Neotropical Genera

of Ferns and Lycophytes

a guide for students
Robbin C. Moran
The New York Botanical Garden

Tropical Plant Systematics

Organization for Tropical Studies
San José, Costa Rica, June-July 2014
 Neotropical Genera
of Ferns and Lycophytes
A guide for students

Robbin C. Moran
The New York Botanical Garden
 Neotropical Genera
of Ferns and Lycophytes
A guide for students

Robbin C. Moran
The New York Botanical Garden

Prepared for Tropical Plants Systematics, OTS 2014-09

Not for sale or redistribution

2
 Phylogeny of the three main groups of vascular plants (Smith et al., 2006).

cover page, Pleopeltis minima

previous page, left to right: Serpocaulon levigatum, Hemionitis palmata y Anemia oblongifolia.

3
 A Phylogeny of the families of ferns.
Interactive key to the neotropical genera of ferns and lycophytes available at
www.plantsystematics.org

4
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).

Petiole vasculature can be extremely useful for identification purposes when faced to an unknown fern.
Refer to Appendix 6 (page 531) for common vascular patterns in ferns.

5
this page has been intentionally left blank

6
 About the author

Robbin C. Moran
Dr. Robbin Moran is the Mary Flagler Cary Curator of Botany at the New
York Botanical Garden. He studies ferns and lycophytes and has published
four books and over 100 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.

7
 Contents
Index to genera.......................................................................................................................................................................... 12
page numbers in bold face indicate genera main entries............................................................................................................................12
Acknowledgements .................................................................................................................................................................. 15
Introduction to Ferns and Lycophytes ................................................................................................................................. 16
The Pteridophyte Life Cycle.................................................................................................................................................... 19
The typical life cycle ........................................................................................................................................................................................... 19
Observing gametangia (antheridia and archegonia)......................................................................................................................... 21
Antheridiogens....................................................................................................................................................................................................... 23
Apogamy .................................................................................................................................................................................................................. 25
Hybridization .......................................................................................................................................................................................................... 28
Branching patterns in Ferns & Lycophytes ............................................................................................................................................ 31
Lycophylls versus Euphylls (or Microphylls versus Megaphylls).................................................................................................. 36
Eusporangia versus Leptosporangia........................................................................................................................................................... 38
Spores ........................................................................................................................................................................................................................ 40
Sterile-fertile leaf dimorphism in ferns .................................................................................................................................................... 45
Tips on Collecting Ferns................................................................................................................................................................................... 47
Overview of land plant phylogeny ......................................................................................................................................... 48
Lycophytes .................................................................................................................................................................................. 50
Lycopodiaceae Mirbel | Club Moss Family............................................................................................................................................. 52
Selaginellaceae Willkomm | Spike-moss Family.................................................................................................................................... 57
Isoëtaceae Reichenbach | Quillwort Family........................................................................................................................................... 62
Ferns............................................................................................................................................................................................. 64
Ophioglossales................................................................................................................................................................................................................66
Ophioglossaceae Agardh | Adder’s-tongue Family............................................................................................................................ 66
Psilotales.............................................................................................................................................................................................................................70
Psilotaceae Kanitz | (Whisk Fern Family)................................................................................................................................................. 70
Equisetales.........................................................................................................................................................................................................................72
Equisetaceae Michx. ex DC. | Horsetail Family.................................................................................................................................... 72
Marattiales.........................................................................................................................................................................................................................75
Marattiaceae Bercht. & J. S. Presl | Marattia Family............................................................................................................................. 75
Leptosporangiate Ferns............................................................................................................................................................ 78
Osmundales......................................................................................................................................................................................................................78
Osmundaceae Bercht. & J. C. Presl | Royal Fern Family.................................................................................................................. 78
Hymenophyllales............................................................................................................................................................................................................82
Hymenophyllaceae Link | Filmy Fern Family.......................................................................................................................................... 82
Gleicheniales....................................................................................................................................................................................................................88
Gleicheniaceae (R. Brown) C. Pres l Forked Fern Family.............................................................................................................. 89

8
 Dipteridaceae Diels | Dipteris family......................................................................................................................................................... 91
Matoniaceae C. Presl | Matonia family...................................................................................................................................................... 92
Schizaeales A. B. Frank in Leunis...........................................................................................................................................................................93
Lygodiaceae M. Roem. | Climbing Fern Family.................................................................................................................................... 93
Anemiaceae Link................................................................................................................................................................................................... 94
Schizaeaceae Kaulf. | Curly Grass Family.................................................................................................................................................. 96
Salviniales | Water Ferns............................................................................................................................................................................................97
Marsileaceae Mirbel | Clover Fern Family............................................................................................................................................... 98
Salviniaceae Dumortier...................................................................................................................................................................................100
Cyatheales A. B. Frank in Leunis | Tree Fern Order................................................................................................................................102
Thyrsopteridaceae C. Presl...........................................................................................................................................................................104
Loxomataceae C. Presl....................................................................................................................................................................................104
Culcitaceae Korall...............................................................................................................................................................................................105
Plagiogyriaceae Bower.....................................................................................................................................................................................105
Cibotiaceae (Nayar) Korall............................................................................................................................................................................107
Cyatheaceae Kaulf. | Scaly Tree Fern Family.........................................................................................................................................107
Dicksoniaceae Bower.......................................................................................................................................................................................111
Metaxyaceae Pic. Serm.....................................................................................................................................................................................112
Polypodiales...................................................................................................................................................................................................................114
Lindsaeaceae Pic. Serm....................................................................................................................................................................................114
Saccolomataceae Doweld..............................................................................................................................................................................115
Pteridaceae Reichenbach | Maidenhair Family...................................................................................................................................117
Eupolypods.....................................................................................................................................................................................................................124
EUPOLYPODS I.........................................................................................................................................................................................................125
Hypodematiaceae Ching................................................................................................................................................................................125
Dryopteridaceae Herter | Wood Fern Family...................................................................................................................................125
Lomariopsidaceae Alston...............................................................................................................................................................................128
Tectariaceae Lellinger........................................................................................................................................................................................129
Oleandraceae (J. Sm.) Ching ex Pichi Serm.........................................................................................................................................131
Davalliaceae Mett. ex Frank | Rabbit’s foot Fern Family................................................................................................................133
Polypodiaceae Berchtold & J. Presl | Polypody Family....................................................................................................................133
Eupolypods II.................................................................................................................................................................................................................137
Cystopteridaceae (Payer) Schmakov.......................................................................................................................................................137
Aspleniaceae Newman | Spleenwort Family......................................................................................................................................139
Hemidictyaceae Christenh. & Schneid.....................................................................................................................................................140
Diplaziopsidaceae X. C. Zhang & Christenh........................................................................................................................................140
Rhachidosoraceae X. C. Zhang...................................................................................................................................................................141
Woodsiaceae Herter........................................................................................................................................................................................141
Athyriaceae Alston | Lady ferns..................................................................................................................................................................142
Thelypteridaceae Ching ex Pichi Sermolli | Beech Ferns, Marsh Fern Family..................................................................142

9
 Blechnaceae Newman | Chain Fern Family; Deer Ferns..............................................................................................................147
Appendices................................................................................................................................................................................516
Appendix 1 | Squash Technique for Fern and Lycophyte Chromosomes..........................................................................516
Appendix 2 | Principal Fern and Lycophyte Floras for the Neotropics..............................................................................519
Appendix 3 | Families and Genera of Neotropical Pteridophytes.........................................................................................521
Appendix 4 | accepted genera of ferns world-wide......................................................................................................................523
Appendix 5 | A phylogeny for 400 species of leptosporangiate ferns.................................................................................526
Appendix 6 | Petioles in cross section....................................................................................................................................................531

10
 Index to Keys
Key to Families of Lycopodiopsida............................................................................................................................................................................51
Key to the main genera of Lycopodiaceae........................................................................................................................................................... 52
Key to the main genera of Ophioglossaceae...................................................................................................................................................... 67
Key to the genera of Psilotaceae................................................................................................................................................................................70
Key to the Neotropical Genera of Marattiaceae.............................................................................................................................................. 77
Key to the Genera of Osmundaceae....................................................................................................................................................................... 79
Key to the traditional Genera of Hymenophyllaceae..................................................................................................................................... 82
Key to the main groups of Hymenophyllaceae in the Americas.............................................................................................................. 83
Key to the Genera of Gleicheniaceae..................................................................................................................................................................... 89
Key to the Genera of Matoniaceae.......................................................................................................................................................................... 92
Key to Main Taxa of Schizaeales..................................................................................................................................................................................93
Key to the Genera of Marsileaceae.......................................................................................................................................................................... 99
Key to the Genera of Salviniaceae..........................................................................................................................................................................101
Key to the Families of Cyatheales............................................................................................................................................................................102
Key to the genera of Loxomataceae......................................................................................................................................................................105
Key to Genera of Cyatheaceae.................................................................................................................................................................................108
Key to Genera of non-scaly Cyatheales...............................................................................................................................................................112
Key to Major Neotropical Genera of Lindsaeaceae......................................................................................................................................115
Key to the Major Neotropical Genera of Dennstaedtiaceae..................................................................................................................116
Key to Major Genera of Vittarioid Ferns (Pteridaceae)..............................................................................................................................119
Key to the Neotropical Genera of Lomariopsidaceae................................................................................................................................129
Key to the Neotropical Genera of Tectariaceae..............................................................................................................................................130
Key to the Genera of Neotropical Polypodiaceae.........................................................................................................................................135
Key to the American Genera and Subgenera of Thelypteridaceae......................................................................................................144
Key to the Neotropical Genera of Blechnaceae.............................................................................................................................................147
Key to the American Genera of Onocleaceae.................................................................................................................................................149

11
 Index to genera
page numbers in bold face indicate genera main entries

Index 522, 523, 524 Dennstaedtia 33, 116, 117, 240, 241, 316, 452,
A Bommeria 18, 23, 24, 26, 27, 44, 119, 204, 205, 521, 523, 524

Abrodictyum 82, 83, 84, 86, 87, 154, 155, 494,

258, 302, 521, 523 Dicksonia 108, 111, 112, 116, 226, 228, 242, 243,
523 Botrychium 44, 66, 67, 69, 76, 170, 206, 521, 523, 336, 490, 521, 524

Acrostichum 42, 96, 117, 118, 128, 141, 156, 157,

524, 525 Dicranoglossum 380, 422, 423, 522, 524
212, 521, 523 C Dicranopteris 89, 244, 245, 256, 288, 468, 521,
Actinostachys 93, 96, 158, 159, 458, 521, 523 Campyloneurum 134, 135, 208, 209, 262, 264, 524

Adiantopsis 119, 160, 161, 162, 214, 521, 523

342, 368, 380, 382, 508, 522, 523, 524 Didymochlaena 125, 246, 247, 522, 524
Ceradenia 134, 136, 210, 211, 218, 276, 290, 374, Didymoglossum 82, 83, 84, 85, 86, 87, 220, 248,
Adiantum 18, 30, 33, 42, 114, 117, 118, 123, 160,
514, 522, 523 249, 250, 251, 426, 524
162, 163, 328, 448, 521, 523, 524
Ceratopteris 23, 24, 117, 118, 156, 212, 213, 521, D. sect. Didymoglossum 248, 250
Alansmia 134, 136, 164, 165, 286, 326, 374, 476,
523, 525
522, 523 D. sect. Microgonium 248, 250
Cheilanthes 18, 25, 26, 30, 117, 119, 160, 174,
Alsophila 107, 108, 166, 167, 228, 242, 294, 336, Diplazium 139, 142, 182, 186, 190, 252, 253, 254,
178, 214, 215, 284, 300, 376, 384, 406, 492, 510,
464, 521, 523, 524 255, 298, 306, 522, 523, 524
521, 523
Ananthocorus 118, 168 D. Callipteris clade 254
Cheiroglossa 66, 67, 396, 523
Anemia 3, 23, 24, 42, 93, 94, 170, 171, 206, 396, Diplopterygium 88, 89, 244, 256, 257, 521, 524
Cibotium 107, 111, 112, 216, 217, 226, 336, 521,
521, 523, 524
523 Doryopteris 117, 119, 204, 258, 259, 492, 521,
Anetium 117, 118, 119, 172, 173, 436, 521, 523 523, 524
Cochlidium 125, 134, 136, 210, 218, 219, 290,
Anogramma 42, 117, 118, 174, 175, 222, 280, 324, 342, 522, 523, 525 Dryopteris 17, 23, 25, 26, 30, 32, 124, 125, 126,
416, 521, 523 143, 176, 224, 232, 260, 261, 322, 402, 432, 434,
Crepidomanes 82, 83, 84, 85, 86, 87, 220, 221,
Arachniodes 126, 176, 177, 232, 260, 322, 358, 438, 478, 502, 522, 524, 525
426, 523
432, 434, 438, 450, 502, 522, 523, 524, 525 E
C. subgen. Crepidomanes 220
Argyrochosma 18, 117, 119, 178, 179, 214, 376, Elaphoglossum 34, 43, 47, 126, 134, 172, 202, 208,
Cryptogramma 117, 118, 222, 223, 330, 521, 523
384, 406, 416, 521, 523 262, 263, 264, 265, 266, 267, 268, 270, 271, 272,
Ctenitis 126, 224, 225, 350, 360, 482, 502, 522, 273, 274, 275, 332, 366, 388, 508, 522, 524, 525
Ascogrammitis 134, 136, 164, 180, 181, 374, 522,
523
523 E. sect. Amygdalifolium 262
Culcita 104, 105, 106, 111, 112, 216, 226, 227,
Asplenium 17, 23, 24, 25, 26, 30, 40, 41, 43, 44, E. sect. Elaphoglossum 264
242, 336, 490, 521, 523
139, 140, 148, 182, 183, 184, 185, 186, 187, 190,
E. sect. Lepidoglossa 266, 268
252, 298, 522, 523, 524, 525 Cyathea 102, 107, 108, 109, 110, 228, 229, 230,
231, 242, 294, 310, 336, 464, 465, 521, 523, 524, E. sect. Polytricha 268
A. sect. Hymenasplenium 186, 187
525 E. sect. Setosa 270, 271
Loxoscaphe 139, 184, 185, 524
C. clade of Cnemidaria 230 E. sect. Squamipedia 272
Astrolepis 18, 26, 119, 178, 188, 189, 214, 406,
Hymenophyllopsis 108, 310, 311, 524 E. sect. Wrightiana 274
521, 523
Cyclodium 126, 232, 233, 358, 424, 432, 438, 470, Enterosora 134, 136, 210, 276, 277, 514, 522, 524
Athyrium 142, 190, 191, 306, 522, 523, 525
522, 523
Azolla 17, 18, 41, 97, 99, 100, 101, 192, 193, 354, Equisetum 17, 33, 40, 49, 64, 72, 73, 74, 278, 279,
Cyclopeltis 128, 129, 224, 234, 235, 378, 522, 523 521, 524
521, 523
Cyclosorus 143, 144, 478, 480, 481, 486, 488, Eriosorus 42, 117, 118, 280, 281, 320, 386, 504,
B
523, 524, 525 524
Blechnum 30, 33, 106, 110, 147, 194, 195, 196,
Cystopteris 24, 26, 30, 32, 137, 174, 222, 236, Eupodium 76, 77, 282, 283, 352, 521, 524
197, 198, 199, 418, 522, 523
237, 292, 510, 522, 524
B. “fragile group” 196 G
D
B. sect. Lomariocycas 198 Gaga 284, 285, 521, 524
Danaea 75, 76, 77, 238, 239, 282, 352, 364, 521,
Blotiella 116, 200, 201, 304, 334, 386, 521, 523 Galactodenia 134, 136, 286, 287, 522, 524
524
Bolbitis 43, 126, 202, 203, 314, 332, 366, 390, 474, Gleichenella 34, 89, 256, 288, 289, 468, 521, 524

12
Grammitis 134, 136, 210, 218, 290, 291, 324, 342, Loxsomopsis 104, 105, 112, 340, 521, 524 522, 524
466, 522, 524 Onocleopsis 149, 356, 392, 394, 395, 522, 524
Luisma 210, 218, 290, 342, 343, 522, 524
Gymnocarpium 17, 137, 292, 293, 522, 523, 524 Ophioglossum 44, 66, 67, 69, 93, 396, 397, 521,
Lycopodiella 52, 53, 55, 56, 308, 344, 345, 346,
Gymnosphaera 108, 166, 294, 295, 521, 524 521 523, 524

H Lycopodium 44, 52, 53, 55, 56, 57, 70, 308, 344, Cheiroglossa 66, 67, 396, 523
346, 347, 521 Osmunda 18, 78, 79, 80, 94, 398, 399, 400, 521,
Hecistopteris 117, 118, 119, 296, 297, 448, 521,
524 Lygodium 17, 30, 32, 42, 93, 94, 348, 349, 454, 524
521, 524 Osmundastrum 78, 79, 80, 398, 399, 400, 401,
Hemidictyum 139, 140, 298, 299, 522, 524

Hemionanthes 300, 301, 524

M 521

Hemionitis 3, 18, 30, 119, 204, 258, 300, 302, 303,

Macrothelypteris 143, 144, 148, 316, 350, 351, P
360, 410, 522, 524 Paltonium 380
446, 521, 524
T. sect. Macrothelypteris 350 Parapolystichum 126, 322, 323, 402, 403, 522,
Histiopteris 32, 33, 116, 200, 240, 304, 305, 334,
386, 521, 524 Marattia 75, 76, 77, 282, 352, 353, 521, 524 525

Homalosorus 140, 141, 148, 306, 524 Marsilea 41, 97, 98, 99, 100, 354, 355, 521, 524 Pecluma 26, 134, 135, 164, 180, 378, 404, 405,
412, 428, 430, 462, 476, 522, 525
Huperzia 18, 32, 42, 52, 53, 54, 55, 308, 309, 344, Matteuccia 27, 149, 151, 356, 357, 392, 394, 522,
346, 521 524 Pellaea 18, 25, 26, 27, 44, 117, 119, 123, 178, 258,
376, 406, 407, 521, 524, 525
Hymenophyllopsis 108, 310, 311, 524 Maxonia 126, 232, 358, 359, 424, 432, 438, 522,
524 Phanerophlebia 24, 126, 408, 409, 522, 525
Hymenophyllum 82, 83, 84, 88, 154, 310, 312,
313, 498, 506, 521, 523, 524, 525 Megalastrum 126, 143, 224, 322, 350, 360, 361, Phegopteris 32, 143, 144, 145, 350, 410, 411, 522,
402, 450, 502, 522, 524 525
Hypoderris 129, 130, 314, 315, 522, 524
Melpomene 134, 136, 180, 210, 286, 362, 363, Phlebodium 41, 134, 135, 412, 413, 428, 522, 525
Hypolepis 32, 33, 35, 116, 118, 200, 240, 304, 314,
372, 374, 522, 524 Pilularia 41, 97, 98, 99, 318, 414, 415, 521, 525
316, 317, 386, 521, 524
Metaxya 112, 113, 238, 364, 365, 521, 524
I Pityrogramma 24, 42, 118, 174, 280, 416, 417,
Mickelia 17, 43, 126, 202, 274, 366, 367, 522, 524 446, 521, 525
Isoëtes 16, 38, 40, 41, 50, 62, 63, 318, 319, 521
Microgramma 17, 33, 35, 134, 135, 208, 262, 264, Plagiogyria 104, 105, 106, 112, 194, 198, 226, 418,
Stylites 63, 318
338, 368, 369, 370, 380, 382, 388, 420, 522, 524, 419, 521, 525
J 525 Pleopeltis 134, 135, 208, 262, 264, 338, 368, 380,
Jamesonia 117, 118, 280, 320, 321, 446, 504, 521, M. subgen. Solanopteris 17, 134, 370 382, 420, 421, 422, 428, 462, 472, 522, 524, 525
524
Moranopteris 134, 136, 372, 373, 522, 524 Dicranoglossum 380, 422, 423, 522, 524
L
Mycopteris 180, 374, 375, 376, 522, 524 Neurodium 380, 381, 522, 524
Lastreopsis 126, 176, 224, 260, 322, 350, 358,
Myriopteris 117, 119, 131, 376, 377, 521, 524 Polybotrya 17, 43, 45, 46, 126, 232, 260, 332, 358,
360, 402, 432, 438, 450, 522, 523, 524
390, 424, 425, 432, 450, 502, 522, 525
Lellingeria 134, 136, 180, 210, 218, 286, 290, 324,
N
Polyphlebium 82, 83, 84, 85, 86, 87, 220, 248, 250,
325, 326, 342, 362, 372, 374, 466, 522, 524 Nephrolepis 128, 129, 234, 378, 379, 408, 522,
426, 427, 494, 500, 506, 525
524
Leucotrichum 134, 136, 164, 326, 327, 466, 522,
Polypodium 18, 33, 34, 107, 125, 133, 134, 135,
524 Neurodium 380, 381, 522, 524
137, 142, 194, 196, 218, 290, 368, 370, 378, 404,
Lindsaea 114, 115, 162, 246, 328, 329, 521, 524 Paltonium 380 412, 420, 428, 429, 430, 431, 462, 472, 522, 525
Llavea 117, 118, 222, 330, 331, 521, 524 Niphidium 135, 208, 368, 380, 382, 383, 522, 524, P. dulce Group 135, 430
525
Lomariopsis 43, 128, 129, 196, 234, 238, 332, 333, Polystichopsis 126, 232, 424, 432, 433, 522, 525
366, 424, 522, 524 Notholaena 18, 117, 119, 178, 188, 214, 384, 521,
Polystichum 19, 30, 126, 176, 260, 402, 408, 432,
524
Lonchitis 114, 115, 118, 200, 304, 334, 335, 521, 434, 435, 450, 522, 523, 524, 525
524 O
Polytaenium 117, 118, 119, 172, 338, 436, 437,
Lophosoria 107, 111, 112, 216, 336, 337, 521, Odontosoria 114, 115, 328, 386, 387, 521, 524 460, 521, 525
524
Oleandra 131, 132, 134, 388, 389, 522, 524 Pradopteris 438, 439
Loxogramme 134, 135, 136, 338, 339, 522, 523,
Olfersia 126, 232, 358, 390, 391, 424, 432, 522, Psilotum 30, 32, 35, 70, 71, 72, 78, 440, 441, 521,
524
524 525
Loxoscaphe 139, 184, 185, 524
Onoclea 89, 149, 151, 356, 392, 393, 394, 410,

13
Pteridium 18, 23, 24, 33, 116, 240, 442, 443, 521, Sticherus 89, 124, 244, 256, 288, 468, 469, 521, 133, 154, 220, 248, 250, 312, 340, 426, 494, 496,
525 525 497, 498, 499, 500, 506, 521, 525

Pteris 25, 26, 27, 30, 32, 33, 117, 118, 122, 156, Stigmatopteris 126, 143, 232, 470, 471, 522, 525 T. subgen. Davalliopsis 494
238, 442, 444, 445, 521, 523, 524, 525
Stylites 63, 318 T. subgen. Feea 83, 86, 87, 494, 496, 500
Pterozonium 117, 118, 446, 447, 504, 521, 525
Synammia 134, 472, 473, 525 T. subgen. Lacostea 83, 494, 498, 500
R T T. subgen. Trichomanes 500
Radiovittaria 117, 118, 119, 124, 168, 296, 448,
Tectaria 30, 129, 130, 202, 314, 366, 474, 475, Triplophyllum 129, 130, 224, 314, 432, 474, 502,
449, 460, 508, 521, 525
502, 512, 522, 523, 524, 525 503, 522, 525
Rumohra 126, 176, 358, 360, 402, 432, 434, 438,
Terpsichore 134, 136, 164, 180, 210, 286, 363, Tryonia 504, 505
450, 451, 522, 525
372, 374, 404, 476, 477, 522, 525
V
S Thelypteris 32, 124, 143, 144, 145, 350, 356, 400,
Vandenboschia 82, 84, 86, 87, 426, 494, 498, 500,
Saccoloma 115, 176, 240, 260, 340, 450, 452, 453, 478, 479, 480, 481, 482, 483, 484, 485, 486, 487,
506, 507, 525
521, 524, 525 488, 489, 512, 522, 523, 524, 525
Vittaria 23, 24, 117, 118, 119, 168, 436, 448, 460,
Salpichlaena 17, 34, 45, 94, 147, 348, 454, 455, Cyclosorus 143, 144, 478, 480, 481, 486, 488,
508, 509, 521, 525
522, 525 523, 524, 525
W
Salvinia 17, 18, 41, 97, 99, 100, 101, 414, 456, 457, T. sect. Macrothelypteris 350
521, 525 Woodsia 17, 141, 142, 174, 236, 510, 511, 522,
T. subgen. Amauropelta 144, 478, 488
524, 525
Schizaea 43, 93, 96, 158, 458, 459, 521, 525
T. subgen. Cyclosorus 144, 480
Woodwardia 24, 34, 147, 392, 512, 513, 522, 523,
Scoliosorus 117, 118, 119, 436, 460, 461, 521,
T. subgen. Goniopteris 144, 482 524, 525
525
T. subgen. Meniscium 144, 484, 485 Z
Serpocaulon 3, 134, 135, 412, 428, 462, 463, 484,
522, 525 T. subgen. Stegnogramma 144, 486 Zygophlebia 134, 136, 210, 276, 374, 514, 522,
525
Sphaeropteris 107, 108, 166, 228, 229, 242, 294, T. subgen. Steiropteris 144, 488
336, 464, 465, 521, 524, 525 Thyrsopteris 111, 112, 490, 491, 521, 525
Stenogrammitis 134, 136, 286, 324, 326, 466, 467, Trachypteris 119, 258, 492, 493, 521, 525
522, 525
Trichomanes 35, 82, 83, 84, 85, 86, 87, 88, 110,

14
 Acknowledgements
It’s humbling to sit down and tally your debts to others after compiling a work such as this. 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 partic-
ipated in my short pteridology courses taught in Costa Rica, Venezuela, Ecuador, and Bolivia, and 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 (there were seven). 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, Barry Hammel, Daniel Janzen, Barbara Lewis, David Neill, Benjamin Øllgaard,
and Nelson Zamora.

Many Latin American botanists have helped with the production of this book. Some revised the descriptions, and oth-
ers corrected my Spanish translations. For their interest, patience, and willingness to help, I thank Mercedes Asanza,
Mauricio Bonifacino, Marta Crisci, Jasivia González, Amanda Grusz, Myra Guzmán-Teare, Francisco Lorea, Blanca León,
Fernando Matos, Leticia Pacheco, Blanca Pérez-García, Katya Romoleroux, Ricardo Rueda, Carmen Ulloa Ulloa, Re-
nato Valencia, and Alejandra Vasco.

Illustrations are a vital part of this book, and I thank those people who have allowed me to use illustrations from their
works. John T. Mickel, my colleague and predecessor at the New York Botanical Garden, generously gave me per-
mission to use illustrations from his book with Alan R. Smith, Pteridophytes of Mexico (New York Botanical Garden
Press, 2004). Most of these illustrations were done by Mr. Haruto Fukuda. Also, 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 Pte-
ridophyta 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.

Robbin C. Moran
The New York Botanical Garden
15 May 2014

15
 Introduction to Ferns and Lycophytes
Why study ferns and lycophytes? There are several reasons, I think. 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.
In the tropics there is a more immediate reason to learn about them. Taken together, ferns and lycophytes number
about 12,000 species worldwide, with the vast majority 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 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 (Psi-
lotaceae); the lycophytes consist of club mosses (Lycopodiaceae), spike mosses (Selaginella), 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 (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. Never-
theless, 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 appear.

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

16
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 Dicksonia-
ceae), forked ferns (Gleicheniaceae), and dwarf polypodies (grammitids). Because the temperate zones harbor fewer
life forms and taxonomic groups, to get an overall view of ferns and lycophytes 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, elon-
gate 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.

17
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, Cheilan-
thes, Hemionitis, Notholaena, and Pellaea. Most of their species reproduce asexually by diploid spores—a mode of
reproduction called apogamy. The spores germinate, grow into a prothallus, and then vegetatively proliferate a new
sporophyte. 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 Polypodium 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 a rich source of nitrogen. Other economically
important ferns are the ornamentals, particulary Adiantum (maindenhair fern), Platycerium (staghorn fern), and Da-
vallia (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.

Some 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.

18
 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 (sing. sorus), which are
clusters of sporangia (sing. 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)

19
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
(sing. 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.

The main groups of vascular plants differ in how they modify their life cycles, and these modifications characterize
the main 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
contrast, mosses and liverworts, although they also disperse by spores, have the sporophyte attached to, and nutri-
tionally dependent upon, the gametophyte. Furthermore, the gametophyte is the dominant generation. Seed plants
(angiosperms and gymnosperms) resemble pteridophytes by dominant sporophyte, 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.

20
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

21
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.

22
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 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.

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,
Pteridaceae, 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.

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. Antherid-
iogens, 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
(Yatskievych, 1993).

Some evidence indicates that the antheridiogen response might actually be caused by two substances. The response
actually has two components: 1) precocious antheridia production, and 2) stunted growth (the prothalli usually re-
maining as protonemata or becoming merely spatulate). Naf (1956) suggested that gametophytes subjected to an-
23
theridiogen divert their potential vegetative growth into antheridia formation; however, some gametophytes 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.
Hamilton, R. G. & R. M. Lloyd. 1991. Antheridiogen in the wild: the development of fern gametophyte communities. Func-
tional 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: 15941601.
_____ & T. A. Ranker. 1985. Differential antheridiogen response and evolutionary mechanisms in Cystopteris. American Journal
of Botany 72: 659665.
Lloyd, R. M. 1974. Reproductive biology and evolution in the Pteridophyta. Annals of the Missouri Botanical Garden 61: 318331.
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: 507532 [especially pages
513521].
_____. 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 435470. 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: 986990.
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: 243249. Voeller, B. R. 1964. Antheridiogens in ferns. Pages 665684. 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.
Fern J. 57: 107112.
_____ & _____. 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. Amer. Fern J. 83: 30–36.

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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.

25
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 (Cheilanthes 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.

26
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.

27
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.

28
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. By using isozyme electrophoresis, hybrids have

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

29
been detected that are scarcely distinguishable morphologically from one or both of their parents, as has especially
been the case in North American Cystopteris. More recently, DNA has been used to detect hybrids. If cpDNA is used,
the female parent can be determined (cpDNA is inherited maternally).

Hybrids may be designated by either a formula or binomial name. 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).

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: 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.Thus the hybrid genus is no longer used. XAsplenosorus is an example:
Camptosorus nests within Asplenium and therefore the hybrid genus is superfluous.

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 platyneuron, 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.

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.
_____, M. Gibby, & C. Fraser-Jenkins. 1984. Recent advances in our knowledge of pteridophyte hybrids. Taxon 33: 256–270.
Moran, R. C. & J. E. Watkins, Jr. 2002. The occurrence and morphology of Adiantum X variopinnatum (Pteridaceae). Brittonia
54: 57–60.
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. Amer. Fern J. 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.

30
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-

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

edly as the branch below it develops.

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

31
between isotomy and lateral branching.

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 Thelypteris 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 ap-
pears to have evolved independently in the dennstaedtioids (e.g., Hypolepis, Dennsteaditia, Histiopteris), thelypteroids
(Phegopteris, Thelypteris palustris), and cystopteroids (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. Rhizome branching types. A. Hypolepis. B. Davallia. C. Thelypteris reticulate. D. Dryopteris (adventitious bud on adaxi-
al surface). E. Pteris. F. Gleichenia (bud at right is a branch bud, the on at left, a leaf bud). (Modified from Hagemann 1989)

32
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 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) to 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.

33
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 has a distinctive type of rhizome branching.The rhizomes are monstichous, with one rank (ortho-
stichy) 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.

34
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 Dichoto-
mie. 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.

35
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, this 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
iciently absorbing carbon dioxide (Kenrick, 2001).

36
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 lat-
eral 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.

37
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 tree). 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.

38
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.

39
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
hapotypic 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).

40
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 or
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. They have no need for stored food as do the female spores.

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.

41
Figure 20. Trilete spores. A. Pityrogramma trifoliata. B. Lygodium microstachyum. C. Huperzia lucidula. D. Anogramma lepto-
phylla, left spore shows proximal face, right, distal. E. Anemia intermedia. F. Acrostichum aureum. G. Adiantum pedatum. H. Erio-
sorus 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 two ways: they remain viable for shorter periods and they germinate faster. Typ-
ically, 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 active, 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

42
Figure 21. Monolete spores. A. Schizaea 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

43
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-pollinated 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 incomplete 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, con-
sequently, 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. American
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 (Adiantaceae).
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.

44
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).

45
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

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.

46
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.

47
 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).

48
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.

49
 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 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).

50
 Key to Families of Lycopodiopsida

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

51
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 (Lycopodiella) or subterranean, mycotrophic, and non-green (Huperzia and Lycopodium). Chromosome num-
bers varies, but is constant with a given subgroup. 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. Habi-
tats extremely diverse. About half (185) of the species occur in the American tropics (Øllgaard, 1979a, 1992, 1995).
Slightly more than half of the Huperzia species are epiphytes (Beitel, 1979), whereas Lycopodium and Lycopodiella are
terrestrial. The gametophytes of Huperzia and Lycopodium are subterranean, acholorophylous, and mycotrophic, but
those of Lycopodiella and Phylloglossum are green and epigeal.

Distinctive is the distribution of each genus. Lycopodiella and Lycopodium are cosmopolitan and contain many wide-
spread species (e.g., Lycopodiella caroliniana and Lycopodium complantatum). Huperzia, 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. Phylloglossum is endemic to
Australia, Tasmania, and New Zealand. (Wikström & Kenrick, 2001).

Genera/species: 4/480. Genera: Huperzia (400 spp.), Lycopodium (40), Lycopodiella (40), Phylloglossum (1).

Key to the main genera of Lycopodiaceae

1. Stems equally forked (isodichotomous), each fork equally thick; sporophylls and vegetative leaves alike or, if differ-
ent, then the sporophylls smaller and persistent; spores foveolate-fossulate..................................................................... Huperzia
1. Stems unequally forked (anisodichotomous), consisting of an elongated indeterminate main stem and shorter lat-
eral determinate branches; sporophylls and vegetative leaves dissimilar, the sporophylls aggregated in terminal strobili;
spores reticulate or rugose.
2. Strobili either pendent and sessile, or erect and terminating simple (rarely forked) stems; cells of sporangium
wall straight; spores rugose............................................................................................................................................................. Lycopodiella
2. Strobili either pendent and stalked, or erect and sessile or stalked, borne on lateral branchlet systems arising
from the main stems; side walls of sporangium epidermis cells sinuate, lignified throughout (use microscope);
spores reticulate..................................................................................................................................................................................... Lycopodium

52
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 sur-
gical gloves and condoms to prevent them from sticking together. An alkaloid in Huperzia serrata, called huperzine, is
reported to slow the onset of Alzheimer’s disease.

Figure 25. Relationships of the main clades within the Lycopodiaceae. Based on Wagner & Beitel (1992) and Wikström et al.
(1999).

Discussion: The Lycopodiaceae is well supported by molecular studies but has only one anatomical synapomorphy:
plectosteles Figure 26E,F; Figure 27D). In the past, it has been recognized as containing only two genera: Lycopodium
and the monotypic Phylloglossum. Molecular and morphological phylogenetic studies show that the family contains
three major clades (Figure 25): the Huperzia clade (containing Phylloglossum) and Lycopodiella and Lycopodium (Wag-
ner & Beitel, 1992; Wikström & Kenrick, 1997, 1999). These clades were recognized as genera by Øllgaard (1987,
1992), the world’s expert on the family, but nowadays the tendency is to split these clades into smaller monophyletic
groups. For instance, Wagner and Beitel (1992) recognized up to 11 genera, 9 of which are in the United States and
Canada. Haines (2002) split two more genera (Dendrolycopodium and Spinulum) from Lycopodium (sens. str.), recog-
nizing 11 genera in the United States and Canada. Because all these genera are monophyletic, it is merely a matter
of opinion at which rank they should be recognized. I here follow Øllgaard because he has written many floristic
treatments for the Neotropics and compiled an index (Øllgaard, 1989) to all names of Lycopodiaceae. But when I
teach field courses, it is helpful to recognize the segregates.

The largest genus is Huperzia, with about 400 species worldwide. It is most diverse in the Andes, where nearly 50 spe-
cies occur, half of which are endemic (Øllgaard, 1995). The stems are isodichotomous, and the sporophylls, although
they may differ in size and shape from the vegetative leaves, remain green and photosynthetic after spore dispersal.
A unique feature of the genus is roots that arise near the apex and travel within the cortex to soil level where they
emerge (Figure 26). In the strict sense, the name Huperzia applies of the “H. selago group,” about 12 species (mostly
temperate) characterized by gemmae borne in the leaf axils and spores with truncate corners and concave, pitted
proximal faces.

If Huperzia is recognized in the strict sense, then the other species fall into Phlegmariurus, which is divided into paleo-
and neotropical clades. Øllgaard (1987) recognized 21 informal species groups in this group, emphasizing they were
preliminary. Wikström et al. (1999) and Wikström and Kenrick, (1999) who found that some of these subgroups were

53
polyphyletic. Epiphytism is the ancestral condition in Phlegmariurus, evolving before the break-up of South America
and Africa. In the Andean páramo, there has been a reversion to the terrestrial habit at least twice (Wikström et al.,
1999; Wikström & Kenrick, 1999).

Sister to Huperzia is Phylloglossum, consisting only of the sole species P. drummondii (Figure 25). Hackney (1950) gave a
general account of its biology.The plants are up to 5 cm tall and occur in seasonally wet habitats, persisting throughout
the dry season by perennial tubers produced at the tip of a dropper branch. Its gametophytes are green and epigeal,
unlike those of Huperzia which are subterranean. Its sperm have 16 flagellae instead of the normal two within the
family.

Figure 26. Lycopodiaceae, Huperzia. A. Huperzia selago, 1) habit and bulbil. 2) gametophyte. B. H. pithyoides, stem cross-section.
C. H. linifolia. D. H. phyllicifolia, modified sporophylls and sporangia. E. H. varia, stem cross-section. F. H. squarrosa, stem cross-sec-
tion. G. H. ecuadoriana, unmodified sporophylls and sporangia.(1999).

54
Sister to the clade that contains Huperzia and Phylloglossum is Lycopodium + Lycopodiella. These two genera are held
together by the synapomorphies of mucilage canals in the leaf base and strobilus, anisotomous branching, fertile stems
determinate, sporophylls stalked and with a decurrent wing, and sporophylls senescing when the sporangia mature or
soon after (Figure 27). They can be distinguished by stem anatomy (Bierhorst, 1971; Jones, 1905; Ogura, 1972), spore
morphology (Tryon & Lugardon, 1991; Wilce, 1972), presence of mucilage canals along the veins (Bruce, 1976a),
branching pattern (Øllgaard, 1979b), shape of the sporangial epidermis cells (Øllgaard, 1975), and gametophyte mor-
phology (Bruce 1976b). Unlike Huperzia, hybrids in Lycopodiella and Lycopodium have normal chromosome pairing
and produce apparently normal, viable spores. In one case (L. X habereri) the spores have been germinated and grown

Figure 27. Lycopodiaceae, Lycopodium. A. L. 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. L. complanatum, gametophyte. C. L. volubile, habit. 1) Anisophyllous branchlet. 2) Strobilus. D. L. tris-
tachyum, cross-section of stem showing plectostele.

55
into gametophytes that produce normal archegonia and antheridia (Whittier & Britton, 1995).

Lycopodium consists of about 40 species worldwide. Nine sections were recognized within by Øllgaard (1987, 1992),
and their monophyly has been supported by sequence data from rbcL and the trnL intron (Wikström & Kenrick,
1999). The genus is extremely 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: L. thyoides.

Lycopodiella also has about 40 species. All have green surficial gametophytes, rugulose spores, erect stems produced
dorsall (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. Based
on morphology, Øllgaard (1987) recognized four sections, the monophyly of which has been supported by sequence
data from rbcL and the trnL intron (Wikström & Kenrick, 1999). These four sections are sometimes recognized as
genera: Lateristachys, Lycopodiella, Palhinhaea, and Pseudolycopodiella.

Unlike the ferns, within the Lycopodiaceae there is a complete absence of apogamy and alloploidy

56
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: 1/450. 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.

Discussion: The Selaginellaceae are a well-supported monophyletic group based on morphology (Kenrick & Crane,
1997; Stevenson & Loconte, 1996) and rbcL sequences (Korall & Kenrick, 2002; Korall et al., 1999). Morphological
synapomorphies are stem steles suspended by endoderm cells in a cavity (Figure 29E), epidermal cells with a reduced
number of chloroplasts (often only one), the presence of strobili, megaspores reduced to 4 per sporangium (some-
times less), and microsporangia globose.

The following discussion of the family’s phylogeny is based on Korall and Kenrick (2002). They found that subgen. Se-
laginella is sister to the rest of the genus (Figure 30). The subgenus consists of only two species: S. selaginoides (Figure
29B), cicrumboreal and type of the genus, and S. deflexa, endemic to Hawaii. These species have helically arranged
monomorphic trophophylls, unlike most other species the genus which are four-ranked. All the roots (less than 8)
are produced from a thickened hypocotyl (the region below the first two sporeling leaves); roots are not produced
elsewhere along the stems, nor are they produced from special root-bearing branches, the rhizophores, as in the
rest of the genus (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.

