and in South Africa and
Madagascar. In the Americas, Woodisa is most diverse
in Mexico (8 spp.). Only one species occurs in South
America and the Antilles (Hispaniola): W. montevidensis
(Spreng.) Hyl. Three subgenera are recognized: subg.
Woodsia, subg. Perrinia Hook., and subg. Cheilanthopsis
(Hieron.) X.C. Zhang & R. Wei. The latter is Asian and
does not occur in the Americas. The two subgenera that
occur in the Americas differ by several characters. Subg.
Woodsia consists of three species (W. alpina, W. glabella,
and W. ilvensis) characterized by concolorous rhizome
scales, articulate petioles, indusial segments uniseriate
throughout and composed of elongate cells. In contrast,
subg. Perrinia has bicolorous rhzome scales, non-articulate
petioles, indusia multiseriate at base, composed of
isodiametric cells or nearly so. The genus name honors
Joseph Woods (1776–1864), British architect and author
of A Tourist’s Flora.
LITERATURE: Brown, D. F. M. 1964. A monographic study of the fern genus Woodsia. Beihefte zur Nova Hedwigia 16:
1–154. Shao,Y. , R. Wei, X.-C. Zhang & Q. Xiang. 2015. Molecular phylogeny of the cliff ferns (Woodsiaceae: Polypodiales) with a
proposed infrageneric classification. PloS One 10(9):e0136318.
574
DESCRIPCIÓN: Terrestres o saxícolas; tallos erectos
o decumbentes, escamosos, las escamas delgadas,
translúcidas; hojas estériles y fértiles monomorfas;
pecíolos con 2 haces vasculares; láminas usualmente
10–35 cm de largo, 1-pinnado-pinnatífdas a 2-pinnado-
pinnatífidas, escamosas (sobre los ejes), usualmente
pubescentes con pelos articulados y a veces con pelos
capitados-glandulosos; pinnas sésiles o casi sésiles; ejes
surcados, los surcos decurrentes; nervaduras libres,
los ápices un poco engrosados (hidatodos); soros
redondos; indusios unidos alrededor de la base del
receptáculo, se hende en varios lobulos o muchos
pelitos que forman una copa suelta alrededor de los
esporangios; x=38, 39, 41.
GÉNEROS PARECIDOS: Cystopteris difiere por el
indusio unido solo al lado proximal del receptáculo.
Cheilanthes difiere por un haz vascular en la base del
pecíolo y soros marginales protegidos por indusios falsos.
COMENTARIOS: Woodsia es el único género de
Woodsiaceae. Consta de casi 40 especies mayormente
en zonas templadas del Hemisfério Norte pero con
extensiones en América tropical, y África del sur y
Madagascar. En las Américas, Woodisa más diverso
en México (8 spp.). Solamente una especie existe
en América del Sur y las Antillas (Hispaniola): W.
montevidensis (Spreng.) Hyl. Se reconoce tres subgéneros:
subg. Woodsia, subg. Perrinia Hook. y subg. Cheilanthopsis
(Hieron.) X.C. Zhang & R. Wei. Este último es Asiático
y no existe en las Américas. Los dos subgéneros que
existen en las Américas difieren por varias características.
El subg. Woodsia consta de tres especies (W. alpina, W.
glabella, and W. ilvensis) caracterizadas por escamas del
rizoma concoloras, pecíolos articulados, segmentos
indusiales uniserados por todo se extensión y consta
de células alargadas. En contraste, el subg. Perrinia tiene
escales del rizoma bicoloras, pecíolos no articulados,
indusios multiseradsos en la base y compuestos de
células isodiamétricas. El nombre genérico honora Joseph
Woods (1776–1864), arquitecto británico y autor de A
Tourist’s Flora.
Figure 284. Woodsia. A–E. W. plummerae. F–K. W. cystopteroides. L–O. W. cochisensis. (From Mickel & Smith, 2004.)

575
BLECHNACEAE
Woodwardia Sm.
DESCRIPTION: Terrestrial; rhizomes creeping,
decumbent or erect, scaly, non-stoloniferous; sterile
and fertile leaves monomorphous; laminae 1-pinnate-
pinnatifid (in Mesoamerica) to 2-pinnate; pinnae
pinnatifid, the segments spinulose (in Mesoamerica) or
entire; aerophores absent; veins areolate, at least along
the costae, without included veins; sori parallel to
the costae, one per areole, in most species deeply
immersed; indusia opening toward the costae, not the
margin. Spores monolete, nongreen. x=31, 34.
SIMILAR GENERA: Anchistea differs by long-creeping
rhizomes and pinnae articulate to the rachis. Lorinseria
differs by dimorphic sterile and fertile leaves, and sterile
pinnae adnate to the rachis.
COMMENTS: Woodwardia is mostly north-temperate
and contains about 13 species. In the Neotropics, it
extends from Mexico to Nicaragua in wet forests above
1500 m. It contains two neotropical species, W. spinulosa
M. Martens & Galeotti and W. martinezii Maxon ex Weath.,
and their hybrid, W. X semicordata Mickel & Beitel. The
center of diversity for the genus is in China, where about
10 species occur. The common name for the genus is
“chain fern,” alluding to the chain-like row of sori along the
costae or costules. Other Blechnaceae, except Anchistea
and Lorinseria, have a single continous sorus parallel to
the midrib. The genus name honors Thomas Jenkinson
Woodward (1745–1820), British phycologist.
LITERATURE: Gasper, A. L. & V. A. O. Dittrich, A. R. Smith & A. Salino. 2016. A classification for Blechnaceae
(Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations. Phytotaxa 275: 191–227. Maxon, W. R.
1919. Notes on American ferns XIV. American Fern Journal 9: 67–73. Weatherby, C. A. 1949. Two Mexican ferns. American
Fern Journal 39: 88–91.
576
DESCRIPCIÓN: Terrestres; rizomas rastreros,
decumbentes o erectos, escamosos, no estoloníferos;
hojas estériles y fértiles monomorfas; láminas 1-pinnado-
pinnatífidas (en Mesoamérica) a 2-pinnadas; pinnas
pinnatífidas, los segmentos espinulosos (en Mesoamérica)
o enteros; aeróforos ausentes en las bases de las pinnas
en el envés y a lo largo del pecíolo; nervaduras areoladas,
al menos a lo largo de las costas, sin nérvulos incluidos;
soros paralelos a las costas o cóstulas, uno por
aréola, en la mayoría de las especies profundamente
inmersos; indusios abriéndose hacia la costa, no el margen;
esporas monoletes, no verdes. x=31, 34..
GÉNEROS PARECIDOS: Anchistea difiere por
rhzomas largamente rastreros y pinnas articuladas
al ráquis. Lorinseria difiere por hojas estériles y fértiles
dimorfas, y pinnas estériles adnatas al ráquis.
COMENTARIOS: Woodwardia es principalmente
en el norte templado y consta de 13 especies. En el
Neotrópico se extiende desde México hasta Nicaragua
en bosques húmedos arriba de 1500 m. Contiene
dos especies neotropicales, W. spinulosa M. Martens &
Galeotti y W. martinezii Maxon ex Weath., y su híbrido W.
X semicordata Mickel & Beitel. El centro de diversificación
está en China, en donde existen ca. 10 especies. El nombre
común para el género es “helecho de cadena,” en alusión
a la hilera de soros que asemejan cadenas por las costas y
cóstulas. Otras Blechnaceae, con excepcion de Anchistea
y Lorinseria, tienen un solo soro continuo paralelo a la
costa. El nombre génerico honora a Thomas Jenkinson
Woodward (1745–1820), ficólogo británico.
Figure 285. Woodwardia. A–E. W. spinulosa. (From Mickel & Smith, 2004.)