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 rhizophore clade consists of two subclades. One contains the taxa Articulatae, Tetragonostachys, and Ericetorum,
the other contains various anisophyllous species. Taxa Articulatae and Tetragonostachys are distinctive by possessing
dorsal rhizophores; that is, the rhizophore is produced by an angle meristem on the dorsal side of the fork.The rhizo-
phore loops over the narrower branch of the fork and grows downward. In the other subclade, the rhizophores are

57
Figure 28. 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
produced on the ventral side of the fork and grow directly downward.

Subgenus Tetragonostachys consists of 40 species that grow in dry habitats or deserts. Nearly all are New World.
Their 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 be-
tween branches along the stem, in contrast to the other subgenera where they occur at the branch forks. 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.

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

58
Figure 29. Selaginellaceae. A1-5. Selaginella lepidophylla. A. Habit when wet. A1. Habit when dry. A2. Heterophylly. A3. Microspo-
rangium. 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.

59
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 28A). 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 28A, 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 28A, 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 30).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 30).

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 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)..

60
Figure 30. Phylogeny of the Selaginellaceae based on rbcL ( Korall & Kenrick, 2002). Strict concensus tree. Branches with boot-
strap support above 80% are thicker.

61
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

62
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. 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-cuticled, 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 31. . 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 stele.

63
 Ferns
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.

64
Figure 32. Phylogeny of the ferns families. About 80% of the extant species belong to the Polypodiales.

65
 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 34A, 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).

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

66
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 typically circumscribed, the Ophioglossaceae constitute three genera: Ophioglossum, Helminthostachys, and Botrychi-
um. A fourth genus, Mankyua, has recently been described (Sun et al., 2001). These genera have distinctive geographic
distributions. 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 classifica-
tion 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 33). A phylogeny of Botrychium was presented
by Hauk et al. (2012). Presl (1845) treated Ophioglossum as four separate genera: Ophioglossum (s.s.), Rhizoglossum,
Ophioderma, and Cheiroglossa.

At the species level, Clausen (1938) published a worldwide monograph that is now badly out-of-date. In the past
two decades many new species of Botrychium (s.s.) have been discovered, especially in western North America, and
reticulate relationships and ranges have become much better known. Nowadays, 27 species are recognized in Botry-
chium subg. 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). 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 expanded 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 wightonii, from the Paleocene of western
Canada (Rothwell & Stockey, 1989). It resembles B. virginianum, an extant species of temperate and tropical Americ

Key to the main genera of 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; roots not proliferous
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.

67
Figure 33. Phylogeny of the Ophioglossaceae. Strict consensus of 12 equally parsimonious trees based on rbcL, trnL–F, and
morphology. Numbers above the branches are bootstrap supports. From Hauk et al. (2003).

68
Figure 34. Ophioglossaceae. A1–3. Botrychium multifidum. A1. Habit. A2, 3. Sporangia. A4–12. Botrychium virginianum. 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. Ophioglossum palmatum. B2.
Ophioglossum vulgatum. C1, 2. Helminthostachys zeylanica. C1. Habit. C2. Sporangia.

69
 Psilotales
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

70
Figure 35. 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/).

71
 Equisetales
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

72
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 an-
cestor present during the Eocene (Figure 36; 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. Equisetum 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

73
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.

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

74
 Marattiales
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.

75
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 37). The pulvinus-like swellings are not
true pulvini because they do not function in reversible movement of the leaf or its segments. The stipules are fleshy
and if cut and placed in soil will often form plantlets along the margin. 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. It appears in the late Carboniferous where it is rep-
resented 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 Botrychium subgen.
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 of the gametophyte.

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.

76
 Key to the Neotropical Genera of 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

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)

77
 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
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 39–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 40).

In agreement with its basal position in fern phylogeny, the family has a long fossil history, extending back to the late
Permian (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).

78
 Key to the Genera of 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

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 39B) 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,

79
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).

Figure 39. Osmundaceae. A1. Osmunda regalis, note winged petiole base. A2–A3. Developing sporangia. A4. Open sporangi-
um 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

80
Figure 40. Phylogeny of the Osmundaceae. Fifty percent majority rule consensus tree from a Bayesian analysis of seven plas-
tid loci (Metzgar et al., 2008).

81
 Hymenophyllales
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

82
 Key to the main groups of Hymenophyllaceae in the Americas
(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

83
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

84
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.

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

85
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).

86
Figure 42. 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).

87
 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. Phylogeny of the Gleicheniales. In this analysis, the filmy ferns (Hymenophyllum and Trichomanes) are sister to the
Gleicheniales. (After Pryer et al. 2004)

88
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).

Key to the Genera of 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

89
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 44). Just below this bud develops

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

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.

90
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).

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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

92
 Schizaeales A. B. Frank in Leunis
This order is characterized by differentiation of the sterile and fertile parts of the leaf, the absence of well defined
sori, and sporangia with a complete subapical annulus. It has been strongly supported as monophyletic (Hasebe & al.,
1995; Pryer & al., 2001a, 2004b; Skog & al., 2002; Wikström & al., 2002). Its fossil record dates back to the early Jurassic
(Collinson, 1996). Vegetatively the order is extremely diverse, and this diversity is reflected by its three main clades,
here recognized at the rank of family.

Wikström et al. (2002) inferred the phylogeny of the Schizaeales using rbcL sequence data (Figure 45). They found
Lygodium was sister to a clade consisting of Schizaea, Actinostachys, and Anemia. The estimated minimum age for the
split between Lygodium and the other genera was mid-Jurassic. 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).

Key to Main Taxa of 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; sporangia in 2 or more rows
each side of the midrib.......................................................................................................................... Schizaeaceae (Actinostachys)
3. Sporangiophores less than 2 cm long, pinnate, curved; sporangia in one row on either side of the midrib......
Schizaeaceae (Schizaea)

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.

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

93
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.

Anemiaceae Link

Terrestrial; rhizomes creeping to suberect, pubescent; leaves determinate, mostly hemidimorphic, some species com-
plete dimorphic; veins free, dichotomous, rarely anastomosing; sporangia usually borne on a basal pair of erect
pinnae; spores 128-256 per sporangium, tetrahedral, 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 a few species in Africa, Madagascar, islands in the Indian Ocean,
and southern India.

Genera/species: 1/120. Sole genus: Anemia.

Economic plants and products: None.

Distribution and ecology: Primarily neotropical, but a few species in Africa, Madagascar, islands in the Indian Ocean,
and southern India.

Genera/species: 1/120. Sole genus: Anemia.

Discussion: Nearly all species of Anemia have two erect basal pinnae that bear sporangia, making identification easy
in most instances. Surprisingly, these dimorphic fertile pinnae are pleisiomorphic for the Anemia + Mohria clade (Wik-
strom et al., 2002). Most of the genus is well known taxonomically thanks to the work of Mickel (1962, 1967, 1982).
The recognition of Mohria (African) leaves Anemia paraphyletic (Dettmann & Clifford, 1992; Reed, 1947; Skog, 1992;
Stevenson & Loconte 1996; Skog et al., 2002).

94
Figure 45. Phylogeny of the Anemiaceae. Courtesy of Paulo Labiak and John Mickel.

95
Schizaeaceae Kaulf. | Curly Grass Family

Plants terrestrial or epipetric. Rhizomes bristly, the bristles multicellular; protostelic, solenostelic or dictyostelic. Pet-
ioles containing a single vascular bundle. Blades simple, linear or fan-shaped, variously cleft; veins free, dichotomous;
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.

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

Economic plants and products: None.

Discussion: The monophyly of the family is supported by rbcL (Hasebe et al., 1995; Pryer et al., 1995; Wikstöm et al.,
2002, Skog et. al., 2002). The family has a fossil record extending back to the Jurassic (Wikström et al., 2002).

Schizaea and Actinostachys are closely related (Actinostachys is often treated as a subgroup of Schizaea). Their leaves
are (mostly) grass-like and bear sorophores apically.The sorophores are either digitate or pinnate with a short, curved
midrib. According to rbcL sequence data, both genera are monophyletic (Wikstöm et al., 2002).

96
 Salviniales | Water Ferns
The Salviniales, a group of about 70 species, are the only heterosporous ferns. They are the only plant group to have
evolved heterospory since the Paleozoic (Nangalingum et al., 2006). Their megasporangiate sori bear a single mega-
spore (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 either
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. Phylogeny of the Salviniales from Nagalingum et al. (2008). Phylogram with average branch lengths obtained by
Bayesian inference using atpB, rbcL, and rps4. Numbers above the branches are MP bootstrap/ML bootstrap/BI posterior prob-
ability. MPBS and MLBS with “*” = 100%; PP with “*” = 1.0.

97
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 colorful variegated fronds.

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

98
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). As documented by Darwin (1869), the four pinnae fold upward, forming a
“packet,” as darkness approaches in the late afternoon. Marsilia is the only nyctinastic fern.

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

Key to the Genera of 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

Figure 47. Spore-bearing structures of Marsiliaceae (A, B) and Salviniaceae (C. D). A, Longitudinal section of a Marsilea sporo-
carp. 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 = microspo-
rangia. Mis = microsporangiate sori. Pl (se) = parenchymatous layer or sporophore envelope. S= sorophore. V=vein. Scale bars
= 1 mm. From Nagalingum et al. (2006).

99
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 48).

Figure 48. Salvinia reussii, a Miocene fossil species with expanded laminae on submerged leaf. (from Collinson, 1991)

100
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.

Key to the Genera of 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

101
 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 the Families of Cyatheales

1. Rhizomes glabrous; leaves dimorphic.................................................................................................................................. Plagiogyriaceae

1. Rhizomes hairy, scaly, or bristly; leaves monomorphic.
2. Sori abaxial (borne on the undersurface of the frond)
3. Stems and leaves scaly...........................................................................................................................................................Cyatheaceae
3. Stems and leaves hairy.
4. Leaves 1-pinnate, not glaucous beneath; plants
usually found below 1000 m.......................................................................................................................................... Metaxyaceae
4. Leaves 2- to 3-pinnate-pinnatifid, glaucous beneath; plants
usually found above 1000 m.................................................................................................................................. Lophosoriaceae
2. Sori marginal.
5. Rhizomes scaly; plants from southern Venezuela....................................................................................................Cyatheaceae
5. Rhizomes hairy; plants from elsewhere.
6. Stems long-creeping, slender, covered by dark bristles; indusia cylindrica....................................Loxomataceae
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
7. Laminae widest at the base................................................................................................................................... Cibotiaceae

102
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 maxi-
mum 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).

103
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).

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).

104
 Key to the genera of Loxomataceae

1. Laminae catadromous, green beneath (not glaucous); costae and veins usually hairy beneath; lower pinnae sessile
or subsessile, asymmetric; sporangia with transverse dehiscence; Neotropics............................................................. Loxsomopsis
1. Laminae anadromous at base, catadromous distally, glaucous beneath; costae and veins glabrous beneath; lower
pinnae stalked, symmetric; sporangia with longitudinal dehiscence; New Zealand................................................................Loxoma

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).

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)

105
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
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).

106
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).

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).

107
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). Sphaeropteris is sister to
a trichotomy containing Cyathea, Alsophila, and Gymnosphaera (Figure 49). 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, Al-
sophila, Cyathea, and Gymnosphaera have marginate scales; that is, with the marginal cells differentiated, usually being
shorter, divergent, and irregular.

As currently construed, Cyathea contains four groups previously recognized as distinct (Korall et al., 2006, 2007). 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 a thorough analysis of 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).

Key to Genera of 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

108
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.

109
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 51, 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).

110
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 spran-
gia. 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 examined 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.

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.

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 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

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.

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Genera/species: 1/2.

Economic plants and products: none.

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).

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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.

Lindsaeaceae Pic. Serm.

Plants terrestrial or (rarely) epiphytic. Stems short- or long-creeping, hairy or scaly, protoselic or solenostelic, if protos-
telic then with internal phloem. Petioles with a single vascular bundle or (in Lonchitis) with two, 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: 6/200. Major Neotropical Genera: Lindsaea (150 spp.), Odontosoria (12), Lonchitis (2), Spheno-
meris (1).

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), but no morphological synapomorphies support it (Mickel, 1973). The
roots have a sclerenchymatous outer cortex combined with an innermost cortical layer six cells wide, but this is
lacking in Cystodium and Lonchitis (Schneider, 1996). In the past, the Lindsaeaceae have usually been classified in the
Dennstaedtiaceae, but molecular studies have shown such a classification renders the Dennstaedtiaceae paraphyletic.

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,

114
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.

Key to Major Neotropical Genera of 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; leaves erect or spreading.............................................................................................................. Ormoloma
2. Laminae 2- to 4-pinnate; leaves scandent.................................................................................................................. Odontosoria

Saccolomataceae Doweld

Terrestrial or saxicolous; rhizomes decumbent to erect, scaly, surrounded by persistant petiole bases; 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, 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: 1/12. Sole genus: Saccoloma (12 spp.).

Economic plants and products: None.

Discussion: This family was tentatively recognized as monophyletic by Smith et al. (2006), and their treatment is
followed here until more sampling can be done. Saccoloma was previously treated in the Dennstaedtiaceae because
of its omega-shaped vascular bundles in the petiole and submarginal sori, but it was atypical there because of its
short rhizome surrounded by persistent petiole bases and lack of epipetiolar buds (dennstaedtioids typically have
long-creeping, scaly rhizomes and epipetiolar buds).

115
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. All genera except Histiopteris have pubescent (not scaly)
rhizomes. The distinctive epipetiolar buds are described by Troop and Mickel (1963) for Dennstaedtia, Hypolepis, Pae-
sia, 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.

The Dennstaedtiaceae is recognized here in a restricted sense, excluding the Lindsaeaceae that has long been includ-
ed. Within the family, Dennstaedtia and Microlepia form a well-supported basal clade basal, with Monachosorum (Asian)
as the next most distal taxon, and the other genera distal to this (Hasebe et al., 1995; Wolf, 1997). DNA sequencing
of the remaining genera is needed before phylogenetic relationships within the family are resolved.

Key to the Major Neotropical Genera of Dennstaedtiaceae

1. Sorus supplied by one vein.

2. Indusia cup-shaped....................................................................................................................................................................... Dennsteadtia
2. Indusia scale-like, formed by the recurved margin of the lamina................................................................................Hypolepis
1. Sorus supplied by two or more veins.
3. Veins anastomosing; rhizomes scaly..........................................................................................................................................Histiopteris
3. Veins free; rhizomes pubescent
5. Rachises flexuous; spores monolete............................................................................................................................................. Paesia
5. Rachises straight; spores trilete............................................................................................................................................... Pteridium

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Figure 53. Dennstaedtiaceae, Dennstaedtia. A1. Rhizome with leaf bases. A2. Leaf. A3. Pinnule with marginal, cup-shaped
sori. A4. Sorus.

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 always 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. Major genera: Subfam. Adiantoideae: Adiantum (200), Ananthacorus (1), Anetium (1),
Antrophyum (20), Haplopteris (20), Hecistopteris (3), Monogramma (1), Polytaenium (8), Radiovittaria (8), Rheopteris (1),
Scoliosorus (3), Vittaria (6). Subfam. Cheilanthoideae: Cheilanthes (180), Doryopteris (30), Pellaea (25), Notholaena
(23), Myriopteris (ca. 20), Argyrochosma (17). Subfam. Cryptogrammoideae: Coniogramme (20), Cryptogramma (3),
Llavea (1). Subfam. Parkerioideae: Acrostichum (4), Ceratopteris (4). Subfam. Taenitidoideae: Actiniopteris (5),
Anogramma (s), Austrogramme (5), Cosentinia (1), Eriosorus (25), “Jamesonia” (19), Nephropteris (1), Onychium (8), Pteris
(250), Pterozonium (13), Syngramma (15), Taenitis (15).

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

117
in unrelated ferns such as Hypolepis (Dennstaedtiaceae) and Lonchitis (Lindsaeaceae).

The Pteridaceae can be divided into at least five major groups, and perhaps these will eventually be recognized at
the family level. The first group 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. If desired, these could be called the Cryp-
togrammaceae or Cryptogrammoideae.

A second group consists of the aquatic genera 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 taenitidoid ferns, or subfamily Taenitidoideae, 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, Ano-
gramma, Austrogramme, Cosentinia, Onychium, Pityrogramma, Pterozonium, Eriosorus (including Jamesonia), 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 taenitidoid clade was studied by Sánchez-Baracaldo (2004b), who produced a cladogram
showing generic relationships (Figure 56). In another study (2004a), she examined the phylogeny Jamesonia and Erio-
sorus, two high-elevation genera in the Neotropics. She found that Jamesonia, characterized by narrow indeterminate
leaves with small rounded pinnae, is 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 vittarioid +Adiantum clade (Figure 55), referred to as the Adiantaceae or Adiantoideae (the
vittarioids were long recognized as a separate family, the Vittariaceae). It consists of Adiantum and the 10 genera of
vittarioid ferns: Ananthacorus, Anetium, Antrophyum, Haplopteris, Hecistopteris, Monogramma, Polytaenium, Radiovittaria,
Scoliosorus, 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 57 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
cells (Wilson, 1959).

Crane (1997) revised the generic taxonomy of the vittarioids based on rbcL data (Figure 54). 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 funnel-shaped terminal cells at the tips of the para-
physes were placed in either Haplopteris (paleotropical) or Radiovittaria (neotropical) depending on whether their

118
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.)

The last group is the cheilanthoids (Siniopteridaceae, or subfamily Cheilanthoideae), which includes about 400 species.
It is subdivided into the myripteroids, pellaeoids, hemionitidoids, and nothlaenoids (Windham et al., 2009).The genera
included are Adiantopsis, Argyrochosma, Astrolepis, Aspidotis, Bommeria, Cheilanthes, Cheiloplecton, Doryopteris, Hemio-
nitis, Myriopteris, Notholaena, Pellaea (in part), Pentagramma, and Trachypteris. Apparent synapomorphies 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 55) by Gastony & Rollo (1998).

Key to Major Genera of Vittarioid Ferns (Pteridaceae)

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

1. Blades entire; veins anastomosing
2. Veins anastomosing to form a single row of aereolae; 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
8. Sporangia present only along the veins; leaves clumped..................................................................... Polytaenium
8. Sporangia scattered on and between veins; leaves distant......................................................................... Anetium

119
Figure 54. Phylogeny of the vittarioid ferns. From Crane (1997).

120
Figure 55. Phylogeny of cheilanthoid ferns (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).

121
Figure 56. Phylogeny of subfam. Taenitidoideae (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 Conio-
gramme and the two Pteris species as an outgroup.

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Figure 57. 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.

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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 vas-
culature 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 58A). 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
Eupolypods 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. Eupoly-
pods II (Thelypteris imbricata; Thelypteridaceae).

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 EUPOLYPODS I
This clade is sister to Eupolypods II and consists of the Hypodematiaceae, Dryopteridaceae, Lomariopsidaceae, Tec-
tariaceae, 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 members 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 are, when viewed in cross-section, hooked.

Hypodematiaceae Ching

Plants terrestrial or rarely epipetric. Rhizomes dictyostelic, erect or short- to long-creeping, scaly. Petioles with 4 or
more vascular bundles at the base. Sterile and fertile leaves monomorphic. Rachises and costae grooved or terete,
the grooves (when present) usually decurrent into those of the next lower order. Veins free. Sori indusiate, round to
(Dicymochlaena) elongate; 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.

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

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

Discussion: This family is supported by molecular evidence (Schuettpelz & Pryer, 2007). Previously, its three genera
have been associated with different families: Didymochlaena with the Dryopteridaceae, Luecostegia with the Davalli-
aceae, and Hypodematium with the Athyriaceae. The genera lack any morphological or anatomical synapomorphies.
Both Leucostegia and Hypodematium have creeping, dorsiventral rhizomes.

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.

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Genera/species: 40–45/1700. Major genera: Elaphoglossum (600 spp.), Dryopteris (275), Polystichum (200), Ctenitis
(100), Arachniodes (50), Bolbitis (45), Megalastrum (35), Polybotrya (35), Parapolystichum (25), Stigmatopteris (24), Lo-
magramma (22), Pleocnemia (19), Lastreopsis (15), Cyrtomium (15), Teratophyllum (11), Mickelia (10), Phanerophlebia
(8), Pradoia (2), 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 is a large and diverse family strongly supported by molecular data (Schuettpelz & Pryer, 2007)
but does not have any morphological synapomorphies. It consists of five main clades. The first is a terrestrial group
with erect or short-creeping rhizomes and monomorphic sterile and fertile leaves. It consists of Ctenitis, Dryopteris,
Polystichum, Phanerophlebia, 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 Bar-
rington (2003) and, in the Andes where it is particularly species-rich, by McHenry & Barrington (2014).

The second group, the “polybotryoid clade,” is entirely neotropical and 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, Pradosia (gen nov.), and Polystichopsis. The latter genus, sister to the others,
is West Indian and atypical by thin rhizomes, terrestrial habit, monomorphic sterile and fertile leaves, and long, straight
whitish hairs on the laminae.

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 fourth group, the “Lastreopsid” clade, is formed by Megalastrum, Lastreopsis, Parapolystichum, and Rumorha (Labiak
et al., in review). A synapomorphy for this group are colored glandular hairs on the leaves (and on the sporangial
stalks in Lastreopsis and Parapolystichum).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 surcurrent
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 59). 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 about 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 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 60). Section Amygdalifolia
is characterized by the unique combination of long-creeping rhizomes, phyllopodia, hydathodes, and reddish young
leaves. It is the only species in the genus that is a primary hemiepiphyte (Lagomarsino et al., 2012). Section Elapho-
glossum 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 aerophores on the rhizome near the

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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. Section 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).

Figure 59. 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 boot-
strap (bellow the branches) equals to 1 and 100%, respectively. (from Moran et al., 2010.)

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Figure 60. 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 boot-
strap (bellow the branches) equals to 1 and 100%, respectively. (from Moran et al., 2010.)

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: 4/70. Genera: Lomariopsis (45), Nephrolepis (23 spp.), Cyclopeltis (5), Thysanosoria (1).

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

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).

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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.

The rhizome system of Nephrolepis is unusual (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 sto-
loniferous part. Unlike other fern rhizomes, the erect rhizomes of Nephrolepis do not produce roots; they produce
stolons, and only the stolons produce the plants’ roots. The stolons are further unusual by not bearing leaves as sto-
lons usually do. An analysis of three plastid sequences, including the maturase matK, resolved Nephorlepis apart from
other lomariopsids, suggesting that it should be recognized in its own family, the Nephrolepidiaceae (Kuo et al., 2011).

Key to the Neotropical Genera of 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. Sori in one row between costa and margin; stolons present........................................................................... Nephrolepis
2. Sori in 2 or 3 rows between costa and margin; stolons absent
3. Leaves simple, entire, with apical bud; veins netted............................................................................ Dracoglossum
3. Leaves 1-pinnate, without apical bud; veins free........................................................................................... Cyclopeltis

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: 10/250. Major genera: Tectaria (200), Triplophyllum (22), Arthropteris (12), Pteridrys (8), Hypoderris
(3).

Economic plants and products: none.

Discussion: The monophyly of the Tectariaceae is supported by DNA sequence evidence (Schuettpelz & Pryer,
2007). An apparent morphological synapomorphy is the puberulence generally found on the upper 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

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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).

Key to the Neotropical Genera of Tectariaceae

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

1. Plants terrestrial or epipetric; widespread
2. Rhizomes erect, with polystichous leaves radially arranged ............................................................................................ Tectaria
2. Rhizomes creeping
3. Spores cristate; veins free................................................................................................................................................. Triplophyllum
3. Spores spiny; veins netted, if free, then rachis winged.......................................................................................... Hypoderris

Figure 61. Phylogeny and synapomorphies of three Eupolypods I families.

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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 (30 spp.)

Economic plants and products: none.

Discussion: The monophylly of Oleandra and its sister relationship to Davalliaceae and Polypodiaceae 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 62A; 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
62D, 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 perispore (see
SEM images on www.plantsystematics.org).

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 62 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 62A). 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

131
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 62. 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 spo-
rangial 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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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).

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, or (in grammitids) erect, radial, with leaves polystichous. Leaves mono-
morphic 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: 50/1000. 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 63). 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 63. . Typical rhizome of Polypodiaceae, showing creeping habit, two rows of phyllopodia, and ventral roots.

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 Key to the Genera of Neotropical 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 pinna,
these 0.5–5 cm wide ......................................................................................................................................Serpocaulon

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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

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 Eupolypods II
Eupolypods II contains nearly one-third of the extant fern species (Figure 63; Rothfels et al., 2012b). It is characterized
by two vascular bundles in the petioles, each bundle elongate as seen in transverse section (Figure 58; also see petiole
vasculatures at end of this book). The recognition of Eupolypod II families below follows Rothfels et al. (2012a, b).

Figure 64. Phylogeny of the families belonging to Eupolypod II (Rothfels et al., 2012).

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: 4/30. Genera: Cystopteris (20), Gymnocarpium (7), Acystopteris (3), Cystoathyrium (1).

Economic plants and products: None.

Discussion: The Cystopteridaceae are sister to the rest of Eupolypods II (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,
Cystoathyrium, and Cystopteris are unique by being attached to the proximal side of the receptacle and arching over
the sporangia. 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.

137
Figure 65. Phylogeny of the Cystopteridaceae. Maximum likelihood tree based on concatenated chloroplast sequence data.
(From Rothfels et al., 2013).

138
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.

139
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.

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).

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.

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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).

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 (8).

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.

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.

141
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).

Athyriaceae Alston | Lady ferns

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: 5/700. Genera: Diplazium (400 spp.), Athyrium (220), Deparia (70); Cornopteris (9); Anisocampium
(4).

Economic plants and products: The fiddleheads of Diplazium esculentum are eaten in Southeast Asia. This species
is also frequently cultivated in greenhouses.

Discussion: A large clade difficult to characterize morphologically. Clear morphological synapomorphies are lacking.
All are terrestrial, and most are medium-sized ferns with x=40, but Diplazium has x=41, a synapomorphy for that
genus (Wei et al., 3013). As for 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).

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 monomorphic or
(less commonly) subdimorphic; petioles with 2 vascular bundles at base; laminae most commonly 1-pinnatifid-pinnat-
ifid, rarely simple or 3- or 4-pinnate, 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 the segment margins; sori round or rarely
elongate; indusia present or absent; sporangial stalks 3-rowed; spores monolete, nongreen. Gametophytes epigeal,
142
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.

Genera/species: 3/950. Genera: Thelypteris (875 spp.), Macrothelypteris (10), Phegopteris (3).

Economic plants and products: none.

Discussion: The Thelypteridaceae can usually be recognized by the presence of acicular hairs; that is, hairs that are
needle-shaped and sharp-pointed. These hairs are dead and maturity and do not contain cytoplasm; thus, they tend
to be whitish or colorless. The hairs are most commonly unicellular but may be multicellular (e.g., Macrothelypteris).
Besides leaves, acicular hairs may occur on the rhizomes, rhizome scales, and sporangial capsules (e.g., Phegopteris).

Many Thelypteridaceae (e.g., subgen. Stieropteris and Amauropelta) bear aerophores (a.k.a., pneumatophores) on the
abaxial surfaces of the rachis-costa junctures, but some 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 mucilaginous
fiddleheads, the aerophores protrude through the mucilage, presumably helping to aerate the developing leaves.

Two other tendencies are notable in the family: Sterile-fertile leaf dimorphy is absent or poorly developed, and the
pinnae, most importantly the basal pair, are equilateral, not prolonged basiscopically (Figure 66). This latter character
helps distinguish similar 1-pinnate-pinnatifid species in genera such as Dryopteris and Megalastrum.

Up to 30 monophyletic groups have been proposed as genera within the family. Holttum (1971, 1982). An interme-
diate number of genera will probably be accepted after the phylogeny of the family is well resolved (Smith & Cranfill,
2002). One study based on three plastid markers and 115 species (representing 27 segregates of the family) recog-
nized seven genera (Figure 67; He & Zhang, 2012).

Within the family are two main clades. First is the phegopteroid clade composed of Macrothelypteris, Pseudophe-
gopteris, and Phegopteris (Figure 67) These have adaxially ungrooved leaf axes and laminae typically more than the
1-pinnate-pinnatifid.The second is the thelypteroid clade, Thelypteris sens. lat., which is predominantly free-veined and
1-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 helps to distinguish them from similar ferns (e.g., Dryopteris, Stigmatopteris). In the strict sense, Thelypteris
has branched veins (Figure 66-A2), which distinguishes it from many proposed segregate genera in the family. Also
helpful is whether the veins meet the laminar margin at the base of the sinus (the cyclosoroid group; Figure 66-A5)
or above it (other groups; Figure 66-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 66-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.

143
Key to the American Genera and Subgenera of 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................................................................................................................................Thelypteris subgen. 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...........................................................................................................................Thelypteris subgen. 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.............................................................................Thelypteris subgen. Goniopteris
6. Laminae more than 1 cell thick between veins, usually iridescent....Thelypteris subgen. 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 subgen. Thelypteris
7. Veins usually simple; proximal pinnae much shorter than the longest or not
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 ......................................................................................... Thelypteris subgen. Amauropelta
8. Lamina without reduced proximal pinnae, or if reduced pinnae present,
then veins united below the sinus
9. Sori elliptic; sporangial capsules setose....................................Thelypteris subgen. 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..............................Thelypteris subgen. Steiropteris
10. Cartilaginous keel lacking; indusia present, persistent; aerophores
absent at pinna bases abaxially............................................................Thelypteris subgen. Cyclosorus

144
Figure 66. Thelypteridaceae. A1. Thelypteris palustris, habit. A2. Segment and veins. A3. Cross-section of the petiole, showing
two vascular bundles. A4. Phegopteris connectilis. A5. Thelypteris ferox, segments and venation. A6. Thelypteris oppositipinna,
pinna and non-indusiate sori. A8. Thelypteris puberula, indusiate sorus.

145
Figure 67. Phylogeny of the Thelypteridaceae based on maximum likelihood analysis of rbcL sequence data. Thick branches
have < 95% bootstrap support. From He & Zhang (2013).

146
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: 10/250. Major genera: Blechnum (220 spp.), Woodwardia (14), Doodia (13), Stenochlaena (8), Sadle-
ria (6), Salpichleana (3).

Economic plants and products: Species of Blechnum, Doodia, Stenochlaena, 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.The indusium is born along a continuous commissural vein parallel to the midribs.
It is often hard to see because it shrivels and is pushed back and obscured by the maturing sporangia. Also distinctive
of the family 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). The monophyly of the Blechnaceae is supported by morphology
and DNA sequences, as is its sister relationship to the Onocleaceae (Figure 68; Cranfill & Kato, 2003; Hasebe, 1995;
Pryer et al., 1995).

Blechnum, the main genus in the family, is paraphyletic as currently delimited (Cranfill & Kato, 2003). It contains
many distinct subgroups that will eventually be recognized as genera. Two examples are the Lomariocycas group,
characterized by arborescent habit, dimorphic sterile and fertile leaves, and narrow, curved petiolar scales with dark
central stripes, and the B. fragile group (B. sect. Lomaria (Willd.) Diels), characterized by climbing rhizomes, meristeles
arranged in a circle (seen in cross-section) and surrounded by dark sclerenchymatous sheaths, adnate pinnae, and
dimorphic sterile and fertile leaves.

Woodwardia (including Anchistea, Chieniopteris, and Lorinseria) contains 14 species scattered throughout the northern
hemisphere. Their biogeography, ecology, and phylogeny were studied by Cranfill & Kato (2003), who suggested that
the group originated at high latitudes in North America during the late Cretaceous. Woodwardia is sister to all other
Blechnaceae.

Key to the Neotropical Genera of Blechnaceae

1. Leaves twining, 2-pinnate................................................................................................................................................................... Salpichlaena

1. Leaves not twining, simple to 1-pinnate-pinnatifid.
2. Veins free; pinnae entire or serrate....................................................................................................................................... Blechnum
2. Veins netted; pinnae pinnatifid.......................................................................................................................................... Woodwardia

147
Figure 68. Phylogeny of the Blechnaceae and Onocleaceae. Strict consensus tree of three most-parsimonious trees obtained
from maximum parsimony analysis of a combined morphological and molecular data set. Macrothelypteris, Asplenium, and Hom-
alosorus are outgroups. From Cranfill and Kato (2003)

148
 Onocleaceae Pichi Sermolli
(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 pinnati-
sect to 1-pinnate. Veins areolate or free. Receptacle conic. Indusium present. 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: The onocleoid ferns are a small, distinctive group whose monophyly has long been inferred on the basis
of morphology. The brown, indurated dimorphic fertile leaves are highly unusual, as are the sori completely enclosed
by the beadlike segments of the fertile pinnules (Figure 70). The family is sister to the Blechnaceae (Figure 68; 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 69).

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..

Key to the American Genera of 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

149
Figure 69. 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).

150
Figure 70. Onocleaceae. A. Matteuccia struthiopteris, habit. A2. Fertile pinnule. A3. Sori. A4. Sorus and indusium. A5. Cross-sec-
tion of fertile pinnule. B1. Onoclea sensibilis. B2. Sterile pinnule showing venation. B3. Fertile pinnule.

151
152
GÉNEROS NEOTROPICALES DE
HELECHOS y LICOFITAS

153
 Abrodictyum C. Presl
DESCRIPTION: Terrestrial; roots many, robust, of- DESCRIPCIÓN: Terrestres; raíces numerosas, robus-
ten stilt-like; rhizomes 1-2 mm wide, erect, ascending or tos, a menudo como raíces zancudos; rizomas 1-2 mm
creeping, unbranched, pubescent, the hairs dark, multicel- de ancho, erectos, ascendentes o reptantes, no ramifi-
lular; sterile and fertile leaves up to 30 × 18 cm, mono- cados, pubescentes, los pelos oscuros, multicelulares; ho-
morphic, polystichous; laminae 1-cell thick between the jas estériles y fértiles hasta 30 × 18 cm, monomorfas,
veins, 2-pinnate to 4-pinnate-pinnatifid; ultimate seg- polistícas; láminas una célula de grosor entre las venas,
ments narrow, only a few rows of cells along the 2-pinnadas a 4-pinnado-pinnatífidas; segmentos últi-
veins, the cells large, usually evident with 10×; veins free, mos estrechos, solamente unas hileras de células
anadromous, false veins absent; sori marginal; indusia por las venas, las células grandes, usualmente evidentes
tubular to campanulate, the mouth entire, truncate; re- con 10×; venas libres, anádromas, falsas venas ausentes;
ceptacles exert; spores green, tetrahaedral-globose; x=33. soros marginales; indusia tubular, la boca entera, tun-
cada; receptáculos exertos; esporas verdes, tetrahédri-
cas-globosas; x=33.

SIMILAR GENERA: Trichomanes subgen. Trichomanes GÉNEROS PARECIDOS: Trichomanes subgen. Tricho-
usually differs by less divided laminae, catadromous vena- manes difiere usualmente por láminas menos divididas,
tion, and sori that tend to be clustered toward the pinna nervación catadrómica y soros que tienden ser agrupa-
tips. Trichomanes subgen. Davalliopsis differs by laminae dos hacia los ápices de los pinnas. Trichomanes subgen.
opaque, 3-cells thick between the veins, and usually irides- Davalliopsis difiere por láminas 3 células de grosor entre
cent. Hymenophyllum differs by epiphytic habit, creeping las venas y usualmente iridiscentes. Hymenophyllum difi-
rhizomes, two-valved indusia, and included receptacles. ere por hábito epifítico, rizomas reptantes, indusios bival-
vados y receptáculos incluidos.

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

about 25 species. The four neotropical species (A. cellu- de 25 especies. Las cuatro especies neotropicales (A. cellu-
losum, A. sprucei, A. rigidum, and A. windischianum) belong losum, A. sprucei, A. rigidum y A. windischianum) pertenecen
to subgen. Pachychaetum. They typically have short erect al subgén. Pachychaetum. Típicamente tienen rizomas cor-
rhizomes supported by prop roots and in aspect resem- tas erectas apoyadas por raíces zancudos, con el aspecto
ble a small tree. The large cells along the narrow margin de un arbolito. Las células grandes por los márgenes es-
of veins are easily visible with a 10× lens, although in T. trechos de venas son visibles con una lupa de 10×, aun-
ridigum (the most widespread neotropical species) the que en A. rigidum (la especie neotropical más difundida)
cells are occluded. The chromosome number of x=33 is las células son ocluidas. El número cromosómico de x=33
derived within the family (Ebihara et al., 2007). Abrodic- se deriva dentro de la familia (Ebihara et al., 2007). Abrod-
tyum is derived from the Greek abros, fine, delicate + dic- ictyum se deriva del griego abros, fino o delicado + dictyon,
tyon, net. The cell walls appear as a net. red. Las paredes celulares aparecen como 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. 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 Lepto-
sporangiées). Adansonia 12: 469–495.

154
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).

155
PTERIDACEAE
Acrostichum L.
DESCRIPTION: Terrestrial or palustral; roots large, DESCRIPCIÓN: Terrestres o palustres; raíces grandes,
fleshy; rhizomes erect or ascending, generally clustered; carnosos; rizoma erecto o ascendente, generalmente
petioles with many vascular bundles; leaves 2–5 m, amacollado; pecíolos con muchos haces vasculares; hojas
1-pinnate, with a terminal segment similar in shape 2–5 m, 1-pinnadas, con una pinna terminal similar
to the lateral pinnae, erect or slightly spreading,; pinnae en forma a las pinnas laterales, erectas o un poco di-
20–60 pairs, 15–40 × 3–6 cm, the margins entire and vergentes; pinnas 20–60 pares, 15–40 × 3–6 cm, los már-
cartilaginously thickened; veins reticulate throughout, genes enteros y cartilaginosos, engrosados; nervaduras
the areoles ca. 1 mm wide, without included veinlets; sori reticuladas a todo lo largo, las aréolas c. 1 mm an-
acrostichoid, paraphysate; spores trilete, whitish to tan; cho, sin nervaduras libres incluidas; soros acrosticoides,
x=30. 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. carnosos y soros lineales protegidos por un indusio falso.

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

156
Figure 72. Acrostichum A, B. A. aureum. C, D. A. danaeifolium. (from Tryon & Stolze, 1989)

157
SCHIZAEACEAE
Actinostachys Wall. ex Hook.
DESCRIPTION: Terrestrial; stem short, creeping or DESCRIPCIÓN: Terrestres; rizomas cortos, reptantes o
erect, pubescent, the hairs orange or brownish, flexuous; erectos, pelosos, los tricomas anaranjados a parduscos,
leaves clumped to spaced; petioles triangular or flattened, flexuosos; hojas cespitosas o espaciadas; pecíolo triangu-
narrowly winged, glabrous, pale green, becoming brown lar o aplanado, estrechamente alado, glabro, verde pálido
near the base; laminae linear, grasslike, or rarely ex- a pardo hacia la base; láminas lineares, graminiformes,
panded, glabrous; veins free; fertile segments (sporan- escasamente a no foliáceas, glabras; segmentos fértiles
giophores) terminal, digitate or subdigitate; sporan- (esporangióforos) terminales, digitados a subdigit-
gia pear-shaped or oblong, with an apical annulus, in 4–6 ados; esporangios pyriformes o oblongos, en 4–6 hil-
rows on the indusiform segments; spores monolete; ga- eras en los segmentos indusiformes; esporas monoletes;
metophytes subterranean, not green, tuberous, brown gametofitos subterraneos, no verdes, tuberosos, par-
hairy; x=134, 140, 168. do-peludos; x=134, 140, 168.

SIMILAR GENERA: Schizaea differs by curved, pinnate GÉNEROS PARECIDOS: Schizaea difiere por los seg-
fertile segments, sterile portions of the leaves often di- mentos fértiles curvados, pinnados, las partes estériles de
chotomous to flabellate, and sporangia in 2–4 rows. las hojas a menudo dicotómas a flabeladas; esporangios
en 2–4 hileras.
COMMENTS: Actinostachys is pantropical has about 20 COMENTARIOS: Actinostachys es pantropical y consta
species worldwide, 2 of which are neotropical (A. pen- de casi 20 especies mundial, 2 de las cuales son neotrop-
nula and A. subtrigujata). It typically grows on sandy soil icales (A. pennula y A. subtrigujata). Crece típicamente en
or decaying logs below 600 m. Many species exhibit a suelos arenosos o sobre troncos podridos bajo de 600
clumped habit that results from repeated branching from m. Muchas especies exhiben un hábito amacollado se
the buds at the base of the petioles. It is often difficult debe a las ramificaciones repetidas de brotes en las bas-
or find the original stem among the numerous petiole es de los pecíolos. Es a menudo difícil encontrar el tallo
bases in the clump. In Actinostachys the fertile segments original entre las bases numerosas en la macolla. En Ac-
(sporangiophores) are actually pseudodigitate with a very tinostachys los segmentos fértiles (esporangióforos) son
short axis. In Schizaea this axis is longer, and the fertile en realidad pseudodigitados con un eje corto. En Schi-
segments have a feathery appearance. The gametophytes zaea este eje es más largo, y los segmentos fértiles tienen
of Actinostachys are subterranean, tuberous, hairy, and re- una apariencia plumosa. Los gametofitos de Actinostachys
semble a short rhizome, whereas those of Schizaea are son subterráneos, tuberosos, peludos y se asemejan a un
superficial, green, and filamentous. Both are mycorrizal. rizoma corto, mientras que los de Schizaea son superfi-
The species of Actinostachys are often recognized under ciales, verdes, y filamentosos. Los dos son micorrizales.
Schizaea. The genus name is derived from the Greek ak- A menudo las especies de Actinostachys se trata bajo de
tis, ray + stachys, spike. The apical segments of the fertile Schizaea. El nombre genérico se deriva del griego aktis,
leaves are digitately arranged. raya + stachys, espiga. Los segmentos apicales de las hojas
fértiles son arrglados digitamente.

LITERATURE: Lellinger, D. B. 1969. Schizaeaceae (Filicales). In: B. Maguire and collaborators.The botany of the Guayana High-
land—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. Takeuchi, M. 1960. O gênero Schizaea no Amazonia. Bol.
Mus. Paraense Emilio Goeldi 5: 1–26.

158
Figure 73. A, B. Actinostachys pennula (Sw.) Hook. (Mickel & Smith, 2004).

159
PTERIDACEAE
Adiantopsis Fée
DESCRIPTION: Terrestrial or rupestral; rhizomes erect, DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto,
decumbent, or short-creeping, scaly, the scales bicolorous decumbente y cortamente rastrero, escamoso, las es-
with a dark central stripe; leaves monomorphic, fascic- camas estrechas, bicolores, con una raya oscura central;
ulate, non-farinose; petioles atropurpureous; laminae hojas monomorfas, fasciculadas, no farinosas; pecíolos
pedate, radiate or 1-4-pinnate; pinnules or segments atropurpúreos; lámina pedata, radiada o 1 a 4-pin-
asymetrical, sessile or short-stalked, glabrous, anadro- nada; pínnulas o segmentos asimétricos, sésiles a
mous; rachises and costa grooved adaxially, with cortamente peciolulados, glabra, anadrómicos; ra-
narrow lateral wings on either side; veins free, simple; quis y costas sulcados o adaxialmente aplanados,
sori marginal, discrete (not in lines), several per segment, con alas estrechas y membranáceas en cada lado;
protected by a scarious false indusium; spores echinate or nervaduras libres, simples; soros marginales, discretos (no
(2 spp.) cristate, tetrahedral-globose, trilete; x=30. en líneas), varios por segmento, protejidos por un indusio
falso escarioso; esporas equinatas o (2 spp.) crestadas,
tetraédrico-globosas, triletes; x=30.

SIMILAR GENERA: Adiantum differs by terete petioles GÉNEROS PARECIDOS: Adiantum difiere por pecío-
and rachises. Cheilanthes has terete rachises and costae los y ráquises teretes. Cheilanthes tiene ráquises y cos-
or, if grooved, then the groove is V-shaped (not flat) and tas teretes o, si surcados, el surco es en forma de V (no
narrow lateral wings on either side are lacking. 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. tropical y consta de 40 especies, 9 de las cuales están
They occur in wet or seasonally dry forests. It is closely en los Caribes. Existe en bosques húmedos o estaciona-
related to Cheilanthes, especially C. microphylla (Sw.) Sw. lmente secos. El género es vinculado a Cheilanthes, es-
It differs only by the rachis-costa characters. The genus pecialmente C. microphylla (Sw.) Sw. Difiere solo por las
is notable for containing species with pinnate, pedate, or características del los ráquises y costas. El género es nota-
digitate lamina architectures. Adiantopsis radiata (L.) Fée is ble por contener especies con láminas pinnadas, pedatas
widespread in the Neotropics and the main species in the o digitatas. Adiantopsis radiata (L.) Fée es muy difundida
genus with digitate laminae. The genus name is derived en el Neotrópico y la especie principal en el género con
from Adiantum and the Greek suffix -opsis, like. láminas digitatas. El nombre genérico se deriva de Adian-
tum 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 Adiantopsis 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 biogeogra-
phy.Taxon 60: 1255–1268. Maxon, W. R. 1908.The Cuban species of Adiantopsis. Contributions from the United States National
Herbarium 10: 485–486.

160
Figure 74. A, B. Adiantopsis seemannii (Hook.) Maxon C, D. A. radiata (L.) Fée (Mickel & Smith, 2004).

161
PTERIDACEAE
Adiantum L.
DESCRIPTION: Terrestrial or rupestral; rhizome short- DESCRIPCIÓN: Terrestres or rupestres; rizoma corta
to long-creeping, rarely suberect, scaly; axes of the leaf a largamente rastrero, raramente suberecto, escamoso;
dark reddish to black, usually lustrous, terete; lami- ejes de la hoja pardo-rojizos a negros, lustrosos,
nae simple to 4-pinnate, often reddish when young, elon- teretes; láminas simples a 4-pinnadas, a menudo ro-
gate spicular cells often present adaxially; ultimate seg- jizas cuando jóvenes, con células espiculares alargadas a
ments sesile to stalked, never adnate; veins free or rarely menudo presentes en el haz; últimos segmentos sésiles
anastomsing; sporangia borne on the inner surface of a pedicelados, nunca adnatos; nervaduras libres o rara-
the false indusium (not below it); paraphyses absent; mente anastomosadas; esporangios nacen en la cara
spores globose-tetrahaedral, trilete; x=29, 30. interna del falso indusio; parafisos ausentes; esporas
globoso-tetrahédricas, triletas; x=29, 30.

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

LITERATURE: 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. Scam-
man, E. 1960. The maidenhair ferns (Adiantum) of Costa Rica. Contributions from the Gray Herbarium of Harvard University
187: 3–22.

162
Figure 75. A, B. Adiantum feei. C, D. Adiantum tricholepis. E, F. A. patens. G, H. A. galeottianum. (Mickel & Smith,
2004).

163
POLYPODIACEAE
Alansmia M. Kessler, Moguel, Sundue & Labiak
DESCRIPTION: Epiphytic; rhizomes setose; rhizome DESCRIPCIÓN: Epífitas; rizomas setosos; escamas
scales usually orange to castaneous, not clathrate, bas- del rizoma usualmente anaranjadas a castañas, no clatra-
ifixed (not cordate), hairy on margins and often both das, basifijas (no cordadas), peludos en los márgenes y
surfaces, or sometimes absent (T. alfarii), the cells tur- a menudo los superficies, o a veces ausentes (T. alfarii),
gid; leaves pendulous, indeterminate; blades lacking las células turgidas; hojas péndulas, indeterminadas;
black clavate fungi; stipes usually less than 1 cm long, the láminas se carecen de hongos claviformes negros; pecíolos
pinnae reduced nearly to the base of the leaf; laminae pin- usualmente menos de 1 cm de largo, las pinnas reducidas
natisect to 1-pinnate, with tendency for paired or clus- casi a la base de la hoja; láminas pinnatisectas a 1-pinnadas,
tered setae; veins simple, free; hydathodes present, often con tendencia tener setas pareadas o agrupadas; venas
cretaceous; sori separate, round, superficial (i.e., not sunk- simples, libres; hidatodos presentes, a menudo cretásicos;
en in the lamina); sporangial capsules setose; spores soros separados, redondos, superficiales (i.e., no hundidos
green, trilete and/or monolete; x=37. en la lámina); cápsulas esporangiales setosos; esporas
verdes, triletes y/o monoletes; x=37.

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

164
Figure 76. A, B. Alansmia cultrata. C–H. A. spathulata. J, K. A. senilis. (Mickel & Smith, 2004).

165
CYATHEACEAE
Alsophila R. Br.
DESCRIPTION: Terrestrial; stems arborescent; sterile DESCRIPCIÓN: Terrestres; tallos arborescentes; ho-
and fertile leaves 1–4.4 m long, monomorphous; petioles jas estériles y fértiles 1–4.4 m de largo, monomorfas;
smooth or with stout black spines, scaly, the scales with pecíolos lisos o con espinas negras fuertes, escamo-
black setae (at least apically), concolorous to bicolor- sos, las escamas con setas negras (a lo menos en
ous, usually the marginal cells different in shape and orien- el ápice), concoloras a bicoloras, las células marginales
tation from the central ones; laminae 1-pinnate-pinnatifid difieren en forma y orientación de las centrales; láminas
to 4-pinnate, the apex pinnatifid or rarely conform; veins 1-pinada-pinatífidas a 4-pinadas, el ápice gradualmente
free; axes pubescent adaxially, the hairs thick, strigose, mul- reducido o raras veces conforme; nervaduras libres; ejes
tiseptate; sori round, the receptacle elevated, subglobose; pubescentes adaxialmente, los pelos estrigosos, gruesos,
indusia present or absent, scalelike or globose, minutely estrigosos, multiseptados; soros redondos, el receptácu-
scaly; spores 16 per sporangium, tetrahedral-globose, lo elevado, subgloboso; indusios presentes o ausentes,
without large equatorial pores; x=69. escuamiformes o globosos, diminutamente escamosos;
esporas 16 por esporangio, tetrahédricas-globosas sin
poros ecuatoriales grandes; x=69.

SIMILAR GENERA: Gymnosphaera differs by the pres- GÉNEROS PARECIDOS: Gymnosphaera difiere por la
ence of small, skeletonized pinnae (aphlebia) near the presencia de pinas pequeñas en esqueletos (aflebias) cer-
petiole base, and 64 spores per sporangium. Other Cy- ca la base del pecíolo y 64 esporas por esporangio. Otros
atheaceae differ by lacking a black seta at the apex of the Cyatheaceae difieren por la ausencia de una seta negras
petiole scale. Sphaeropteris further differs by conform en el ápice de la escama del pecíolo. Sphaeropteris difiere
petiole scales (cells with the same shape and orientation). 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 casi 200 especies. Existe en bosques sombreados. Nephe-
that has often been used for many species of Alsophi- lea, un nombre que ha sido usado a menudo en los trópi-
la in the American tropics is Nephelea It here placed in cos americanos, está ubicado aquí en sinonimia por que
synonymy because there are no constant differences no hay diferencias constantes entre él y Alsophila y varios
between it and Alsophila and several hybrids are known híbridos y especies de origen híbrido son conocidos en-
between the two genera. The 2-pinnate-pinnatifid spe- tre los dos. Las especies 2-pinnadas de Alsophila tienen
cies of Alsophila have black spines on the petioles. They espinas negras en los pecíolos. Producen ramas laterales
produce narrow lateral branches at the base of the trunk angostas en la base del tronco que crecen hacía el suelo
that grow into the soil where they eventually form erect donde eventualmente forman tallos erectos. Estas espe-
stems. Thus the plants form colonies. The genus name is cies forman colonias. El nombre genérico se derive del
derived from the Greek alsos, grove + philein, to love. It griego alsos, arboleda + philein, amar. Se refiere a al hecho
refers to the shade-loving habit of the genus. 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.

166
Figure 77. A–C. Alsophila cuspidata. F–H, J–L, A. firma. M–R. A. tryoniana (A-C, @R. C. Moran 2010; F–R from
Mickel & Smith, 2004)

167
PTERIDACEAE
Ananthocorus Maxon & Underwood
DESCRIPTION: Epiphytic or epipetric; rhizomes DESCRIPCIÓN: Epífitas o epipétricas; rizomas cor-
short- creeping solenostelic, short-creeping, dor- tamente rastreros, solenostélicos, dorsiventrales; escam-
siventral; rhizome scales clathrate; leaves 15–45 × as del rizoma clatradas; hojas 15–45 × 0.9–1.5 cm,
0.9–1.5 cm, simple, entire, glabrous, coriaceous, dis- simples, enteras, glabras, coriáceas, dísticas, ama-
tichous, clumped; petioles absent or very short and colladas; pecíolos ausentes o muy cortos y achata-
flattened; costae present; veins areolate, 2 or 3 rows of dos; costas presentes; nervaduras areoladas, en 2 o 3
areoles between the costa and margin; so r i sub- hileras de areolas entre la costa y el margen;
marginal on each side of the costa, nearly continuous; so r o s submarginales en cada lado de la costa,
indusia absent; paraphyses abundant, slender, without casi continuous; indusios ausentes; parafisos abundantes,
a swollen apical cell; spores monolete, nongreen; game- delgados, sin una célula apical hinchada; esporas
tophytes lacking gemmae; x=120. monoletes, no verdes; gametofitos sin yemas; x=120.

SIMILAR GENERA: Radiovittaria and Vittaria differ by GÉNEROS PARECIDOS: Radiovittaria y Vittaria
having only one row of areoles between the costa and difieren por tener una sola hilera de areolas entre
margin (vs. 2 or 3) and by the presence of gemmae la costa y el margen (vs. 2 o 3) y por la presencia de
on the gametophytes. gemas en el gametofito.

COMMENTS: Consists of only one species, Anan- COMENTARIOS: Consta de una sola especie, Anan-
thacorus angustifolius (Sw.) Underw. & Maxon, wide- thacorus angustifolius (Sw.) Underw. & Maxon,, difundido
spread from Mexico to Bolivia. It occurs as an desde México hasta Bolivia. Ocurre como epífito
epiphyte in wet or semi-deciduous forests from 50– en bosques húmedos o semi-deciduous de 50–1500
1500 m. Ananthacorus is the only vittarioid fern whose m. Ananthacorus es el único helecho vittarioide cuy-
gametophytes lack gemmae. Its gametophytes bear an- os gametofitos faltan de yemas. Sus gametofitos llevan
theridia scattered over the surface instead of being pre- anterídios salpicados por la superficie en lugar de estar
dominantly on small plants or on germinating principalmente en plantas pequeñas o en yemas germi-
gemmae as in other vittarioid ferns (Farrar, 1974). The nantes como en otros helechos vittarioides (Farrar,
rows of aeroles between the midrib and margin are 1974). Las hileras de areolas entre la costa y el margen
often hard to see because of the thick lamina. The ge- son a menudo difícil ver a causa de la lámina gruesa.
nus name is derived from the Greek an, without + El nombre genérico se derive del griego an, sin +
anthos, flower + Acorus, Sweetflag. Resembling Acorus anthos, flor + Acorus. Se asemeja Acorus per sin flores.
but without flowers.

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.

168
Figure 78. A–D. Ananthacorus angustifolius. (Mickel & Smith, 2004)

169
PTERIDACEAE
Anemia Sw.
DESCRIPTION: Terrestrials or saxicolous; rhizomes DESCRIPCIÓN: Terrestres o saxícolas; rizomas com-
compact, short-creeping or ascending, densely pu- pactos, cortamente rastreros o ascendentes, densa-
bescent, the hairs brown or orange; sterile and mente pubescentes, los pelos pardos o naranja-
fertile leaves hemidimorphic, rarely holodimorphic; dos; hojas estériles y fértiles hemidimórfas, raras veces
laminae pinnatifid to 4-pinnate; basal pinnae typical- holodimórfas; láminas pinnatífidas a 4-pinnadas; pinnas
ly modified into 2 erect, long-stalked fertile basales típicamente modificadas en 2 espigas fér-
spikes; veins free or areolate; sporangia cylindrical tiles lárgamente pedunculadas, erectas, (la hoja
or pear- shaped with an apical annulus; spores raras veces es completamente fértile); nervaduras
tetrahedral- globose, striate with parallel smooth or libres o areoladas; esporangios cilíndricos o piriformes
spiny ridges; x = 38. con un anillo apical; esporas tetraédricas-globosas,
estriadas con filas paralelas lisas o espinosas; x = 38.

SIMILAR GENERA: Botrychium differs by erect GÉNEROS PARECIDOS: Botrychium difiere por
rhizomes, fleshy leaves, a single fertile spike, and glo- rizomas erectos, hojas carnosas, única espiga fértil e es-
bose sporangia. Ornithopteris differs by longer-creeping, porangios globosos. Ornithopteris difiere por rizomas
solenostelic rhizomes, blackish rhizome hairs, and más largamente rastreros, solenostélicos, pelos de los
distichous leaves. rizomas negruzcos y hojas dísticas.

COMMENTS: Anemia contains about 125 spe- COMENTARIOS: Anemia consta de alrededor de
cies, mostly in the Neotropics and Africa. It is most 125 especies principalmente en el Neotrópico y
diverse in Brazil (80 species) and Mexico (20 species). África. Es más diverso en Brasil (80 especies) y
Nearly all species occur in dry, open or semi-open hab- México (20 especies). Casi todas las especies ex-
itats. They often hybridize when growing together. isten en habitats secos, abiertos, o semi-abiertos. A
Ploidy levels in the genus range from 2x to 14x. Or- menudo se hibridizan cuando en poblaciones mixtas.
nithopteris was formerly included in Anemia as subgen. Niveles de poliplodea en el género varian desde 2x
Anemiorrhiza, but unpublished molecular phylogenetic hasta 14x. Ornithopteris fue anteriormente incluído en
evidence (Labiak & Mickel, pers. comm..) suggests it is Anemia como el subgén. Anemiorrhiza, pero evidencia
best treated at the rank of genus. Mohria, an African ge- molecular filogenético sugiere que se trata mejor
nus with scales instead of hairs, is sister to Anemia al rango de género. Mohria, un género Africano
(Skog et al., 2002). The genus name is derived from con escamas en lugar de pelos, está hermana a
the Greek aneimon, naked, and alludes to the exposed Anemia (Skog et al., 2002). El nombre genérico se
sporangia. deriva del griego aneimon, desnudo, y refiere a los
esporangios expuestos.

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. 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. Skog, J., E. A. Zimmer & J. T. Mickel. 2002. Additional support for two
subgenera of Anemia (Schizaeaceae) from data for the chloroplast intergenic spacer region trnL-F and morphology. American
Fern Journal 92: 119–130.

170
Figure 79. A, B. Anemia munchii. C, D. A. phyllitidis. E, F. A. hirsuta × A. phyllitidis. (Mickel & Smith, 2004)

171
PTERIDACEAE
Anetium Splitg.
DESCRIPTION: Epiphytic or rarely rupicolous; roots DESCRIPCIÓN: Epífitas o raramente rupícolas; raíces
numerous, densely hairy, often golden; rh i z o m e s numerosas, densamente pelosas, a menudo doradas; ri-
long- creeping, dorsivental, densely scaly, the scales zomas largamente rastreros, dorsiventrales, densamente
clathrate; petioles absent or very short; laminae oblan- escamosos, las escamas clatradas; pecíolos ausentes
ceolate to narrowly elliptic, entire simple, pendulous, o muy cortos; láminas oblanceoladas a estrechamente
slightly succulent, glabrous, with spicular idoblasts elípticas, simples, enteras, péndulas, algo suculen-
(false veins) adaxially; ve i n s areolate, with many tas, glabras, con idioblastos cortos, espiculares, visibles
rows or areoles between the costa and margin, the are- en el haz; ner v adur as anastomosadas, con muchas
oles without included veinlets; sori subacrostichoid, hileras de aréolas entre la costa y el margen, sin nér-
the sporangia dispersed individually or in small vulos libres incluidos; soros subacrosticoides, los espor-
isolated groups on and between the veins; indusia angios dispuestos individualmente o en pequeños
absent; paraphyses absent; spores trilete; gameto- grupos aislados sobre y entre las nervaduras;
phytes with paired gemmae; x=60. inducíos ausentes; parafisos ausentes; esporas triletes; ga-
metofitos con yemas pareadas; x=60.

SIMILAR GENERA: Polytaenium differs by erect GÉNEROS PARECIDOS: Polytaenium difiere por
or short-creeping rhizomes and sori in distinct lines, not rizomas cortamente rastreros y soros en líneas dis-
subacrostichoid. Elaphoglossum differs by non-clathrate tintos, no subacrostichoides. Elaphoglossum difiere por
rhizome scales, free veins, and (usually) the presence of escamas del rizoma no clatradas, venas libres y (usual-
phyllopodia. Many simple-leaved Polypodiaceae differ mente) la presencia de filopodios. Muchos Polypodia-
by the presence of phyllopodia, included veinlets in ceae con hojas simples difieren por la presencia de filo-
the areoles, and sori round. podios, venas incluidas en las areolas y soros redondos.

COMMENTS: Anetium consists of a single species, A. COMENTARIOS: Anetium consta de una sola espe-
citrifolium (L.) Splitg., which occurs from Mexico cie, A. citrifolium (L.) Splitg., la cual existe desde México
to southern Brazil and Bolivia, and the Greater An- hasta el sur de Brasil y Bolivia, y en las Antillas Mayores.
tilles. It grows as an low-trunk epiphyte in wet forests Crece como un epífito en las partes inferiores de los
generally below 500 m. The rhizome scales are troncos de bosques húmedos generalmente bajo de
a beautiful example of the clathrate condition and are 500 m. Las escamas del rizoma son un ejemplo bellísimo
often highly iridescent. Anetium is sister to Polytaenium de la condición clatrada y a menudo son altamente
(Crane, 1997). The genus name is derived from iridiscente. Anetium es hermana a Polytaenium (Crane,
the Greek a, without + notos, heap. The sporangia 1997). El nombre genérico se deriva del griego a,
are scattered, not in sori. sin + notos, pila. Los esporangios están difundidos, no
en soros.

LITERATURE: 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.

172
Figure 80. A–D. Anetium citrifolium. (Mickel & Smith, 2004)

173
PTERIDACEAE
Anogramma Link
DESCRIPTION: Terrestrial; rhizomes inconspicuous, DESCRIPCIÓN: Terrestres; rizomas inconspicuos, erec-
erect, hairy (not scaly); leaves 5–25 cm long, fasciculate, tos, pelosos (no escamosos); hojas 5–25 cm de largo,
annual; petioles ca. 1 mm wide, as long or slightly lon- fasciculadas, anuales; pecíolo c. 1 mm de ancho, tan largo
ger than the lamina, green to stramineous; laminae elon- como la lámina o escasamente más largo que ella, verdes
gate-triangular, 2- to 4-pinnate, not farinose, anadromic, o pajizos; lámina alargado-triangular, 2- a 4-pinnada, no
thin; terminal segments 0.5–2 mm wide; rachises grooved farinosas, anádroma, delgada, glabra; segmentos termina-
adaxially; veins free; sori borne along the veins, without les 0.5–2 mm de ancho; ráquises surcados adaxialmente;
parafises; indusia absent; spores tetrahedral-globose, nervaduras libres; soros a lo largo de las nervaduras,
trilete, with an equatorial ridge and two parallel flanges; sin parafisos; indusio ausente; esporas tetraédrico-glo-
x=29. bosas, triletes, con una fila ecuatorial y dos costillas para-
lelas; x=29.
SIMILAR GENERA: Pityrogramma differs by scaly rhi- GÉNEROS PARECIDOS: Pityrogramma difiere por ri-
zomes and laminae usually farinose beneath. Cystopteris zomas escamosos y las láminas usualmente farinosas en
and Woodsia have round sori covered by indusia. el envés. Cystopteris and Woodsia tienen soros redondea-
dos cubiertos por indusios.

COMMENTS: Anogramma consists of two pantropical COMMENTARIOS: Anogramma consta de dos espe-
species: A. leptophylla (L.) Link and A. lorentzii (Hieron.) cies 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 Anogramma lugares semi-abiertas de 1000–2800 m. Dos especies
are now placed in Pityrogramma (i.e., P. chaerophylloides previamente en Anogramma ahora están ubicadas en
(Desv.) Domin and P. novogaliciana (Mickel) comb. ined.) Pityrogramma (i.e., P. chaerophylloides (Desv.) Domin y P.
Anogramma is sister to Cosentinia, a monotypic genus of novogaliciana (Mickel) comb. ined.). Anogramma es her-
the Mediterranean- Himalayan region, and these two mana a Cosentinia, un género monotípico de la región
genera are sister to Pityrogramma (Nakazato & Gastony, Mediterráneo a las Amalhayas, y estos dos géneros son
2003). The gametophytes of Anogramma have several un- hermana a Pityrogramma (Nakazato & Gastony, 2003).
usal characteristics. The young ones are spathulate, not Los gametofitos tienen varias características inusuales.
cordate, and meristematic activity is from a lateral meri- Los jóvenes son espatulados, no cordados, y la actividad
stem, not an apical one. These characters are shared with meristamática es de un meristemo lateral, no apical. Es-
Cosentinia (Pangua & Vega, 1996). If fertilization or spo- tos rasgos se compartan con Cosentinia (Pangua & Vega,
rophyte production does not occur, the gametophytes 1996). Si no toma lugar la fecundación ni producción del
can survive an unfavorable growing season by means of esporofito, los gametofitos pueden sobrevivir por una
a tubercle formed by elaborating the thickened archego- temporada desfavorable por medio de un tubérculo
nial region or cushion. When favorable growth conditions desarrollado por elaborar la región aquegonial o “cojín”.
return, the tubercle regenerates new lobes. The game- Cuando condiciones favorables regresan, el tubérculo
tophytes cannot further develop if a new sporophyte regenera lóbulos nuevos gametofíticos. Los gametofitos
is produced. Thus, the gametophytes perennate only in a no pueden desarrollarse más si un nuevo esporofito se
limited sense. The genus name is derived from the Greek produce. Por eso, los gametofitos perenan solo en un
ano, upward + gramme, line. It refers to the elongate sori sentido limitado. El nombre genérico se deriva del griego
on the terminal segments. ano, hacia arriba + gramme, línea. Refiere a los soros alar-
gados en los segmentos terminales.

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. 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.

174
Figure 81. A–C. Anogramma leptophylla. D, E. A. lorentzii. (A–C from Mickel & Smith, 2004; D, E, © R. C. Moran, 2010)

175
DRYOPTERIDACEAE
Arachniodes Adans.
DESCRIPTION: Terrestrial; rhizomes creeping; leaves DESCRIPCIÓN: Terrestres; rizomas rastreros es-
monomorphic; petioles with more than three vascu- camosos; hojas monomorfas; pecíolo con tres o más
lar bundles; laminae widely deltate or pentagonal, 2- to haces vasculares; lámina ampliamente deltada o pentag-
5-pinnate; medial pinnae prolonged acroscopically, the onal, 2-5-pinnada; pinnas medias prolongadas acroscópi-
basal basiscopic side cuneate; basal pinnules anadrom- camente, el lado basal basiscópico cuneado; pínnulas
ically arranged, stalked; rachises and costae with dispuestas anadrómicamente, pediculadas; raquis y
grooves glabrous, without proliferous buds; veins free; costas con surcos glabros, sin yemas prolíferas; ner-
sori round; indusium orbicular-reniform; spores mono- vaduras libres; soros redondos; indusio orbicular-reni-
lete; x=41. forme; esporas monoletes; x=41.

SIMILAR GENERA: Dryopteris differs by rhizomes GÉNEROS PARECIDOS: Dryopteris difiere por rizo-
erect or ascending, pinnules catadromic, and (in the mas erectos o ascendentes, pínnulas catadrómicas y (en
Neotropics) leaves often capitate-glandular. Polystichum el Neotrópico) hojas a menudo glanduloso-capitadas.
differs by the basal acroscopic pinnules slightly elongate, Polystichum difiere por las pínnulas basales basiscópicos
and generally the margins of the pinnules are spinulose. alargados, y generalmente los márgenes de las pínnulas
Lastreopsis and Rumohra differ by a thickened marginal son espinulosos. Lastreopsis y Rumohra difieren por una
ridge of the costa or costules surcurrent along the green fila marginal engrosada de la costa o cóstula surcurrente
laminar margin of the segment of the next highest order por el margen del tejido laminar verde del segmento del
(not adaxially along the midrib of the costule). Saccoloma próximo orden más alto (no adaxialmente por la cóstula
differs by marginal sori and a single, continuous (not divid- del segmento). Saccoloma difiere por soros marginales y
ed) omega-shaped vascular bundle in the petiole. Pradoia un haz vascular continuo (no dividido) en forma de ome-
differs by adaxial grooves of rachises and costae pubes- ga en el pecíolo. Pradoia difiere por los surcos adaxiales
cent, as least at the junctures. de los raquises y costas pubescentes, al menos en las
uniones.
COMMENTS: Arachniodes is pantropical with about 50 COMENTARIOS: Arachniodes es pantropical con casi
species, most of which are Asian. In the Neotropics it con- 50 especies, la mayoría de las que son Asiáticas. En el
tains about 4 species, and these occur mostly above 2000 Neotrópico contiene alrededor de 4 especies, e estas
m. The most widespresd species is A. denticulata (Sw.) existen principalmente arriba de 2000 m. La especie
Ching), which highly variable. Christensen (1920) listed más difundida es A. denticulata (Sw.) Ching), la cual es
numerous forms and varieties of this species, and many of muy variable. Christensen (1920) listó numerosas formas
these probably represent distinct taxa, but biosystemat- y variedades de él, y muchos de estos probablemente
ic work is needed. The genus name is derived from the representan taxones distintos, pero se necesita trabajo
Greek arachnion, spider’s web + odes, having the form or biosistemático. El nombre genérico se deriva del griego
nature of. What it refers to is unknown. arachnion, tela de araña + odes, con la forma o la carac-
terística de algo. Se desconoce a lo que se refiere.

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.

176
Figure 82. A, B. Arachniodes denticulata. (Mickel & Smith, 2004)

177
PTERIDACEAE
Argyrochosma (J. Sm.) Windham
DESCRIPTION: Terrestrial or rupestral; rhizomes DESCRIPCIÓN: Terrestres o rupícolas; rizomas cor-
creeping or erect, scaly, the scales glandular, commonly tamente rastreros o erectos, escamosos, las escamas
not sclerotic, concolorous or slightly bicolorous; leaves glandulares, comúnmente no escleróticas, concoloras o
2-4 pinnate, with separate segments all the way to ligeramente bicoloras; hojas 2-4-pinnadas, 2-4-pinnadas,
the apex (not becoming confluent in a pinnatifid con segmentos separatas hacia el ápice (no llegan-
apical portion), monomorphic; petiole with a single vas- do a ser confluentes en una parte apical pinnatífi-
cular bundle; pinnules short-stalked or narrowed toward da), monomorfas; pecíolo con un haz vascular; pínnulas
the base, typically white or yellow farinose beneath; cortamente pediculadas o basalmente constrictas, típica-
veins free, the tips not or only weakly thickened; sori sub- mente blancas o amarillo-cerosas en el envés; ner-
marginal along the distal 1/4-2/3 of each vein, partially vaduras libres, las puntas débilmente o no engrosadas;
hidden by the weakly to strongly revolute margins of the soros submarginales a lo largo del 1/4-2/3 apical de cada
pinnules; spores trilete; x=27. nervadura, ocultos parcialmente por los márgenes débil
a fuertemente reflejos de las pínnulas; esporas triletes;
x=27.

SIMILAR GENERA: Pellaea differs by nonglandular rhi- GÉNEROS PARECIDOS: Pellaea difiere por escamas
zome scales and nonfarinose laminae. Cheilanthes also del rizoma no glandulares. Cheilanthes difiere también por
differs by nonfarinose lamina. Notholaena differs by bi- láminas nofarinosas. Notholaena difiere por escamas del
colorous (centrally sclerotic) rhizome scales. rizoma bicoloras (escleróticas en la parte central).

COMMENTS: Argyrochosma is exclusively American COMENTARIOS: Argyrochosma es género exclusiv-

genus of about 20 species. It center of diversity is the amente americano de casi 20 especies. Su centro de
Chihuahuan desert region of northern Mexico and ad- diversidad es la región del Desierto Chihuahuense del
jacent Texas, where it has 8 species. Most of the species norte de México y el área adyacente de Texas, donde
are apogamous and grow on or among rocks. The base se encuentra 8 especies. La mayoría de las especies son
chromosome number of x=27 is unique among cheilan- apógamas y crecen sobre o entre rocas. El número cro-
thoid ferns. Argyrochosma is sister to Pellaea and Astrolepis. mosómico básico de x=27 es único entre los helechos
The gametophytes of Argyrochosma lack farina, where as cheilantoides. Argyrochosma es hermana a Pellaea y Astro-
those of Notholeana have it (Windham, 1987). The gener- lepis. Los gametófitos de Argyrochosma carecen de una fa-
ic name comes from the Greek argyros, silver, + chosma, rina, mientras que los de Notholaena la tienen (Windham,
poweder. It refers to the white (silvery) powder on the 1987). El nombre genérico se deriva del griego argyros,
abaxial surface of the laminae. plata, + chosma, polvo. Se refiere al polvo blanco (platea-
do) en el envés de las láminas.

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 revi-
sion of the American species of Nothlaena. Contributions from the Gray Herbarium of Harvard.

178
Figure 83. A-C. Argyrochosma incana. D-G. A. delicatula. E-G. A. delicatula. H, J. A. incana. K, L. A. fendleri. (Mickel & Smith,
2004).

179
POLYPODIACEAE
Ascogrammitis M. Sundue
DESCRIPTION: Plants epiphytic, rarely saxicolous; DESCRIPCIÓN: Plantas epífitas, raras veces saxícolas;
rhizomes short-creeping, dorsiventral, unbranched; rhi- rizomas cortamente rastreros, dorsiventrales, no ram-
zome scales usually clathrate, the cells flat (not turgid ificados; escamas del rizoma usualmente clatradas,
or inflated), margins usually reddish setose, sometimes las células planas (no túrgidas o hinchadas), márgenes
entire or provided only with a single apical seta; leaves usualmente setosos-rojizos, a veces enteros o con
pubescent and setose, the fertile portions more densely una seta apical; hojas pubescentes y setosas, las partes
setose than sterile portions, usually bearing a black, fértiles más setosas que las estériles, usualmente por-
clavate fungus ca. 1 mm long; laminae 1-pinnatisect tando un hongo negro clavado pequeño (ca. 1 mm
to 1-pinnate; veins greenish (not dark); hydathodes long); láminas 1-pinnatisectas a 1-pinnadas; venas
present, usually cretaceous or not; sori slightly eliptic, verdosas (no oscuros); hidatodos presentes, usual-
sometimes provided with receptacular setae; paraphyses mente cretásicos; soros levemente elípticos, a veces con
absent; sporangial capsules glabrous; spores trilete, green; setas receptaculares; cápusulas esporangiales glabras es-
x=37? poras triletes, verdes; x=37?

SIMILAR GENERA: Lellingeria differs by rhizome scales GÉNEROS PARECIDOS: Lellingeria difiere por las es-
with hyaline marginal setae, fronds that lack reddish se- camas del rizoma con setas marginales hialinas, hojas que
tae, costae not visible on the abaxial side of the blade, faltan setas rojizas, costas no visibles en el envés de la lámi-
and hairs with an acicular rather than setiform branch. na y pelos con una rama acicular en lugar de setiforme.
Mycopteris differs by concolorous, orange to castaneous Mycopteris difiere por escamas de rizoma concoloras,
rhizome scales with turgid (swollen) cells, leaves not more anaranjadas a castaneas con células túrgidas (hichadas),
densely setose in the fertile parts, and blackish veins. hojas no más densamente setosas en las partes fértiles
Melpomene differs by reddish-brown scales with entire y venas negruzcas. Melpómene difiere por escamas ro-
margins and frequently with multiple apical papillate cells. jizas-pardas enteras y frequentamente con varias células
Dried specimens have a sweet, spicy odor. Pecluma differs apicales papilosas. Ejemplares secas tienen un olor dulce
by monolete yellowish spores and lack of foliar setae. especiosa. Pecluma difiere por esporas monoletes amaril-
lentosas y la falta de setas foliares.

COMMENTS: Ascogrammitis contains about 17 species. COMENTARIOS: Ascogrammitis consta de ca. 17 epe-
It is entirely neotropical, occurring in the Greater Antil- cies. Es completamente neotropical y existe en las Antillas
les, Guadeloupe, and Mexico to Bolivia. The species occur Mayores, Guadeloupe, México hasta Bolivia. Las especies
in cloud forests usually from 1800–3200 m. The fungus inhabitan bosques nublados usualmente desde 1800–
Acrospermum maxonii is a nearly constant associate of the 3200 m. El hongo Acrospermum maxonii es casi constan-
genus. Its mycelia are whitish and the spore cases (as- temente asociado con el género. Sus micelias son blan-
comes) are black, clavate, and ca. 1 mm long. The fungus quecinas y las esporangios (ascomas) son negras, clavadas
does not appear to harm the fern in any way. The genus y ca. 1 mm de largo. El hongo no parece dañar el helecho
name refers to the ascomes (Greek, asco-, bag or blad- en ninguna manera. El nombre genérico refiere a las as-
der) of the fungal associate. 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.

180
Figure 84. Ascogrammitis david-smithii A. Habit. B. Rhizome scale. C. Detail of abaxial lamina surface, note ascomes of Acros-
permum. Ascogrammitis colombiensis D. Habit. E. Rhizome scale. F. Detail of abaxial lamina surface including ascomes of Acros-
permum. Ascogrammitis cuencanana G. Habit. H. Detail of adaxial lamina surface. J. Detail of abaxial lamina surface, note ascome
of Acrospermum. (From Sundue, 2008).

181
ASPLENIACEAE
Asplenium L.
DESCRIPTION: Epiphytic, terrestrial or saxicolous; DESCRIPCIÓN: Epífitas, terrestres o saxícolas; tallos
stems erect to creeping, scaly, the scales clathrate; sterile erectos a rastreros, escamosos, las escamas clatradas;
and fertile leaves monomorphous, rarely subdimorphous; hojas estériles y fértiles monomorfas, raras veces subdi-
petioles with 2 vascular bundles basally which unite to- morfas; pecíolos con 2 haces vasculares en la base los
ward the distally to form a single X-shaped vascular bun- cuales se unen distalmente formando un solo haz vas-
dle; laminae simple to 4-pinnate, anadromic; axes of the cular en forma de “X”; láminas simples a 4-pinnadas,
lamina grooved, the grooves decurrent, glabrous; veins anádromas; ejes de la lámina surcados, los surcos decur-
free; sori elongate to linear, borne singly (not double) rentes, glabros; venas libres; soros alargados a lineares,
per vein indusiate; annulus of the sporangia consisting nacen uno por vena (no doblados) indusios; anillo del
of 20-28 cells; spores bilateral, brown; x=36. esporangio consiste de 20-28 células; esporas bilate-
rales, pardas; x=36.

SIMILAR GENERA: Asplenium and Diplazium can be GÉNEROS PARECIDOS: Asplenium y Diplazium
difficult to distinguish; nevertheless, three characteristics pueden ser difíciles de distinguir; sin embargo, tres car-
consistently separate them.The first is the number of cells acterísticas los separan consistentemente. La primera
in the annulus: Asplenium has 20-28, Diplazium has 15-20. es el número de células en el anillo: Aplenium tiene 20-
The second is the number of rows of cells in the stalk 28, Diplazium tiene 15-20. La segunda es el número de
of the sporangium: Asplenium has only 1, Diplazium has células en el pecíolo del esporangio: Asplenium tiene 1,
2 or 3. The third is the form of the vascular bundle(s) in Diplazium tiene 2 o 3. La tercera es la forma del haz
the distal part of the petiole: in Asplenium it is X-shaped, vascular en la parte distal del pecíolo: en Asplenium forma
in Diplazium the two strands are separate and elliptic. una ”X”, en Diplazium forma dos haces distintos, elípti-
Unfortunately, the characteristics cited above are often cos. Lamentablemente, las características arribas citadas
difficult to observe. There are two other characteristics son a menudo difíciles de observar. Hay otras dos car-
that, although not always as constant as the above, are acterísticas que aunque no son tan constantes (como las
often helpful. The rhizome scales of Asplenium are usu- anteriores), a menudo son útiles. Las escamas del rizoma
ally clathrate; those of Diplazium are never clathrate. The de Asplenium usualmente son clatradas; las de Diplazium
sori of Asplenium are almost always borne singly along nunca son clatradas. Los soros de Asplenium casi siempre
a single vein; those of Diplazium are often paired back nacen únicamente por una sóla vena; las de Diplazium
to back along a single vein. See Diplazium for additional a menudo nacen pareados, en par en par, por una sóla
comments. 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- de aproximadamente 700 especies, haciéndolo uno de
est fern genera in the world. Section Hymenasplenium is los más grandes géneros de helechos en el mundo. A vec-
sometimes segregated as distinct because it is sister to es la sección Hymenasplenium se reconoce como distinto
the rest of Asplenium. The genus name comes from the por que es hermana el resto de Asplenium. El nombre
Greek splen, spleen. Dioscorides thought a fern with this genérico se deriva del griego splen, esplín. Dioscorides
name was useful in treating diseases of the spleen. 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.

182
Figure 85. Asplenium. Note linear sori and clathrate rhizome scales. Fijase en los soros lineares y escamas del rizoma clatra-
das. (©Robbin Moran, 2009)

183
ASPLENIACEAE
Asplenium (Loxoscaphe T. Moore)
DESCRIPTION: Epiphytic; rhizomes short-creeping DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros
to suberect, the scales clathrate; leaves 5–15 cm a suberectos, las escamas clatradas; hojas 5–15 cm de
long, clumped, monomorphic; petioles scaly, with two largo, amacolladas, monomorfas; pecíolos escamosos,
vascular bundles at the base; blades finely dissected, con 2 haces vasculares en la base; láminas finamente
2-4-pinnate; segments slender, linear, fleshy; veins free; disectadas, 2-4-pinnadas; segmentos delgados, lineares,
sori near the margin, protected by a pouch-like indusia carnosos; margen de la lámina engrosado y decurrente
that opens toward the margin; spores bilateral, brown; por el raquis; venas libres; soros cerca del margen,
x=35. 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: Loxoscaphe contains a single species COMENTARIOS: Loxoscaphe contiene una sola
in the Neotropics: L. thecifera (Kunth) T. Moore. (=A. especie en el Neotrópico: L. thecifera (Kunth) T. Moore.
theciferum (Kunth) Mett. It is an epiphyte in wet forests Es un epífito en bosques húmdos de 1200–1900 m. La
from 1200–1900 m. The species is represented in Africa especie está representada en África por la var. concinnum
by var. concinnum (Schrad.) Schelpe. The only difference (Schrad.) Schelpe. La única diferencia es el lóbulo apical
is the short or absent apical lobe that extends beyond corto o ausente que extiende más alla el soro en las
the sorus in the American plants; this lobe is absent or plantas Americanas; este lóbulo es ausente o muy corto
very short in the African plants.Loxoscaphe belongs to en las plantas Africanas.Loxoscaphe pertenece a Asplenium
Asplenium subg. Daraea T. Moore, a group primarily of subg. Daraea T. Moore, un grupo principalmente de África
Africa and Asia whose species have finely dissected fronds y Asia, cuyas especies tienen láminas finamente disectadas
with narrow segments and a single sorus parallel to the con segmentos angostos y un solo soro parallelo a la vena
vein and margin, often reaching the latter. Loxoscaphe y el margen, a menudo alcanzando el margen. Loxoscaphe
thecifera differs only by pocket-like indusia, as wide as thecifera difiere solo por los indusios en forma de bolsa,
long, and adnate on both sides to the laminar tissue.The tan ancho como largo, y adnato en ambos lados al tejido
name is derived from the Greek loxos, oblique + skaphe, laminar. El nombre se deriva del griego loxos, oblicuo
a small boat or bowl. The bowl-shaped sori are oblique + skaphe, una barca pequeña o escudilla. Los soros en
to the costae. forma de escudilla son oblicuos a la costa.