577
POLYPODIACEAE
Zygophlebia L.E. Bishop
DESCRIPTION: Epiphytic; rhizome scales not clathrate,
shiny, cilate or glandular-papillose on the margins;
phyllopodia present, swollen; petiole equalling or longer
than the lamina; laminae pinnatisect, with or without setae,
with simple or branched glandular hairs; hydathodes
absent; veins 1—2(—3) forked, connivent at the apices
or rarely free; sori separate, round, paraphyses presnt,
consisting of a 1-seriate stalk with 2 or 3 brown glands
distally; spores green.
DESCRIPCIÓN: Epífitas; escamas del rizoma no
clatradas, lustrosas, ciliadas o glanduloso-papilosas en los
márgenes; filopodios presentes, hinchados; pecíolo tan
largo como la lámina o más largo; láminas pinnatisectas, con
o sin setas, con tricomas glandulosos simples o bifurcados;
hidatodos ausentes; nervaduras 1—2(—3) bifurcadas,
conniventes en los ápices o raramente libres; soros
separados, redondos, parafisos presentes, constituidos
por un pedicelo 1-seriado con 2 o 3 glándulas pardas
distalmente; esporas verdes.

DIAGNOSIS: Rhizome scales not clathrate, shiny, cilate DIAGNOSIS: Escamas del rizoma no clatradas, lustrosas,
or glandular-papillolse on the margins; phyllopodia present, ciliadas o glanduloso-papilosas en los márgenes; filopodios
swollen; veins areolate; hydathodes absent. presentes, hinchados; nervaduras areoladas; hidatodos
ausentes.

SIMILAR GENERA: Several species of Ceradenia GÉNEROS PARECIDOS: Varias especies de Ceradenia
resemble Zygophlebia but can usually be distinguished by se parecen a Zygophlebia pero usualmente pueden
having free veins and whitish globose glands in the sori distinguirse por tener venas libres y glándulas blanquecinas
(i.e., as paraphyses). globosas en el soro (i.e., como parafisos).

COMMENTS: Unfortunately, the two important COMENTARIOS: Desafortunadamente, las dos

characterisitcs of Zygophlebia (phyllopodia and areolate
veins) are often difficult to observe due to the covering
of scales on the rhizome and the thickness of the lamina.
The phyllopodia are evident as a swollen base with
a darker color than the petiole. The change in color is
usually abrupt. The absence of hydathodes on the upper
surface of the lamina will separate Zygophlebia from the
other neotropical genera of Grammitidaceae except
for Ceradenia and Enterosora (which see). The leaves of
Zygophlebia have a peculiar habit, but this characteristic
is found in other grammitid ferns: the petiole is nearly
horizontal, i.e., perpendicular to the branch or trunck on
which the plant is epiphytic and is sharply geniculate at its
apex so that the lamina is pendulous.
características importantes de Zygophlebia (filopodios
y nervaduras areoladas), son a menudo difíciles de
observar debido a la cobertura de escamas del rizoma y
al grosor de la lámina. Los filopodios son evidentes como
una base hinchada con un color más oscuro que el del
pecíolo. El cambio de color es típicamente abrupto. La
ausencia de hidatodos en el haz separará Zygophlebium
de los otros géneros neotropicales de Grammitidaceae
con excepción de Ceradenia y Enterosora (véase). Las
hojas de Zygophlebia tienen un hábito peculiar, pero esta
característica la poseen otros helechos gramitidáceos por
igual: el pecíolo es casi horizontal, i.e., casi perpendicular
a la rama o tronco en el que la planta es epífita y es
marcadamente geniculado en el ápice, de tal modo que
las láminas son péndulas.

LITERATURE: Bishop, L. E. 1989. Zygophlebia, a new genus of Grammitidaceae. American Fern Journal 79: 103–118.

578
Figure 286. Zygoplebia werfii. Note in C the anastomosing veins.These are often difficult to see because the laminae are thick.
(From Bishop, 1989.)

579
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_____, _____, & J. C. Vogel. 2002. Schizaeaceae: a phylogenetic approach. Review of Palaeobotany and Palynology 119: 35–50.
_____ & K. M. Pryer. 2005. Incongruence between primary sequence data and the distribution of a mitochondrial atp1 group
II intron among ferns and horsetails. Molecular Phylogenetics and Evolution 36: 484–493.
Wilce, J. H. 1965. Section Complanata of the genus Lycopodium. Beihefte Nova Hedwigia 19: i–ix, 1–233, plate 40.
_____. 1972. Lycopod spores, I. General spore patterns and the generic segregates of Lycopodium. American Fern Journal 62:
65–79.
Wilson, K. A. 1959. Sporangia of the fern genera allied with Polypodium and Vittaria. Contributions from the Gray Herbarium
of Harvard University 185: 97–127.
Windham, M. D. & G. Yatskievych. 2003. Chromosome studies of cheilanthoid ferns (Pteridaceae: Cheilanthoideae) from the
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Windisch, P. G. 1978. Synopsis of the genus Sphaeropteris (Cyatheaceae) with a revision of the neotropical species of the
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Wolf, P. G. 1995. Phylogenetic analysis of rbcL and nuclear ribosomal RNA gene sequences in Dennstaedtiaceae. American
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_____, S. D. Sipes, M. R. White, M. L. Martines, K. M. Pryer, A. R. Smith & K. Ueda. 1999. Phylogenetic relationships of
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 Appendices
Appendix 1 | Squash Technique for Fern and Lycophyte Chromosomes

Pre-treatments

To find sporangia that are undergoing meiosis, look for sori that are white. Any hint of brown means that meiosis has
already taken place and that the spore walls are beginning to form. At this stage it is too late.