LITERATURE: Morton, C. V. & D. B. Lellinger. 1966. The Polypodiaceae subfamily Asplenioideae in Venezuela. Memoirs of
the New York Botanical Garden 15: 1–49.

184
Figure 86. A. Asplenium (Loxoscaphe) theciferum. (Mickel & Beitel, 2004)

185
ASPLENIACEAE
Asplenium sect. Hymenasplenium (Hayata) Iwatsuki
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: Asplenium sect. Hymenasplenium is pan- COMENTARIOS: Asplenium secc. Hymenasplenium es

tropical and contains ca. 40 species, of which 11 are neo- pantropical y consta de ca. 40 especies, de las cuales 11
tropical. In the Americas it extends from Mexico and the son neotropicales. En las Américas se extiende desde
Antilles to southern Brazil. It occurs in wet forests, gen- México y las Antillas hasta Brasil del sur. Existe en bosques
erally below 2000 m, and is remarkable for the variety of húmedos, generalmente abajo de 2000 m, y es destacado
growth habits among its species (terrestrial, saxicoulous, por la variedad de hábitos de crecimiento entre sus espe-
hemiepiphytic) and diversity of spore types.Section Hy- cies (terrestre, saxícola, hemiepífita) y diversidad de tipos
menasplenium is sister to the rest of Asplenium; therefore, de las esporas. La secc. Hymenasplenium es hermana al
sometimes ranked as a genus. The leaves resemble those resto de Asplenium; por eso, a veces se reconoce al nivel
typical of sect. Asplenium, but the creeping rhizomes with genérico. Las hojas se asemejan a ellas que son típicas
distichous leaves and swollen petiole bases are striking- de secc. Asplenium, pero obviamente diferente son los
ly different. The chromosome number of the section is rizomas reptantes con hojas dísticas y bases hinchados
x=38 or 39, in contrast to that in sect. Asplenium which de los pecíolos. El número cromosómico en la sección
is uniformly x=36. The section name comes from the es x=38 o 39, mientras que es uniformemente x=36 en
Greek hymen, membrane + Asplenium. The type species la secc. Asplenium. El nombre de la sección se deriva del
has membranous leaves. griego hymen, membrana + Asplenium. 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.

186
Figure 87. Asplenium sect. Hymenasplenium. A. A. repandulum. B. A. delitescens. C. A. obtusifolium. D. Vasculature of dor-
siventral rhizome; note distichous leaves with swollen bases. E. A. ortegae. F. A. laetum. (Murakami & Moran, 1993)

187
PTERIDACEAE
Astrolepis D. M. Benham & Windham
DESCRIPTION: Rhizomes compact, horizontal to erect, DESCRIPCIÓN: Rizomas compactos, horizontales a
occasionally branched; rhizome scales stramineous to cas- erectos, ocasionalmente ramificados; escamas del rhi-
taneous, concolorous to weakly bicolorous, linear, ciliate zoma estramineas a castañas, concoloras a debilmente
to entire; leaves monomorphic, clumped, 7-80 cm long; bicoloras, lineares, ciliadas a enteras; hojas monomorfas,
petioles with 2 vascular bundles, 1/6–1/3 the leaf amacolladas, 7–80 cm de largo; pecíolos con 2 haces
length, terete; blades linear to linear-oblong, pinnate vasculares, 1/6–1/3 el largo de la hoja, teretes; láminas
to pinnate-pinnatifid, coriaceous, abaxially clothed with lineares a lineares-oblongas, pinnadas a pinnada-pi-
imbricate, ciliate scales with underlying scurf of substel- natífidas, coriáceas, con escamas imbricadas ciliadas en
late scales, adaxially with substellate to long-ciliate el envés arriba de una caspilla de escamas subesteladas,
scales, often glabrescent, dull; pinnae 15–42 pairs, en el haz con escamas subesteladas a largo-ciliadas,
short-stalked to subsessile, articulate, ovate to oblong or a mendo glabrescentes, opaco; pinnas 15–42 pares,
narrowly deltate, cordate to truncate or rounded at base, corte-pedicelado a subsessiles, articuladas, ovadas a ob-
coriaceous; pinna margins undifferentiated, not recurved; longas o estrechamente deltadas, cordatas a truncatas o
veins obscure, free; sporangia along terminal 1/3 of veins; redondeadas; margenes de las pinnas no diferenciadas,
spores dark brown, tetrahedral-globose, rugose; x=29. 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- COMENTARIOS: Todas son epipétricas o terrestres en
tively dry regions. The genus occurs from California and regiones secas. El género existe en California y Oklahoma
Oklahoma south to Argentina, and is disjunct in Alabama, al sur hasta Argentina, y es disjunto en Alabama, Geor-
Georgia, and Hispaniola. All species occur in Mexico. The gia y Española. Todas las especies existen en México. El
genus consists of five diploid species which form auto- género consta de cinco especies diploides que forman
polyploids (6 taxa) and allopolyploids (13). The diploids autopolyploideas (6 taxones) y allopolyploideas (13). Los
are sexual, with 64 spores per sporangium, the spores diploides son sexuales con 64 esporas por esporangio,
less than 50 mμ long. All polyploids are apomictic, with las esporas menos de 50 mμ de largo. Todos los polip-
32 spores per sporangium. Triploids have spores 50–70 loideas son apomícticas, con 32 esporas por esporangio.
mμ long, tetraploids more than 71 mμ long. Astrolepis to Los triploides tienen esporas 50–70 mμ de largo, tetra-
be monophyletic with a chromosome base number of ploides más de 71 mμ de largo. Astrolepis es un grupo
29, fronds pinnate to pinnate-pinnatifid with many pinna monofilético con un número cromosómico base de 29,
pairs, two vascular bundles in the petiole, imbricate scaly hojas pinadas a pinada-pinatífidas con muchos pares de
abaxially, and substellate to ciliate scales on adaxial blade pinas, dos haces vasculares en el pecíolo, escamas imbri-
surface. The genus name is derived from the Greek astro, cadas en el envés, y escamas subesteladas, ciliadas en el
star + lepis, scale. Star-shaped scales occur on the upper haz. El nombre genérico se deriva del griego astro, estrella
surface of the pinnules. + 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.

188
Figure 88. 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. (Mickel & Smith, 2004).

189
ATHYRIACEAE
Athyrium Roth.
DESCRIPTION: Terrestrial; rhizomes erect to creep- DESCRIPCIÓN: Terrestres; tallos erectos o rastreros,
ing, scaly, the scales not clathrate; sterile and fertile leaves escamosos, las escamas no clatradas; hojas estériles y
monomorphous; petioles with 2 vascular bundles ba- fértiles monomorfas; pecíolos con dos haces vascu-
sally, these uniting distally to form a “U”; laminae 1-pin- lares en la base, estos uniéndose distalmente en forma
nate to 4-pinnate; veins free; rachis and costa grooved, the de “U”; lámina 1-pinnada a 3-pinnado-pinnatífida; ner-
grooves decurrent; sori shortly linear or J-shaped or vaduras libres; raquis y costas surcados, los surcos de-
horseshoe-shaped, with the distal portion hooked curentes; soros cortamente lineares o en forma de
around the tip of the vein; indusia membranous, the “J” a hipocrepiformes con la porción distal uncina-
margins ciliate or lacerate; sporangial stalks with 3 rows of da alrededor de la punta de la nervadura; indusio
cells; spores bilateral, brown; x=40. membranáceo, los márgenes glandulosos, ciliados o lacer-
ados; pedículos de los esporangio con 3 hileras de células;
esporas reniformes, pardas; x=40.

SIMILAR GENERA: Diplazium differs by having straight GÉNEROS PARECIDOS: Diplazium difiere por nunca
(not J-shaped) sori. Asplenium differs by having straight tener soros rectos (no en forma de J), Asplenium difiere
(not J-shaped) sori, and the 2 vascular bundles in the pet- por tener soros rectos (no en forma de J), y los dos haces
iole uniting distally to form an “X.” Athyrium is always ter- vasculares en el pecíolo unen distalmente para fomar un
restrial, whereas Asplenium can be terrestrial, epiphytic or “X”. Athyrium es siempre terrestre, mientras que Aspleni-
saxicolous. um puede ser terrestre, epífita, o saxícolo.

COMMENTS: Athyrium contains about 100 species and COMENTARIOS: Athyrium consta de casi 100 especies
is cosmopolitan, but most species occur in eastern Asia. y es cosmopolito, pero la mayoría de las especies existen
It has few species in the Neotropics. Athyrium filix-femina en Asia oriental. Tiene pocas especies en el Neotrópico.
(L.) Roth, the type species of the genus, is represented in Athyrium filix-femina (L.) Roth, la especie tipa del género,
the Neotropics by a complex of species, probably none está representada en el Neotrópico por un complejo de
of which are conspecific with the European type. Names especies, probablemente ninguna es conespecífica con el
that apply to this complex in Central America are A. arcua- tipo de Europa. Nombres que aplican a este complejo
tum Liebm. and A. bourgaei Fourn., and in South America en America Central son A. arcuatum Liebm. y A. bour-
the plants are often called A. dombeyi Desv. These species gaei Fourn., y en America del Sur las plantas se llaman A.
occur above 1500 m. Two species at lower elevations are dombeyi Desv. En el sentido amplio, esto es la única espe-
A. palmense (H. Christ) Lellinger and A. skinneri (Baker) C. cies que existe arriba de 1500 m. Dos especies en eleva-
Chr. They both have creeping rhizomes, in contrast to the ciones más bajas son A. palmense (H. Christ) Lellinger y A.
previous ones that have erect rhizomes. The genus name skinneri (Baker) C. Chr. Los dos tienen rizomas reptantes,
is derived from the Greek athyros, doorless. It apparently en contraste a las especies previas que tienen rizomas
refers to the sporangia only tardily pushing back the outer erectos. El nombre genérico se deriva del griego athyros,
edge of the indusium. sin una puerta. Aparentamente refiere a los esporangios
que empujan atrás lentamente el margen exterior del in-
dusio.

LITERATURE: Stolze, R. G. 1994. Athyrium. In: Harling, G. & L. Andersson, editors. Flora of Ecuador, 14 (5B). Polypodiace-
ae-Dryopteridoideae-Physematieae 49: 1–108.

190
Figure 89. A. Athyrium arcuatum. B. pinnule with J-shaped sori. C. Athyrium bourgeaui. (Mickel & Smith, 2004)

191
SALVINIACEAE
Azolla Lam.
DESCRIPTION: Floating aquatics; plants typically 0.5– DESCRIPCIÓN: Acuáticas flotantes; plantas típi-
2 cm long, more or less triangular; roots present, slender, camente 0.5–2 cm de largo, más o menos triangulares;
unbranched; stems hidden by the imbricate leaves; leaves raíces presentes, delgadas, no ramificadas; tallos ocultos
0.4–2 mm, distichous, appressed, sessile, with an upper por las hojas imbricadas; hojas 0.4–2 mm, dísticas, ad-
and lower lobe, the lower lobe translucent and floating, presas, sésiles, con un lóbulo superior e inferior, el lóbulo
the upper lobe green or reddish, arched-erect, papillose; inferior translúcido y flotante, el superior verde o rojizo,
sori generally paired, one megasporangiate (smaller) and arqueado-erecto, papiloso; soros generalmente pareados,
one microsporangiate (larger), borne on the underside of uno megasprangiodo (más pequeño) y el otro microspo-
the first leaf of a lateral branch, covered by a membrana- rangiado (más grande), portados sobre el envés de la pri-
ceous indusium; x=22. mera hoja de una rama lateral, envueltos por un indusio
membranáceo; x=22.
SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Azolla consists of abut 7 species world- COMENTARIOS: Azolla consta de casi 7 especies
wide, mostly in tropical and subtropical regions of the mundial, principalmente en las zonas tropicales y sub-
world.They grow on the surface of stagnant or slow-mov- tropicales del mundo. Crecen sobre la superficie de agua
ing water but may become stranded on mud. In the tem- estancada o en arroyos lentos pero pueden estar sobre
perate zones, the plants turn reddish in the fall. Azolla lodo. En las zonas templadas, las plantas se tornan rojizas
is well known for its symbiosis with the nitrogen-fixing en el otoño. Azolla es bien conocida por su simbiosis con
cyanobacteria Anabaena azollae Strasb. The cyanobacte- la cianobacteria fijadora de nitrógeno Anabaena azollae
ria can be observed by crushing a leaf on a microscope Strasb. La cianobacteria puede ser observada por trit-
slide and examining it with a compound microscope. The urar una hoja sobre un portaobjetos y examinándolas
plants are a source of nitrogen, and are used as fertilizer bajo un microscopio compuesto. Las plantas son una rica
in the rice paddies of southeast Asia. Four species have fuente de nitrógeno y se emplean como fertilizante en
long been recognized in the New World, but according to los campos de arroz de sureste Asia. Cuatro especies ha
Evrard & Van Hove (2004), there are only two: A. cristata sido reconocidas en el Nuevo Mundo, pero conforme a
Kaulf. (leaf hairs 2(-3)-celled, glochidia mostly 2 to pluri- Evrard & Van Hove (2004), hay solamente dos especies: A.
septate; megaspores variously granular) and A. filiculoides cristata Kaulf. (pelitos de la hoja 2(-3) celulares, glochidias
Lam. (leaf hairs unicellular, glochidia mostly septate; mega- 2 a pluriseptadas, megasporas granulares variosamente)
spores warty). A molecular phylogenetic analysis suggests y A. filiculoides Lam. (pelitos de la hoja unicelulares, las
there are three species in the New World (Reid et al., glochidias septadas; megaesporas verrugosas). Un análisis
2006). The genus name is derived from the Greek azo, to filogenético molecular sugiere que hay tres especies en
dry + olluo, to kill. The fern is presumably killed by drying. 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.

192
Figure 90. 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 megaspo-
rangiate. F, I. Mature megaspores. G. Massula with microspores and septate glochidia. H. Massula attached to megaspore. (D–I
from Eames, 1936)

193
BLECHNACEAE
Blechnum L.
DESCRIPTION: Terrestrial, rarely hemiepiphytic or ep- DESCRIPCIÓN: Terrestres, hemiepífitas, o raras veces
iphytic; stems scaly, creeping or erect, in some species epífitas; tallos escamosos, rastreros a erectos, en algunas
massive and arborescent; sterile and fertile leaves mono- especies masivos y arborescentes; hojas estériles y fér-
morphous or dimorphous; petioles with several vascular tiles monomorfas o dimorfas; pecíolos con varios haces
bundles, the two adaxial ones enlarged; laminae pinna- vasculares, los dos adaxiales agrandodos; láminas pinna-
tisect to 1-pinnate, rarely simple; pinnae or segments tisectas o 1-pinnadas, raras veces simples; pinnas o
entire or serrate, never lobed; aerophores absent or segmentos enteros a serrados, nunca lobados; aeróforos
present at the base of the pinnae abaxially, sometimes ausentes o presentes en las bases de las pinnas abaxial-
also along the petioles; veins free, bifurcate; sori linear, mente y algunas veces a lo largo del pecíolo; venas libres,
parallel to the costae; indusia opening toward the cos- bifurcadas; soros lineares, paralelos a las costas; indu-
tae (not towards the margin); spores bilateral, nongreen; sios abriéndose hacia la costa (no hacia el margen); es-
2n=28, 29, 31, 32, 33, 34, 36. 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 150 species COMENTARIOS: Blechnum consta de casi 150 espe-
and is pantropical. In the future it will probably be divid- cies y es pantropical. En el futuro se espera que será di-
ed into smaller monophyletic genera. In the dimorphic vidido en géneros más pequeños monofiléticos. En las
species of Blechnum with narrow fertile pinnae, it is often especies dimorfas de Blechnum con pinnas fértiles angos-
difficult to interpret the soral condition. The sporangia, tas, a menudo es difícil interpretar la condición soral. Los
when mature, cover the entire abaxial surface of the pin- esporangios, cuando maduros, cubren toda la superficie
nae, obscuring the costae and indusia. This imparts the abaxial de las pinnas fértiles, obscureciéndo las costas y
false impression of an acrostichoid sorus. The only solu- los indusios. Da la impresión falsa de un soro acrosti-
tion to the correct interpretation is careful observation. coide. La sola solución para la interpretación correcta es
Usually a vestige of the indusium can be seen near the la observación cuidadosa. Usualmente se puede ver un
margin of the pinna. In the Blechnum occidentale group véstigio del indusio cerca del margen de la pinna. En el
(those species with stolons and monomorphous sterile grupo de Blechnum occidentale (las especies con hojas
and fertile leaves) hybrids are frequent. Although they estériles y fértiles monomorfas) híbridos son frequentes.
have abortive spores, the hybrids can become abundant Aunque tienen esporas abortivas, los híbridos pueden
locally through reproduction by stolons. The genus name ser abundantes localmente via reproducción por estolo-
is derived from the Greek blechnon, an ancient name for nes. El nombre genérico se deriva del griego blechnon, un
ferns in general. 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. Murillo, M. T. 1968. Blechnum subgénero Blechnum en Sur América, con especial ref-
erencia 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. 2006. Revisión de los grupos de especies del género
Blechnum (Blechnaceae-Pteridophyta): el grupo B. penna-marina. Acta Botánica Malac. 31: 7–50.

194
Figure 91. A. Blechnum appendiculatum. B, C. B. occidentale. D. B. polypodioides. E, F. B. serrulatum. G. B. gracile.
(Mickel & Beitel, 1988).

195
BLECHNACEAE
Blechnum “fragile group”
Lomaridium C. Presl
DESCRIPTION: Hemiepiphytes; rhizomes DESCRIPCIÓN: Hemiepífitas; rizomas largamente
long-creeping, with 8-15 vascular bundles arranged rastreros, con 8-15 haces vasculares dispuestos en
in a circle (as seen in cross-section), scaly, the scales un circulo (visto en sección transversal), escamo-
with a dark central stripe; sterile and fertile leaves sos, las escamas con una raya oscura central; hojas
dimorphous; petioles with 4 or more vascular bundles; estériles y fértiles dimorfas; pecíolos con 4 o más haces
laminae pinnatisect; segments adnate to the rachis, entire; vasculares; láminas pinnatisectas; segmentos adnatos al ra-
aerophores absent at the base of the pinnae abaxially; quis, enteros; aeróforos ausentes en las bases de las pin-
veins free; sterile and fertile leaves dimorphic; sori nas abaxialmente; venas libres; hojas estériles y fértiles
linear, parallel to the costae; indusia opening toward the dimorfas; soros lineares, paralelos a las costas; indusios
costae (not towards the margin); spores bilateral, non- abriéndose hacia la costa (no hacia el margen); esporas
green; x=29. bilaterales, no verdes; x=29.

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: The group of Blechnum fragile consists of COMENTARIOS: El grupo de Blechnum fragile consta
about 7 species and occurs in the Neotropics, Africa, and de 7 especies y ocurre en el Neotrópico, África y Mad-
Madagascar. All the species grow in wet shaded forests at agascar. Todas las especies existen en bosques húmedos
middle elevations. The most common, widespread ones sombreados en elevaciones medias. Las más comunes y
in the Neotropics are B. ensiforme (Liebm.) C. Chr. and B. difundias en el Neotrópico son B. ensiforme (Liebm.) C.
fragile (Liebm.) C. V. Morton & Lellinger. The sporophytes Chr. and B. fragile (Liebm.) C.V. Morton & Lellinger. Usual-
usually begin growth as epiphytes on the base of trees mente las esporofitas empiecen crecer en las bases de
and then climb. Thick roots are sent from the basal por- los árboles y luego trepan. Se envian raíces gruesas des-
tion of the rhizome into the soil, thus the plants become de la parte basal del rizoma al suelo, por eso tornando
hemiepiphytes.The juvenile leaves are quite different than en hemiepífitos. Las hojas juveniles son diferentes que
the mature ones, being simple or pinnatifid basally. Be- las maduras, siendo simples a pinnatífidas basalmente.
cause the fertile pinnae are contracted, it can be difficult A causa de las pinnas fértiles son contraídas, puede ser
to interpret the typical blechnoid soral condition (i.e., sori difícil interpretar la condición típica soral blechnoide (i.e.,
along the costa and indusium opening toward the costae). los soros por la costa y los indusios abriéndose hacia las
Careful observation with a hand lens is required. Lomarid- costas). Se requiere la observación esmera con una lupa.
ium means resembling Lomaria. Lomaridium significa semejante a Lomaria.

LITERATURE: Cranfill, R. & M. Kato. 2003. Phylogenetics, biogeography, and classification of the woodwardioid ferns
(Blechnaceae). Pages 25–48. In: S. Chandra & M. Srivastava, editors. Pteridology in the New Millennium. Kluwer Academic Publish-
ers, Netherlands. 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. Rojas-Alvarado, A. F. 2006. Two new species of Blechnum (Blechnaceae)
from the Neotropics. Brittonia 58: 388–394.

196
Figure 92. A–E. Blechnum fragile. F. B. ensiforme. (A, F, courtesy of John Mickel; B–E by Susanah Graedel).

197
BLECHNACEAE
Blechnum sect. Lomariocycas (J. Sm.) C. V. Morton
DESCRIPTION: Terrestrial, rarely epiphytic; trunks up DESCRIPCIÓN: Terrestres, raras veces epífitas; tron-
to 4 m tall, arborescent, scaly, the scales 2–4 cm long, cos hasta 4 m de altura, arborescentes, escamosos,
curved, bicolorous with a dark central stripe; sterile las escamas 2–4 cm de longitud, curvadas, bicolores con
and fertile leaves dimorphous; petioles with several una raya oscura central; hojas estériles y fértiles di-
vascular bundles, the two adaxial ones enlarged; lami- morfas; pecíolos con varios haces vasculares, los dos
nae 1-pinnate basally, pinnatisect distally; pinnae or adaxiales agrandodos; láminas 1-pinnadas basalmente,
segments entire or serrate, never lobed, adnate distally; pinnatisectas distalmente; pinnas o segmentos en-
costae sulcate adaxially; veins free, bifurcate; sori linear, teros a serrados, nunca lobados; costas surcadas adaxial-
parallel to the costae; indusia opening toward the costae mente; venas libres, bifurcadas; soros lineares, paralelos a
(not towards the margin); spores bilateral, nongreen; x=? las costas; indusios abriéndose hacia la costa (no hacia el
margen); esporas bilaterales, no verdes; x=?

SIMILAR GENERA: Plagiogyria differs by absence of GÉNEROS PARECIDOS: Plagiogyria difiere por la au-
scales, three vascular bundles in the petiole base, pinnae sencia de escamas, tres haces vasculares en la base del
adnate throughout, and adaxially non-sulcate costae. pecíolo y pinnas adnatas y costas no surcadas adaxial-
mente.
COMMENTS: Blechnum sect. Lomariocycas consists of COMENTARIOS: Blechnum sect. Lomariocycas consta
about 20 species and is mainly Southern Hemisphere. de casi 20 especies y es principalmente del Hemisferio
About 5 species occur in the Americas. They general- Sur. Hay ca. 5 especies en las Américas. Generalmente
ly grow at high elevations, above 2000 m, especially in existen en elevaciones altas, arriba de 2000 m, especial-
páramos and peat marshes. The plants tolerate ground mente en páramos y tuberas. Las plantas soportan fuegos
fires because their stems are insulated by a mass of sur- porque sus tallos están protegidos por una masa grue-
rounding petiole bases and roots. The typical blechnoid sa de bases de los pecíolos y raíces. La condición típica
soral pattern, consisting of a linear sorus on either side soral blechnoide, la cual consta de un soro lineal en los
of the costa, is often difficult to detect because of the ambos lados de las costas, es a menudo difícil detectar
narrow fertile pinnae. The sporangia, when mature, cover a causa de las pinnas fértiles estrechas. Los esporangios,
the entire abaxial surface of the pinnae, obscuring the cuando maduros, cubren toda la superficie abaxial de las
costae and indusia.This imparts the false impression of an pinnas fértiles, obscureciéndo las costas y los indusios.
acrostichoid sorus. The only solution to the correct inter- Da la impresión falsa de un soro acrosticoide. La sola
pretation is careful observation. Usually a vestige of the solución para la interpretación correcta es la observación
indusium can be seen near the margin of the pinna. The cuidadosa. Usualmente se puede ver un véstigio del indu-
genus name is derived from Lomaria, a dimorphic fern sio cerca del margen de la pinna. El nombre genérico se
genus, + Cycas. Refers to the sterile-fertile leaf dimorphy deriva Lomaria, un género dimórfico de los helechos, +
and cycad-like 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. Murillo, M. T. 1968. Blechnum subgénero Blechnum en Sur América, con especial referen-
cia 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.

198
Figure 93. . Blechnum auratum. At right is a bicolorous rhizome scale. (©Robbin Moran, 2004)

199
DENNSTAEDTIACEAE
Blotiella R. M. Tryon
DESCRIPTION: Terrestrial; rhizomes creeping, pubes- DESCRIPCIÓN: Terrestres; rizomas rastreros, pubes-
cent; leaves up to 3 m long, distant, pendulous to sub- centes; hojas hasta 3 m de largo, distantes, péndulas a
scandent; petiole bases in cross-section with a vascular subescandentes; la base del pecíolo en sección cruz con
bundle in the form of a omega (Ω); laminae pubescent un haz vascular en forma de omega (Ω); láminas pubes-
on both sufaces, 1- o 2-pinnate-pinnatifid,; pinnae sub- centes en ambas superficies, 1- a 2-pinnada-pinnatífi-
opposite; pinnules sessile, mostly adnate; rachises and das; pinnas subopuestas; pínnulas sésiles, a menudo
costae moderately to densely pubescent, the hairs 3–10 adnatas; requises y costas moderada a densamente
mm long, septate, spreading; veins netted, the areoles pubescentes, los pelos 3–10 mm de largo, septados,
polygonal, without included veinlets; sori marginal, divergentes; venas anastomosadas, las areolas poli-
continuos at the base of the sinus, supplied by more than gonales, sin venas incluídas; soros marginales, contin-
one vein, blade margin recurved and differentiated as an uos por las bases de los senos, suministrados por más de
indusium (false indusium); inner indusium absent; spores una vena, margen de la lámina recurvado y diferenciado
bilateral, monolete; x=38. como indusio (indusio falso); un indusio interior ausente;
esporas bilaterales, monoletes; x=38.

SIMILAR GENERA: Histiopteris differs by laminae gla- GÉNEROS PARECIDOS: Histiopteris difiere por lámi-
brous and glaucous, with stipule-like pinnules at the base nas glabras y glaucas, con pínnulas basales que asemejan
of the pinnae. Hypolepis differs by sori supplied by a single como estípulas en la base de las pinnas. Lonchitis difiere
vein. Lonchitis differs by free veins. por venas libres.

COMMENTS: Blotiella contains about 15 species, all in COMENTARIOS: Blotiella consta de alrededor de 15
Africa and Madagascar except for one American species, especies, todas en África y Madagascar con excepción de
B. lindeniana (Hook.) R. M. Tryon. This species grows in una especie Americana, B. lindeniana (Hook.) R. M. Tryon.
wet forests from 1000–2000 m in the Greater Antilles Esta especie inhabita bosques húmedos de 1000–2000
and from Costa Rica to eastern Brazil. Its leaves, soft and m en las Antillas Mayores y desde Costa Rica a Brasil
pliable, are often supported on the surrounding vegeta- oriental. Sus hojas, suaves y flojas, están a menudo apoy-
tion. Blotiella is sister to Histiopteris. Both have laminae adas por la vegetación circundante. Blotiella es hermana
with intermittent apical growth, opposite pinnae, and a Histiopteris. Los dos tienen láminas con crecimiento
areolate veins. The genus name honors Marie-Louise Tar- apical intermitente, pinnas opuestas y venas areoladas.
dieu-Blot, (1902–), French pteridologist El nombre genérico honora a Marie-Louise Tardieu-Blot,
(1902–), pteridólogo francés.

LITERATURE: Tryon, R. M. 1962. Taxonomic fern notes. III. Contributions from the Gray Herbarium 191: 91–107.

200
Figure 94. . 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)

201
DRYOPTERIDACEAE
Bolbitis Schott
DESCRIPTION: Terrestrial, saxicolous, or hemi-epiphyt- DESCRIPCIÓN: Terrestres, saxícolas, o hemiepífitas; ri-
ic; rhizome creeping, scaly, in cross section with a sin- zomas reptantes, escamosos, en corte transversal con
gle, wide, ventral vascular bundle and 3 or 4 dorsal ones; un sólo haz vascular ancho ventral y con 3 o 4 haces
sterile and fertile leaves dimorphous, the fertile ones dorsales; hojas estériles y fértiles dimorfas, las hojas
more erect, longer petiolate, and with narrower pinnae; fértiles más erectas, con pecíolos más largos y pinnas más
laminae simple and entire to 1-pinnate-pinnatifid, glabrous angostas; láminas simples y enteras a 1-pinnado-pinnatí-
or sparsely scaly, often with buds at or near the apex; fidas, glabras o escasamente escamosas, a menudo con
pinnae continuous (not articulate to) the rachis; rachis brotes en o cerca del ápice; pinnas continuas (no artic-
and costae above without decurrent grooves, glabrous; uladas) al raquis; raquis y costas en la superficie superior
veins reticulate (free in some Old World species), with sin surcos decurrentes, glabros; venas reticulate (libres
or without included veinlets; sori acrostichoid; indusia en algunas especies del Viejo Mundo), con o sin venillas
absent; spores monolete; x=41. incluidas; soros acrosticoides; indusios ausentes; espo-
ras monoletes; x=41.

SIMILAR GENERA: Mickelia differs by (usually) scan- GÉNEROS PARECIDOS: Mickelia difiere por ser
dent habit, laminae imparipinnate (not pinnatifid at the (usualmente) hábito escandente, láminas imparipinadas
apices), and buds (when present) at the base of the medi- (no pinatífidas en los apices) y brotes (si presentes) en la
al pinnae. Tectaria differs by the pubescent upper surfaces base de las pinnas mediales. Tectaria difiere por la pubes-
of its axes and non-acrostichoid (usually round) sori. cencia en la superficie superior de los ejes y usualmente
soros no acrostichoides (usualmente redondos).

COMMENTS: Bolbitis contains about 45 species, with COMENTARIOS: Bolbitis consta de casi 45 especies,
about 10 in the Neotropics. It is primarily a genus of low- con casi 10 en el Neotrópico. Es principalmente un géne-
to middle-elevation forests. Most of the species grow on ro de bosques de elevaciones bajas a medias. La mayoría
the forest floor or among on rocks in streams. Bolbitis is de las plantas existen en el sotobosque o entre rocas de
sister to the rest of the bolbitidoid ferns; that is, Arthro- riachuelos. Bolbitis es hermana al resto de los helechos
botrya, Elaphoglossum, Lomagramma, Mickelia, and Terato- bolbitidoides; i.e., Arthrobotrya, Elaphoglossum, Lomagram-
phyllum, It greatly resembles Anapausia, with which many ma, Mickelia y Teratophyllum. Se parece mucho a Anapau-
of its species were formerly classified, and no single char- sia, con que muchas de sus especies fueron clasificadas
acter distinguishes them. There are, however, tendencies anteriormente, y no carácter por su mismo distingue los
to differ. Most species of Bolbitis bear buds at the lamina dos. Pero hay, no obstante, tendencias diferir. La mayoría
apices, whereas only two species of Mickelia have buds, de las especies de Bolbitis llevan brotes en los ápices de
and these occur at the bases of the medial pinnae. The las láminas, mientras que solo dos especies de Mickelia
lamina apex of Bolbitis is usually pinnatifid, whereas that of llevan brotes, estos en las bases de las pinnas mediales. El
Mickelia is conform, similar to the lateral pinnae. The ge- ápice de la lámina de Bolbitis es usualmente pinnatífido,
nus name is derived from the Greek bolbition, diminutive mientras que lo de Mickelia es conforme, similar a las pin-
of bolbos, bulb. It alludes to the small buds formed on the nas laterales. El nombre genérico se derive del griego bol-
leaves of some species. bition, 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. In Press. Phylogeny and character evolution in bolbitidoid
ferns (Dryopteridaceae). International Journal of Plant Sciences. Moran, R. C., P. Labiak & M. Sundue. In review. Synopsis of
Anapausia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae). Brittonia

202
Figure 95. A–C. Bolbitis aliena (Sw.) Alston D, E. B. serratifolia (Mert. ex Kaulf.) Schott. F–H. B. umbrosa (Liebm.) Ching.
(Mickel & Smith, 2004)

203
PTERIDACEAE
Bommeria E. Fourn.
DESCRIPTION: Terrestrial; rhizomes short- to DESCRIPCIÓN: Terrestres; rizomas corto a largamente
long-creeping, scaly and sometimes hairy with unicellular rastreros, escamosos y a veces pelosos con tricomas
hairs; leaves monomorphous; petioles castaneous to unicelulares; hojas monomorfas; pecíolos castaño a at-
atropurpureous, terete or adaxially grooved; laminae ropurpúreos, teretes a sulcados adaxialmente; lámina
1-16 cm, generally as long as wide, pentagonal, pe- 1-16 cm, generalmente tan larga como ancha, pen-
date, deeply lobed, the indument of the upper sur- tagonal, pedata, profundamente lobada, el indumento
face consisting of unicellular acicular hairs, on the del haz de tricomas unicelulares, aciculares, el indu-
lower surface of scales and unicellular acicular hairs or mento del envés de escamas y tricomas unicelulares, acic-
occasionally enrolled, occasionally with 2-cell hairs with ulares o a veces enrollados, ocasionalmente con tricomas
a glandular head; basal segments generally connected to 2-celulares con la punta glandulosa; segmentos basales
the apical ones by a wing along the rachis. the basiscopic conectados generalmente a los apicales por un ala angos-
lobe elongated or bipinnatifid; veins free or netted, the ta que corre a lo largo del raquis, el lobo basiscópico alar-
areoles without included veinlets; sporangia spreading gado a bipinnatífido; nervaduras libres o anastomosadas,
along the veins, covering 2/3-3/4 the distance from the las aréolas sin nérvulos incluidos; esporangios difundi-
margin to the costa, occasionally restricted toward the dos a lo largo de las nervaduras, que cubren 2/3-3/4
margins; indusium absent; paraphyses absent; spores de la distancia del margen a la costa, ocasionalmente se
globose; x = 30. 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 COMENTARIOS: Bommeria tiene 5 especies y ocurre
the southwestern United States to Costa Rica. The ge- desde el suroeste de los Estados Unidos hacia Costa Rica.
nus typically grows in dry, rocky habitats. The genus name El género crece típicamente en hábitats secas, rocosas.
honors Joseph Edouard Bommer (1829–1895), Belgian El nombre genérico honora Joseph Edouard Bommer
pteridologist. (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.

204
Figure 96. A–C. Bommeria pedata (Sw.) E. Fourn. D–G. B. subpaleacea Maxon. (Mickel & Smith, 2004)

205
OPHIOGLOSSACEAE
Botrychium Sw.
DESCRIPTION: Terrestrial; stems subterranean, fleshy, DESCRIPCIÓN: Terrestres; tallos subterráneos, carno-
erect, usually unbranched, without scales or hairs; roots sos, erectos, usualmente no ramificados, sin escamas o
fleshy, without hairs; leaves one per plant, divided into pelos; raíces carnosas, sin pelos; hojas una por planta, divi-
a green, horizontal or spreading lamina and a non- didas en una lámina verde, horizontal o divergente
green, erect, sporangia-bearing spike (sorophore); y una espiga no verde, erecta llevando esporganios
petiole base sheathing buds at the stem apex; laminae (el soróforo); base del pecíolo envainando los brotes
1-4-pinnate, fleshy, usually tringular; veins free; so- en el ápice del tallo; láminas 1-4-pinnadas, carnosas,
rophore arising from the adaxial surface of the petiole; usualmente triangulares; nervaduras libres; soróforos
sporangia globose, opening transverely, after dehiscence surgiendo desde la superficie adaxial del pecíolo; espo-
appearing clamlike; x=44–47. rangios globosos, abriéndose transversamente, después
de la dehiscencia se parecen un bivalva; x=44–47.

SIMILAR GENERA: Anemia differs by 2 erect fertile GÉNEROS PARECIDOS: Anemia difiere por 2 espigas
spikes (sorophores) and densely pubescent rhizomes. fértiles (soróforos) erectas y rizomas densamente pubes-
centes.

COMMENTS: Of the approximately 45 species of COMENTARIOS: De las aproximadamente 45 espe-

Botrychium worldwide, only 5 occur in the Neotropics. cies de Botrychium mundialmente, solo 5 existen en el
They represent 2 of the 4 subgenera in the genus. Sub- Neotrópico. Representan 2 de los 4 subgéneros en el
genus Sceptridium (Lyon) Claussen is represented by the género. El subgénero Sceptridium (Lyon) Claussen está
other four species and can be recognized by the fertile representa por las otras cuatros especies y se puede
spike born near the petiole base, usually well below the reconocer por la espiga fértil que nace cerca de la base
soil (see A). Subgenus Osmundopteris (Milde) Claussen is del pecíolo (vease A), por lo común bajo del suelo. El
represented by B. virginianum (L.) Sw. and can be recog- subgénero Osmundopteris (Milde) Claussen está repre-
nized by the fertile spike born at the base of the lamina sentado por B. virginianum (L.) Sw. y se puede reconocer
(see B). Botrychium typically grows in open or semi-open por la espiga fértil que nace cerca de la base de la lámi-
habitats such as pastures and second-growth woodlands. na (vease B). Botrychium típicamente crece en hábitats
Usually, when one plant is found, others are nearby. The abiertos o semi-abiertos tales como potreros y bosques
genus name is derived from the Greek botrys, cluster, al- secondarios. Usualmente, cuando se encuentra una
luding to the grape-like clusters of sporangia. planta, otras están cerca. El nombre genérico se deriva
del griego botrys, en referenica a los macollos de espor-
angios que aparecen como un racimo de uvas.

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 mor-
phology. Molecular Phylogenetics and Evolution 28: 131–151.

206
Figure 97. A. Botrychum decompositum. B. B. virginianum. (Mickel & Smith, 2009).

207
POLYPODIACEAE
Campyloneurum C. Presl
DESCRIPTION: Epiphytic, occasionally terrestrial or DESCRIPCIÓN: Epífitas, ocasionalmente terrestres o
rupestral; rhizomes short- to long-creeping scaly, the saxícolas; rizomas cortamente a largamente rastreros, es-
scales usually lanceolate, clathrate or nonclathrate, gla- camosos, las escamas usualmente lanceoladas, clatradas o
brous; sterile and fertile leaves monomorphous; no clatradas, glabras; hojas estériles y fértiles mono-
petioles articulate to the stem or to short phyllopodia; morfas; pecíolos articulados al tallo o a las filopódios
laminae simple, entire (but 1-pinnate in 2 species), ob- cortos; láminas simples, enteras, (pero 1-pinnadas en
long to narrowly elliptic, glabrous or with a few scales 2 especies), oblongas a angostamente elípticas, glabras
on the rachis; venation areolate with included veinlets, o con pocas escamas sobre el raquis; venación areo-
these usually between prominent, lateral veins; sori in lada, con venillas incluídas, estas usualmente entre venas
2–4 rows between the main lateral veins, borne on laterales prominentes; soros en 2–4 hileras entre las
the tips of included veinlets, round, without indusia, venas laterales principales, se nacen en los ápices
and several rows between the rachis and margin of the de venillas incluídas, redondos, sin indusios, en varias
lamina; sporangia glabrous; paraphyses absent; spores bi- hileras entre el raquis y margen de la lámina; esporangios
lateral, yellow; x=37. glabros; parafises ausentes; esporas bilaterales, amarillas;
x=37.

SIMILAR GENERA: Niphidium differs by one row of GÉNEROS PARECIDOS: Niphidium difiere por una
sori between the lateral veins. Other genera of Polypo- hilera de soros entre las venas laterales. Otros géneros
diaceae with simple, entire leaves, such as Microgramma de Polypodiaceae con hojas simples y enteras, tales como
and Pleopeltis, are easily distinguished by having only one Microgramma y Pleopeltis, son distinguidos facilmente por
row of sori between the rachis and margin of the lamina. tener una sola hilera de soros entre el raquis y margen de
Elaphoglossum differs by acrostichoid sori and (almost al- la lámina. Elaphoglossum difiere por soros acrostichoides
ways) free veins. y (casi siempre) venas libres.

COMMENTS: Campyloneurum consists of about 50 spe- COMENTARIOS: Campyloneurum consta de alrededor

cies and is entirely neotropical. Its species grow primarily de 50 especies y es completamente neotropical. Sus es-
in wet forests.Two species are atypical by having 1-pinnate pecies se encuentra principalmente en bosques húmedos.
laminae: C. decurrens (Raddi) C. Presl and C. magnificum T. Dos especies son atípicas por tener láminas 1-pinnadas:
Moore. They are included in the genus because they have C. decurrens (Raddi) C. Presl y C. magnificum T. Moore. Es-
venation like that of the simple-leaved species. The genus tán incluídas dentro del género porque tienen venación
name is derived from the Greek kampylos, arched + neu- parecida a éso en las especies con hojas simples. El nom-
ron, vein. It refers to the secondary veins that are arched bre genérico se deriva del griego kampylos, arqueado +
between the main lateral ones. neuron, vena. Se refiere a las venas secundarias que son
arcuadas entre las venas laterales principales.

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. Dissertation.
Aarhus University, Denmark. Rojas, A. F. 2005. El complejo de Campyloneurum angustifolium (Sw.) Fée en Costa Rica. Lankes-
teriana 5: 41–48. Vasques, D. T. & J. Prado. 2011. Campyloneurum C. Presl (Polypodiaceae) in São Paulo State, Brazil. Hoehnea
38: 147–163.

208
Figure 98. A, B. Campyloneurum amphostenon. C, D. C. angustifolium. E–G. C. tenuipes. H. C. xalapense. J, K. C. repens. L–N.
C. phyllitidis. (Mickel & Beitel, 1988).

209
POLYPODIACEAE
Ceradenia L.E. Bishop
DESCRIPTION: Typically epiphytic; rhizome scales non- DESCRIPCIÓN: Típicamente epífitas; escamas del
clathrate, often ciliate and with whitish, marginal glands rizoma no clatradas, a menudo ciliadas y con glándulas
when young; petiole often articulate; laminae simple, marginales blanquecinas cuando jóvenes; pecíolo a menu-
pinnatifid or peripinnate, often pectinate; veins often do articulado; láminas simples, pinnatífidas o peripinnadas,
1-forked, sometimes simple; hydathodes absent; setae a menudo pectinadas; nervaduras a menudo 1-bifurcadas,
reddish; trichomes simple or branched, sometimes glan- a veces simples; hidátodos ausentes; setas rojizas; tri-
dular; sori round or oblong, superficial or subimpressed; comas simples o ramificados, a veces glandulares; soros
paraphyses glandlike, waxy, globose or pyriform, redondeados u oblongos, superficiales o subimpresos;
whitish, brown-yellow to yellowish, multicellular; spores paráfisis como glándulas, cerosos, globosos o piri-
green, tetrahedral; x=37. formes, blanquecinos, pardo-amarillentos a amarillentos,
multicelulares; esporas verdes, tetraédricas; x=37.

SIMILAR GENERA: Zygophlebia and Enterosora differ GÉNEROS PARECIDOS: Zygophlebia e Enterosora di-
by anastomosing veins, but this is difficult to see because fieren por venas anastomosadas, pero esto es difícil de
of the thickness of the lamina. They lack glands in the sori, distinguir debido al grosor de la lámina. Faltan glándulas
or if present (Zygophlebia), then they are brown and not en los soros, y si son presentes (Zygophlebia), entonces
waxy or whitish. Many other genera of Grammitidace- son pardos y no cerosos blanquecinos. Se puede distin-
ae, such as Lellingeria, Melpomene y Terpsichore, can be guir muchos otros géneros de las Grammitidaceae, tales
distinguished by the presence of hydathodes. Cochlidium como Lellingeria, Melpomene y Terpsichore, por la presen-
Grammitis, Lellingeria, and Luisma differ by the absence of cia de hidatodos. Cochlidium Grammitis, Lellingeria y Luis-
reddish setae. 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 tropical con casi 60 especies, con 2 en los Azores y Liberia,
in Africa-Madagascar. Nearly all grow in cloud forests. The y 6 en África-Madagascar. Casi todos existen en bosques
most distinctive characteristic of this genus is the whitish nublados. El carácter más distintivo del género es las
glands in the sorus, from which the genus takes its name. glándulas blanquecinas en el soro, de las cuales el género
These can be so numerous that they obscure the young toma su nombre. Estas pueden san tan numerosas que
sporangia and can be mistaken for spores in more mature esconden los esporangios jóvenes, y pueden ser malin-
specimens. Some species also have branched, gland-bear- terpretados como esporas en ejemplares más maduros.
ing hairs on the lamina, axes, and margins of the rhizome Algunas especies también tienen pelitos glandulíferos por
scales. In these locations, the hairs are often branched, las laminas, ejes y márgenes de las escamas del rizoma.
with 2–5 glands per hair. Sometimes the glands are diffi- En estas locaciones, las pelitos son a menudo ramificados
cult to observe in specimens collected in alcohol or dried con 2–5 glándulas por pelito. A veces las glándulas son
with high heat. Ceradenia is monophyletic and sister to difíciles de observar en ejemplares colectados en alcohol
Enterosora (Ranker et al., 2004). The genus name is de- o secados con alto calor. Ceradenia es monofilético y her-
rived from the Greek keras, wax, and aden, gland, referring mano a Enterosora (Ranker et al., 2004). Ceradenia deriva
to the distinctive glands in the sorus. 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 Cerad-
enia 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. 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 (Grammit-
idaceae): a case of rampant morphological homoplasy. Taxon 53: 415–428. Smith, A. R. 1993. New species and combinations of
Ceradenia (Grammitidaceae). Novon 3: 182–185.

210
Figure 99. Ceradenia. In G and K, note whitish glands among sporangia. En G y K, fijase las glándulas blanquecinas entre los
esporangios. (Smith, 1993).

211
PTERIDACEAE
Ceratopteris Brongn.
DESCRIPTION: Plants free-floating or rooting DESCRIPCIÓN: Plantas flotantes o echando raíces
in the mud; rhizomes inconspicuous, short-creeping, en el lodo; rizomas inconspícuos, cortamente rasteros,
sparsely scaly; sterile and fertile fronds dimorphic, escamosos; las hojas estériles y fértiles dimorfas, las
the fertile ones contracted and more erect compared to fértiles contraídas y más erectas en comparación con las
the sterile; sterile fronds pinnatifid to 3-pinnate, glabrous; estériles; láminas estériles pinatifídas a 3-pinnadas, glabras;
veins areolate; buds present in the pinna sinuses; sori venas areoladas; brotes presentes en los sinuses de las
marginal, with the margin recurved over the sori; annulus pinnas; soros marginales, con los márgenes recurvados
weak and ill-defined; spores tetrahedral-globose, 16 or 32 por los soros; anillos débiles y mal definidos, 16 o 32 por
per sporangium; x=39. esporangio; esporas tetraédricas-globosas; x=39.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Ceratopteris is pantropical and subtropical COMENTARIOS: Ceratopteris es pantropical y sub-

usually occurs in standing or slow-moving water. The ge- tropical y ocurre usualmente en aguas estancadas o len-
nus was monographed by Lloyd (1974), who recognized tas. El género fue monografiado por Lloyd (1974) quién
three species in the Neotropics: C. pteridoides (Hook.) reconoció tres especies en el Neotrópico: C. pteridoides
Hieron., C. richardii Brongn. and C. thalictroides (L.) Brongn. (Hook.) Hieron., C. richardii Brongn. y C. thalictroides (L.)
The spores, which are 70–150 micrometers long, are Brongn. Las esporas, cuales son 70–150 micrometros
among the largest of any homosporous fern. They bear de largo, son unas de los más grandes entre helechos
have parallel ridges that can easily be seen under a dis- homospóricos. Tienen crestas paralelas fácilmente vistas
secting microscope at 30X. The chromosome number of con un microscopio de disección a 30X. El número cro-
x=39 is unique in the family. The genus is frequently used mosómico de x=39 es único en la familia. El género se
for genetic research because it has the shortest life cycle usa frecuentemente para investigacíon genética porque
of any homosporous fern (three months), and the plants tiene el ciclo de vida más corto de cualquier helecho
grow easily in the laboratory. Reproduction by buds in homospórico (tres meses), y las plantas crecen fácilmente
the sinuses of the pinnae is extremely common. The spe- en el laboratorio. La reproducción por brotes en los si-
cies are frequently sold for use in aquaria. Ceratopteris is nuses de las pinnas es muy común. Se venden las especies
the sister genus to Acrostichum, which looks quite differ- con frecuencia para uso en los acuarios. Ceratopteris es el
ent but also grows in wet soils. The two are classified in género hermana a Acrostichum, que parece muy diferente
the subfamily Parkerioideae. The generic name is derived pero también inhabita suelos mojados. Los dos se clasif-
from the Greek keras, horn + pteris, fern. It alludes to the ican en la subfamilia Parkerioideae. El nombre genérico
antler-like fertile leaves. se deriva del griego keras, cuerno + pteris, helecho. Se
refiere a las hojas fértiles como cuernos.

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.

212
Figure 100. A–D. Ceratopteris pteridoides. D–G. C. thalictroides (Mickel & Smith, 2004).

213
PTERIDACEAE
Cheilanthes Sw.
DESCRIPTION: Terrestrial or rupicolous; rhizomes DESCRIPCIÓN: Terrestres o rupícolas; rizoma postra-
prostrate or ascending, scaly; plants generally less than do o ascendente, escamoso; plantas generalmente
30 cm long; leaves monomorphic or subdimorphic, her- menores de 30 cm de largo; hojas monomorfas o
baceous to coriaceous; petioles with 1 vascular bun- subdimorfas, herbáceas a coriáceas; pecíolo con un haz
dle, often castaneous to blackish y shiny, terete or sulcate vascular, a menudo castaño a negruzco y lustroso, terete
adaxially; laminae 1-5-pinnate, not farinose; veins free, o sulcado adaxialmente; lámina 1-5-pinnada, no farinosa;
the tips not claviform; indusium reflexed, scarious; nervaduras libres, los ápices no claviformes; esporangios
paraphyses absent; spores tetrahedral-globose to glo- dispuestos sobre o cerca de los ápices de las nervaduras;
bose, trilete, 64 or, more commonly, 32 per sporangium. indusio generalmente reflexo, escarioso; parafisos
x=29, 30. 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 de casi 150 especies. Existe principalmente en hábitats
most of its species are apogamous with 32 spores per secas, y la mayoría de sus especies son apógamas con 32
sporangium. In the New World, its center of diversity and esporas por esporangio. Su centro de diversificación y
endemism is Mexico. Even with the removal of Adian- endemismo en el Nuevo Mundo es en México. Aun con
topsis, Aleuritopteris, Argyrochosma, Astrolepis, Cheiloplecton, la exclusión de Adiantopsis, Aleuritopteris, Argyrochosma,
Mildella, and Notholaena, Cheilanthes remains a large and Astrolepis, Cheiloplecton, Mildella y Notholaena, Cheilanthes
heterogeneous group. Future studies will probably result permanece como un grupo numeroso y heterogéneo. Es
in the recognition of additional segregate genera, such as probable que estudios futuros resulten en la designación
Physapteris. The genus name is derived from the Greek de géneros adicionales segregados, tales como Physapter-
cheilos, lip + anthos, flower. It alludes to the indusia and is.El nombre genérico se deriva del griego cheilos, labio +
the sori. 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 Uni-
versity, Tempe. Tryon, R. M. 1956. A revision of the American species of Notholaena. Contributions to the Gray Herbarium of
Harvard University 179: 1–106.

214
Figure 101. A–C. Cheilanthes kaulfussii. D, E. C. longipila. F–H. C. lendigera. J–P. C. myriophylla. (Mickel & Smith, 2004)

215
CIBOTIACEAE
Cibotium Kaulf.
DESCRIPTION: Terrestrial; trunks erect, to 1.5 m tall DESCRIPCIÓN: Terrestres; troncos erectos, hasta 1.5
(to 15 m in Asia and Pacific), densely lanose, the hairs soft, m de altura (hasta 15 m en Asia y el Pacífico), densa-
golden; leaves 2–4 m long; petioles densely lanose like the mente lanosos; los pelos suaves, dorados; hojas 2–4 m
trunk; laminae 2-pinnate-pinnatifid to 3-pinnate, glabrous, de longitud; pecíolos densamente lanosos como el tron-
glaucous or green abaxially; veins free; sori marginal; in- co; láminas 2-pinnado-pinnatífidas a 3-pinnadas, glabras,
dusial bivalvate, the lower valve narrower and fitting glaucas o verdes abaxialmente; nervaduras libres; soros
into the recurved upper one, both valves differing from marginales; indusios bivalvados, la valva inferior más
the lamina tissue in color; spores non-green, tetrahedral, angosta y cabe a dentro lo de arriba recurvada, las
with a prominent equatorial flange. x=68. dos valvas difieren del tejido de la lámina en color; espo-
ras no verdes, tetrahédricas, con un cíngulo prominen-
te ecuatorial. x=68.

SMILAR GENERA: Culcita differs by non-arborescent GÉNEROS PARECIDOS: Culcita difiere por troncos
trunks, broadly deltate laminae that are green abaxially no arborescentes, láminas anchamente deltadas que son
(never glaucous), and valves of the indusial of equal size. verdes abaxialmente (nunca glaucas) y valvas del indusio
Lophosoria differs by abaxial, non-indusiate sori. de tamaño igual. Lophosoria difiere por soros abaxiales
no indusiados.

COMMENTS: Culcita consists of 9 species, of which 2 COMENTARIOS: Culcita consta de 9 especies, de las
occur in northern Central America, 3 in Asia, and 4 in cuales 2 existen en América Central septentrional, 3 en
Hawaii. Our species typically grow in forests or thickets Asia y 4 en Hawaii. Nuestras especies crecen típicamente
from (–750)1000–2500(–3000) m. The two American en bosques o matorrales desde (–750)1000–2500(–
species are C. schiedei Schldtl. & Cham. (Veracruz and 3000) m. Las dos especies Americanas son C. schiedei
Oaxaca) and C. regale Verschaff. & Lem. (s. Mexico to Schldtl. & Cham. (Veracruz and Oaxaca) and C. regale
Honduras and El Salvador). The genus name is derived Verschaff. & Lem. (s. Mexico to Honduras and El Salva-
from the Greek kibotos, a small chest or casket, alluding dor).El nombre genérico se deriva del griego kibotos, una
to the shape of the indusium. caja pequeña o arquita.

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.

216
Figure 102. Cibotium. A–C. Cibotium schiedei. D. C. regale (Mickel & Smith, 2004).

217
POLYPODIACEAE
Cochlidium Kaulf.
DESCRIPTION: Epiphytic or saxícolous; rhizome scales DESCRIPCIÓN: Epífitas o rupícolas; escamas del rizoma
not clathrate, eciliate, dull or somewhat lustrous, brown-yel- no clatradas, sin cilias, opacas o algo lustrosas, pardo-am-
low to brown; leaves less than 25 cm long, typically simple, arillentas a pardas; hojas menos de 25 cm de longitud,
linear, entire or deeply serrate, glabrous or with simple, típicamente simples, lineares, enteras o profundamente
2—8-celled hairs (without dark setae); veins free or con- serradas, glabras o con pelos simples 2—8-celulares
nivent basally en a commissural vein, sterile veins simple (sin setas oscuras); nervaduras libres o conniventes
or 1-forked; hydathodes present; sori grouped in the basalmente en una nervadura comisural, nervaduras
distal part of the lamina; separate or linearly confluen; estériles simples o 1-bifurcadas; hidátodos presentes; so-
paraphyses absent; spores green, tetrahedral. x=33, 34?, ros agrupados en la parte distal de la lámina, separa-
35. dos o linearmente confluentes; paráfisis ausentes; esporas
verdes, tetrahédricas. x=33, 34?, 35.

SMILAR GENERA: Grammitis differs by having a dark GÉNEROS PARECIDOS: Grammitis difiere por un
laminar margin. Luisma differs by two vascular bundles margen laminar oscuro. Luisma difiere por dos haces
in the petiole. Several species of Lellingeria resemble C. vasculares en el pecíolo. Varias especies de Lellingeria se
serrulatum (Sw.) L. E. Bishop, but can be distinguished by parecen a C. serrulatum (Sw.) L. E. Bishop, pero pueden
clathrate rhizome scales and at least a few small hairs distinguírseles por escamas del rizoma clatradas y al
(especially on the petiole and raquis) unequally forked. 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, Cochlid-
ium, 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.

218
Figure 103. C–E. Cochlidium serrulatum (Sw.) L. E. Bishop. G. Cochlidium rostratum (Hook.) Maxon ex C. Chr.; note sorus
sunken in the central groove (Mickel & Beitel, 1988).

219
 Crepidomanes (C. Presl) C. Presl subgen. Crepidomanes
DESCRIPTION: Mostly epiphytes; roots absent; rhi- DESCRIPCIÓN: Mayormente epífitas; raíces ausentes;
zomes 0.1–0.8 mm wide, long-creeping, frequent- rizomas 0.1–0.8 mm de ancho, largamente rastre-
ly branched, densely pubescent, the hairs reddish ros, ramificados con frecuencia, densamente pu-
brown or (in ours) black; sterile and fertile mono- bescentes, los pelos pardo-rojizos o (en nuestra)
morphic; petioles to 8 cm long, pubescent basally and neguzcos; hojas estériles y fértiles monomorfas; pecíolos
along margin of wing, the hairs like those on the hasta 8 cm de largo; pubescentes en la base y por el
rhizomes; sterile laminae 1-cell thick between the veins, margen de la ala del pecíolo, los pelos como los del
1-pinnate to 4-pinnate, the apices not proliferous; veins rizoma; láminas estériles una célula de grosor entre las
anadromous, free, false veins often present; receptacles venas, 1-pinnadas a 4-pinnadas, los ápices no prolíferos;
exert; spores green, tetrahaedral-globose; x=36. venas anádromas, libres, falsas venas a menudo pre-
sentes; receptáculos exertos; esporas verdes, tetrahédri-
cas-globosas; x=36.

SIMILAR GENERA: Polyphlebium differs by reddish GÉNEROS PARECIDOS: Polyphlebium difiere por pe-
brown rhizome hairs, presence of roots, glabrous peti- los de los rizomas pardo-rojizos y presencia de raíces,
ole bases and petiole wing margins, and lack of folds in bases de los pecíolos y márgenes de las alas de los pecío-
the lamina parallel the veins. Didymoglossum differs by the los glabros, y falta de pliegues en la lámina paralelos a las
presence of false veins in the laminae and smaller leaves venas. Didymoglossum difiere por la presencia de venas
(< 6 cm). falsas en las láminas y hojas más cortas (< 6 cm).

COMMENTS: Crepidomanes consists of about 30 spe- COMENTARIOS: Crepidomanes consta de casi 30 es-
cies that grow in wet forests. All of its species are paleo- pecies que existen en bosques húmedos.Todos sus espe-
tropical except for C. pyxidiferum (L.) Dubuisson & Ebiha- cies son paleotropicales con excepción de C. pyxidiferum
ra, which is widespread in the Neotropics. It is distinctive (L.) Dubuisson & Ebihara, la cual está muy difundida en
by bearing folds in the lamina parallel to the veins. Perhaps el Neotrópico. Es distinctiva por llevando pliegues en la
not distinct from C. melanotrichum of Africa and Mada- lámina paralelas a las veins.Tal vez lo mismo como C. mel-
gascar. It resembles Polyphlebium diaphanum and P. hy- anotrichum de Africa y Madagascar. Se asemeja a Polyphle-
menophylloides and is often misidentified as these species. bium diaphanum y P. hymenophylloides, y es a menudo mal
Crepidomanes pyxidiferum can be distinguished from these identificada con estas especies. Crepidomanes pyxidiferum
species by black (not red-brown) rhizome hairs, petiole se puede distinguir por los pelos negruzcos de los rizom-
bases pubescent on all sizes (the hairs like those of the as (no pardo-rojizos), bases de los pecíolos pubescentes
rhizome), and wings of the petioles pubescent on mar- en todos lados (los pelos como los en el rizoma) y alas
gins. The rhizomes branch frequently to produce short de los pecíolos pubescentes por el margen. Los rizomas
branches that resemble roots (Schneider, 2000). This ramifican con frecuencia y producen ramas cortas que
character is shared with Didymoglossum and Polyphlebium. parecen a raíces (Schneider, 2000). Esta característica se
The genus name is derived from the Greek crepis, slipper, comparta con Didymoglossum y Polyphlebium. El nombre
and manes, cup. It alludes to the shape of the sorus. 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 phy-
logeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of the Linne-
an 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.

220
Figure 104. Crepidomanes pyxidiferum. Note dark hairs at petiole base and along wing –charcteristic of the genus. (from
Mickel & Smith, 2004.)

221
PTERIDACEAE
Cryptogramma R. Br.
DESCRIPTION: Saxicolous; stems short-creeping ,de- DESCRIPCIÓN: Saxícolas; tallos cortamente rastreros,
cumbent to erect, scaly; sterile and fertile leaves di- decumbentes a erectos, escamosos; hojas estériles y
morphic, with the fertile leaves longer-petiolate, taller and fértiles dimorfas, con la fértil más largamente pecioladas,
more erect than the sterile; sterile leaves 5-20 cm long; más largas y erectas que las estériles; hojas estériles 5-20
petioles grooved on the upper surface, with 1 vascular cm de largo; pecíolos surcados en la superficie superior,
bundle; laminae 2-4-pinnate, glabrous on both surfaces, con 1 haz vascular; láminas 2-4-pinnado, glabras en am-
not farinose beneath, scales rare or absent, the apex pin- bas superficies, no farinosas en el envés, escamas raras
natifid, tapered; ultimate segments sessile or short-stalked; o ausentes, el ápice pinnatífido, reducido gradualmente;
veins free; false indusia present; sporangia borne along segmentos últimos sésiles o cortamente peciolulados;
the veins, often intermixed with farina-producing glands, nervaduras libres; inducíos falsos presentes; esporan-
64-spored; spores yellow, trilete, verrucate, without gios nacen a lo largo de las venas, a menudo mezcladas
equatorial flange; x=30. con glándulas farinosas, con 64 esporas; esporas amarillas,
triletes, verrucatas, sin un borde ecuatorial; x=30.

SIMILAR GENERA: Cystopteris differs by round sori, GÉNEROS PARECIDOS: Cystopteris difiere por soros
presence of a true indusium (not marginal), and two vas- redondeados y un indusio verdadero (no marginal) y dos
cular bundles in the petiole. Anogramma differs by sori not haces vasculares en el pecíolo. Anogramma difiere por
protected by a reflexed marginal indusium. 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.

222
Figure 105. A, B. Cryptogramma (sect. Cryptogramma) acrostichoides. C, D. C. (sect. Homopteris) stelleri. (©R. C. Moran,
2011)

223
DRYOPTERIDACEAE
Ctenitis (C. Chr.) C. Chr.
DESCRIPTION: Terrestrial or rarely rupicolous; rhi- DESCRIPCIÓN: Terrestres o raramente rupícolas;
zomes erect to obliquely ascending; petioles with more pecíolos con más de tres haces vasculares; rizomas erec-
than three vascular bundles; leaves monomorphous; lam- tos a oblicuamente ascendentes; láminas 1-pinnado-pin-
inae 1-pinnate-pinnatifid to 4-pinnate-pinnatifid, cata- natífidas a 4-pinnado-pinnatífidas, catádromas, a menudo
dromic, often glandular, the glands cylindrical, appressed; glandulosas, las glándulas cilíndricas, adpresas; raquis y
rachis and costae flat to rounded (not grooved) on costas aplanados a redondeados (no surcados) en la
the upper surface, puberulent, often scaly beneath, superficie superior, puberulentos, a menudo escamo-
the scales typically clathrate; veins free, the tips narrow sos abaxialmente, las escamas típicamente clatradas;
(not hydathodous), ending at the margin; sori round; in- nervaduras libres, las puntas delgadas (no como un hid-
dusia present or absent; sporangial stalks with unicellular atodo), terminando en el margen; soros redondeados;
cylindrical glands; x=41. indusios presentes o ausentes; pedículos esporangiales
con glandulos unicelulares, cilíndricos; x=41.

SIMILAR GENERA: Megalastrum differs by strigose, GÉNEROS PARECIDOS: Megalastrum difiere por pe-
whitish hairs on the upper surface of the axes, and hy- los estrigosos, blanquecinos en la superficie superior de
dathodes on the upper surface of the lamina. Lastreopsis los ejes y hidatodos en la superficie superior de la lámina.
has a completely different kind of architecture on the up- Lastreopsis tiene un tipo diferente de arquitectura en el
per surface of the rachis and costas. haz de la los ráquises y costas.

COMMENTS: Ctenitis is pantropical and contains about COMENTARIOS: Ctenitis es pantropical y consta de
75 species. It grows on wet forest floors mostly from alrededor de 75 especies. crece sobre los suelos de
0–2000 m. It appears to contain two species groups: one bosques húmedos principalmente de 0–2000 m. Parece
with laminae lanceolate and 1-pinnate-pinnatifid (e.g., C. contener dos grupos de especies: uno con láminas lance-
subincisa (Langsd. & Fischer) Ching), the other with lam- oladas y 1-pinnado-pinnatífidas (e.g., C. subincisa (Langsd.
inae deltate to ovate and 2-pinnate-pinnatifid or more & Fischer) Ching), el otro con láminas deltadas a ovatas
finely divided (e.g., C. excelsa (Desv.) Proctor). Ctenitis has y 2-pinnado-pinnatífidas (e.g., C. excelsa (Desv.) Proc-
distinctive hairs, called appropriately “Ctenitis hairs,” on tor). Ctenitis tiene pelos distinctivos, llamados apropri-
the upper surface of its lamina axes. These hairs are short adamente “pelos de Ctenitis, en la superficie superior de
(0.2–0.5 mm long), often reddish, several celled, and often los ejes de las láminas. Estos pelos son cortos (0.2–0.5
appear jointed because the cells twist at right angles to mm de largo), a menudo rojizos, multicelulares, y a menu-
each other upon drying. These hairs are useful in identi- do aparecen artículados a causa de las células torcen en
fying the genus, but they are also found in other genera angulos rectos a cada uno cuando se secan. Estos pelos
such as Cyclopeltis, Lastreopsis, and Triplophyllum. Ctenitis son útiles en identificar el género, pero ocurren en otros
is derived from the Greek kteis, comb, and refers to the géneros como Cyclopeltis, Lastreopsis y Triplophyllum. Cte-
pectinate pinnae of certain species. nitis génerico 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-bipinnatifid
species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 7, 10: 55–282. Christensen, C. 1920. A monograph of
the genus Dryopteris. Part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvi-
densk. Math. Afd. ser. 8, 6: 1–132.

224
Figure 106. A–C. Ctenitis submarginalis. D, E. C. nigrovenia. F–H. C. melanosticta. J–M. C. excelsa. (Mickel & Smith, 2004)

225
CULCITACEAE
Culcita C. Presl
DESCRIPTION: Terrestrial; rhizome subterranean, DESCRIPCIÓN: Terrestres; rizoma subterráneo, cor-
stout, short, not forming a trunk, the apex with dense, to y grueso, no formando un tronco, ápice con pelos
long, golden hairs. Leaves monomorphic; stipes hairy, with densos, largos, y dorados. Hojas monomorfas; pecíolos
an omega-shaped vascular bundle; blades 4-5-pinnate, pelados, con una haz vascular en forma de una omega;
deltate, glabrous to slightly hairy, coriaceous; rachises láminas 4-5-pinnadas, deltadas, glabras a ligeramente
and costae grooved on the upper surface; veins free; peludas, coriáceas; ráquises y costas surcados en el
sori marginal; indusia bivalvate, the lower valve fitting haz; venas libres; soros marginales; indusios bivalvos,
inside the outer (upper), outer valve green like the lami- la valva inferior cabe adentro del superior, la valva supe-
nar tissue, inner valve whitish to tan, differentiated; spores rior verde como el tejido láminar, la inferior blanceciña a
trilete, nongreen; x = 66. leonada, diferenciada; esporas triletes, no verdes; x = 66.

SIMILAR GENERA: Dicksonia and Cibotium differ by GÉNEROS PARECIDOS: Dicksonia y Cibotium difieren
arborescent trunks, laminae to only 3-pinnate-pinnatifid, por troncos aborescentes, láminas hasta 3-pinadas-pin-
and non-grooved rachises and costae adaxially. Dicksonia natífidas, y ráquises y costas no surcados adaxialmente.
further differs by laminae strongly tapered toward the Dennsteadtia tiene también los soros marginales, pero los
base. Dennsteadtia also has marginal sori, but the indusia indusios son en forma de una copa, no bivalvos. Carece
are cup-shaped, not bivalvate. It lacks long, golden hairs de pelos largos dorados en el rizoma y bases de los
on the rhizome and petiole bases. pecíolos.

COMMENTS: Culcita consists of two species: C. coniifo- COMENTARIOS: Culcita consta de dos especies: C.
lia (Hook.) Maxon of the Neotropics, and C. macrocarpa coniifolia (Hook.) Maxon en el Neotrópico, y C. macrocar-
C. Presl, of Spain, the Azores, Madeira, and Tenerife The pa C. Presl en España, los Azores, Madeira, y Tenerife. La
neotropical species grows above 2000 m in cloud forests, especie neotropical existe arriba de 2000 m en bosques
generally in semi-open places. Previously Culcita was con- nubosos, generalmente en lugares semi-abiertos. Previa-
sidered related to Calochleana, a genus of 8 species from mente Culcita se consideró relacionado a Calochleana, un
southeastern Asia and the South Pacific. Recent molecu- género de 8 especies del sureste de Asia y el Sur Pacíf-
lar phylogenetic studies show Culcita is sister to Plagiogyria ico. Estudios filogenéticos moleculares recientes mues-
(Korall et al., 2006), with which it seems to have little in tran que Culcita es hermana a Plagiogyria (Korall et al.,
common morphologically or anatomically.The name Cul- 2006), con que tiene poco común morfológicamente o
cita means bed or cushion is Latin. It might refer to the anatómicamente. El nombre Culcita significa cama o cojín
arched, cushion-like indusia, or perhaps the soft, hairy rhi- en Latín.Tal vez se refiere a los inducíos arqueados, como
zome apex, or the use of the hairs to stuff pillows. cojín, o el ápice del rizoma suave y peludo, o el uso de los
pelos en almohadas.

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. 1922.The genus Culcita. Journal of the Washington Academy of Science 12: 454–460. White, R. A. & M. D. Turner. 1988.
Calochleana, a new genus of Dicksonioid ferns. American Fern Journal 78: 86: 95.

226
Figure 107. A–D. Culcita coniifolia. (Mickel & Smith, 2004).

227
CYATHEACEAE
Cyathea J. E. Sm.
DESCRIPTION: Usually terrestrial; stems typically ar- DESCRIPCIÓN: Usualmente terrestres; tallos típica-
borescent; petioles scales bicolorous or concolorous, mente arborescentes; escamas de los pecíolos bicolor-
lacking long black setae, the borders with cells of as o concoloras, sin setas largas negras, los bordes
different size, shape, and orientation from those con células de tamaño, forma y orientación de las
in the medial part; sterile and fertile leaves 1–4(–6) en la parte medial; hojas estériles y fértiles 1–4(–6) m
m long, monomorphous; laminae 1-pinnate to 4-pinnate de largo, monomorfas; láminas 1–pinnadas a 4-pinnadas
(most species 2-pinnate-pinnatisect), the apex pinnatifid (la mayoría de las especies son 2-pinnada-pinnatisectas),
or rarely conform; veins free or anastomosing; sori round, el ápice pinnatífido o raras veces conforme; nervaduras
receptacles elevated, usually paraphysate; indusia present libres o anastomosadas; soros redondos, receptáculos el-
or absent, globose or cup-shaped or scalelike; spores tet- evados, usualmente con paráfises; indusios presentes o
rahedral, 64 per sporangium; x=69. 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
150 species. It is sister to Alsophila, the two sharing the de alrededor de 150 especies. Es hermana a Alsophila,
synapomorphy of marginate petiole scales (i.e., scales los dos compartiéndo la sinapomorfia de escamas pecio-
with cells of the borders differentiated from those of the lares marginadas (i.e., escamas con las células por los bor-
medial portions. Cnemidaria, long recognized as a genus, des diferentes de las de la parte mediana de la escama).
is nested within Cyathea. It is herein treated as Cyathea, Cnemidaria, reconocido por mucho tiempo como género,
Cnemidaria group (which see). Trichipteris differs from Cy- es anidado dentro de Cyathea. Se trata aquí como el gru-
athea only by the absence of indusia, a structure that has po de Cyathea, Cnemidaria (vease). Trichipteris difiere de
been lost independently several times in Cyathea. Lehnert Cyathea sólo por la ausencia del indusio, una estructura
revised the species of Cyathea with globose indusia and que ha sido perdido varias veces en Cyathea. Lehnert
white petiole scurf (Lehnert, 2008) those with globose revisó las especies de Cyathea con indusios globosos y
indusia and dark petiole scurf (Lehnert, 2009). The genus caspilla peciolar blanquecina (Lehnert, 2008) y las con
name is derived from the Greek kyathos, wine cup. It al- indusios globosos y caspilla peciolar oscura (Lehnert,
ludes 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. Leh-
nert, 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. Lellinger, D. B. 1988.
The disposition of Trichopteris (Cyatheaceae). 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 Uni-
versity 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.

228
Figure 108. Cyathea caracasana var. boliviensis. A. Pinna. B. Sphaeropteroid (globose) indusia. C. Marginate petiole scale; note
cells not all of same shape as in Sphaeropteris. (Tryon & Stolze, 1989)

229
CYATHEACEAE
Cyathea, clade of Cnemidaria C. Presl
DESCRIPTION: Terrestrial; stems non-arborescent, DESCRIPCIÓN: Terrestres; tallos no arborescentes,
or if so (rarely), up to 1.5 m tall; sterile and fertile leaves o si (raras veces), hasta 1.5 m de alto; hojas estériles y
1-3.5 m long, monomorphous; petioles smooth or spiny, fértiles 1-3.5 m de largo, monomorfas; pecíolos lisos o
scaly, the scales bicolorous with a dark brown center espinosos, escamosos, las escamas bicoloras con un cen-
and white margins, lacking setae; laminae 1-pinnate to tro oscuro y márgenes blanquecinos, carecente de setas;
1-pinnate-pinnatisect (rarely 2-pinnate), with con- láminas 1-pinnadas a 1-pinnada-pinnatisectas (raras
form or pinnatifid apices; pinnae equilateral, sessile or veces 2-pinnadas), con los ápices conformes o pinnatí-
short-stalked; veins free or anastomosing; rachis and fidos; pinnas equiláteras, sésiles o cortamente peciolula-
costae glabrous adaxially (except in C. suprastrigosa das; nervaduras libres o anastomosadas; raquis y costas
R.C. Moran); sori round, the receptacle elevated, subglo- glabros adaxialmente (con excepción de C. suprastrigo-
bose; indusia scalelike, attached to the base of the recep- sa R.C. Moran); soros redondos, con receptáculos eleva-
tacle, either semicircular and attached on the costular side dos, subglobosos; indusios escamiformes, únidos a la base
of the receptacle, or circular and completely encircling del receptáculo, semicirculares y únidos al lado costular
the receptacle; spores trilete, with three large equa- del receptáculo, o circular y completamente rodeando
torial pores and often smaller ones; x=69. el receptáculo; esporas tetrahédricas-globosas, con
tres poros grandes ecuatoriales y a menudo unos
más pequeños; x=69.
SIMILAR GENERA: Cyathea (s.s.) may have laminae GÉNEROS PARECIDOS: Cyathea (s.s.) peude tener
1-pinnate to 1-pinnate-pinnatisect; however, the axes are láminas 1-pinnadas a 1-pinnada-pinnatisectas; no obstan-
always pubescent adaxially with thick strigose hairs. Most te, los ejes siempre son pubescentes adaxialmente con
species of Cyathea differ by arborescent trunks. pelos gruesos estrigosos. La mayoría de especies de Cy-
athea difiere por troncos arborescentes.

COMMENTS: The Cnemidaria clade is entirely neotrop- COMENTARIOS: El clado de Cnemidaria es completa-
ical and contains about 27 species. It inhabits wet forests mente neotropical y consta de casi 27 especies. Inhabita
from low to middle elevations. Cnemidaria is monophy- bosques húmedos de elevaciones bajas a medias. Cnemi-
letic, nested within Cyathea (Korall et al., 2006).Besides daria es monofiletico y anidado dentro de Cyathea (Korall
the characteristics in boldface in the description, many et al., 2006). Además de las características en negritas da-
species of Cnemidaria have anastomosing veins, conform das la descripción, muchas especies de Cnemidaria tienen
or subconform lamina apices, and short stems. It is rarely venas anastomosadas, ápices de las hojas conformes o
arborescent. When a trunk is present, it is usually less than subconformes y tallos cortos. Raras veces es arborescen-
one meter tall.The genus name is derived from the Greek te, pero cuando si, el tronco es usualmente menos de un
knemis, spoke of a wheel + aris, resembling. The lowest metro de altura. El nombre genérico se deriva del griego
veins radiate like spokes of a wheel. 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. Lehnert,
M. 2011. A synopsis of the species of Cyathea (Cyatheaceae–Polypodiopsida) with pinnate to pinnate-pinnatifid fronds. Phytotaxa
61: 17–36. Stolze, R. G. 1974. A taxonomic revision of the genus Cnemidaria (Cyatheaceae). Fieldiana, Botany 37: 1–98.

230
Figure 109. A–E. Cyathea (Cnemidaria) grandifolia. F–H. C. horrida. (original, © Moran & Mickel, 2010)

231
DRYOPTERIDACEAE
Cyclodium C. Presl
DESCRIPTION: Terrestrial, hemiepiphytic, or rheoph- DESCRIPCIÓN: Terrestres, hemiepífitas o reofíticas; ri-
ytic; rhizomes creeping, scaly; leaves monomorphic or zomas reptantes, escamosos; hojas monomorfas a algo
slightly dimorphic; petioles with 4 or more vascular bun- dimorfas; pecíolos con 4 o más haces vasculares; lámina
dles; laminae pinnate to 2-pinnate-pinnatifid, glabrous or pinnada a 2-pinnado-pinnatífida, glabra o casi así, anádro-
nearly so, anadromous, chartaceous to coriaceious, the ma, cartácea a coriácea, el ápice abruptamente reducido
apex pinnatifid, abruptly tapered or similar in form to the o pinnatífido similar en forma a las pinnas laterales; seg-
lateral pinnae; longest segments on the acroscopic side mentos más largos en el lado acroscópico de las pinnas;
of the pinnae; rachis and costae grooved adaxially, the raquis y costas sulcados adaxialmente, los surcos más o
grooves more or less continuous with those of the lower menos confluentes de un eje al siguiente, pelosos por
order, puberulent within the groove; costae abaxially with dentro; costas abaxialmente con tricomas uniseriados,
uniseriate trichomes, reddish or brownish, appressed; rojizos oparduscos, adpresos; nervaduras libres o anas-
veins free or anastomosing, ending at the margins, not en- tomosadas, terminándose antes del margen, no hichados;
larged; sori round; indusium round, peltate or attached soros redondeados; indusio redondeado, peltado o
at a narrow sinus, often deciduous before the sporangia unido a un seno estrecho, a menudo deciduo antes de la
mature; spores monolete; x=41. madurez del esporangio; esporas monoletes; x=41.

SIMILAR GENERA: Arachniodes differs by more finely GÉNEROS PARECIDOS: Arachniodes difiere por lámi-
divided blades. Polybotrya differs by (mostly) climbing rhi- nas anadrómicas y rizomas reptantes. Polybotrya es (prin-
zomes and strongly dimorphic sterile and fertile leaves. cipalmente) un helecho trepador y tiene hojas estériles y
Dryopteris differs by erect or decument rhizomes. Stig- fértiles dimorfas. Dryopteris difiere por rizomas erectos o
matopteris differs by thinner blades with internal punctu- decumentes. Stigmatopteris difiere por laminas más delga-
ate glands, and veins ending before the margins in clavate das con glándulas internas punctatas y venas terminando
hydathodes. ante del margen en hidatodos clavados.
COMMENTS: Cyclodium is entirely neotropical and con- COMENTARIOS: Cyclodium es completatmente neo-
tains about 10 species, these mostly in the Guianas. It is tropical y consta de casi 10 especies, estas principalmente
primarily a lowland genus, growing only up to 1500 m. en las Guianas. Es un genero de las tierras bajas, alca-
It forms a clade with other dryopteroid ferns that have ncando solo hasta 1500 m. Forma un clado con otros
creeping rhizomes and (often) dimorphic sterile and fer- helechos dryopteroides que tienen rizomas reptantes y
tile leaves, such as Maxonia, Olfersia, Polybotrya, and Polys- (a menudo) hojas estériles y fértiles dimorfas, tales como
tichopsis. The genus name is derived from the Greek kyk- Maxonia, Olfersia, Polybotrya, and Polystichopsis.El nombre
lodes, ring-like. It alludes to the circular indusia. genérico se deriva del griego kyklodes, como un anillo.
Alude a los indusios circulares.