Put the leaf tissue with sori into small bottles or vials filled with a saturated solution of paradichlorobenzene (i.e.,
“PDB” or moth crystals) made by adding distilled water to a vial containing a few layers of crystals. Keep at refriger-
ator temperature, around 5°C for 4 to 20 hours.

Remove material from the PDB solution, blot off water, and drop into the fixative (either Newcomer's or Farmer’s
solution). Leave at room temperature for two to six hours, and then place in freezer (not refrigerator) where it will
keep indefinitely. The bottles can be removed from freezer and left at room temperature for as long as six hours if
necessary while making squashes.

Squashing

To make a chromosome squash (Figure 287), place a small drop of ACH (50% acetocarmine and 50% Hoyer's solu-
tion mixed in dropping bottle) on a slide, and at one end of the slide a fragment of leaf containing sori or sporangia.
Blot the leaf tissue lightly and quickly with a kim wipe or paper towel, and under a dissecting microscope remove the
sporangia with a dissecting needle and place them in the drop of ACH. With two needles, spread apart the sporangia
in the drop and remove any excess material such as bits of indusia, leaf tissue, or scales. Cover with a square cover
slip.Tap gently with a needle to squeeze the spore mother cells from the sporangia. Heat gently over an alcohol lamp
until slide feels warm; be careful not to boil the acetocarmine, which is easy to do. Immediately place the warm slide
on hard smooth surface covered with absorbent paper such as smooth filter paper. Place a second piece of filter
paper on top of the slide, and holding the edge of the cover slip with fingers or thumb of left hand, press mightily,
straight down on cover slip with right thumb (if right-handed, or reverse, if left-handed). Keep your arm straight and
steady because if the cover slip moves, the spore mother cells will be smeared.

Under low power of the compound microscope, survey the slide systematically, up and down, viewing fields in con-
secutive frames instead of continuous motion (this is easier on the eyes and helps prevent headaches). If you find a
cell you want to examine later under oil immersion, note the vernier reading. Less than perfect chromosome figures
can be resquashed by heating slightly and squashing as above with thumb directly above area of the slide containing
the figure.

Slides with good chromosome figures should be sealed with EUKITT, or other recommended material for sealing
slides. (Nail polish, formerly used, does not form an adequate seal.) If chromosomes are near the edge of the slide,
do not view under oil until the slide is sealed. Good figures can be circled on the slide using a fine ball-point pen.
Label slide.

Slides made with ACH are best viewed with a phase contrast microscope. If none is available, the procedure above
can be followed using acetocarmine without Hoyer's. There are several techniques for making acetocarmine slides
permanent (CO2, dry ice, acid-alcohol transfer, etc.) but the simplest procedure is to ring one side of the cover glass
with Hoyer's. Within a day or so the Hoyer's will replace the acetocarmine. When the Hoyer's solution is dry, the
excess can be carefully removed with water-dampened tissue, and the cover glass sealed with EUKITT.

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Figure 287. Chromosome squashing in ferns. (Drawn by F. A. Lang, 1983).

600
Recipes for Fixatives

There are three main solutions for fixing chromosomes: Farmer’s, Carnoy’s, and Newcomer’s solutions.

Farmer’s solution

ingredients amount
Glacial acetic acid 1 part
Absolute alcohol 3 parts

This is a good general fixative for all kinds of tissues, but it must be mixed in the field just before use (use plastic squirt
bottles). Fix for at last 15 minutes and up to 24 hours at room temperature. If the objects are to be stored, they can
be kept indefinitely in the solution in a freezer. For angiosperms (not ferns), if cold storage is unavailable, transfer to
70% alcohol.

Carnoy’s solution

ingredients amount
Glacial acetic acid 1 part
Absolute alcohol 6 parts
Chloroform 3 parts

A modification of the aceto-alcohol fixative and used the same way. Some cytologists prefer it because it hardens the
cells less and this makes it easier to spread the chromosomes. It also leaves the cytoplasm clearer and the specimens
can be kept in the solution longer. The problem using it is carrying chloroform in the field.

Newcomer’s solution

ingredients amount
Acetone 1 part
Dioxane (spectroquality) 1 part
Petroleum ether 1 part
Propionic acid 3 parts
Isopropyl alcohol 6 parts

This is the most stable fixative. For some material it is possible to get better chromosome spreading and somewhat
darker staining than in Farmer’s solution. The results are generally similar to material fixed in Farmer’s.

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Hoyer’s Solution

ingredients amount
Gum Arabic (USP flake, do not use powder) 60 g
Distilled water 100 ml
Chloral hydrate 400 g
Glycerin 40 ml

Dissolve the gum arabic in water for 24 hours. It goes into solution slowly. Flakes are best, although crystals will work.
Avoid powder because it is difficult to work with. Use any available mechanical means of mixing. The mixture might
seem to need filtering, but this is not necessary. After a few hours the bubbles will disappear and the solution will
clear. Add chloral hydrate. Let stand another 24 hours. Add glycerin. Do not use heat. Bubbles should disappear after
several hours.

When squashing and staining with ACH (Aceto-carmine Hoyer: 50% aceto-carmine, 50% Hoyer’s solution), the
stained chromosomes often become pale-colored because of the clearing effect of the chloral hydrate. For this
reason, the slide containing the squash is best viewed under a phase-contrast microscope. If only a non-phase light
microscope is available, it is best to squash in 100% aceto-carmine, although the resulting slide will not be permanent.
Its cover slip can be ringed with nail polish enamel to retard drying, but it will not be as durable as a squash prepared
in ACH, where Hoyer’s solution acts as a mounting medium.

For further information about the use of Hoyer's solution for mounting spores and leaf fragments see Anderson
(1954).

Literature cited

Anderson, L. W. 1954. Hoyer’s solution as rapid permanent mounting medium for bryophytes. Bryologist 57: 242–244.