LITERATURE: Smith, A.R. 1986. Revision of the neotropical fern genus Cyclodium. American Fern Journal 76: 56–98.

232
Figure 110. Cyclodium meniscioides. A. Habit. B. Venation. C. Fertile pinnae. D. Detail of sori. (©Robbin Moran, 2009)

233
LOMARIOPSIDACEAE
Cyclopeltis J. Sm.
DESCRIPTION: Terrestrial; rhizomes erect or decum- DESCRIPCIÓN: Terrestres; rizomas erectos o decum-
bent, scaly; sterile and fertile leaves monomorphous; bentes, escamosos; hojas estériles y fértiles monomórfi-
laminae 1-pinnate with a subconform apical pinna; cas; láminas 1-pinnadas con una pinna apical subcon-
pinnae articulate to the rachis, entire or nearly so, forme; pinnas articuladas al raquis, enteras o casi
the basiscopic side auriculate and overlapping the enteras, el lado basiscópico auriculado y traslapa
upper surface of the rachis; rachis and costae round- la superficie superior del raquis; raquis y costas en la
ed or flattened above (not grooved), puberulent adaxially, superficie superior redondeados o planos (no surcados),
these 0.1–0.2 mm long, reddish, articulate; veins free; sori puberulentos adaxialmente, estos 0.1–0.2 mm de largo,
round, in 2 or 3 rows between the costa and mar- rojizos, articulados; nervaduras libres; soros redondea-
gin; indusia peltate. Spores monolete. dos, dispuestos en 2 o 3 hileras entre la costa y el
margen; indusios peltatos. Spores monolete.

SIMILAR GENERA: Nephrolepis differs by having sto- GÉNEROS PARECIDOS: Nephrolepis difiere por ten-
lons, pinnatifid (not conform) lamina apices, and only one er estolones, ápices de la láminas pinnatífidos (no con-
row of sori between the costa and margin. Lomariopsis formes) y solamente una hilera de soros entre la cos-
differs by epiphytic or hemiepiphytic habit, long-creeping ta y el margen. Lomariopsis difiere por hábito epifítico
rhizomes, sterile-fertile leaf dimorphy, and acrostichoid o hemiepifítico, rizomas largamente rastrero, dimorfismo
sori. entre las hojas estériles y fértiles y soros acrosticoides.

COMMENTS: Cyclopeltis has one species in the New COMENTARIOS: Cyclopeltis tiene una especie en el
World (C. semicordata (Sw.) J. Sm.) and five in southeast- Nuevo Mundo (C. semicordata (Sw.) J. Sm.) y cinco en
ern Asia and the western Pacific. The American species el sureste de Asia y el Pacífico occidental. La especie
is common and widespread in wet forests mostly below americana es común y difundida y occure en bosques
600 m. The articulations of the pinnae can be seen as húmedos principalmente bajo los 600 m. Las articula-
dark line around the base of the costa where it joins the ciones de las pinnas puede ser observadas como líneas
rachis.They appear to be nonfunctional; that is, the pinnae oscuras alrededor de la base de la costa donde se une
do not disarticulate. The presence of articulations is the al raquis. Aparecen ser no funcional; o sea, las pinnas no
only morphological synapomorphy known for the Lom- se desprenden. La presencia de las articulaciones es la
ariopsidaceae.The genus name is derived from the Greek única sinapomorfia conocida para las Lomariopsidaceae.
kyklos, circle + peltis, shield, and refers to the indusium. El nombre genérico se deriva del griego kyklos, circulo +
peltis, shield, a se refiere al indusio.

LITERATURE: Lu, J. M. & D. Z. Li. 2006. A study on systematic position of Cyclopeltis. Acta Botanica Yunnan 28(4): 337–340.
[in Chinese]

234
Figure 111. . A, B. Cyclopeltis semicordata (Sw.) J. Sm. (Mickel & Smith, 2004)

235
CYSTOPTERIDACEAE
Cystopteris Bernh.
DESCRIPTION: Terrestrial or saxicolous; rhizomes DESCRIPCIÓN: Terrestres o saxícolas; rhizomas rep-
creeping, scaly, the scales thin, translucent; sterile and fer- tantes, escamosos, las escamas delgadas, translúcidas;
tile leaves dimorphous, thin-textured; petioles with 2 hojas estériles y fértiles dimorfas, de textura delgada;
vascular bundles; laminae usually 15–40 cm long, 2-pin- pecíolos con 2 haces vasculares; láminas usualmente
nate-pinnatifid to 4-pinnate, anadromic; axes grooved 15–40 cm de largo, 2-pinnado-pinnatífidas a 4-pinnadas,
on the upper surface, the grooves decurrent; veins free, anadrómicas; ejes surcados en la superficie superior, los
the tips slender (not enlarged); sori round, medial or in- surcos decurrentes; nervaduras libres, los ápices delgados
framedial; indusia hyaline, scalelike, attached to the (no engrosados); soros redondeados, mediales o infra-
proximal side of the receptacle and (when young) mediales; indusios hialinos, como una escama, unidos
arching over the sporangia, evanescent, often hidden al lado proximal del receptáculo y (cuando jóvenes)
by the mature sporangia; spores monolete, bilateral; x=42. arqueando sobre los esporangios, evanescentes, a
menudo escondido por los esporangios maduros; espo-
ras monoletes, bilaterales; x=42.

SIMILAR GENERA: Woodsia resembles Cystopteris but GÉNEROS PARECIDOS: Woodsia se parece a
differs by having the indusium attached completly around Cystopteris pero difiere por tener el indusio unido com-
the base of the receptacle. At maturity, the indusium pletamente alrededor de la base del receptáculo. Al mad-
splits into lobes or hairs that form a loose cup around urar, el indusio se hende en lobulos o pelos que forman
the sporangia. In sterile specimens, the veins tips are diag- una copa suelta alrededor de los esporangios. En ejem-
nostic: those of Woodsia are enlarged, whereas those of plares estériles, las venas son diagnósticas: las de Woodsia
Cystopteris are slender. son engrosadas, mientras que las de Cystopteris son del-
gadas.

COMMENTS: Cystopteris contains about 10 species COMENTARIOS: Cystopteris consta de alrededor de

and is primarily temperate. The common and widespread 10 especies y es principalmente templado. La especie
species in the neotropics is C. fragilis (L.) Bernh. It is usually común y muy difundida en el neotrópico es C. fragilis (L.)
found above 2500 m. Throughout Latin America this spe- Bernh. Se encuentra usualmente sobre los 2500 m. Por
cies has diploid, tetraploid, and hexaploid cytotypes. Some todo América Latín esta especie tiene citotipos diploides,
specimens have misshapen spores, suggesting hybridiza- tetraploides y hexaploides. Algunos ejemplares tienen es-
tion.Two notes about observing identifying the genus:The poras malformadas, sugiriendo hibridación. Dos notas so-
indusium is often pushed back and hidden by the sporan- bre identificar el género: El indusio es a menudo empuja
gia. Therefore, it is difficult to see in mature sori. The two hacia atrás y escondido por los esporangios. Por eso es
vascular bundles in the petioles can usually be seen in difícil verlo en soros maduros. Los dos haces vasculares
specimens where the petiole has been broken.The genus en el pecíolo se pueden ver usualmente en ejemplares
name is derived from the Greek kystos, bladder + pteris, donde el pecíolo ha sido quebrado. El nombre genérico
fern. It refers to the inflated indusium. se deriva del griego kystos, vejiga + pteris, helecho. Se
refiere al indusio inflado.

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.

236
Figure 112. A–D. Cystopteris fragilis. G, H, J. C. membranifolia. (Mickel & Smith, 2004)

237
MARATTIACEAE
Danaea Sm.
DESCRIPTION: Terrestrials (rarely saxicolous), fleshy; DESCRIPCIÓN: Terrestres (raras veces saxícolas), car-
rhizomes creeping or erect; sterile and fertile leaves di- nosas; rizomas reptantes o rectos; hojas estériles y fértiles
morphic, 1-pinnate (rarely simple), with a conform ter- dimorfas, 1-pinnadas (raras veces simples), con una pinna
minal pinna or this is replaced by a bud; petioles with terminal conforme o en su lugar un brote; pecíolos con o
or without swollen nodes, basally with rounded, earlike, sin nudos hinchados, basalmente con estípulas redondea-
fleshy stipules; pinnae opposite, entire or with serrate das parecidas a orejas; pinnas opuestas, enteras o con
apices; rachis swollen-nodose at the insertion of the ápices serrados; raquis nodoso-hinchado en la unión de
pinnae; veins free, simple or 1-forked near the base, las pinnas; nervaduras libres, simples o 1-furcadas cerca
parallel to each other; fertile leaves long-petiolate and de la base, paralela entre sí; hojas fértiles lárgamente pe-
therefore taller than the sterile, with narrower pinnae; cioladas y por eso más largas que las estériles, con pinnas
sporangia fused into an elongate, double-rowed synan- más angostas; esporangios fusionados en sinangios elon-
gium, with 20–100 locules, opening by terminal pores; gados de doble hilera, con 20100 lóculos, abriéndose
synangia covering most of the distance between the costa por poros terminales; sinángios cubren la mayoría de la
and margin; x=40. distancia entre la costa y el margen; x=40.

SIMILAR GENERA: Lomariopsis differs by its long-creep- GÉNEROS PARECIDOS: Lomariopsis difiere por sus
ing and climbing stems, alternate pinnae, and lack of swol- tallos largamente reptantes y rastreros, pinnas alternas
len nodes along the rachis. Metaxya differs by hairs at y la ausencia de nudos hichados por el raquis. Metaxya
the base of the petioles, the absence of swollen nodes at difiere por pelos en la base de los pecíolos, la aucencia
the rachis-pinna junctures, and round sori. Once-pinnate de nudos hinchados en la unión del raquis y pinnas, y so-
species of Pteris differ by areolate veins and sori along the ros redondos. Especies de Pteris 1-pinnadas difieren por
margins. 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 mente americano con alrededor de 50 especies. Se en-
to middle elevations. When dried, the swollen nodes on cuentran en bosques húmedos en elevaciones bajas a
the petioles and rachises shrink and darken. In several medias. Cuando es secado, los nudos hinchados de los
species, the apical pinna is often or always replaced by a pecíolos y ráquises se encogen y oscuren. En varias espe-
bud. After the leaf has died, the petiole bases and stipules cies, la pinna apical es a menudo o siempre replazado por
persist on the stem and contribute substantially to its di- un brote. Después de que la hoja se muere, las bases de
ameter. The genus totally lacks hairs; the only indument los pecíolos persisten sobre el tallo y contribuyen sub-
consists of peltate, irregular scales that provide few dis- stancialmente a su diámetro. El género carece totalmente
tinctions between the species. When collecting Danaea, it de pelos; el único indumento consiste de escamas pelta-
is important to note whether the rhizome is creeping or das iregulares que provienen pocas distinciones entre las
erect. The very young leaves of some species are irides- especies. Cuando se colecta a Danaea, hay que notar si el
cent. The genus is named for Giovanni Pietro Maria Dana rizoma es reptante or erecto. Las hojas muy jóvenes de
(1736–1801), professor of botany and friend of J. E. Smith. algunas especies son iridescentes. El género se 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. Underwood, L. M. 1902. American Ferns–V. A review of the genus
Danaea. Bulletin of the Torrey Botanical Club 29: 699–679. Tuomisto, H. & R. C. Moran. 2001. Marattiaceae. In: G. Harling
(editor). Flora of Ecuador 66: 23–68.

238
Figure 113. Danaea elliptica. Note in C the rounded stiples, and in B the poricidal synangia. Fijase en C las estipulas redon-
deadas y en B los sinángios poricidales. (from Proctor, 1985; drawn by Peter J. Edwards)

239
DENNSTAEDTIACEAE
Dennstaedtia Bernh.
DESCRIPTION: Terrestrial or epiphytic; rhizomes DESCRIPCIÓN: Terrestres o epífitas; rizomas rastre-
creeping, pubescent, without scales; leaves 1–4(-7) ros, pubescentes, sin escamas; hojas 1–4(-7) m de largo,
m long, erect or supported by surrounding vegetation; erectas o apoyándose sobre la vegetación circundante;
petiole bases frequently with buds that give rise to bases de los pecíolos frecuentemente portando
new stems, in transverse section with a vascular bundle yemas que dan lugar a nuevos tallos, en corte trans-
en the form of an omega; laminae 2-pinnate-pinnatifid versal con un haz vascular en forma de omega; láminas
to 4-pinnate, rarely (in D. wercklei (H. Christ) R.M. Tryon) 2-pinnado-pinnatífidas a 4-pinnadas, raras veces (en D.
1-pinnate, some species with buds in the axils of the distal wercklei (H. Christ) R.M. Tryon) 1-pinnatas, algunas espe-
pinnae; pinnules arranged anadromically; veins free, end- cies con yemas en las axilas de las pinnas distales; pínnulas
ing behind the margin in slender or clavate apices; sori arregladas anadrómicamente; venas libres, terminando
marginal, deflexed, terminal on the vein apices, in most antes del margen, con las puntas delgadas o claviformes;
species borne in the sinuses between lobes; indusia cup- soros marginales, deflexos, terminales en las puntas de
shaped; spores tetrahaedral-globose, trilete, nongreen. las venas, naciendo en la mayoría de las especies en los
x=30, 33, 34, 46, 47. senos entre los lobulos; indusios en forma de copa;
esporas tetrahédricas-globosas, triletes, no verdes. x=30,
33, 34, 46, 47.
SIMILAR GENERA: Hypolepis differs by false indusia. GÉNEROS PARECIDOS: Hypolepis difiere por false
Paesia and Pteridium differs by elongate sori supplied by indusia. Paesia y Pteridium tienen soros alargados y sum-
several to many veins. Saccoloma differs by persistant pet- inistrados con varias a muchas venas. Saccoloma difiere
iole bases around a short, thick rhizome. por los bases de los pecíolos persistentes alrededor de
un rizoma corto y grueso.
COMMENTS: Dennstaedtia is mostly pantropical with COMENTARIOS: Dennstaedtia es mayormente pan-
about 20 species in the New World. With the excep- tropical con alrededor de 20 especies en el Nuevo
tion of D. bipinnata and D. globulifera, all other species in Mundo. Con excepción de D. bipinnata y D. globulifera,
the genus have epipetiolar buds. These are borne on the todas las otras especies del género tienen ramas epipe-
sides of the petiole near the base and may number more ciolares. Estas provienen de los lados del pecíolo cerca
than one. They are an integral part of the branching sys- de la base y pueden ser varias en número. Son una parte
tem of the plant. Epipetiolar branches are found in other integral de la sistema de ramificación de la planta. Ramas
dennstaedtioid genera such as Histiopteris, Hypolepis, and epipeciolares se encuentran en otros géneros de Denn-
Pteridium. The genus name is named for August Wilhelm staedtiaceae como Histiopteris, Hypolepis y Pteridium. El
Dennstaedt (1776–1826), German botanist and author. 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 char-
acters, new species and a new combination. Nordic Journal Botany 319–346. Tryon, R. M. 1960. A review of the genus Denns-
taedtia in America. Contributions from the Gray Herbarium of Harvard University 187: 23–52.

240
Figure 114. A–J: Dennstaedtia cornuta. Q–K. D. bipinnata. R–U. D. globulifera. (Mickel & Smith, 2004).

241
DICKSONIACEAE
Dicksonia L’Hér.
DESCRIPTION: Terrestrial; stems arborescent, to 10 DESCRIPCIÓN: Terrestres; tallos arborescentes, has-
m tall, densely covered by adventitious roots; petioles ta 10 m de longitud, densamente cubierto por raíces ad-
with dense, wool-like, golden brown hair, lacking venticias; pecíolos cubiertos por pelos como lana, densos,
spines; sterile and fertile leaves 1–3.5 m long, monomor- largos, pardo-dorados, espinas ausesntes; hojas estériles y
phous; laminae 2-pinnate-pinnatifid to 4-pinnate, cori- fértiles 1–3.5 m de largo, monomorfas; láminas 2-pinna-
aceous, gradually reduced basally and apically; axes do-pinnatífidas a 4-pinnadas, coriáceas, gradualmente
adaxially not grooved, pubescent; veins free; sori mar- reducidas basalmente y apicalmente; ejes adaxial-
ginal, reflexed; indusia bivalved, with a scarious inner mente no surcados, pubescentes; nervaduras libres; so-
(abaxial) lobe and a thicker green outer (adaxial) lobe, ros marginales, reflejados; indusios bivalvados, con una
the two lobes appearing clam-shaped; spores tetrahe- valva interior (abaxial) escariosa y una valva exterior
dral-globose; x = 65. (adaxial) más gruesa y verde, las dos valvas semejan a una
almeja; esporas tetrahédricas-globosas; x = 65.
SIMILAR GENERA: Alsophila, Cyathea, and Sphaerop- GÉNEROS PARECIDOS: Alsophila, Cyathea y Sphaer-
teris differ by scaly petioles, abaxial sori (not marginal), opteris difieren por pecíolos escamosos, soros abaxiales
and varied shapes of indusia but never bivalved (in some (no marginales), e indusios variados pero nunca bivalva-
species the indusia are lacking). Culcita differs by lacking dos (en algunas especies los indusios están ausentes). Cul-
a trunk, deltate laminae, and rachises and costa grooved cita difiere por faltar un tronco, laminas deltatas y ráquises
adaxially. y costas surcados adaxialmente.

COMMENTS: Dicksonia contains about 20 species, pri- COMENTARIOS: Dicksonia consta de alrededor de 20
marily in the Malesian-Australian region. Three species especies principalmente en la región Malesian-Australiana.
occur in the New World: D. berteroana (Colla) C. Chr. Three especies existen en el Nuevo Mundo: D. berteroa-
(endemic to Juan Fernández Islands), D. sellowiana Hook. na (Colla) C. Chr. (endémica a las islas Juan Fernández),
and D. spruceana Kuhn. Only D. sellowiana is common and D. sellowiana Hook. and D. spruceana Kuhn. Solamente D.
widespread. It grows above 1500 m in cloud forests. It sellowiana es común y difundida. Existe arriba de 1500 m
can usually be recognized from a distance by its basally en bosques nubosos. Usualmente se puede reconocer
tapering laminae, short petiole, and dead leaves that per- de una distancia por sus láminas reducidas basalmente,
sist around the stem to form a “skirt.” Dicksonia is named pecíolos cortos, y hojas muertas que persisten alrededor
for James Dickson (1738–1822), Scottish physician and del tallo formando una “falda”. Dicksonia se nombra para
botanist. 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. 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 sequenc-
es. Plant Systematics and Evolution 219: 263–270.

242
Figure 115. A–C. Dicksonia sellowiana. (Mickel & Smith, 2004).

243
GLEICHENIACEAE
Dicranopteris Bernh.
DESCRIPTION: Terrestrial; rhizomes long-creeping, DESCRIPCIÓN: Terrestre; rizomas largamente rastre-
pubescent, the hairs stiff, multicellular, bearing leaves in ros, pubescentes, los pelos tiesos multicelulares, llevando
one row on the dorsal surface; leaves indeterminate, of hojas en una hilera en la superficie dorsal; hojas indeter-
periodic growth; pinnae opposite, forked pseudodichoto- minadas, de crecimiento periódico; pinnas opuestas, fur-
mously, the bud in the angle of each fork pubescent cadas pseudodicotómicamente, el brote en el ángulo
by dark bristles; penultimate segments pectinate; veins de cada bifurcación pubescente por cerdas oscuros;
2–4-forked; sori round, without indusia; sporangia usu- segmentos penúltimos pectinados; venas 2-4-furcadas;
ally 8–15 per sorus, sessile; spores trilete; x=39. soros redondos, sin indusios; esporangios usualmente
8–15 por soro, sésiles; esporas triletes; x=39.

SIMILAR GENERA: Sticherus differs by scaly buds in the GÉNEROS PARECIDOS: Sticherus difiere por brotes
pinna forks, scales (never bristles) present on the costae escamosos en las furcaciones de las pinnas, y escamas
and costules, veins 1-forked, and 3–6 sporangia per sorus. (nunca cerdas) usualmente presentes en las costas y
Diplopterygium differs by unforked pinnae and the pres- cóstulas. Diplopterygium difiere por pinnas no bifurcadas
ence of scales (not bristles) on the leaf. y la presencia de escamas (no cerdas) en la hoja.

COMMENTS: Dicranopteris is pantropical and consists COMENTARIOS: Dicranopteris es pantropical y consta

of about 10 species. It is found in open or semi-open hab- de ca. 10 especies. Se encuentra en habitats abiertos o
itats, especially roadsides. Their leaves grow periodically, semi-abiertos, especialmente en taludes. Sus hojas crecen
forming a dormant bud on the tip of the rachis while the periódicamente y forman un brote latente en el ápice
two opposite pinnae develop beneath. When the pinnae del raquis mientras las dos pinnas opuestas se desarrol-
finish their development, the bud at the tip of the rachis lan abajo. Cuando las pinnas terminan su desarrollo, el
continues growth. This process may repeat itself many brote latente apical del raquis continua su crecimiento.
times, resulting in leaves up to 20 m long. The pinnae are Este proceso puede repetirse muchas veces resultando
also forked and have a dormant bud in the axil(s). These en hojas de hasta 20 cm de largo. Las pinnas son tam-
buds usually do not develop unless the rachis bud is dam- bién furcadas y tienen un brote latente en la(s) axila(s).
aged or destroyed. Two species are common and wide- Estos brotes usualmente no se desarrollan a menos que
spread in the Neotropics: Dicranopteris flexuosa (Schrad.) el brote del raquis sea dañado o destruido. Dos especies
Underw. and D. pectinata (Willd.) Underw. The genus son comunes y muy difundias en el Neotrópico: Dicran-
name is derived from the Greek dikranos, two-pronged + opteris flexuosa (Schrad.) Underw. y D. pectinata (Willd.)
pteris, fern. The leaves and pinnae are repeatedly forked. Underw.El nombre genérico se deriva del griego dikranos,
dos-cuernos + pteris, helecho. Las hojas y pinnas se bifur-
can repetidamente.

LITERATURE: see Sticherus; véase Sticherus.

244
Figure 116. A-D: Dicranopteris flexuosa. (Mickel & Smith, 2004)

245
HYPODEMATIACEAE (EUPOLYPODS I)
Didymochlaena Desv.
DESCRIPTION: Terrestrial; leaves 1–2 m long, the ster- DESCRIPCIÓN: Terrestres; hojas 1–2 m de largo, las es-
ile and fertile monomorphous; laminae 2-pinnate, im- tériles y fértiles monomorfas; láminas 2-pinnadas, im-
paripinnate; pinnae 15–20 pares, sessile; pinnules 1–2.5 paripinnadas; pinnas 15–20 pares, sésiles; pínnulas 1–2.5
× 0.5–1 cm, subdimidiate, sessile, obtuse, entire or cren- × 0.5–1 cm, subdimidiadas, sésiles, obtusas, enteras o
ate, straight along the basiscopic side; basal acroscop- crenuladas, rectas a lo largo del lado basiscópico; pín-
ic pinnule overlapping the rachis; ridges of the axes nulas basales basiscopicas sobrelapando el raquis;
with spine-like projections on the upper surface of the crestas de los ejes con proyecciones espiniformes en
junctures of the rachis, costae, and costules; axes scaly; la superficie adaxial en las uniones del raquis, costas y
veins free, hydathodes present; sori elongate, along cóstulas; ejes escamosos; nervaduras libres, hydatodos
both sides of the vein and curved around its apex; indusia presentes; soros alargados, a lo largo de ambos lados
round distally, entire; spores monolete; x=41. de la nervadura y curvados alrededor de su ápice; indu-
sios redondeados en el extremo distal, enteros; esporas
monoletes; 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 COMENTARIOS: Un género pantropical que consiste
species D. truncatula (Sw.) J. Sm. (a second species might de una sola especie D. truncatula (Sw.) J. Sm. (una segunda
occur in Madagascar). It grows in wet forests at low to especie posiblemente existe en Madagascar). Crece en
mid elevations and is common and widespread through- bosques húmedos en elevaciones bajas o medias, es es
out the Neotropics. Didymochlaena has two characters común y muy difundida por todo el Neotrópico. Didy-
unique among the Eupolpods I: elongate sori and minute mochlaena tiene dos características únicas entre los Eu-
spine-like projections on the upper surface of the costa polypods I: soros alargados y proyecciones espiniformes
where it joins the costule. The young unfurling leaves are minutas en la superficie superior de las costa donde se
reddish brown, giving rise to the common name of the une con las pínnulas. Las hojas jóvenes en el proceso
plant in horticulture: mahogany fern. The leaves emit a de desplagarse son pardo rojizas, dando lugar al nom-
skunk-like odor when cut or crushed, similar to that of bre común en la horticultura de helecho de mahoga-
Phanerophlebium. The genus name is derived from the ny. Cuando cortadas o estrujadas, las hojas emiten un
Greek didymos, double + chlaena, cloak. It refers to the olor como zorrillo, similar a eso en Phanerophlebium. El
indusia that bends around the vein apex and covers both nombre genérico se deriva del griego didymos, double +
sori, thus appearing double. chlaena, capa. Se refiere al los indusios que doblan alred-
edor del ápice de la vena para cubrir los dos soros, por
eso apareciéndose doble.

LITERATURE: None; ninguna.

246
Figure 117. Didymochlaena truncatula (Tryon & Stolze, 1991)

247
 Didymoglossum sect. Didymoglossum
DESCRIPTION: Epiphytic; roots absent; rhizomes ca. DESCRIPCIÓN: Epífitas; raíces ausentes; rizomas a 1
1mm wide, long-creeping, branched, densely pubescent, mm de ancho, largamente rastreros, ramificados, densa-
the hairs blackish; leaves to 6 cm long, entire to bipinnat- mente pubescentes, los pelos negruzcos; hojas a 6 cm de
ifid, margins pubescent, the hairs dark, simple or paired largo, enteras a bipinnatífidas, márgenes pubescentes;
or arranged in stellate groups of 3 or 4; petioles short or pecíolos cortos o ausentes, a menudo pubescentes con
absent, often pubescent with dark hairs like those on the pelos oscuros como los en el rizoma; costas presentes o
rhizomes; costae present or absent, if absent; veins cata- ausentes, si ausentes; venas catádromas, a menudo flabel-
dromous, often flabellate; false veins present as short, ladas; venas falsas presentes como rayos cortos débiles
faint streaks between the true veins, continuous sub- entre las venas verdaderas, falsa vena continua sub-
marginal false vein absent; sori one to several, margin- marginal ausente; soros uno a varios, marginales en la
al in the distal part of the leaf; indusia funnel-shaped or parte apical de la hoja; indusios infundibuliformes o tubu-
tubular, bilabiate, the lobes often with dark margins; lares, bilabiados, los lóbulos con márgenes oscuros;
receptacles exert; spores green, trilete; x=34. receptáculos exertos; esporas verdes, triletes; x=34.

SIMILAR GENERA: Didymoglossum sect. Microgonium GÉNEROS PARECIDOS: Didymoglossum sect. Microgo-
differs by the presence of a continuous submarginal false nium difiere por la precencia de una falsa vena continua
vein, marginal hairs absent, indusia truncate (not bival- submarginal, pelos marginales ausentes, indusios trunca-
vate) without a dark margin. Polyphlebium differs by false dos (no bilobados) sin un margen oscuro. Polyphlebium
veins absent and larger leaves. difiere por venas falsas ausentes y hojas más grandes.

COMMENTS: Section Didymoglossum is mainly neotrop- COMENTARIOS: La secc. Didymoglossum es principal-

ical and contains about 20 species. Wessels Boer (1962) mente neotropical y consta de ca. 20 especies. Wessels
recognized 15 species in his monograph of the neotropi- Boer (1962) reconoció 15 especies en su monografía de
cal taxa.They grow in wet forests up to ca. 1500 m, usually los taxones neotropicales. Crecen en bosques húmedos
as low-trunk epiphytes and sometimes on moss-covered hasta 1500 m, usualmente como epífitos en las partes
rocks. Often they are overlooked because of their small basal de troncos y a veces sobre rocas cubiertas con
size. An unusual character of Didymoglossum (both subge- musgos. Usualmente están pasados por alto a causa de
nera) are false veins. These are elongate collenchymatous su tamaño pequeño. Una característica poco común de
cells that appear as faint streaks between and parallel to Didymoglossum (ambos subgéneros) es falsas venas. Estos
the main veins. Section Microgonium also has a continuous son células colenquímatas alargadas y se aparecen como
submarginal false vein parallel to the margin (this lacking rayas finas entre (y paralelas a) las venas principales. La
in sect. Didymoglossum). The dark hairs on the rhizome sección Microgonium tiene también una falsa vena conti-
are specialized for adhesion to the substrate, branching nua submarginal (esto ausente en secc. Didymoglossum).
when they touch the surface (Schneider, 2000). The ge- Los pelos oscuros en el rizoma son especializados para
nus name is derived from the Greek didymos, double + adherir al sustrato, ramificando en ápice cuando tocan la
glossa, tongue. The mouth of the indusium typically con- superficie (Schneider, 2000). El nombre genérico se deri-
sists of two widely spreading lobes. va del griego didymos, doble + glossa, lengua. La boca del
indusio típicamente consta de dos lóbulos ampliamente
divergentes.

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. 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.

248
Figure 118. A. Didymoglossum kraussii. B. D. membranaceum. C, E. D. bucinatum. D. D. ovale. F. D. hymenoides. (modified
from Mickel & Smith, 2004)

249
 Didymoglossum sect. Microgonium
DESCRIPTION: Epiphytic; roots absent; rhizomes ca. DESCRIPCIÓN: Epífitas; raíces ausentes; rizomas a 1
1mm wide, long-creeping, branched, densely pubescent, mm de ancho, largamente rastreros, ramificados, densa-
the hairs blackish; leaves to 6 cm long, entire to bipin- mente pubescentes, los pelos negruzcos; hojas a 6 cm
natifid, margins glabrous; petioles short or absent, of- de largo, enteras a bipinnatífidas, márgenes glabros;
ten pubescent with dark hairs like those on the rhizomes; pecíolos cortos o ausentes, a menudo pubescentes con
costae present or absent, if absent; veins catadromous, pelos oscuros como los en el rizoma; costas presentes o
often flabellate; false veins present as short, faint streaks ausentes, si ausentes; venas catádromas, a menudo flabel-
between the true veins, continuous submarginal false ladas; venas falsas presentes como rayos cortos débiles
vein present; sori one to several, marginal in the distal entre las venas verdaderas, falsa vena continua sub-
part of the leaf, usually completely immersed in the lami- marginal presente; soros uno a varios, marginales en
na; indusia funnel-shaped or tubular, truncate (not bila- la parte distal de la hoja, usualmente inmersados en la
biate), without dark margins; receptacles exert; spores lámina; indusios infundibuliformes o tubulares, truncados
green, trilete; x=34. (no bilabiados), sin márgenes oscuros; receptáculos
exertos; esporas verdes, triletes; x=34.

SIMILAR GENERA: Didymoglossum sect. Didymoglos- GÉNEROS PARECIDOS: Didymoglossum sect. Didymo-
sum differs by the absence of a continuous submarginal glossum difiere por la ausencia de una falsa vena conti-
false vein, the indusia bilabiate (not truncate) with dark nua submarginale, indusios bilabiados (no truncados) con
margins, and marginal hairs present. Polyphlebium differs bordes oscuros, y pelos marginales presentes. Polyphlebi-
by false veins absent and larger leaves. um difiere por venas falsas ausentes y hojas más grandes.

COMMENTS: Section Microgonium is mainly paleotrop- COMENTARIOS: La sect. Microgonium es principal-

ical and consists of about 10 species. They occur as ep- mente paleotropical y consta de casi 10 especies. Exis-
iphytes or on mossy rocks in wet tropical forests. In his ten como epífitas o sobre rocas musgosos en bosques
monograph of the neotropical taxa, Wessels Boer (1962) húmedos. En su monografía de taxones neotropicales,
recognized 4 species: T. ekmanii, T. godmanii, T. hookeri, and Wessels Boer (1962) reconoció 4 especies: T. ekmanii, T.
T. kapplerianum. Often they are overlooked because of godmanii, T. hookeri y T. kapplerianum. Muchas veces están
their small size. Both subgenera of Didymoglossum lack pasados por alto a causa de su tamaño pequeño. Los
roots but have root-like rhizomes (Schneider, 2000). dos subgéneros de Didymoglossum faltan de raíces pero
These are short, leafless rhizomes that are densely pubes- tienen rizomas que asemejan a raíces (Schneider, 2000).
cent with adhesive hairs. They help anchor the plant to Estos son rizomas cortas sin hojas que son densamente
the substrate. (For comments about adhesive hairs and pubescentes con pelos adhesivos. Ayudan anclar la planta
false veins, see subgen. Didymoglossum.) The chromosome al sustrato. (Para comentarios sobre pelos adhesivos y fal-
number x=34 is considered derived within the family.The sas venas, vease subgén. Didymoglossum.) El número cro-
name Microgonium is derived from the Greek mikros, mosómico x=34 se considera derivado dentro de la fa-
small + gonos, progeny, referring to the size of these ferns. milia. El nombre Microgonium se deriva del griego mikros,
pequeño + gonos, progenie, refiriéndose al tamaño de
estos helechos.

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.

250
Figure 119. A. Didymoglossum godmanii. B. D. ekmanii. (modified from Mickel & Smith, 2004)

251
ATHYRIACEAE (EUPOLYPODS II)
Diplazium Sw.
]DESCRIPTION: Terrestrial; rhizomes creeping to erect, DESCRIPCIÓN: Terrestres; rizomas rastreros a erec-
scaly, the scales not clathrate; sterile and fertile leaves tos, escamosos, las escamas no clatradas; hojas estériles
monomorphous; petioles with 2 vascular bundles ba- y fértiles monomorfas; pecíolos con 2 haces vascu-
sally, these uniting distally to form a “U”; laminae lares basalmente, éstos uniéndose distalmente
simple to 4-pinnate; veins free or areolate, the areoles para formar un haz vascular en forma de “U”;
without included veinlets; sori elliptic to linear, usually láminas simples a 4-pinnadas; venas libres o areoladas
paired back to back along a single vein, indusiate,; an- sin venillas incluidas; soros elípticos o lineares, usual-
nulus consisting of 15–20 cells; spores bilateral, brown; mente pareados de par en par por una sóla vena,
x=41. con indusios,; anillo consiste de 15–20 células; esporas
bilaterales, pardas; x=41.

SIMILAR GENERA: Asplenium differs by more than 20 GÉNEROS PARECIDOS: Asplenium difiere por más
cells per annulus and, typically, unpaired sori. Whereas de 20 células por anillo y, típicamente, soros no pareados.
Diplazium is always terrestrial, Asplenium can be terrestri- Diplazium es siempre terrestre, pero Asplenium puede
al, epiphytic or saxicolous, Another difference is that the puede ser terrestre, epífita, o saxícola. Otra diferencia es
annulus of Asplenium generally remains extended after que el anillo de Asplenium generalmente se queda exten-
dehiscence, and often the distal part of the capsule splits dido después de dehiscencia, y a menudo la parte distal
divaricately, resembling two small wings. The annulus of de la cápsula se hunde divaracatamente, pareciendo dos
Diplazium returns to its original position after dehiscence, pequeñas alas. El anillo en Diplazium regresa a su posición
forming an elongate “C,” and the distal part of the cap- original después de la dehiscencia, formando una “C” alar-
sule walls do not split apart. Callipteris differs by rhi- gada, y la parte distal de la cápsula no se hunde. Callipteris
zome (and often laminar) scales with black margins and difiere por escamas del rizoma (y a menudo las de las
black, bifid teeth. láminas) con márgenes negros y dientes bífidos negros.

COMMENTS: Diplazium consists of about 350 species, COMENTARIOS: Diplazium consta de casi 350 espe-
of which about 150 are neotropical. It grows primarily cies, de las cuales casi 150 son neotropicales. Crece prin-
in wet forests at low to middle elevations. In the Neo- cipalmente en bosques húmedos en elevaciones bajas a
tropics, few fern genera have received as little study as medias. En el Neotrópico, pocos géneros de los helechos
Diplazium. Consequently, there are numerous taxonomic han recibidos tan poco estudio como Diplazium. En con-
and nomenclatural problems, and these require mono- secuencia, hay numerosos problemas taxonómicos y
graphic study to solve. The sori arranged back-to-back nomenclaturales, y requieren estudio monográfico para
along the same vein are characteristic of the genus and resolverlos. Los soros arreglados espalda a espalda por
are called diplazioid sori. Many species have distinctive la misma vena son característicos del género y se llaman
hairs on the grooves of the rachises, and a peculiar flange soros diplazioides. Muchas especies tienen pelitos ca. 0.1
of tissue where the costules join the pinna rachises. The mm de largo en los surcos de los ráquises, y una pliega de
genus name is derived from the Greek diplazios, double. tejido donde las cóstulas se unen con los ráquises de las
The indusia are often double, or back-to-back along the pinnas. El nombre genérico se deriva del griego diplazios,
same vein. doble. A menudo los inducíos son dobles, o espalda a
espalda por la misma vena.

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.

252
Figure 120. . Diplazium lonchophyllum. A: leaf (hoja) B: pinna with sori (pinna con soros). C: diplazioid sori (soros diplazioi-
des).

253
ATHYRIACEAE (EUPOLYPODS II)
Diplazium (Callipteris clade)
DESCRIPTION: Terrestrial; rhizomes erect or decum- DESCRIPCIÓN: Terrestres; rizomas erectos a decum-
bent, the scales with black borders and black, bifid bentes, las escamas del rizoma con márgenes negros
marginal teeth; sterile and fertile leaves monomor- y dientes bífidos negros; hojas estériles y fértiles mono-
phous; petioles with 2 vascular bundles basally, these morfas; pecíolos con 2 haces vasculares basalmente,
uniting distally to form a U-shaped strand; laminae simple éstos uniéndose distalmente para formar un haz vascular
to 1-pinnate-pinnatifid, broadly cut, never finely divided; en forma de “U”; láminas simples a 1-pinnada-pinnatífidas,
veins areolate or rarely free, the areoles without includ- cortadas ampliamente (nunca finamente divididas); venas
ed veinlets; sori linear, indusiate, usually paired back to areoladas or raras veces libres, las areolas sin venillas in-
back along a single vein; annulus consisting of 15–20 cells; cluidas; soros lineares, con indusios, usualmente parea-
spores bilateral, brown; x=41. dos de par en par por una sóla vena; anillo consiste de
15–20 células; esporas bilaterales, pardas; x=41.

SIMILAR GENERA: The other species of Diplazium are GÉNEROS PARECIDOS: Los otros especies de Dipla-
distinguished by rhizome scales without dark borders. zium se distinguen por las escamas del rizoma sin bordes
Also, most species of Diplazium have free veins, where- oscuras. También, la mayoría de las especies de Diplazium
as the Callipteris group almost always has anastomosing tienen venas libres, mientras que los en el grupo de Cal-
veins. lipteris usualmente son anastomosantes.

COMMENTS: The Callipteris group consists of about 20 COMENTARIOS: El grupo Callipteris consta de casi 20
species in the New World and 70 in the Old. It grows especies en el Nuevo Mundo y 70 en el Viejo. Existe en
on wet, shaded, forest floors from 0–2000 m. The group sotobosques húmedos y sombreados de 0–2000 m. El
appears to be monophyletic but nested within Diplazium. grupo se parece ser monofilético pero anidado dentro
The laminae are often fleshy, and the teeth of the rhizome de Diplazium. Las láminas a menudo son carnosas, y los
scales are dark and bifid. The name Callipteris is derived dientes de las escampas del rizomas son oscuros y bífi-
from the Greek callos, beautiful, + pteris, fern. dos. El nombre Callipteris se deriva del griego callos, her-
moso, + pteris, helecho.

LITERATURE: Pacheco, L. & R. C. Moran. 1999. Monograph of the neotropical species of Callipteris (Woodsiaceae) with
anastomosing veins. Brittonia 51: 343–388.

254
Figure 121. Diplazium (Callipteris group) chimborazense (Pacheco & Moran, 1999).

255
GLEICHENIACEAE
Diplopterygium (Diels) Nakai
DESCRIPTION: Terrestrial; rhizomes long-creeping, DESCRIPCIÓN: Terrestres; rizomas largamente ras-
scaly; leaves indeterminate, of periodic growth, bearing treros, escamosos; hojas indeterminadas, de crecimiento
pairs of opposite pinnae, the distalmost pair with a scaly periódico, llevando pares de pinnas opuestas; el par más
resting bud between them; pinnae not forked, pin- distal con una yema latente escamosa entre sí; pin-
nate-pinnatisect or 2-pinnate; penultimate segments nas no surcadas, pinadas-pinnatisectas o 2-pinnadas;
pectinate; veins once-forked; sori round, not indusiate; segmentos penúltimos pectinados; venas una vez fur-
sporangia 2–5 per sorus; spores trilete; x=56. 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- COMENTARIOS: Diplopterygium tiene casi 10 especies
cies worldwide, all in Asia (India to Japan and Malesia). mundialmente, todos en Asia (India a Japón y Malesia).
Only one species occurs in the Neotropics: D. bancroftii Solamente una especie existe en el Neotrópico: D. ban-
(Hook.) A. R. Sm. This species generally grows in road- croftii (Hook.) A. R. Sm. Esta especie se encuentra usual-
sides and steep banks at middle elevations (1000–2600 mente en taludes y al lado de los caminos en elevaciones
m). This genus is sometimes treated in Gleichenia, but it is medias (1000–2600 m). Algunas veces este género se
maintained here as distinct because of its differences in trata en Gleichenia, pero aquí se mantiene distinto debido
pinna architecture and chromosome number. The genus a sus diferencias en la arquitectura de las pinnas y en el
name is derived from the Greek diploos, double + pterygi- número cromosómico. El nombre genérico se deriva del
um, little wing. The opposite pinnae resemble wings. 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.