602
Appendix 2 | Principal Pteridophyte Floras for the Neotropics

Mexico & Mesoamerica

Gómez, L. D. & A. L. Arbeláez. 2009. Helechos. In: W. D. Stevens, O. M. Montiel & A. Pool (eds.), Flora de Nicaragua. Vol. IV.
Monographis in Systematic Botaniy from the Missouri Botanical Garden 116: 1–348.
Knobloch, I. W. & D. S. Correll 1962. Ferns and fern allies of Chihuahua, Mexico. Contributions from the Texas Research
Foundation 3: 1–198.
Lellinger, D. B. 1989.The ferns and fern allies of Costa Rica, Panama, and the Chocó. (Part 1: Psilotaceae through Dicksoniaceae).
Pteridologia 2A: 5–364.
Mickel, J. T. 1992. Pteridophytes. In: R. McVaugh, editor, Flora Novo-Galiciana 17: 120–455.
Mickel, J. T. & J. Beitel 1988. Pteridophyte flora of Oaxaca, Mexico. Memoirs of the New York Botanical Garden 48: 1–568.
_____, & A. R. Smith. 2004. The pteridophytes of Mexico. Memoirs of the New York Botanical Garden 88: 1–1055.
Monterrosa S., J. A., M. del Carmen Peña-Chocarro, S. Knapp & R. Escobar L. 2009. Guía de identificación de helechos
de El Salvador. Jardín Botánico La Laguna y The Natural History Museum. La Libertad, El Salvador.
Moran, R. C. & R. Riba (editores.) 1995. Psilotaceae a Salviniaceae. In: G. Davidse, M. Sousa S. & S. Knapp (editores generales),
Flora Mesoamericana. Volumen 1. Univ. Nacional Autónoma de México, Ciudad Universitaria. [available on the web: http://
www.mobot.org/mobot/fm/ ]
Smith, A. R. 1981. Pteridophytes. In: D. E. Breedlove, editor. Flora of Chiapas 2: 1–370. California Academy of Science, San Fran-
cisco.
Stolze, R. G. 1976. Ferns and fern allies of Guatemala, part I. Ophioglossaceae through Cyatheaceae. Fieldiana, Botany 39: i–v
+ 1–130.
_____. 1981. Ferns and fern allies of Guatemala, part II. Polypodiaceae. Fieldiana, Botany, new series, 6: i–ix + 1–522.
_____. 1983. Ferns and fern allies of Guatemala, part III. Marsileaceae, Salviniaceae, and the fern allies. Fieldiana, Botany, new series,
12: i–iii + 1–91.

South America

Kramer, K. U. 1962. Flora of the Netherlands Antilles. Pteridophyta. In: A. L. Stoffers, editor. Uitgaven “Natuurwetenschappelijke
Studiekring voor Suriname en de Nederlandse Antillen,” Utrecht, no. 25. vol. 1: 1–84.
_____. 1978. The pteridophytes of Suriname. Uitgaven. Natuurwetenschappelijke Studiekring voor Suriname Nederlandse An-
tillen 93: 1–198.
Lellinger, D. B. 1989.The ferns and fern allies of Costa Rica, Panama, and the Chocó. (Part 1: Psilotaceae through Dicksoniaceae).
Pteridologia 2A: 5–364.
Lüer, H. G. 1984. Helechos de Chile. Monografías Anexas a Los Anales de la Universidad de Chile, Núm. 1.
Mickel, J. T. 1985. Trindad Pteridophytes. Published privately by the author, New York Botanical Garden.
Mori, S. A., G. Cremers, C. Gracie, J.-J. De Granville, M. Hoff, J. D. Mitchell. 1997. Guide to the vascular plants of Central
French Guiana. Part I. Pteridophytes, Gymnosperms, and Monocotyledons. Memoirs of the New York Botanical Garden 76:
1–422.
Navarrete, H. 2001. Helechos communes de la Amazonia baja ecuatoriana. Editorial Simbioe, Quito.
Prado, J., R. Y. Hirai & R. C. Moran. 2017. Fern and lycophyte flora of Acre State, Brazil. Biota Neotropica 17(4): [1–59].
e20170369. http://dx.doi.org/10.1590/1676-0611-BN-2017-0369
Rodríguez R., R. 1995. Pteridophyta. Pages 119–309. In: C. Marticorena & R. Rodríguez, Flora de Chile, Vol. I, Pteridophyta—
Gymnospermae. Universidad de Concepción, Chile.

603
Sehnem, A. 1967–1979. (18 familias de pteridofitas), In: R. Reitz, editor. Flora illustrada catarinense. Itajai, Santa Catarina.
Smith, A. R. 1985. Pteridophytes of Venezuela, an annotated list. Published privately by the author.
_____. 1992. 16. Thelypteridaceae. In: R. M. Tryon & R. G. Stolze, Pteridophyta of Peru. Fieldiana, Botany new series 29: iii + 1–80.
_____. 1995. Pteridophytes. In: Steyermark et al., general editors. Flora of the Venezuelan Guayana, volume 2. Pages 1–334. Tim-
ber Press, Portland, Oregon.
Sota, E. R. de la 1977. Pteridophyta. In: A. L. Cabrera, editor. Flora de la Provincia de Jujuy, República Argentina, parte 2: 1–275.
Colección Científica INTA, Buenos Aires.
Tryon, R. M. & R. G. Stolze. 1989. Pteridophyta of Peru, Part 1, 1. Ophioglossaceae-12. Cyatheaceae. Fieldiana, Botany new
series 20: iii + 1–145.
_____. 1989. Pteridophyta of Peru, Part II, 13. Pteridaceae-15. Dennstaedtiaceae. Fieldiana, Botany new series 22: iii + 1–128.
_____. 1991. Pteridophyta of Peru, Part IV, 17. Dryopteridaceae. Fieldiana, Botany new series 27: iii + 1–176.
_____.1993. Pteridophyta of Peru, Part V, 18. Aspleniaceae—21. Polypodiaceae. Fieldiana, Botany new series 27: iii + 1–176.
_____.1994. Pteridophyta of Peru, Part VI, 22. Marsileaceae—28. Isoetaceae. Fieldiana, Botany new series 32: iii + 1–190.
Vareschi, V. 1969. Helechos. Flora de Venezuela, 1(1 & 2), Edición Especial del Instituto Botánico, Caracas. 1033 Pages
Zuquim, G., F. R. C. Costa, J. Prado & H. Tuomisto. 2008. Guide to the Ferns and Lycophytes of REBIO Autuma, Central
Amazonia. Áttema Design Editorial, Manaus.

West Indies

Greuter, W. (ed). 2003. Aspleniaceae, Cyatheaceae, Cycadaceae, Zamiaceae. Fasículo 8. Flora de la República de Cuba. Serie A.
Plantas Vasculares. Koeltz Scientific Books. Königstein, Germany.
Greuter, W. (ed). 2006. [13 fern families by various authors]. Fasículo 11. Flora de la República de Cuba. Serie A. Plantas Vascu-
lares. Koeltz Scientific Books. Königstein, Germany
Proctor, G. R. 1977. Pteridophyta. In: R.A. Howard, editor, Flora of the Lesser Antilles, vol. 2: 1–414. Arnold Arboretum of Har-
vard University, Jamaica Plain, Massachusetts.
_____. 1985. Ferns of Jamaica. British Museum (Natural History), London.
_____. 1989. Ferns of Puerto Rico and the Virgin Islands. Memoirs of the New York Botanical Garden 53: 1–389.
Sánchez, C. 2000. Hymenophyllaceae Fasículo 4. In: W. Greuter, ed. Flora de la República de Cuba. Serie A. Plantas Vasculares.
Koeltz Scientific Books. Königstein, Germany.

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Appendix 3 | Families and Genera of American Pteridophytes

LYCOPHYTES Dicksonia
1. Lycopodiaceae 10. Matoniaceae Lophosoria
Dendrolycopodium (Old World)
Diphasium 25. Cyatheaceae
Diphasiastrum 11. Dipteridaceae Alsophila
Huperzia (Old World) Cyathea
Lycopodiella Gymnosphaera
Lycopodium 12. Gleicheniaceae Sphaeropteris
Palhinhaea Dicranopteris
Phlegmariurus Diplopterygium 26. Saccolomataceae
Pseudolycopodiella Gleichenella Saccoloma
Spinulum Sticherus
27. Cystodiaceae
2. Isoëtaceae 13. Lygodiaceae (Old World)
Isoëtes Lygodium
28. Lonchitidaceae
3. Selaginellaceae 14. Schizaeaceae Lonchitis
Selaginella Actinostachys
Schizaea 29. Lindsaeaceae
Lindsaea
FERNS 15. Anemiaceae Odontosoria
4. Equisetaceae Anemia Sphenomeris
Equisetum
16. Salviniaceae 30. Pteridaceae
5. Psilotaceae Azolla Acrostichum
Psilotum Salvinia Adiantopsis
Adiantum
6. Ophioglossaceae 17. Marsileaceae Allosorus
Botrychium Marsilea Ananthacorus
Botrypus Pilularia Anogramma
Cheiroglossa Regnellidium Antrophyum
Ophioglossum Argyrochosma
Sceptridium 18. Thyrsopteridaceae Aspidotis
Thyrsopteris Astrolepis
7. Marattiaceae Bommeria
Danaea 19. Loxsomataceae Ceratopteris
Eupodium Loxsomopsis Cheilanthes
Marattia Cheiloplecton
20. Culcitaceae Cryptogramma
8. Osmundaceae Culcita Doryopteris
Osmunda Gaga
Osmundastrum 21. Plagiogyriaceae Hecistopteris
Plagiogyria Hemionitis
9. Hymenophyllaceae Jamesonia
Abrodictyum 22. Cibotiaceae Llavea
Crepidomanes Cibotium Mildella
Didymoglossum Myriopteris
Hymenophyllum 23. Metaxyaceae Notholaena
Polyphyllum Metaxya Pellaea
Trichomanes Pentagramma
Vandenboschia 24. Dicksoniaceae Pityrogramma

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 Polytaenium Telmatoblechnum 47. Lomariopsidaceae
Pteris Woodwardia Cyclopeltis
Pterozonium Dracoglossum
Radiovittaria 41. Athyriaceae Lomariopsis
Scoliosorus Athyrium
Trachypteris Deparia 48. Tectariaceae
Tryonia Diplazium (incl. Callipteris) Arthropteris
Vittaria Ephemeropteris Hypoderris
Malaifilix
31. Dennstaedtiaceae 42. Thelypteridaceae Pterydrys
Blotiella Amauropelta Tectaria
Dennstaedtia Amblovenatum Triplophyllum
Histiopteris Christella
Hypolepis Coryphopteris 49. Oleandraceae
Microlepia Cyclosorus Oleandra
Paesia Goniopteris
Pteridium Macrothelypteris 50. Davalliaceae
Meniscium (Old World)
32. Cystopteridaceae Oreopteris
Cystopteris Parathelypteris 51a. Polypodiaceae
Gymnocarpium Phegopteris Adetogramma
Stegnogramma Campyloneurum
33. Rhachidosoraceae Steiropteris Loxogramme
(Old World) Thelypteris Microgramma
Niphidium
34. Diplaziopsidaceae 43. Didymochlaenaceae Pecluma
Homalsosorus Didymochlaena Phlebodium
Platycerium
35. Desmophlebiaceae 44. Hypodematiaceae Pleopeltis
Desmophlebium (Old World) Polypodium
Serpocaulon
36. Hemidictyaceae 45. Dryopteridaceae Synnamia
Hemidictyum Arachniodes
Atalopteris 51b. Grammitid clade
37. Aspleniaceae Bolbitis Alansmia
Asplenium Ctenitis Ascogrammitis
Cyclodium Ceradenia
38. Woodsiaceae Dryopteris Cochlidium
Woodsia Elaphoglossum Enterosora
Lastreopsis Galactodenia
39. Onocleaceae Lomagramma Grammitis
Matteuccia Maxonia Lellingeria
Onoclea Megalastrum Leucotrichum
Onocleopsis Mickelia Lomaphlebia
Olfersia Luisma
40. Blechnaceae Parapolystichum Melpomene
Anchistea Phanerophlebia Micropolypodium
Austroblechnum Polybotrya Moranopteris
Blechnum Polystichopsis Mycopteris
Cranfillia Polystichum Stenogrammitis
Lomaridium Rumohra Terpsichore
Lomariocycas Stigmatopteris Zygophlebia
Lorinseria
Parablechnum 46. Nephrolepidiaceae
Salpichlaena Nephrolepis
Struthiopteris

606
Appendix 4 | accepted genera of ferns and lycophytes world-wide
The genera of ferns and lycophytes and the families to which they belong. .The family numbers follow PPG1
(2016).

FAMILY NUMBER Thyrsopteridaceae (18) Hemidictyaceae (36)

Lycopodiaceae (1) Loxomataceae (19) Aspleniaceae (37)
Isoetaceae (2) Culcitaceae (20) Woodsiaceae (38)
Selaginellaceae (3) Plagiogyriaceae (21) Onocleaceae (39)
Equisetaceae (4) Cibotiaceae (22) Blechnaceae (40)
Psilotaceae (5) Metaxyaceae (23) Athyriaceae (41)
Ophioglossaceae (6) Dicksoniaceae (24) Thelypteridaceae (42)
Marattiaceae (7) Cyatheaceae (25) Didymochlaenaceae (43)
Osmundaceae (8) Saccolomataceae (26) Hypodematiaceae (44)
Hymenophyllaceae (9) Cystodiaceae (27) Dryopteridaceae (45)
Matoniaceae (10) Lonchitidaceae (28) Nephrolepidiaceae (46)
Dipteridaceae (11) Lindsaeaceae (29) Lomariopsidaceae (47)
Gleicheniaceae (12) Pteridaceae (30) Tectariaceae (48)
Lygodiaceae (13) Dennstaedtiaceae (31) Oleandraceae (49)
Schizaeaceae (14) Cystopteridaceae (32) Davalliaceae (50)
Anemiaceae (15) Rhachidosoraceae (33) Polypodiaceae (51)
Salviniaceae (16) Diplaziopsidaceae (34)
Marsileaceae (17) Desmophlebiaceae (35)