256
Figure 122. A-F. Diplopterygium bancroftii (Hook.) A. R. Sm. (Mickel & Smith, 2004)

257
PTERIDACEAE
Doryopteris Sm.
DESCRIPTION: Terrestrial or rupestral; rhizomes erect, DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto,
scaly, the scales bicolorous with a dark central strip and escamoso, las escamas bicoloras estrechamente lance-
wide pale borders; leaves monomorphic or slightly di- oladas, con un centro oscuro y ancho, los bordes par-
morphic; petioles brown or black, with 1 or 2 vascular do claro; hojas monomorfas a casi dimorfas; pecíolos
bundles, terete or flattened adaxially, if flattened then with pardos a negros, con 1 o 2 haces vasculares, terete o
narrow wings on either side; laminae glabrous, gener- aplanado adaxialmente, en este caso acompañado por
ally pedate and pentagonal, rarely cordiform, deltate bordes o emarginaciones laterales estrechas; láminas
or reniform; veins netted or (in 2 species) free, often hard glabras generalmente pedadas y pentagonales, rara-
to see because of the thick lamina, the areoles without mente cordiformes, deltadas o reniformes; nervaduras
included veinlets; sori continuous along the margin; generalmente oscurecidas por la gruesa lámina; areoladas
indusium formed by the recurved margin; x=30. o (en dos especies) libres, las aréolas sin nérvulos inclu-
idos; soros continuos a lo largo del margen; indusio
formado por el margen foliar reflexo; x=30.

SIMILAR GENERA: Bommeria differs by pubescent GÉNEROS PARECIDOS: Bommeria difiere por lámi-
laminae and free veins (only two species of Doryopteris nas pubescentes y venas libres (solo dos especies de Do-
have free veins). Hemionitis differs by sori spreading be- ryopteris tienen venas libres). Hemionitis difiere por los
tween the midrib and margin. soros difundidos entre la costa y el margen.

COMMENTS: Doryopteris has 33 species, 28 of which COMENTARIOS: Doryopteris tiene 33 especies, 28 de

occur in the Neotropics. Most occur in the Planalto re- las cuales existen en los Neotropics. La mayoría existen
gion of southern Brazil. The genus typically grows in dry, en la región Planalto del sur de Brasil. El género crece
rocky habitats. Phylogenetic studies have shown that Do- típicamente en ambientes secas y rocosas. Estudios mo-
ryopteris is most closely related to Trachypteris of South leculares han demostrado que Doryopteris es más cer-
America and Pellaea calomelanos of Africa (Gastony & canamente relacionado a Trachypteris de América del Sur
Rollo, 1995, 1998). The genus name is derived from the y Pellaea calomelanos de África (Gastony & Rollo, 1995,
Greek dory, lance + pteris, fern. It alludes to the shape of 1998). El nombre genérico se deriva del griego dory, lanza
the blade in some species. + 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. Contribu-
tions 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.

258
Figure 123. Doryopteris. A–C. Doryopteris palmata. D, E. Doryopteris concolor. (from Mickel & Smith, ined.)

259
DRYOPTERIDACEAE (EUPOLYPODS I)
Dryopteris Adans.
DESCRIPTION: Terrestrial or more rarely epiphytic or DESCRIPCIÓN: Terrestres o menos comúnamente
rupestral; rhizomes compact, erect or ascending, scaly; epífitas o rupícolas; rizomas compactos, erectos o as-
leaves monomorphic; petioles with more than three vas- cendentes, escamosos; hojas monomorfas; pecíolos con
cular bundles, these arranged in a U with the two adaxial más tres haces vasculares, estas arregladas en forma de U
ones the largest; laminae 2- to 3-pinnate-pinnatifid, less con los dos adaxiales lo más grandes; láminas 2- o 3-pin-
commonly 1-pinnate-pinnatifid, the apices gradually ta- nadas, menos común 1-pinnada-pinnatífidas, los ápices
pered and pinnatifid (not imparipinnate); pinnules of the gradualmente reducidas y pinatífidas (no imparipinadas);
lower pinnae arranged isodromically or catadromically; pínnulas de las pinnas inferiores arregladas isodrómica-
rachises and costae grooved adaxially, the grooves more mente o catadrómicamente; ráquises y costas sulcados
or less confluent at their junctures; veins free; sori round; adaxialmente, los surcos más o menos confluentes en
indusia present, attached at the base of a narrow las uniones; venas libres; soros redondos; indusios pre-
sinus, not peltate, reniform or orbicular; spores mono- sentes, unidos en la base de un sinus estrecho, no
lete, winged; x=41. peltados, reniformes o orbiculares; esporas monoletes,
aladas; x=41.

SIMILAR GENERA: Polystichum usually differs by 1- or GÉNEROS PARECIDOS: Polystichum usualmente difi-
2-pinnate laminae, the base of the pinnae or pinnules ere por láminas 1- a 2-pinnadas, la base de las pinnas y
slightly elongate on the acroscopic side, and the spinulose pínnulas alargada en el lado acroscópico, y los márgenes
lamina margins. Arachniodes differs by anadromic blades espinulosos de las láminas. Arachniodes difiere por láminas
and creeping rhizomes. Lastreopsis differs by the upper anadrómicas y rizomas reptantes. Lastreopsis difiere por
surfaces of the rachises and costae rounded or flat (not el haz de las ráquises y costas redondeado o achatado
grooved). Polybotrya is (mostly) a climbing epiphytic fern (no surcado). Polybotrya es (principalmente) un helecho
and has dimorphic sterile and fertile leaves. Saccoloma dif- trepador epífito y tiene hojas estériles y fértiles dimorfas.
fers by an omega-shaped vascular bundle in the base of Saccoloma difiere por una haz vascular en forma de una
the petiole, and marginal or submarginal sori. omega en la base del pecíolo y soros marginales o sub-
marginales.
COMMENTS: Dryopteris consists of about 225 species COMENTARIOS: Dryopteris consta de casi 225 espe-
worldwide and is well represented in the northern tem- cies mundial y es bien representado en las zonas temp-
perate zones, especially in Asia. In the Neotropics it grows ladas del norte, especialmente en Asia. En el Neotrópico
in middle to high elevation forests and contains about crece en bosques de elevaciones medias y altas y con-
12 neotropical species, most all of which occur in Me- tiene alrededor de 12 especies neotropicales, la mayoría
soamerica. A subgeneric classification of the genus was de las cuales existen en Mesoamérica. Una clasificación
proposed by Frasier-Jenkins (1989). The genus name is subgenérica del género fue propuesta por Frasier-Jenkins
derived from the Greek drys, tree + pteris, fern. (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-bipinnatifid
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). Bull. Brit. Mus. (Nat. Hist), Bot. 14(3): 183–218. Frasier-Jenkins, C. R.
1989. A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. Bull. Brit. Mus. (Nat. Hist.), Bot. 18:
323–477.

260
Figure 124. A, B. Dryopteris muenchii. C, D. D. arguta. E, F. D. futura. G, H. D. filix-mas. (Mickel & Smith, 2004)

261
DRYOPTERIDACEAE
Elaphoglossum sect. Amygdalifolium (H. Christ) Mickel & Atehortúa
DESCRIPTION: Plants epiphytic; rhizomes long- DESCRIPCIÓN: Plantas epífiticas; rizomas largamente
creeping, the scales linear to lanceolate; aerophores rastreros, las escamas lineares a lanceoladas; aeróforos
absent(?); phyllopodia present; blades glabrous or ausentes(?); filopódios presentes; láminas glabras a
nearly so, entire, thin, sometimes reddish when young, subglabras, enteras, delgadas, a veces rojizas cuando
the scales (when present) minute, non-setose, laciniate- jóvenes, las escamas diminutas, no setosas, lacinadas-
pectinate to stellate, minute glandular hairs absent; veins pectinadas a estrelladas, pelos diminutos glandulares
free; hydathodes present; spores monolete, brown or ausentes; venas libres; hidatodos presentes; esporas
blackish; x=41. monoletes, pardas o negruzcas; x=41.

SIMILAR GENERA: The simple-leaved species of GÉNEROS PARECIDOS: Las especies de las
Polypodiaceae (e.g., Campyloneurum, Microgramma, Polypodiaceae con hojas enteras (e.g., Campyloneurum,
Pleopeltis) differ by reticulate veins and round sori. The Microgramma, Pleopeltis) difieren por nervadura reticulada
other sections of Elaphoglossum differ by having at least y soros redondos. Las otras secciones de Elaphoglossum
one of the following characters: hydathodes absent, difieren por tener a lo menos una de las características
conspicuously scaly laminae, or phyllopidia absent. 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. y consta de una sola especie: E. amygdalifolium (Mett.) H.
It occurs from 0–1500 m in Cuba and Central America Christ. Existe desde 0–1500 m en Cuba y América Central
to Colombia and Ecuador (restricted to the western side hasta Colombia y Ecuador (restingada al lado occidental
of the Andes). No other species of Elaphoglossum has de los Andes). Ninguna otra especie de Elaphoglossum
the combination of characters given in boldface in the tiene la combinación de caracteres dados en negritos en
description above. The species is sister to the rest of the la descripción arriba. La especie es hermana al resto del
genus (Rouhan et al., 2004). Plants in Central America género (Rouhan et al., 2004). Plantas de América Central
have reddish young leaves—a characteristic unique in tienen hojas jóvenes rojizas—una característica única
the genus—but plants from Ecuador apparently lack this en el género—pero plantas de Ecuador aparentemente
color (pers. obs.). Elaphoglossum amygdalifolium might faltan este color (pers. obs.). Elaphoglossum amygdalifolium
eventually prove to be hemiepiphytic, a pleisiomorphic puede ser hemiepifítica, una condición pleisiomórfica.
condition. Plants appear to start near the bases of trunks Plantas aparecen empezar cerca las bases de los troncos
and then climb. Such plants might produce at their bases y luego suben. Es posible que tales plantas produzcan en
long, straight roots that grow into the soil where they sus bases raíces largas rectas que van directamente al
ramify. This growth form, which occurs in the outgroup suelo donde ramifican. Esta forma de crecimiento, que
genera (Anapausia, Lomagramma, Teratophyllum), needs existe en los grupos externos (Anapausia, Lomagramma,
to be searched for. The sectional name is derived from Teratophyllum), se necesita buscar. El nombre genérico se
the Latin amygdalus, almond + folium, leaf. The leaves deriva del latín amygdalus, almendra + folium, hoja. Las
resemble that of an almond. 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.

262
Figure 125. Elaphoglossum amygdalifolium. A. Habit. B. Long-creeping rhizome and dark phyllopodium. C. Venation, showing
hydathodes. (©Moran, 2009)

263
DRYOPTERIDACEAE
Elaphoglossum Schott ex J. Sm., sect. Elaphoglossum
DESCRIPTION: Plants epiphytic; rhizomes long- DESCRIPCIÓN: Plantas epífiticas; rizomas largamente
creeping, the scales linear to lanceolate; aerophores rastreros, las escamas lineares a lanceoladas; aeróforos
absent(?); phyllopodia present; blades glabrous or ausentes(?); filopódios presentes; láminas glabras a
nearly so, entire, thin, sometimes reddish when young, subglabras, enteras, delgadas, a veces rojizas cuando
the scales (when present) minute, non-setose, laciniate- jóvenes, las escamas diminutas, no setosas, lacinadas-
pectinate to stellate, minute glandular hairs absent; veins pectinadas a estrelladas, pelos diminutos glandulares
free; hydathodes present; spores monolete, brown or ausentes; venas libres; hidatodos presentes; esporas
blackish; x=41. monoletes, pardas o negruzcas; x=41.

SIMILAR GENERA: The simple-leaved species of GÉNEROS PARECIDOS: Las especies de las
Polypodiaceae (e.g., Campyloneurum, Microgramma, Polypodiaceae con hojas enteras (e.g., Campyloneurum,
Pleopeltis) differ by reticulate veins and round sori. The Microgramma, Pleopeltis) difieren por nervadura reticulada
other sections of Elaphoglossum differ by having at least y soros redondos. Las otras secciones de Elaphoglossum
one of the following characters: hydathodes absent, difieren por tener a lo menos una de las características
conspicuously scaly laminae, or phyllopidia absent. 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. y consta de una sola especie: E. amygdalifolium (Mett.) H.
It occurs from 0–1500 m in Cuba and Central America Christ. Existe desde 0–1500 m en Cuba y América Central
to Colombia and Ecuador (restricted to the western side hasta Colombia y Ecuador (restingada al lado occidental
of the Andes). No other species of Elaphoglossum has de los Andes). Ninguna otra especie de Elaphoglossum
the combination of characters given in boldface in the tiene la combinación de caracteres dados en negritos en
description above. The species is sister to the rest of the la descripción arriba. La especie es hermana al resto del
genus (Rouhan et al., 2004). Plants in Central America género (Rouhan et al., 2004). Plantas de América Central
have reddish young leaves—a characteristic unique in tienen hojas jóvenes rojizas—una característica única
the genus—but plants from Ecuador apparently lack this en el género—pero plantas de Ecuador aparentemente
color (pers. obs.). Elaphoglossum amygdalifolium might faltan este color (pers. obs.). Elaphoglossum amygdalifolium
eventually prove to be hemiepiphytic, a pleisiomorphic puede ser hemiepifítica, una condición pleisiomórfica.
condition. Plants appear to start near the bases of trunks Plantas aparecen empezar cerca las bases de los troncos
and then climb. Such plants might produce at their bases y luego suben. Es posible que tales plantas produzcan en
long, straight roots that grow into the soil where they sus bases raíces largas rectas que van directamente al
ramify. This growth form, which occurs in the outgroup suelo donde ramifican. Esta forma de crecimiento, que
genera (Anapausia, Lomagramma, Teratophyllum), needs existe en los grupos externos (Anapausia, Lomagramma,
to be searched for. The sectional name is derived from Teratophyllum), se necesita buscar. El nombre genérico se
the Latin amygdalus, almond + folium, leaf. The leaves deriva del latín amygdalus, almendra + folium, hoja. Las
resemble that of an almond. 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.

264
Figure 126. A, B. Elaphoglossum seminudum. C-E. E. lonchophyllum. F-J. E. sartorii. K, L. E. glabellum.  M. E. parduei. (from Mickel &
Beitel, 1988)

265
DRYOPTERIDACEAE
Elaphoglossum J. Sm. sect. Lepidoglossa
DESCRIPTION: Primarily epiphytic, occasionally DESCRIPCIÓN: Principalmente epífitas, a veces
terrestrial or rupestral; stem short- to long-creeping, rarely terrestres o rupestres; tallos cortamente a largamente
erect, scaly, in transverse section with an elongated ventral rastreros, raras veces erectos, escamosos, en sección
vascular bundle; leaves simple, entire, dimorphic; petioles transversal con un haz ventral vascular alargado; hojas
articulate to dark phyllopodia; laminae subglabrous to simples, enteras, dimorfas; pecíolos articulados a filopódios
densely scaly, scales with acicular, one-celled, non- oscuros; láminas subglabras a densamente escamosas, las
glandular teeth; rachis grooved adaxially; veins free; sori escamas con dientes aciculares, unicelulares, no
acrostichoid; spores bilateral, typically nongreen; x=41. 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. pantropical e consta de ca. 350 especies mundialmente.
Most of the species are neotropical and epiphytic. They La mayoría de las especies son neotropicales y epifíticas.
occur in a wide variety of habitats, from rainforests to Ocurren un una amplia variedad de hábitats, desde
páramos. The species are poorly known and in need of bosques lluviosos hasta páramos. Las especies se
taxonomic study. They can be identified without fertile conocen pobremente y necesitan estudio taxonómico.
leaves because they are defined primarily on the basis of Se puede identificarlas sin hojas fértiles porque son
leaf shape, venation, presence or absence of hydathodes definidas principalmente en base de la forma de las hojas,
and phyllopodia, and characteristics of the laminar scales. venación, presencia o ausencia de hidatodos y filopódios,
The laminae of sect. Lepidoglossa are often conspicuously y características de las escamas laminares. Las láminas
scaly, but in some species the scales are reduced to en secc. Lepidoglossa son típicamente escamosas, pero
reddish dots and the laminae therefore appear glabrous. en algunas especies las escamas son reducidas a puntos
The genus name is derived from the Greek elaphos, stag rojizos y pore so las láminas aprecen glabras. El nombre
+ glossa, tongue. The leaves resemble a deer’s tongue. genérico se deriva del griego elaphos, venado + glossa,
Lepido- means scaly. 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..

266
Figure 127. A–F. Elaphoglossum muscosum. G–P. E. paleaceum. Q. E. vestitum. R–W. E. pringlei. (Mickel & Beitel, 1988)

267
DRYOPTERIDACEAE
Elaphoglossum sect. Polytricha H. Christ
DESCRIPTION: Plants generally epiphytic; rhizomes DESCRIPCIÓN: Plantas generalmente epifítas, rizomas
short- to (rarely) long-creeping, the scales linear to cortamente a (raras veces) largamente rastreros, las
lanceolate; phyllopodia present; petioles glabrous or with escamas lineares a lanceoladas; filopódios presentes;
minute capitate-glandular hairs; blades simple, entire, pecíolos glabros o con pelitos gladulosos-capitados;
with or without glandular hairs, scaly particularly along láminas simples, enteras, con o sin pelos glandulares,
the costae, the scales subulate with a slightly widened escamosas particularmente por las costas, las escamas
enrolled base, patent; veins free; hydathodes absent; sori subuladas con una base enrollada, patentes; venas
acrostichoid; spores monolete; x=41. 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. COMENTARIOS: Como todos los grupos de
Polytrichia is primarily neotropical. The exact number Elaphoglossum, la secc. Polytrichia es principalmente
of species belonging to the group is unknown. Rouhan neotropical. El número exacto que pertenece al grupo
et al. (2004) found the section to be monophyletic. In se desconoce. Rouhan et al. (2004) encontraron que
Elaphoglossum, subulate scales are highly distinctive and el grupo es monofilético. En Elaphoglossum las escamas
found only in sects. Polytrichia and Setosa. These scales subuladas son altamente distintivas y se encuentran solo
are enrolled longitudinally and en masse have a bristly en las seccs. Polytrichia y Setosa. Estas escamas se enrollan
appearance. They are found only on the leaves, not the longitudinalmente y conjunto tienen una apariencia
rhizomes. In some species, flat scales or scales enrolled cerdosa. En algunas especies, escamas planadas o
basally and flat distally, occur along the lamina margins. solamente enrolladas basalmente y planadas distalmente,
In such species, typical subulate scales are found on the ocurren por los márgenes de las láminas. En tales
petioles and costae. Some widespread species of sect. especies, escamas subuladas típicas se encuentran por los
Polytrichia are E. apodum (Kaulf.) Schott ex J. Sm., E. crinitum pecíolos y las costas. Algunas especies muy difundidas de
(L.) Christ, E. decoratum (Kunze) T. Moore, E. erinaceum sect. Polytrichia son E. apodum (Kaulf.) Schott ex J. Sm.,
(Fée) T. Moore, E. hybridum (Bory) Brack., and E. pallidum E. crinitum (L.) Christ, E. decoratum (Kunze) T. Moore, E.
(Baker ex Jenm.) C. Chr. The section name comes from erinaceum (Fée) T. Moore, E. hybridum (Bory) Brack. y E.
the Greek poly, many, and trichia, hair, referring to the pallidum (Baker ex Jenm.) C. Chr. El nombre seccional se
many hair-like subulate scales on the leaves. 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. In press. The
Brazilian species of Elaphoglossum section Polytrichia (Dryopteridaceae). Brittonia 00: 00–00. 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.

268
Figure 128. Elaphoglossum sect. Polytrichia. A–F. E. scolopendrifolium. (Matos & Mickel, in press)

269
DRYOPTERIDACEAE
Elaphoglossum sect. Setosa (H. Christ) Mickel & Atehortúa
DESCRIPTION: Plants generally epiphytic; rhizomes DESCRIPCIÓN: Plantas generalmente epifíticas, rizomas
short- to long-creeping, the scales linear; phyllopodia cortamente a largamente rastreros, las escamas lineares;
absent; petioles with minute capitate-glandular hairs; filopódios ausentes; pecíolos con pelitos gladulosos-
blades sparsely puberulent and conspicuously scaly, capitados; láminas esparcidamente puberulentas y
the hairs minute and glandular (handlens needed), conpicuosamente escamosas, los pelos minutos
the scales subulate with a slightly widened enrolled y glandulares (se necesita una lupa), las escamas
base, spreading to erect, orange to brown; veins free, subuladas con la base poco ensanchada y enrollada,
spaced well apart; hydathodes present; sori acrostichoid; patentes a erectas, narangas a pardas; venas libres, bien
spores monolete; x=41. espaciadas; hidatodos presentes; soros acrosticoides;
esporas monoletes; x=41.

SIMILAR GENERA: Similar species of Elaphoglossum GÉNEROS PARECIDOS: Especies similares de

can usually be distinguished by non-subulate scales, lack Elaphoglossum se pueden distinguir usualmente por
of minute glandular hairs, the presence of phyllopodia, or escamas no subuladas, el carecer de pelitos glandulosos,
the absence of hydathodes. la presencia de filopodios, o la ausencia hidatodos.

COMMENTS: Like all groups of Elaphoglossum, sect. COMENTARIOS: Como todos los grupos de
Setosa is primarily neotropical. The exact number of Elaphoglossum, la secc. Setosa es principalmente
species belonging to the group is unknown. Rouhan neotropical. El número exacto que pertenece al grupo
et al. (2004) found the group to be monophyletic. In se desconoce. Rouhan et al. (2004) encontraron que
Elaphoglossum, subulate scales are highly distinctive. They el grupo es monofilético. En Elaphoglossum las escamas
are slender and en masse often have a bristly appearance. subuladas son altamente distintivas. Son delgadas y a
With magnification it can be seen that the scales are menudo en mases parecen cerdosas. Con magnificación,
rolled at the base to form a short cylinder-like part on se puede ver que las escamas son enrolladas en la base y
which supports the rest of the scale. Some widespread forman un breve cilíndrico que apoya el resto de la escama.
species of sect. Elaphoglossum Setosa are E. lindenii (Bory Algunas especies muy difundidas de Elaphoglossum sect.
ex Fée) T. Moore, E. piloselloides (C. Presl) T. Moore, y E. Setosa son E. lindenii (Bory ex Fée) T. Moore, E. piloselloides
villosum (Sw.) J. Sm. The section name comes from the (C. Presl) T. Moore, y E. villosum (Sw.) J. Sm. El nombre
Latin seta, bristle, and refers to the stiff, spreading subulate seccional se deriva del latín seta, cerdo, y se refiere a las
scales. escamas subuladas rígidas y patentes.

LITERATURE: Mickel, J. T. & L. Atehortúa G. 1980. Subdivision of the genus Elaphoglossum. American Fern Journal 70:
47–68.

270
Figure 129. Elaphoglossum sect. Setosa. A–E. E. pilosius. F–G. E. setosum. J–L. E. jaliscanum. M, N. E. pusillum. Q. E. monicae.
(Mickel & Smith, 2004).

271
DRYOPTERIDACEAE
Elaphoglossum sect. Squamipedia Mickel & Atehortúa
DESCRIPTION: Plants epiphytic, generally less than DESCRIPCIÓN: Plantas epífiticas, generalmente
25 cm tall; rhizomes long-creeping, the scales usually menos de 25 cm de alto; rizomas largamente
pale brown, concolorous; aerophores 1-3 mm long, rastreros, las escamas usualmente pardo pálido,
peg-like, found slightly behind the base of the petiole; concoloras; aeróforos 1-3 mm de largo, cilíndricos,
phyllopodia absent; blades entire or either flabellately atrás de la base del pecíolo; filopódios ausentes; láminas
or pinnately dissected, subglabrous to moderately scaly, enteras or disectadas dicotómamente o pinnatamente,
the scales non-setose, often hastate or slightly so at base; subglabras a escamosas, las escamas sin setas, a menudo
veins free; hydathodes absent; sori acrostichoid; spores hastatas or casi así en la base; venas libres; hidatodos
minutely spiny; x=40. 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 casi 17 especies, todos de las cuales son neotropicales
in Madagascar. The section is entirely epiphytic and con excepción de una en Madagascar. La sección es
generally occurs above 1000 m. The peglike aerophores completamente epifítica y generalmente existe arriba
on the rhizomes are often hidden by the scales and arriba de 1000 m. Los aeróforos como dedos en las
must be carefully searched for. They are situated about rizomeas a menudo están ocultos por las escamas y hay
1 mm behind the petiole. Three species with dissected que buscarlas con cuidado. Están ubicados casi 1 mm atrás
laminae occur in the section: Elaphoglossum peltatum del pecíolo. Dos especies con láminas disectadas existe en
(laminae orbicular and flabellately dissected), E. tripartitum el sección: Elaphoglossum peltatum (láminas redondeadas
(laminae tripartite), and E. moorei (laminae lanceolate y discectadas dicotómamente), E. tripartitum (láminae
pinnately dissected, often with the pinnae forked at tripartitas) y E. moorei (láminas disectadas pinatamente,
the apex). Because of their dissected laminae, they are a menudo con las pinnas furcadas en el ápice). A causa
sometimes placed in Peltapteris Link, but molecular de sus láminas disectadas, están ubicadas en Peltapteris
phylogenetic results indicate that lamina division evolved Link., pero resultados moleculares filogenéticos indican
three times separately within the section. Thus Peltapteris, que división de la lámina se evolucionó tres veces dentro
defined as such, is polyphyletic. Wagner (1980) found dentro de la sección. Por eso Peltapteris, definido tal, es
n=40 in E. peltatum, which differs from n=41 in other polifilético. Wagner (1980) encontró n=40 en E. peltatum,
Dryopteridaceae. This might be a synapomorphy for the lo cual difiere de n=41 en otros Dryopteridaceae. Es
section. The name Squamipedia refers to E. squamipes posible esto es una sinapomofia para la sección. El nombre
which belongs to the section. 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.

272
Figure 130. A–F. Elaphoglossum squamipes. G–L. E. aff. squamipes. M. E. deltoideum. N–R. E. anthracinum. S, T. E. lloense.
(From Vasco et al., 2013).

273
DRYOPTERIDACEAE
Elaphoglossum sect. Wrightiana Lóriga et al.
DESCRIPTION: Plants climbering, with long-creeping DESCRIPCIÓN: Plantas trepadoras, con rizomas
rhizomes that begin growth on the ground and eventually largamente rasteros que empiezan crecimiento en el
climb trunks to heights of 1–2 m, maintaining the suelo y eventualmente suben troncos hasta alutras de
connection to the ground by the rhizome and roots 1–2 m, mantaniéndose la coneccion con el suelo por
emitted from the lower portions of the climbing rhizome; el rizoma y raíces emitidas de las porciones proximales
aerophores present, finger-like; phyllopodia present; del rizoma trepador; aeróforos presentes, con forma de
hydathodes absent; laminar scales flat (not subulate), with dedo; filopodios presentes; hidatodos ausentes; escamas
marginal processes or teeth ending in a slightly swollen foliares planas (no subuladas), con dientes marginales
cell (i.e., scales never with acicular marginal cells as in E. terminándose en una célula levamente hinchada (i.e.,
sect. Lepidoglossa); x=?. 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 COMENTARIOS: Sección Wrightiana consta de una
species endemic to Cuba: E. wrightii (Mett. ex D. C. Eaton) sola especie endémica a Cuba: E. wrightii (Mett. ex D.
T. Moore. It grows in wet montane forests and is the sister C. Eaton) T. Moore. Existe en bosques húmedos y es la
species to all others in the genus except E. amygdalifolium. especie hermano a las demas especies del género con
Its growth form is unique in the genus: young sporophytes excepción a E. amygdalifolium. Su forma de crecimiento
begin on the forest floor and creep around until they es único en el género: esporofitos jóvenes empiezan en
encounter a trunk, which is then climbed by means of el sotobosque y trepan hasta que encuentran un tronco,
adhesive roots. Fertile leaves are produced on the lo cual es subido por medios de raíces adhesivos. Hojas
climbing portion of the rhizome. This growth form is fértiles están producidas de la porción reptante del rizoma.
present in the bolbitioid outgroups to Elaphoglossum, Esta forma de crecimiento es presente en los grupos
such as Mickelia, Lomagramma, and Teratophyllum. The externos bolbitidoides, tales como Mickelia, Lomagramma
sectional name Wrightiana refers to E. wrightii, the type y Teratophyllum. El nombre seccional Wrightiana refiere a
species of the genus, which is named for Charles Wright E. wrightii, la especie tipo de la sección, que es nombrado
(1811–1885), a plant collector. 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]

274
Figure 131. A–F. Elaphoglossum (sect. Wrightiana) wrightii. D. Heteroblastic series, youngest leaf at right. A. Habit. B. Petiole
scale from fertile leaf. C. Rhizoe scale. (Modified from Vasco et al., 2013)

275
POLYPODIACEAE
Enterosora Baker in Thurn
DESCRIPTION: Epiphytes, occasionally saxicolous; DESCRIPCIÓN: Epífitas, ocasionalmente saxícolas;
rhizome scales not clathrate, yellowish-brown to golden escamas del rizoma no clatradas, pardo-amarillento a
brown, sometimes ciliate or glandular; laminae simple pardo-doradas, a veces ciliadas o glandulares; láminas
to lobed 4/5 of the way to the midrib, not pectinate, simples a lobadas 4/5 de la longitud a la costilla
generally thick and spongy, mesophyll completely media, no pectinada, generalmente gruesa y esponjosa,
composed of stellate parenchyma (pallisade mesofilio consta completamente de parénquima
parenchyma lacking); veins simple when free, forked estrellado (parénquima palisade ausente); nervaduras
when fertile, but often variously areolate, especially near simples cuando libres, furcadas cuando fértiles, pero a
the midrib; hydathodes absent; setae present, dark menudo variadamente areoladas, especialmente cerca de
reddish, often surrounding the sorus; hairs often la costilla media; hidatodos ausentes; setas presentes,
branched and glandular at the apex; sori slightly to rojizas oscuras, a menudo rodeando el soro; pelitos
deeply sunken in the lamina tissue, round to elongate; típicamente ramificados, a menudo glandulares en el ápice;
spores green, trilete; x=37. soros ligeramente a profundamente embebidos en
el tejido laminar, redondeados a alongados, esporas
verdes, triletes; x=37.

SIMILAR GENERA: The absence of hydathodes will GÉNEROS PARECIDOS: La ausencia de hidátodos
separate Enterosora from all the other neotropical genera separa Enterosora de todos los otros géneros
except Ceradenia and Zygophlebia. Ceradenia differs neotropicales con excepción de Ceradenia y Zygophlebia.
by the waxy glands in the sori, and Zygophlebia by the Ceradenia difiere por las glándulas cerosas en los soros,
petioles articulate to the rhizome and pinnatisect to y Zygophlebia por los pecíolos articulados al rizoma y
pinnate laminae. 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 neotropicales y 2 Africanas. En el Neotrópico, se existe
Mexico to Bolivia and Suriname and in the Antilles. The desde México hasta Bolivia y Surinam y en las Antillas. Las
species grow mainly in cloud forests. The thick, spongy especies crecen principalmente en bosques nublados. El
mesophyll can be seen where the tissue has been torn mesofilio grueso, esponjoso se puede ver donde el tejido
or deteriorated. It cells bear long “arms” and between ha estado roto o deteriorado. Sus células llevan “brazos”
them are large intercellular air spaces. A palisade layer is largos e entre si están espacios intercelulares grandes.
lacking. Enterosora is monophyletic and probably sister to Un estrato palisade se carece. Enterosora es monofilético
Ceradenia (Ranker et al., 2004), a genus that can also have y probablemente hermana a Ceradenia (Ranker et al.,
spongy mesophyll, although less developed. The genus 2004), un género que puede tener mesofilio esponjoso,
name is derived from the Greek enteron, intestine + soros. aunque menos desarrollado. El nombre genérico se
The sori are not superficial, but embedded in pockets deriva del griego enteron, intestino + soros. Los soros no
within the leaf tissue. 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.

276
Figure 132. Enterosora. A–C. E. trifurcate. D–F. E. insidiosa. G–I. E. trichosora. J–L. E. parietina. (from Bishop & Smith, 1992)

277
EQUISETACEAE
Equisetum L.
DESCRIPTION: Stems jointed, green, longitudinally DESCRIPCIÓN: Tallos articulados, verdes, surcados
grooved, usually hollow in the center. Branches (if longitudinalmente, usualmente huecos en el centro.
present) whorled at the nodes. Leaves whorled, fused Ramas (cuando presentes) verticilados en los nudos.
into a sheath, the tips free, tooth-like, often deciduous. Hojas verticiladas, fusionadas en una vaina, los ápices
Sporophylls aggregated in terminal strobili, peltate, tan libres, como dientes, a menudo deciduos. Sporofilas
or yellowish, bearing 5–10 sporangia on the inner surface; agregados en estróbilos terminales, peltados, leonados
spores green, with 4 arm-like appendages (elaters); x=108. 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 COMENTARIOS: Equisetum tiene 15 especies mundial
and occurs on all continents exceopt Australia and y existe en todos los continentes con excepción de
New Zealand. It exhibits a reverse latitudinal gradient Australia y Nueva Zelanda. Exhibe un gradiente latitudinal
in diversity, being most species-rich in north-temperate de diversidad en revés, siendo más rico en especies en
regions between 40° and 60°. It occurs in many habitats, las zonas templadas norteñas entre 40° y 60°. Existe en
but most typically in sunny places and wet soils. The muchos hábitats, pero típicamente en lugares soleados y
stems are covered by a layer of silica modified into suelos húmedos. Los tallos están cubiertos por un estrato
spines, tubercles, or teeth. These structures are often de sílice modificado en espinas, tubérculos o dientes.
distinctive for the species. In Equisetum giganteum they are Estas estructuras son a menudo distintivas para la especie.
quadrangular in profile, whereas in E. myriochaetum they En Equisetum giganteum son cuadrangulares en perfil,
are angled. The nodes remain meristematic and capable mientras que en E. myriochaetum son angulados. Los
of generating new stems if placed in soil. Fragmentation of nudos se quedan meristemáticos y capaces de generar
stems enables abortive-spored hybrids to persist at their tallos nuevos si planteados en el suelo. Fragmentación
site of origin and even spread beyond the range of their de los tallos facilita los híbridos con esporas abortivas
parents.The spores bear hygroscopic appendages called persistir en su sitio de origen y también difundirse más
“elaters.” When moist, the elaters coil tightly around the allá el rango de sus padres. Las esporas llevan brazos
spore, and when dry they uncoil and spread, catching the higroscópicos que se llaman “elateres.” Cuando mojados,
wind and creating drag which help disperse the spore. los elateres se enrollan alrededor de la espora, y cuando
The spores are green and have short viability (about one se secan, se extienden, recogiendo los vientos. Las
week). They are unique by lacking haptotypic markings esporas son verdes y tienen una corta viabilidad (casi una
(scars from their position in the tetrad). Thus, they are semana). Son únicas por carece de marcas haptotípicas
neither monolete nor trilete. The genus name is derived (cicatrices de su posición en la tetrada). Por eso no son
from the Greek equis, horse + seta, bristle. The shoots of monoletes ni triletes. El nombre genérico se deriva del
some species resemble a horse’s tail. 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.

278
Figure 133. A–D. Equisetum hyemale. E–J. E. myriochaetum (Mickel & Smith, 2004).

279
PTERIDACEAE
Eriosorus Fée
DESCRIPTION: Terrestrial, rupestral, or rarely epiphytic; DESCRIPCIÓN: Terrestres, rupícolas o raramente
rhizomes short- to long-creeping, covered by dense epifíticas; rizoma corta a largamente rastrero,
dark trichomes or bristles; leaves erect, pendent or cubierto densamente por tricomas o cerdas
scandent; petiole and rachis atropurpueous, sparsely to oscuras; hojas erectas, pendientes o escandentes;
densely hairy; laminae 2- to 5-pinnate; pinnae hairy on pecíolo y raquis atropurpúreos, esparcida a densamente
one or both surfaces, the hairs multicellular, frequently pelosos; láminas 2- a 5-pinnadas; pinnas pelosas en
with narrow cell walls; margins of the pinnules flat, not una o ambas superficies, los tricomas pluricelulares,
involute or covering the sporangia; veins free, ending frecuentemente con paredes celulares adelgazadas;
before the margin in a notch; indusia absent; sporangia márgenes de las pínnulas aplanados, no involutos o
spreading along the veins; indusia absent; spores cubriendo los esporangios; nervaduras libres, terminando
tetrahedral-globose; x=87. antes de los márgenes o en una escotadura; indusio
ausente; esporangios difundiéndose a lo largo de las
nervaduras; indusios ausentes; esporas tetrahédricas-
globosas; x=87.

SIMILAR GENERA: Pityrogramma has an erect or GÉNEROS PARECIDOS: Pityrogramma tiene un

decumbent, scaly rhizome and lamina farinose below. In rizoma erecto, escamoso, y una lámina farinosa en el
Anogramma the rhizome is poorly developed and lacks envés. En Anogramma el rizoma no es bien desarrollado
old petiole bases from previous years. y no porta viejas bases peciolares de los años anteriores.

COMMENTS: Eriosorus is found in forests and open COMENTARIOS: Eriosorus se encuentra en en bosques
places above 1200 m. It contains 25 species and is y lugares abiertos arriba de 1200 m. Contiene 25 especies
entirely neotropical, with one population on Tristan y es completamente neotropical, con una población en
da Cunha. Eriosorus flexuosus (Humb., Bonpl. & Kunth) Tristan da Cunha. Eriosorus flexuosus es muy difundida
Copel. is widespread and common throughout much of y común por la mayoría de las Américas. Se puede
the Americas. It can be easily distinguished by its zig-zag distinguirlo fácilmente por su raquis en forma de zig-zag
rachis and scandent, indeterminate leaves.The recognition y hojas escandentes y indeterminadas. La recognición
of Jamesonia leaves Eriosorus paraphyletic. Jamesonia is de Jamesonia deja Eriosorus parafilético. Jamesonia es un
a páramo morphotype that has evolved at least three morfotipo de los páramos que ha evolucionado desde
times within Eriosorus (Sánchez-Baracaldo, 2004). Hybrids Eriosorus al lo menos tres veces (Sánchez-Baracaldo,
between Eriosorus and Jamesonia are relatively common 2004). Los híbridos entre Eriosorus y Jamesonia son
where the two grow together in páramos (Gómez, relativamente comunes donde estos dos crecen juntos
1979).The genus name is derived from the Greek erion, en páramos (Gómez, 1979). El nombre genérico se
wooly + soros, sori. It refers to the wool-like hairs among deriva del griego erion, lanoso + soros. Se refiere a los
the sporangia. pelos como lana entre los esporangios.

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. Scamman, E. 1962. The genus Eriosorus in Costa Rica. Contr. Gray Herb. 191: 81-89.
Tryon, A. F. 1970. A monograph of the fern genus Eriosorus. Contributions from the Gray Herbarium of Harvard University
200: 54–174.

280
Figure 134. A, B. Eriosorus hirsutus. C, D. E. flexuosus. E, F. E. flexuosus var. galeanus (Mickel & Smith, 2004).

281
MARATTIACEAE
Eupodium J. Sm.
DESCRIPTION: Terrestrial; rhizomes erect or nearly so, DESCRIPCIÓN: Terrestres; rizomas erectos o casi así,
each bearing only 1 leaf (rarely 2) and paired rounded cada uno llevando una sola hoja (raras veces 2) y
fleshy stipules at the base of each leaf; fleshy; sterile and estípulas pareadas, redondeadas y carnosas en la base
fertile leaves monomorphic; petioles with polycyclic steles; de cada hoja; hojas estériles y fértiles monomorfas;
laminae 2-4-pinnate, triangular, the apex pinnatifid, axes pecíolos con estelas policíclicos; láminas 2-4-pinnadas,
of the distal segments usually winged; pinnae opposite or triangulares, los ápices pinatífidas, los ejes de los segmentos
nearly so; pulvini present at the junctures of the rachises, distales usualmente aladas; pinnas opuestas o casi así;
costae, and costules; veins free, adaxial surfaces with pulvini presentes en las junciones de las ráquises, costas y
greenish awns; receptacles without scales; synangia cóstulas; nervaduras libres, en superficies adaxiales con
stalked, bilateral, splitting longitudinally, with 2 rows arístas verdosas; receptáculos con escamas; sinángios
of opposing sporangia, each sporangium dehiscing by peciolados, bilaterales, rendiéndose longitudinalmente,
a vertical slit on the inner surface; spores monolete, con 2 hileras de esporangios opuestos, cada esporangio
spinulose; x=? abriéndose por una hendidura vertical en la superficie
interior; esporas monoletes; x= ?