GENUS (FAMILY NUMBER) Anetium = Polytaeniuim (30) Blechnum (40)

Abrodictyum (9) Angiopteris (7) Blotiella (31)
Aconiopteris = Elaphoglossum Anisocampium (41) Bolbitis (45)
Acropelta = Polystichum Anisogonium = Diplazium Bommeria (30)
Acrophorus (45) Anogramma (30) Botrychium (6)
Acrorumohra = Dryopteris (45) Anopteris = Pteris Botrypus (6)
Acrosorus (51) Antigramma = Asplenium Brainea (40)
Acrostichum (30) Antrophyopsis (30) Bryodesma = Selaginella
Actiniopteris (30) Antrophyum (30) Calciphilopteris (30)
Actinostachys (14) Arachniodes (45) Callipteris = Diplazium
Acystopteris (41) Araiostegia (50) Callistopteris (9)
Adenoderris = Polystichum Araiostegiella (50) Calciphilopteris (30)
Adenophorus (51) Archangiopteris = Angiopteris Calochlaena (24)
Adetogramma (51) Archigrammitis (51) Calymmodon (51)
Adiantopsis (30) Argyrochosma (30) Campium = Bolbitis
Adiantum (30) Arthrobotrya (45) Camptodium = Tectaria
Aenigmopteris (45) Arthrobotrya = Dryopteris (45) Campyloneurum (51)
Afropteris = Pteris Arthromeris (51) Caobangia = Lemmaphyllum
Aglaomorpha (51) Arthropteris (48) Cardiomanes = Hymenophyllum
Alansmia (51) Ascogrammitis (51) Cassebeera (30) = Doryopteris
Aleuritopteris (30) Aspidium = Tectaria Cephalomanes (9)
Allantodia = Diplazium Aspidotis (30) Ceradenia (51)
Allosorus (30) Aspleniopsis = Austrogramme Ceratopteris (30)
Alsophila (25) Asplenium (312) Cerosora (30)
Amauropelta (34) xAsplenosorus = Asplenium Ceterach = Asplenium
Amblia = Phanerophlebia Astrolepis (30) Ceterachopsis = Asplenium
Amblovenatum (34) Atalopteris (45) Cheilanthes (30)
Ampelopteris (34) Ataxipteris = Ctenitis Cheilanthopsis = Woodsia
Amphiblestra = Tectaria Athyriopsis = Deparia Cheilolepton = Lomagramma
Amphineuron (34) Athyrium (41) Cheiloplecton = Notholaena (30)
Amphoradenium = Adenophorus Austroblechnum (40) Cheiroglossa (6)
Ananthacorus (30) Austrogramme (30) Cheiropleuria (11)
Anapausia = Bolbitis Azolla (16) Chieniopteris = Woodwardia
Anarthropteris = Loxogramme Belvisia = Lepisorus Chingia (34)
Anchistea (40) Blechnidium (40) Chlamidogramme = Tectaria
Anemia (15) Blechnopsis (40) Christella (34)

607
Christensenia (7)
Christiopteris = Aglaomorpha
Chrysogrammitis (51)
Cibotium (22)
Cionidium = Tectaria
Cleistoblechnum (40)
Cnemidaria = Cyathea
Cochlidium (51)
Colysis (51)
Coniogramme (30)
Coptodipteris (31)
xCornoathyrium = Athyrium
Cornopteris (41)
Coryphopteris (34)
Cosentinia (30)
Costaricia = Dennstaedtia
Coveniella = Lastreopsis
Cranfilia (40)
Crepidomanes (9)
Crypsinus = Selliguea
Cryptogramma (30)
Cryptosorus = Prosaptia
Ctenitis (45)
Ctenitopsis = Tectaria
Ctenopterella (51)
Ctenopteris (51)
Culcita (20)
Currania = Gymnocarpium
Cyathea (25)
Cyclodium (45)
Cyclogramma (34)
Cyclopeltis (47)
Cyclosorus (34)
Cyrtogonellum = Polystichum
Cyrtogonium = Bolbitis
Cyrtomidictyum = Polystichum
Cyrtomium (45)
Cystoathyrium = Cystopteris
Cystodium (23)
Cystopteris (32)
Danaea (7)
Dasygrammitis (51)
Davallia (50)
Davallodes (50)
Dendroconche (51)
Dendrolycopodium (1)
Dennstaedtia (31)
Deparia (41)
xDepazium = Deparia
Desmophlebia (35)
Diacalpe = Dryopteris
Dichasium = Dryopteris
Dicksonia (24)
Diclisodon Dryopteris (45)
Dicranoglossum = Pleopeltis
Dicranopteris (12)
Dictymia (51)
Dictyodroma = Deparia
Dictyoglossa = Elaphoglossum
Dictyoxiphium = Tectaria
Didymochlaena (43)
Didymoglossum (9)
Diellia = Asplenium
Diphasium (1)
Diphasiastrum (1)
Diplaziopsis (34)
Diplazium (41)
Diploblechnum (40)
Diplopterygium (12)
Diplora = Asplelnium
Dipteris (11)
Doodia (40)
Dorcapteris = Olfersia
Doryopteris (30)
Dracoglossum (47)
Draconopteris (48)
Drymoglossum = Pyrrosia
Drymotaenium = Lepisorus
Drynaria = Agloamorpha
Dryoathyrium = Deparia
xDryopolystichum (45)
xDryostichum (45)
Dryopsis = Dryopteris
Dryopteris (45)
Edanyoa = Bolbitis
Egenolfia = Bolbitis
Elaphoglossum (45)
Enterosora (51)
Ephemeropteris (41)
Equisetum (4)
Eriosoriopsis = Woodsia
Eriosorus = Jamesonia
Eupodium (7)
Fadyenia = Tectaria (48)
Filix = Dryopteris (45)
Fourniera = Sphaeropteris
Gaga (30)
Galactodenia (51)
Glaphyropteridopsis (34)
Gleichenella (12)
Gleichenia (12)
Goniophlebium (51)
Goniopteris (34)
Grammitis (51)
Gymnocarpium (32)
Gymnogramma = Hemionitis
Gymnopteris = Hemionitis
Gymnogrammitis (51)
Gymnosphaera (25)
Hemesteum = Polystichum
Haplopteris (30)
Hecistopteris (30)
Helminthostachys (6)
Hemidictyum (39)
Hemigramma = Tectaria
Hemionanthes (30)
Hemionitis (30)
Hemitelia = Cyathea
Heterogonium = Tectaria
Heteroneuron = Bolbitis
Hewardia = Adiantum
Hicriopteris = Dicranopteris
Himalayopteris = Sillegua
Hippochaete = Equisetum
Histiopteris (31)
Holcochlaena (30)
608
Holodictyum = Asplenium
Holostachyum = Aglaomorpha
Homalosorus (34)
Hovenkampia (51)
Humata = Davallia
Huperzia (1)
Hyalotrichopteris = Campyloneurum
Hymenasplenium (312)
Hymenocystis = Woodsia
Hymenodium = Elaphoglossum
Hymenoglossum = Hymen-ophyllum
Hymenophyllopsis = Cyathea
Hymenophyllum (9)
Hypodematium (43)
Hypoderris (48)
Hypolepis (31)
Hypopeltis = Polystichum
Icarus (40)
Ithycaulon = Saccoloma
Isoëtes (2)
Isoloma = Lindsaea
Jamesonia (30)
Japanobotrychium (6)
Jenkinsia = Bolbitis
Kaulinia (51)
Kontumia (51)
Kuniwatsukia = Athyrium
Lastreopsis (45)
Lecanopteris (51)
Lellingeria (51)
Lemmaphyllum (51)
Lepidomicrosorum (51)
Lepisorus (51)
Leptochilus (51)
Leptolepia (31)
Leptopteris (8)
Leptorumohra = Arachniodes
Leucostegia (43)
Leucotrichum (51)
Lindsaea (23)
xLindsaeosoria (23)
Lithostegia = Arachniodes
Litobrochia = Pteris
Llavea (30)
Lomagramma (45)
Lomaphlebia (51)
Lomaria (40)
Lomaridium (40)
Lomariocycas (40)
Lomariopsis (47)
Lonchitis (23)
Lophodium = Dryopteris (45)
Lophosoria (24)
Lorinseria (40)
Loxogramme (51)
Loxoma (19)
Loxoscaphe = Asplenium
Loxsomopsis (19)
Luisma (51)
Lunathyrium = Deparia
Lycopodiella (1)
Lycopodium (1)
Lygodium (13)
Lytoneuron (30) Onocleopsis (39) Pleurosorus = Asplenium
Macrothelypteris (34) Onychium (30) Pneumatopteris (34)
Malaifilix (48) Ophioderma (6) Podopeltis = Tectaria
Mankyua (6) Ophioglossum (6) Podosorus (51)
Marattia (7) Oreogrammitis (51) Poecilopteris = Bolbitis
Marsilea (17) Oreopteris (34) Polybotrya (45)
Matonia (10) Ormoloma = Lindsaea Polyphlebium (9)
Matteuccia (39) Ormopteris (30) Polypodiastrum = Goniophlebium
Maxonia (45) Ornithopteris = Anemia (15) Polypodioides = Goniophlebium
Mecodium = Hymenophyllum Orthiopteris = Saccoloma Polypodiopteris (51)
Megalastrum (45) Osmolindsaea (23) Polypodium (51)
Melpomene (51) Osmunda (8) Polystichopsis (45)
Meniscium (34) Osmundastrum (8) Polystichum (45)
Menisorus (34) Osmundopteris (6) Polytaenium (30)
Merinthosorus = Aglaomorpha Pachypleuria (50) Pradoia (45)
Mesophlebion = Cyclosorus Paesia (31) Pronephrium (34)
Metapolypodium = Goniopteris Palhinhaea (1) Prosaptia (51)
Metathelypteris (34) Papuapteris = Polystichum Protowoodsia = Woodsia
Metaxya (23) Parablechnum (40) Psammiosorus (48)
Mickelia (45) Parathyrium = Deparia Pseudocolysis = Pleopeltis
Microgonium = Didymoglossum Paraceterach (30) Pseudocyclosorus (34)
Microgramma (51) Paragramma (51) Pseudocystopteris = Athyrium
Microlepia (31) Paragymnopteris (30) Pseudodrynaria = Aglaomorpha
Micropolypodium (51) Parahemionitis (30) Pseudolycopodiella (1)
Micropteris = Cochlidium Parapolystichum (45) Pseudophegopteris (34)
Microsorum (51) Paraselliguea (51) Pseudotectaria = Ctenitis
Microstaphyla = Elaphoglossum Parkeria = Ceratopteris Psilotum (5)
Microtrichomanes = Hymenophyllum Parathelypteris (34) Psomiocarpa = Tectaria
Mildella (30) Pecluma (51) Pteridium (31)
Mohria = Anemia Pelazoneuron = Cyclosorus Pteridoblechnum (40)
Monachosorum (31) Pellaea (30) Pteridrys (48)
Monogramma = Haplopteris (30) Peltapteris = Elaphoglossum Pteris (30)
Monomelangium = Diplazium Pentagramma (30) Pterozonium (30)
Moranopteris (51) Pentarhizidium (39) Ptilopteris = Polystichum
Mycopteris (51) Peranema = Dryopteris Ptisana (7)
Myriopteris (30) Pessopteris = Niphidium Pycnodoria = Pteris
Nannothelypteris (34) Phanerophlebia (45) Pycnopteris = Dryopteris
Nematopteris = Scleroglossum Phanerophlebiopsis = Arachniodes Pyrrosia (51)
Neoathyrium = Athyrium Phanerosorus (10) Quercifilix = Tectaria
Neocheiropteris (51) Phegopteris (34) Radiogrammitis (51)
Neolepisorus (51) Phlebodium (51) Radiovittaria (30)
Neolindsaea (23) Phlegmariurus (1) Regnellidium (17)
Nephelea = Alsophila Photinopteris = Aglaomorpha Revwattsia = Dryopteris
Nephrodium = Dryopteris Phyllitis = Asplenium Rhachidosorus (312c)
Nephopteris = Jamesonia Phymatopteris = Sillegua Rheopteris (30)
Neoblechnum (40) Phymatosorus (51) Rhipidopteris = Elaphoglossum
Nephrolepis (49) Physematium = Woodsia Rhizoglossum (6)
Neurocallis = Pteris Pichisermollodes = Silleguea Rosenstockia = Hymenophyllum
Neurodium = Pleopeltis Pilularia (17) Rumohra (45)
Niphidium (51) Pityrogramma (30) Saccoloma (29)
Niphobolus = Pyrrosia Plagiogyria (21) Sadleria (40)
Notholaena (30) Platycerium (51) Salpichlaena (40)
Nothoperanema = Dryopteris Platygyria = Lepisorus Salvinia (16)
Notogrammitis (51) Platyloma = Pellaea Sceptridium (6)
Oceanopteris (40) Platyzoma = Pteris Schaffneria = Asplenium
Ochropteris = Pteris Plecosorus = Polystichum Schizaea (14)
Odontosoria (23) Plectopteris = Calymnodon Scleroglossum (30)
Oenotrichia (31) Pleocnemia (48) Scoliosorus (30)
Oenotrichia p.p (45) Pleopeltis (51) Scyphularia = Davallia
Oleandra (49) Plesioneuron (34) Selaginella (3)
Oleandropsis (51) Pleuroderris = Tectaria Selliguea (51)
Olfersia (45) Pleurogramme = Cochlidiuim Serpocaulon (51)
Onoclea (39) Pleurosoriopsis (51) Serpyllopsis = Hymenophyllum

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Sinephropteris = Asplenium
Sinopteris = Aleuritopteris
Solanopteris = Microgramma
Sorolepidium = Polystichum
Soromanes = Polybotrya
Sphaerocionium =Hymenophyllum
Sphaeropteris (25)
Sphaerostephanos (34)
Sphenomeris (23)
Spinulum (1)
Stenisioblechnum = Blechnum
Stegnogramma (34)
Steiropteris (34)
Stenochlaena (40)
Stenogrammitis (51)
Stenolepia (45)
Stenosemia = Tectaria
Sticherus (12)
Stigmatopteris (45)
Stromatopteris (12)
Struthiopteris (40)
Stylites (2)
Synammia (51)
Syngramma (30)
Taenitis (30)
Tapeinidium (23)
Tectaria (48)
Tectaridium = Tectaria
Teratophyllum (45)
Terpsichore (51)
Thamnopteris = Asplenium
Thelypteris (34)
Themelium (51)
Thylacopteris (51)
Thyrsopteris (18)
Thysanosoria (47)
Tmesipteris (5)
Todea (8)
Telmatoblechnum (40)
Thysanosoria (47)
Tomophyllum (51)
Trachypteris (30)
Triblemna = Deparia
Trichipteris = Cyathea
Tricholepidium (51)
Trichomanes (9)
Trigonospora (34)
Triplophyllum (48)
Trismeria = Pityrogramma
Tryonia (30)
Urostachys = Phlegmariurus
Vaginularia (30)
Vandenboschia (9)
Vittaria (30)
Wagneriopteris (51)
Wibellia (50)
Woodsia (311)
Woodsiopsis = Woodsia
Woodwardia (40)
Xiphopterella (51)
Xiphopteris = Cochlidium
Xyropteris (23)
Zygophlebia (51)

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Appendix 5 | A phylogeny for 400 species of leptosporangiate ferns
taken from: Schuettpelz, E. H. & K. M. Pryer. 2007. Phylogeny of ferns inferred from 400 leptosporangiate
species and three plastid genes. Taxon 56: 1037–1050.

(Figure 289)

Figure 288. Leptosporangiate fern phylogeny resulting from maximum likelihood analysis of plastid rbcL, atpB, and atpA data,
presented both as a phylogram (left) to reveal branch lengths and a cladogram (right) to clarify relationships and allow for the
presentation of maximum likelihood bootstrap percentages (only percentages > 50 are given; if > 70%, branches are bolded;
* = 100%). Note that the five eusporangiate fern outgroups have been pruned. Major clades discussed in text are indicated
in circles on trees: co, core leptosporangiates; ff, filmy ferns; gl, gleichenioids; hf, heterosporous ferns; hy, hymenophylloids; le,
leptosporangiates; of, osmundaceous ferns; po, polypods; sc, scaly tree ferns; sh, schizaeoids; tf, tree ferns; tr, trichomanoids. Families
recognized in the most recent classification of extant ferns (Smith & al., 2006b) are indicated in boxes between trees: Ane,
Anemiaceae; Cib, Cibotiaceae; Cul, Culcitaceae; Cya, Cyatheaceae; Die, Dickso niaceae; Dip, Dipteridaceae; Gle, Gleicheniaceae;
Hym, Hymenophyllaceae; Lox, Loxomataceae; Lyg, Lygodiaceae; Mar, Marsileaceae; Mat, Matoniaceae; Met, Metaxyaceae; Osm,
Osmundaceae; Pia, Plagiogyriaceae; Sal, Salviniaceae; Sch, Schizaeaceae;Thy,Thyrsopteridaceae. Phylogeny continues in Figure 289.

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 (Figure 290)

Figure 289. Continued from Figure 288. Major clades discussed in text are indicated in circles on trees: ad, adiantoids; ce, cera
topteridoids; ch, cheilanthoids; cr, cryptogrammoids; de, dennstaedtioids; eu, eupolypods; li, lindsaeoids; pd, pteridoids; po, polypods;
pt, pteroids; vi, vittarioids. Families recognized in the most recent classification of extant ferns (Smith & al., 2006b) are indicated
in boxes between trees: Den, Dennstaedtiaceae; Lin, Lindsaeaceae; Pte, Pteridaceae; Sac, Saccolo mataceae. Phylogeny continues
in Figure 290.

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 (Figure 291)

Figure 290. Continued from Figure 289. Major clades discussed in text are indicated in circles on trees: as, asplenioids; at,
athyrioids; bl, blechnoids; cs, cyclosoroids; e1, eupolypods I; e2, eupolypods II; eu, eupolypods; on, onocleoids; th, thelypteroids.
Families recognized in the most recent classification of extant ferns (Smith & al., 2006b) are indicated in boxes between trees: Asp,
Aspleniaceae; Ble, Blechnaceae; Ono, Onocleaceae; The, Thelypteridaceae; Woo, Woodsia ceae. Phylogeny continues in Figure
291.

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 (Figure 292)

Figure 291. Continued from Figure 290. Major ciades dis cussed in text are indicated in circles on trees: dc, di morphic
climbers; dr, dry opteroids; e1, eupolypods I; fl, former lomariopsids. Family recognized in the most recent classification of extant
ferns (Smith & al, 2006b) is indicated in box between trees: Dry, Dryopteridaceae. Phylogeny continues in Figure 292.

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Figure 292. Continued from Figure 291. Major clades discussed in text are indicated in circles on trees: da, davallioids; gr, gram
mitids; pg, polygrammoids; te, tectarioids. Families recognized in the most recent classification of extant ferns (Smith & al., 2006b)
are indicated in boxes between trees: Dav, Davallia ceae; Lorn, Lomariopsidaceae; Ole, Oleandraceae; Pol, Polypodiaceae; Tec,
Tectariaceae.

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Appendix 6 | Petioles in cross section
More patterns exist in nature but these are the most common found. In order to observe these patterns you would need
to produce a clean cut of the base of the petiole (using a sharped knife or scissor). When faced to an unknown Fern, it is a
good practice to produce first a clean cut of the petiole base and look for the vascular pattern.

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 We steal as in a castle, cocksure;
we have the receipt of fern-seed,
we walk invisible.

Gadshill to Chamberlain
Henry IV, Part 1, Act II, Scene 1
William Shakespeare, 1564-1616

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