SIMILAR GENERA: Marattia differs by more than one GÉNEROS PARECIDOS: Marattia difiere por más
leaf per plant, lacking scattered awns along the veins de una hoja por planta, la falta de aristas por las venas
adaxially, and sessile synangia. Danaea differs by 1-pinnate adaxialmente y sinángios sésiles. Danaea difiere por
laminae, dimorphic sterile and fertile leaves, and synangia láminas 1-pinadas, hojas estériles y fértiles dimorfas y
that open by circular pores. sinángios que abren por poros circulares.

COMMENTS: Eupodium is entirely neotropical and COMENTARIOS: Eupodium es completamente

occurs in wet forests. It contains three species.The first, E. neotropical y existe en bosques húmedos. Contiene
laeve (Sm.) Murdock, occurs in the Greater Antilles. The dos espeies. La primera, E. laeve (Sm.) Murdock, existe
second, E. kaulfussii (J. Sm.) J. Sm., occurs in southern Brazil en las Antilles Mayores. La segunda, E. kaulfussii (J. Sm.)
and northern Argentina. The third, E. pittieri (Maxon) J. Sm., existe en el sur de Brasil y Argentinal al norte. La
Christenh., occurs from Costa Rica to Venezuela and tercera, E. pittieri (Maxon) Christenh., existe desde Costa
Bolivia. The second species differs from the other two Rica hasta Venezuela y Bolivia. La segunda especie difiere
by the presence of awns on the abaxial surfaces of the de las otras por la presencia de arístas en las superficies
veins, scales absent from the laminar surfaces, pulvini not abaxiales de las venas, escamas ausentes de las superficies
tuberculate, and synangia varying from elongate to circular. de las láminas, pulvini no tuberculados y sinángios que
The genus name is derived from the Greek eu-, good, varian desde elongados hasta circulares.
true + pous, foot. It refers to the stalk of the synangium. El nombre genérico se deriva del griego eu-, bueno,
verdad + pous, pie. Refiere al pecíolo del sinángio.

LITERATURE: Christensen, 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.

282
Figure 135. Eupodium. Note in B the characteristic stalked synangia. A-C: E. laevis. D-E: E. kaufussii. (from Tryon & Stolze,
1989).

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PTERIDACEAE
Gaga Pryer, Fay-Wei Li & Windham
DESCRIPTION:Terrestrial; rhizomes short and compact; DESCRIPCIÓN: Terrestres; rizomas cortos y
petioles castaneous to black, typically without hairs; compactos; pecíolos castaños a negros, típicamente sin
rachises grooved adaxially; laminae 2- to 4-pinnate, pelos; ráquises surcados adaxialmente; láminas 2- a
the basal basiscopic pinnules usually exaggerated; costae 4-pinnadas, las pínnulas basales basiscópicas usualmente
and costules grooved adaxially; veins endings forming exageradas; costas y cóstulas surcados adaxialmente;
prominent hydathodes; sori protected by strongly nervaduras terminando en hidátodos prominentes;
differentiated inframarginal (i.e., behind the leaf soros protegidos por pseuodindusios inframarginales
edge) pseudoindusia, these decurent along costae (i.e., atrás del margen de la lámina) fuertemente
and costules; sporangia clustered at vein tips; sporangia diferenciados, estos decurrentes por las costas y
32- or 64-spored; spores globose, trilete, tan, brown or cóstulas; esporangios agrupados en los puntos de las
black. x=30 venas; esporangios con 32 o 64 esporas; esporas globosas,
triletes, leonadas, pardas o negras. x=30

SIMILAR GENERA: Aspidotis differs by lamina segments GÉNEROS PARECIDOS: Aspidotis difiere por las
more acute or mucronate and adaxial lamina surfaces láminas con segmentos más agudos o mucronados y
striate and shiny. Cheilanthes differs by less differentiated superficies adaxiales estriadas y estriadas. Cheilanthes
pseudoindusia, production of 32 small or 16 large spores difiere por pseudoindusio menos diferenciados,
per sporangium, and often scaly laminae. producción de 32 esporas pequeñas por esporangio o
16 esporas grandes por esporangio, y a mendo laminas
escamosas.

COMMENTS: Gaga consists of 19 species, all neotropical. COMENTARIOS: Gaga consta de 19 especies, todas
It occurs from southwestern United States (Arizona neotropicales. Existe desde el suroeste de los Estados
and Texas) to southern Bolivia. Mexico is the center of Unidos (Arizona y Texas) hasta el sur de Bolivia. México es
diversity, with 17 species. Generally in open or semi-open el centro de diversidad, con 17 especies. Generalmente en
habitats, (1500–)2000–4000 m. Gaga is sister to Aspidotis, lugares abiertos o semi-abiertos, (1500–)2000–4000 m.
from which it differs only slightly. Both genera are sister Gaga es hermana a Aspidotis, de lo cual difiere solamente
to Cheilanthes, from which Gaga is nearly inseparable un pocito. Ambos géneros son hermana a Cheilanthes,
morphologically. Gaga is named for the American pop de la cual Gaga es casi morfológicamente inseparable
singer, songwriter, and performer, “Lady Gaga,” stage name inseparable.. Gaga se nombre para la Americana cantante
of Stefani Joanne Angelina Germanotta (1986– ). de “pop”, escritora de cancines y artista “Lady Gaga,”
nombre de teatro de 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.

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Figure 136. Gaga. A–C. G. marginata. D–G. G. hirsuta. H,J,K. G. marginata. In C and G, note the distinctive decurrent pseudo-
indiusia. (from Mickel & Smith, 2004)

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POLYPODIACEAE
Galactodenia Sundue & Labiak
DESCRIPTION: Epiphytes, sometimes epipetric; DESCRIPCIÓN: Epífitas, a veces epipéticas; rizomas
rhizomes short-creeping, the scales concolorous, non- cortamente rastreros, las escamas concoloras, no
clathrate, orange, golden-brown or castaneous, marginally clatradas, anaranjadas a doradas-pardas o castáñas,
ciliate by glandular hairs; leaves pendent, setose or not, ciliadas en el margen por pelos glandulares; hojas setosas
with glandular hairs, these clavate, hyaline, milky o no, con pelos glandulares, estos clavados, hialinos,
white to gray or golden brown; laminae 1-pinnatisect lechosos a grisáceos o pardo-dorados; hojas péndulas,
to 1-pinnate-pinnatisect; rachises blackish, visible on both láminas 1-pinnatisectas a 1-pinnada-pinnatisectas, ráquises
sides of the laminae; pinnae with blackish costae, these negruzcos, visibles en ambos lados de la lamina; pinnas
often visible on both sides of the laminae; veins simple, con costas negruzcas, a mendo visibles en ambos lados
free, not blackish, and not visible without transmitted light; de las láminas; nervaduras simples, libres, no negruzcas,
hydathodes present, non-cretaceous, sometimes no visibles sin luz transmitida; hidatodos presentes,
poorly developed and difficult to see; sori round; sporangia sin puntos blanquecinos calizos, a veces pobremente
glabrous, or when young glandular, the glands similar to desarrollados y difíciles ver; soros redondos; esporangios
those found on the lamina; spores green, trilete. x=? glabros, o, cuando jóvenes glandulares, las glándulas como
las de las láminas; esporas verdes, triletes. x=?

SIMILAR GENERA: Alansmia also has pendent leaves GÉNEROS PARECIDOS: Alansmia también tiene hojas
but differs by petioles and laminae with stellate or péndulas pero difiere por los pecíolos y láminas con setas
clustered setae and, in most species, setulose sporangial estelladas o agrupadas y, en la mayoría de las species,
capsules. Lellingeria, Melpomene, and Stenogrammitis differ cápsulas esporangiales setulosas. Lellingeria, Melpomene y
by lacking glandular hairs on the laminae. Stenogrammitis difieren por falta de pelos glandulares en
las laminas.

COMMENTS: Galactodenia consists of five species, all COMENTARIOS: Galactodenia consta de cinco
neotropical. It occurs from southern Mexico and the especies, todas neotropicales. Existe desde México del
West Indies to Bolivia, (1500–)2000–4000 m. The most sur y las Antillas a Bolivia, (1500–)2000–4000 m. Las
distinctive characters of the genus are its pendent laminae características más distinctivas del género son sus laminas
covered by glandular hairs. Galactodenia is sister to a clade péndulas cubiertas por pelos glandulares. Galactodenia es
composed of Lellingeria, Melpomene, and Stenogrammitis. hermana a un clado formado por Lellingeria, Melpomene
It is not closely related to Alansmia or Terpsichore, the y Stenogrammitis. No es cercanamente relacionado a
two genera in which two of its species were previously Alansmia o Terpsichore, los dos géneros donde dos de sus
classified. Its species are G. delicatula, G. pumila, G. parriseae, species han sido clasificadas préviamente. Sus especies
G. subscabra, and G. vareschii. A key to these species is son G. delicatula, G. pumila, G. parriseae, G. subscabra y G.
given in Sundue et al. (2012).Galactodenia is derived from vareschii. Una clave a estas species se presenta en Sundue
the Greek galacto, milk + adeno, gland, and refers to the et al. (2012). Galactodenia se deriva del Griego galacto,
milky white glandular hairs characteristic of these plants. leche + adeno, glándula, y se refiere a los pelos glandulares
lechoso-blancos característicos de estas plantas.

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.

286
Figure 137. 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; used with permission.)

287
GLEICHENIACEAE
Gleichenella Ching
DESCRIPTION: Terrestrial; rhizomes solenostelic, long- DESCRIPCIÓN: Terrestre; rizomas solenostélicos,
creeping, densely pubescent, the hairs stiff, multicellular, largamente rastreros, densamente pubescentes, los pelos
unbranched; leaves born in a single row along the dorsal tiesos, multicelulares, no furcados; hojas nacidas en una
surface of the rhizome, up to 3 m or more long, erect sola hilera en la superficie dorsal, hasta 3 m o más de largo,
or scandent, indeterminate, of periodic growth; pinnae erectas o escandentes, indeterminadas, de crecimiento
opposite, unequally branched, with the larger branch periódico; pinnas opuestas, desigualmente furcadas,
alternately to one side of the axis and then the other, con la rama más grande alternadamente a un lado y luego
the minor branches also unequally branched; pinna axes en el otro, las ramas menores desigualmente ramificadas
with a narrow lateral ridges on either side; dormant también; ejes de las pinnas con una fila estrecha
buds in the forks pubescent, the hair dark, stiff; penultimate lateral en cada lado; brotes en los furcos pubescentes,
divisions pectinate, glabrous and often glaucous beneath; los pelos oscuros, tiesos; divisiones penúltimos pectinadas,
veins free, 2-4-forked; sori round, non-indusiate; sporangia glabras y a menudo glaucas en el envés; venas libres,
(6–)8–16(–25), sessile, with oblique annuli; spores 2-4-furcadas; soros redondos, sin indusios; esporangios
monolete, ellipsoid; x=43. (6–)8–15(–25), séssiles, con anillo oblicuo; esporas
monoletes, elipsoides; x=43.

SIMILAR GENERA: Dicranopteris differs by equally GÉNEROS PARECIDOS: Dicranopteris difiere por
forked pinnae and (often) accessory pinnae at the forks. pinnas surcadas igualmente y (a menudo) pinnas accesorias
Sticherus differs by scaly buds in the pinna forks, veins en los furcos. Sticherus difiere por brotes escamosos en
usually 1-forked, and sporangia 3–6 per sorus. Gleichenella las furcaciones de las pinnas, venas usualmente 1-furcadas
is the only genus of Gleicheniaceae with solenostelic y esporangios 3–6 por soro. Gleichenella es el único
rhizomes and a narrow lateral ridge on either side of the género de Gleicheniaceae con rizomas solenostélicos y
axes of the pinnae. una fila estrecha en cada lado de los ejes de las pinnas.

COMENTARIOS: Gleichenella es monotípico, con su

COMMENTS: Gleichenella is monotypic, with its sole única especie ocurriéndose desde México hasta el sur de
species occurring from Mexico to southern Brazil and Brasil y Bolivia, y en las Antillas. Como los otros géneros
Bolivia, and in the West Indies. Like other genera in the en la familia, crece en lugares abiertos, soleados, a menudo
family, it grows in open, sunny, often disturbed habitats. It en ambientes perturbados. Cubre con frecuencia taludes
frequently covers steep roadsides with dense colonies. de caminos con colonias densas. Es la especies de
It is the most frequent species of Gleicheniaceae at low Gleicheniaceae más frecuente en tierras bajas y sus hojas
elevations, and its light yellowish green leaves makes amarillentosas verde claros lo hace fácil distinguir en la
them easy to identify at a distance. Gleichenella is sister distancia.
to Dicranopteris, the genus in which it was formerly Gleichenella es hermana Dicranopteris, el género en
included; therefore, its separation as a distinct genus from lo que fue incluido anteriormente; por eso, su separación
Dicranopteris is only a matter of opinion. It differs from de Dicranopteris es solo un asunto de opinión. Difiere
Dicranopteris by pinna branching pattern, solenostelic de Dicranopteris por el patrón de ramificación de las
rhizomes, chromosome number, and spore morphology. pinnas, rizomas solenostélicos, número cromosómico y
The generic name comes from Gleichenia + -ella, a morfología de las esporas.
suffix denoting the diminutive. El nombre genérico se deriva de Gleichenia + -ella, un
sufijo denotando el diminutivo.

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.

288
Figure 138. Gleichenella pectinata. Note unequal bifurcation of the pinna in A. (From Mickel & Smith, 2004)

289
POLYPODIACEAE
Grammitis Sw.
DESCRIPTION: Epiphytic, rarely saxícolous or terresrial; DESCRIPCIÓN: Epiftíicas, raramente saxícolas o
rhizome scales not clathrate, entire; leaves up to 20 cm terrestres; escamas del rizoma no clatradas, enteras;
long, simple, entire, glabrous, pubescent or setulose, hojas hasta 20 cm de longitud, simples, enteras, glabras,
with a thick, dark sclerotic margin; hydathodes present pubescentes o setulosas, con un margen grueso,
or absent; veins simple or 1-forked, free; sori borne on esclerosado, oscuro; hidátodos presentes o ausentes;
a short acroscopic veinlet, round or slightly elongate, nervaduras simples o 1-bifurcadas, libres; soros sobre una
generally inframedial and grouped near the costa when venilla corta acroscópica, redondeados o ligeramente
mature, without paraphyses; spores green. x= ? alargados, generalmente inframedios y aglomerándose en
la costa cuando maduros, sin parafisos; esporas verdes.
x= ?

SIMILAR GENERA: All other genera of grammitid GÉNEROS PARECIDOS: Todos los otros géneros de
ferns differ by green (not black) lamina margins. Many helechos grammitidoides difieren por márgenes de las
others differ further by pinnatifid to 1-pinnate laminae láminas verdes (no negros). Muchos otros difieren además
and the prescence of reddish setae on the laminae (only por láminas pinnatífidas a 1-pinnadas y la presencia
Cochlidium, Grammitis, Lellingeria and Luisma lack setae). de setas rojizas en las láminas (solamente Cochlidium,
Grammitis, Lellingeria y Luisma faltan setas).

COMMENTS: Grammitis consists of about 22 species COMENTARIOS: Grammitis consta de casi 22 especies
of both the New and Old World tropics and subtropics. de los trópicos y subtrópicos del Nuevo yViejo Mundo. Las
The species usually grow in cloud forests. Molecular data especies usualmente crece en bosques nublados. Datos
indicate that Grammitis as defined here (i.e., restricted moleculares sugieren que Grammitis como definido aquí
to only those species with black laminar margins) are (i.e., restringido a las especies con márgenes laminares
monophyletic (Ranker et al. 2004). Although the genus negros) son monofíleticos (Ranker et al. 2004).A pesar
has been revised three times this century (Bishop, 1977; de que el género ha sido revisado tres veces en este siglo
Copeland, 1952; Maxon, 1915), relatively few specimens (Bishop, 1977; Copeland, 1952; Maxon, 1915), se disponía
were available for study. The many recent collections, de relativamente pocos especímenes para los estudios.
some of which have been described as new species, make La gran cantidad de colecciones recientes, algunas de las
necessary a new revision of the genus. The genus name cuales han sido descritas como especies nuevas, hacen
is derived from the Greek gramme, line, referring to the necesaria una nueva revisión del género. El nombre
elongate to linear sori. 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.

290
Figure 139. A–C. Grammitis bryophila (Maxon) F. Seym. D–F. G. marginella (Sw.) Sw. G­J. G. leptopoda (C. H. Wright) Copel..
K–M. Lomaphlebia graminea (Sw.) comb. ined.? (© R. C. Moran, 2011)

291
CYSTOPTERIDACEAE
Gymnocarpium Newman
DESCRIPTION:Terrestrial; rhizomes long-creeping, scaly; DESCRIPCIÓN: Terrestres; rhizomas largamente
sterile and fertile leaves monomorphous, thin-textured; rastreros, escamosos; hojas estériles y fértiles
petioles with 2 vascular bundles; laminae usually monomorfas, de textura delgada; pecíolos con 2 haces
10–30 cm long, broadly deltate, 2-3-pinnate-pinnatifid; vasculares; láminas usualmente 10­ 30 cm de largo,
pinnae with a slightly swollen point of attachment to the anchamente deltadas, 2-3-pinnado-pinnatífidas; pinnas
rachis; axes grooved on the upper surface, the grooves con un punto de unión con el raquis levemente hinchado;
not decurrent; veins free, the tips slender (not enlarged); ejes surcados en la superficie superior, los surcos no
indument lacking or of minute (0.1 mm long) capitate- decurrentes; indumento ausente o consta de pelitos
glandular hairs; sori round, medial or inframedial; indusia capitados glandulares diminutos (0.1 mm de largo);
absent; spores monolete, bilateral; x=40. 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 y es principalmente de distribución templada en América
Eurasia. All species are terrestrial in shaded forests or del Norte y Eurasia. Todas las especies son terrestres
on shaded limestone. They tend to form diffuse colonies en bosques sombreados o en piedra caliza sombreada.
by the action of their long-creeping rhizomes. Molecular Tienden formar colonias difusas por acción de sus rizomas
phylogenetic studies indicate that Gymnocarpium is largamente rastreros. Estudios filogenéticos molecular
sister to Cystopteris, and the two are sister to the rest of indican que Gymnocarpium es hermana a Cystopteris, y los
Eupolypods II. Thus, the two genera deserve their own dos son hermana al resto de Eupolypods II. Por ende los
family, Cystopteridaceae. This is a deviation from Smith dos géneros merecen su propia familia, Cystopteridaceae
et al. (2006) generally followed here. The genus name is .Esto es una desviación de la clasificación de Smith et al.
derived from the Greek gymnos, naked + karpos, fruit. It (2006) generalmente seguido aquí. El nombre genérico
refers to the absence of an indusium. 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.

292
Figure 140. A–D. Gymnocarpium dryopteris. (©R. C. Moran, 2008)

293
CYATHEACEAE
Gymnosphaera Blume
DESCRIPTION: Stems arborescent; sterile and fertile DESCRIPCIÓN: Tallos arborescentes; hojas estériles
leaves monomorphic or slightly dimorphic; petioles with y fértiles monomorfas o levemente dimorfas; pecíolos
4–10 pairs of small, skeletonized pinnae (aphlebia) con 4–10 pares de pinas pequeñas en esqueletos
near the base and distant from the normal pinnae; (aflebias) cerca la base del pecíolo y distantes
petiole scales with single black apical setae, the de las pinnas normales; escamas peciolares
marginal cells different in shape and orientation from con una seta apical negra, las células marginales
the central ones (i.e., marginate); laminae 1-pinnate- difieren en forma y orientación de las centrales (i.e.,
pinnatifid to 4-pinnate, the apex pinnatifid or rarely marginadas); láminas 2-pinada-pinatífidas a 3-pinadas,
conform; veins free; axes pubescent adaxially, the hairs el ápice gradualmente reducido; nervaduras libres; ejes
thick, strigose, multiseptate; sori round, the receptacle pubescentes adaxialmente, los pelos gruesos, estrigosos,
elevated, subglobose; indusia absent or (in A. capensis) multiseptados; soros redondos, el receptáculo elevado,
present and hemitelioid; spores tetrahedral-globose, 64 subgloboso; indusios ausentes o (en A. capensis) presentes
per sporangium, ridged, without pores; x=69. y hemitilioides; esporas tetrahédricas-globosas, 64 por
esporangio, con crestas bajas, sin poros; x=69.

SIMILAR GENERA: Alsophila differs by 16 spores GÉNEROS PARECIDOS: Alsophila differs by 16

per sporangium and lack of aphlebia. Cyathea and spores per sporangium y la falta de aflebias. Cyathea y
Sphaeropteris differ by lacking an aerophore pit on the Sphaeropteris difieren por la falta de un aeróforo como
rachis at the pinna juncture. Sphaeropteris further differs hueco por el raquis en la unión de la pinna. Sphaeropteris
by conform petiole scales (i.e., all cells of the same shape difiere además por escames peciolares conformes (i.e.,
and orientation). todas la células del mismo forma y orientación).

COMMENTS: Gymnosphaera contains about 16 species COMENTARIOS: Gymnosphaera consta de casi 16

and is entirely Old World except for G. capensis (SE Brazil, especies y es completamente del Viejo Mundo con
Africa) and G. salvinii (Central America, Peru). It occurs excepción de G. capensis (SE Brasil, África) y G. salvinii
in shaded forests. In a phylogenetic analysis (Korall et al., (América Central, Peru). Existe en bosques sombreados.
2007), Gymnosphaera formed a trichotomy with Alsophila En un análisis filogenético (Korall et al., 2007),
and Cyathea. All three genera have marginate petiole Gymnosphaera formó una tricotomía con Alsophila y
scales. As in Alsophila, the aerophore in Gymnosphaera is Cyathea. Los tres géneros tienen escamas peciolares
a sunken pit in the rachis at the juncture of the costa, marginadas. Como en Alsophila, el aeróforo de
and there is a single black apical seta on the petiole Gymnosphaera es un hueco hundido en el raquis cerca de
scales.Gymnosphaera has dark rachises and costae, and la unión de la costa, y hay una sola seta negra apical en las
this will distinguish it from many (not all) scaly tree escamas peciolares. Gymnosphaera tiene ráquises y costas
ferns. Many Old World species have the fertile pinnae oscuras, y esto lo distinguirá de muchos otros (no todos)
contracted compared to the sterile. Only G. capensis has helechos arborescentes con escamas. Muchas especies
an indusium; the others in the subgenus are non-indusiate. del Viejo Mundo tienen las pinnas fértiles contraídas en
The name is derived from the Greek gymnos, naked + comparación de las estériles. Solo G. capensis tiene un
sphaera, ball. It refers to the non-indusiate sori. 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.

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.

294
Figure 141. A–B, H–M. Gymnosphaera salvinii, in note aphlebiae (skeletonized pinnae) on petiole base. C–G. G. capensis ssp.
polypodioides (© R. C. Moran, 2011).

295
PTERIDACEAE (VITTARIOID CLADE)
Hecistopteris J. Sm.
DESCRIPTION: Epiphytic or rupicolous; rhizomes DESCRIPCIÓN: Epífitas o rupícolas; rizomas
short-erect, proliferous roots present; rhizome scales cortamente erectos, raíces prolíferas presentes;
clathrate, not obscured by densely clumped pubescent escamas del rizoma clatradas, no oscurecidas por raíces
roots; petioles absent or very short; laminae less than 4 cespitosas densamente pelosas; pecíolos ausentes o muy
cm long, incised, narrowly cuneate, flabellate, glabrous, reducidos; láminas menos de 4 cm de largo, incisas,
with spicular idioblasts (false veins) adaxially; veins free; estrechamente cuneada en la base, flabelada, glabra, con
sori elongate along the veins in the distal portion of the idioblastas spiculares (venas falsas) en el haz; nervaduras
lamina; paraphyses simple or branched, with an enlarged libres; soros alargados junto a las nervaduras en la
apical cell; spores trilete, nongreen; gametophytes with porción distal de la lámina; parafisos simples o ramificados
unpaired gemmae; x=? con una célula apical agrandada; esporas triletes, no
verdes; gametofitos con yemas no pareadas; x=?

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ningunos.

COMMENTS: Hecistopteris is entirely neotropical COMENTARIOS: Hecistopteris es completamente

and consists of three species. They grows primarily neotropical y consta de tres especies. Crecen
as an epiphyte in wet, shaded forests below 600 m. It principalmente como epífito en bosques húmedos y
sometimes grows on rotting logs.The plants form colonies sombreados bajo de 600 m. Algunas veces crece sobre
by elongate rootlets that bear buds, and it is typical to find troncos podridos. Las plantas forman colonias por
several plants united by a single rootlet. For over 150 raicillas alargadas que llevan yemas adventicias, y es típico
years Hecistopteris was considered monotypic, containing encontrar varias plantas unidas a una raicilla única. Por
the sole species H. pumila (Spreng.) J. Sm., which occurs mas de 150 años se considera a Hecistopteris monotípico,
from Mexico to southern Brazil and Bolivia, and the conteniendo la única especie H. pumila (Spreng.) J. Sm.,
Antilles. But recently a species was described from la que ocurre desde México hasta el sur de Brasil y
Guyana (H. kiaeteurensis Kelloff & G. McKee), and another Bolivia, y las Antillas. Pero recientemente una especie fue
from Ecuador (H. pinnatifida R. C. Moran & Øllgaard). descrita de Guyana (H. kiaeteurensis Kelloff & G. McKee)
Hecistopteris is sister to Radiovittaria (Crane, 1997) and y otra de Ecuador (H. pinnatifida R. C. Moran & Øllgaard).
differs from that genus, and all other vittarioid ferns, by Hecistopteris es hermana a Radiovittaria (Crane, 1997),
incised leaves, proliferous roots, and free veins. The genus y difiere de ese género y todos los demás helechos
name is derived from the Greek hekistos, least + pteris, vittarioides por sus hojas incisas, raíces prolíferas y venas
fern, referring to the small size of H. pumila. libres. El nombre genérico se deriva del griego hekistos,
menos + pteris, helecho, refiriéndose al tamaño pequeño
de 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.

296
Figure 142. 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)

297
HEMIDICTYACEAE
Hemidictyum L.
DESCRIPTION: Terrestrial; leaves ca. 3 m, erect-arching; DESCRIPCIÓN: Terrestres; hojas c. 3 m, erectas-
sterile and fertile leaves monomorphous; the base of the arqueadas; hojas estériles y fértiles monomorfas; la base
petiole with 2 vascular bundles, these uniting distally in del pecíolo con 2 haces vasculares, estos uniéndose
a U-shape; laminae 1-pinnate with a terminal pinnae distalmente en forma de “U”; láminas 1-pinnadas
similar in form to the lateral ones; pinnae 23–50 X con una pinna terminal similar en forma a las
6–12 cm, linear-oblong, entire, opposite to subopposite, pinnas laterales; pinnas 23–50 X 6–12 cm, linear-
sessile, slightly cordate and overlapping the rachis; veins oblongas, enteras, opuestas o subopuestas, la base
areolate toward the marings, a submaringal connecting sésil, ligeramente cordiformes y traslapadas al raquis;
vein present; indusia narrow, linear membranaceous; nervaduras anastomosadas hacia los márgenes, una
spores bilateral, brown; x=31. vena colectora submarginal presente; indusios angostos,
lineares, membranáceos; esporas bilaterales; x=31.

SIMILAR GENERA: Asplenium differs by smaller leaves GÉNEROS PARECIDOS: Asplenium difiere por hojas
(never more than 1 m long). Diplazium differs by at least más pequeñas (nunca más de un metro de longitud).
some of the sori (usually the basal ones) back to back Diplazium difiere por tener a lo menos unos soros
(diplazioid); none of its species have entire pinnae and (usualmente los basales) espalda a espalda (diplazioide);
anastomosing veins. ninguna de sus especies tienen pinnas enteras y venas
anastomosadas.

COMMENTS: Hemidictyum consists of only one species, COMENTARIOS: Hemidictyum consta de una sola
H. marginatum (L.) C. Presl, which occurs throughout especie, H. marginatum (L.) C. Presl, la cual existe por
most of the Neotropics. It generally grows in wet forests la mayoría de los Neotropicos. Generalmente existe
or forest margins from 0–1300 m. The Hemidictyaceae en bosques húmedos o por los bordes de bosques de
is monotypic. Although sister to Aspleniaceae, it 0–1300 m. Hemidictyaceae es monotípico. Aunque es
resembles certain species of the Asian Diplaziopsis in the hermana a Aspleniaceae, se asemeja ciertas especies
Diplaziopsidaceae, the closest outgroup family. The genus del Asiático Diplaziopsis en Diplaziopsidaceae, la familia
name is derived from the Greek hemi, half + diktyon, net. grupo afuera más relacionada. El nombre genérico se
The veins anastomose in the distal half of the pinnae. 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.

298
Figure 143. Hemidictyum marginatum. A. leaf apex. B. sori and venation. C. pinna margin. (from Stolze, Fern & Fern Allies of
Guatemala, 1981).

299
PTERIDACEAE
Hemionanthes Mickel
DESCRIPTION: Terrestrial; rhizomes compact, DESCRIPCIÓN: Terrestres; rizomas compactos,
horizontal, the scales linear, bicolorous with a dark central horizontales, las escamas lineares, bicolores con una raya
stripe; petioles stramineous, mostly glabrous, 3-grooved, central oscura; pecíolos estramineosos, principalmente
ca. half the length of the leaf; blades lanceolate to deltate, glabros, 3-surcados, ca. Una mitad del largo de la hoja;
pinnate-pinatifid to 2-pinnate basally; pinnae 5–7 pairs, láminas lanceoladas a deltadas, pinnadas-pinnatífidas
the lobes obtuse, basiscopically somewhat exaggerated, a 2-pinnadas basalmente; pinnas 5–7 pares, los lobos
the adaxial surfaces glabrous, abaxial surfaces with obtusos, basiscopicamnte also exagerado, superficies
sparse straight jointed hairs; rachises with sparse narrow, adaxiales con pelos articulados esparsos; ráquises con
bicolorous scales; veins free with a few casual anastomoses escamas bicoloras angostas esparsas; venas libres con
(2–5 per pinna); sporangia extending along the distal unas pocas anastomoses casuales (2–5 por pinna);
2–4 mm of the veins; indusial absent, blade margins not esporangios extendiéndose por el distal 2–4 mm de
reflexed or modified; spores tetrahedral-globose, crested; las venas; indusios ausentes, márgenes de las láminas
x=30. 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: COMENTARIOS: Hemionanthes consta de tres species:
H. gryphus (Mickel) Mickel, H. lozanoi (Maxon) comb. ined. H. gryphus (Mickel) Mickel, H. lozanoi (Maxon) comb. ined.
y H. seemannii (Hook.) comb. ined. All are endemic to y H. seemannii (Hook.) comb. ined. Todas son endémicas
Mexico. They grow in deciduous woodlands and moist a México. Crecen en bosques decíduos en arroyos
shaded gullies. Hemionanthes gryphus has the appearance sombreados húmedos. Hemionanthes gryphus tiene
of a hybrid between Cheilanthes skinneri and Hemionitis la apariencia de un híbrido entre Cheilanthes skinneri y
subcordata, which can grow together with it; however, Hemionitis subcordata, que pueden crecer juntos con
Hemionanthes is not a hybrid. It has normal spores and is él; no obstante, Hemionanthes no es un híbrido. Tiene
only distantly related to the two species it resembles.The esporas normales y es solo relacionado distantamente a
genus name is derived from the intermediate appearance las dos especies que se asemeja. El nombre genérico se
of the species between Cheilanthes skinneri and Hemionitis deriva de la apariencia intermedia de la especies entre
subcordata. 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.

300
Figure 144. Hemionanthes gryphus. A. Habit. B. Rhizome scale. C. Pinnule. (Mickel & Smith, 2004)

301
PTERIDACEAE
Hemionitis L.
DESCRIPTION: Terrestrial or rupestral; rhizomes erect DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto a
to short-creeping, scaly, the scales (at least those toward cortamente rastrero, escamoso, las escamas concoloras
the apex of the rhizome) concolorous, but those on the hacia el ápice del rizoma, aquellas en las porciones más
older part of the rhizome bicolorous with a dark central viejas del rizoma bicolores, con una franja media castaño
strip or concolorous (in H. palmata); leaves monomorphic o concolora (en H. palmata); hojas monomorfas a
or dimorphic; petioles castaneous, dark brown or straw- dimorfas; pecíolo castaño, pardo oscuro o pajizo, con un
colored, with 1 vascular bundle; laminae simple, orbicular haz vascular; lámina simple, orbicular, pedata, pinnatífida
to 1-2-pinnate, densely to sparsely hairy, occasionally o 1-2-pinnada, densa a esparcidamente pilosa,
on the lower surface with two-celled, gland-tipped hairs; ocasionalmente con tricomas 2-celulares con glándulas
veins netted or free, if netted then the areoles without en las puntas en el envés; nervaduras o anastomosadas, las
included veinlets; sori running along the veins, aréolas sin nérvulos incluidos; esporangios cubriendo
sometimes (in H. subcordata ) restricted to the distal one- las nervaduras a todo lo largo o a veces restringidos al
quarter of near the margin; indusium absent; x=30. 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. COMENTARIOS: Hemionitis tiene 10 especies,
The genus typically grows in mesic habitats such as shaded todas Neotropicales. El género crece típicamente en
rocks, tall forests, and along rivers. Two clades occur hábitats mesofíticas como rocas sombreadas, selvas
within the genus: one with palmate laminae and netted altas perennifolias, o comunidades raparías. Dos clados
veins (Hemionitis s.s.) and another with pinnate lamina existen dentro del género: uno con láminas palmadas y
and free veins (formerly distinguished as Gymnopteris venas anastomosadas (Hemionitis s.s.) y otra con laminas
Bernh.).Hemionitis arifolia (Burm. f.) T. Moore, a commonly 1-pinnadas y venas libres (anteriormente distinguido
cultivated Asian species, was resolved sister to the four como Gymnopteris Bernh.). Hemionitis arifolia (Burm. f.) T.
neotropical Hemionitis species sampled by Zhang et Moore, una especie Asiática comúnmente cultivada, fue
al. (2010). The genus name is derived from the Greek resuelta hermano a los cuatro especies neotropicales de
hemionis, mule. Ancient name for a fern thought to be a Hemionitis muestradas por Zhang et al. (2010). El nombre
hybrid. 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.

302
Figure 145. A: Hemionitis pinnatifida. B: H. levyi. C: H. palmata. (Mickel & Smith, 2004)

303
DENNSTAEDTIACEAE
Histiopteris (J. G. Agardh) J. Sm.
DESCRIPTION: Terrestrial; rhizomes long-creeping, 2-5 DESCRIPCIÓN: Terrestres; rizomas largamente
mm wide, with castaneous hairs and scales; leaves large, rastreros, 2-5 mm de ancho, con pelos y escamas castaños;
distant, erect to subscandent; petiole base in cross-section hojas grandes, distantes, erectas a subescandentes; la
with a vascular bundle in the form of a corrugated “U”; base del pecíolo en sección cruz con un hace vascular
blades 2- to 3-pinnate, often with large stipule-like en forma de un “U” corrugado; láminas 2- a 3-pinnadas,
pinnules at the pinna base; pinnae and pinnules a menudo con pínnulas que asemejan a estípulas;
opposite, light pea-green, herbaceous; lamina glabrous pinnas y pínnulas opuestas, verde claras, herbáceas,
or with crisped, multicellular hairs, often glaucous; veins glabras o con pelos multicelulares crespadas, a menudo
netted, without included veinlets; sori marginal, glaucas; venas anastomosadas, sin venas incluídas;
continuous, supplied by more than one vein, blade soros marginales, contínuos; suministrados por más de
margin recurved and differentiated as an indusium (false una vena, margen de la lámina recurvado y diferenciado
indusium); inner indusium absent; spores bilateral; x=96. como indusio (indusio falso); un indusio interior ausente;
esporas bilaterales; x=96.

SIMILAR GENERA: Blotiella differs by shorter sori, GÉNEROS PARECIDOS: Blotiella difiere por soros más
pubescent laminae and absence of stipule-like pinnules cortos, láminas pubescentes y la ausencia de pínnulas que
at the base of the pinnae. Hypolepis differs by sori asemejan como estípulas en la base de las pinnas. Lonchitis,
supplied by a single vein. Lonchitis, which differs by free lo cual difiere por venas libres, láminas pubescentes, y la
veins, pubescent laminae, and the presence of an inner presencia de un indusio interior.
indusium.
COMENTARIOS: Histiopteris incsa (Thunb.) J. Sm. es
COMMENTS: Histiopteris incsa (Thunb.) J. Sm. is pantropical y la única especie del género que existe en el
pantropical and the only species of the genus to occur in Neotrópico. Es una especie de las montañas, ocuriendo
the Neotropics. It is a montane plant, occurring primarily principalmente desde 1000–3000 m. Cuatro otras
from 1000–3000 m. Four other species occur in Southeast especies existen en el sureste de Asia y el Pacífico del sur.
Asia and the South Pacific. The leaf grows intermittently. La hoja crece intermitente. El ápice descansa mientras
The apex rests while the pinnae pair beneath develop. que el par de pinnas abajo se desarrollan. Después de que
After the pinnae are nearly fully developed, the apex las pinnas son casi completamente desarrolladas, el ápice
resumes growth. Thus the leaves almost always have a resume crecimiento. Por eso, las hojas casi siempre tienen
fiddlehead at their apex, this usually protected by the un cayado en sus ápices, esto usualmente protegido por
enlarged basal pinnules. The genus name is derived from las pínulas basales agrandados. El nombre genérico se
the Greek histion, sail + pteris, fern. It refers to the small deriva del griego histion, vela + pteris, helecho. Se refiere
sail-like basal pinnules. a la pínnula pequeña basal que asemeja una vela.

LITERATURE: none; ninguna

304
Figure 146. Histiopteris incsa. A. Pinnae. B. Creeping rhizome and petiole base with epipetiolar bud. C. Fertile pinnule and
net veins. (Mickel & Smith, 2004)

305
DIPLAZIOPSIDACEAE
Homalosorus Small ex Pic. Serm.
DESCRIPTION: Terrestrial; rhizomes creeping to erect, DESCRIPCIÓN: Terrestres; rizomas rastreros a erectos,
scaly, the scales not clathrate; sterile and fertile leaves escamosos, las escamas no clatradas; hojas estériles y
subdimorphic, ceaspitose; petiole bases with 2 vascular fértiles subdimorphic; bases de los pecíolos con 2 haces
bundles, these uniting distally to form a “U”; laminae vasculares, éstos uniéndose distalmente para formar
1-pinnate, hairs absent, scales sparse to absent; pinnae un haz vascular en forma de “U”; láminas 1-pinnadas,
sessile, equilateral, truncate or nearly so at base; rachis pelos ausentes, escamas esparcidas al ausentes; ráquises
sulcate adaxially, lacking hairs within the groove; veins free, surcados adaxialmente, sin pelos en los surcos; pinnas
usually ending before the margin in clavate hydathodes; séssiles, equiláterales, truncadas o casi así en la base;
fertile leaves longer petiolate with narrower pinnae; sori venas libres, usualmente terminándose antes del margen
elongate, approximate, filling most of the laminar en hidatodos clavados; hojas fértiles más largamente
tissue between the costa and margin, occasionally pecioluladas con pinnas más estrechas; soros alargados,
paired back to back along a single vein; indusia present; aproximados, llenando la mayoría del tejido láminar
annulus of 15–20 cells; spores bilateral, brown, cristate; entre la costa y márgen, a veces pareados de par en
x=40. 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 GÉNEROS PARECIDOS: Deparia difiere por pelos
hairs on the leaves. Athyrium and Diplazium differs by plateados articulados en las hojas. Athyrium y Diplazium
rachises and costae grooved adaxially with the grooves difieren por ráquises y costas surcados adaxialmente con
decurrent into one another, and usually by having some los surcos decurrentes, y usualmente con algunos pelos
hairs on the leaves, at least in the grooves adaxially. In both foliares, al menos en los surcos adaxialmente. En ambos
genera, the sori are rarely as crowded and filling so much géneros, los soros raras veces son tan congestionados y
space between the costa and margin as in Homalosorus. llenando tanto espacio entre la costa y el márgen como
en Homalosorus.

COMMENTS: Homalosorus consists of a single species: COMENTARIOS: Homalosorus consta de una sola
H. pycnocarpon (Spreng.) Pic. Serm. It is native to especie: H. pycnocarpon (Spreng.) Pic. Serm. Es nativa a
eastern North America where it grows in wet forests. América del Norte oriental donde existe en bosques
Morphologically, it most resembles Diplazium flavoviride húmedos. Morfológicamente, se asemeja más a Diplazium
Alston (Kato & Darnaedi, 1988), a species of eastern flavoviride Alston (Kato & Darnaedi, 1988), una especie de
Malesia that should be transferred to Homalosorus. The Malesia oriental que debe ser transferida a Homalosorus.
sori are single along the veins; rarely are they back-to- Los soros son simples por las venas; raras veces son
back as in Diplazium. The young indusia are inflated and espaldas a espaldas como en Diplazium. Los indusios
silvery, turning brown with age. Homalosorus is sister jóvenes son infladas y plateadas, tornándose pardos con
to Diplaziopsis, an Asian genus. Together they form edad. Homalosorus es hermana a Diplaziopsis, un género
the Diplaziopsidaceae, a family closely related to the asiático. Juntos, forman la Diplaziopsidaceae, una familia
Hemidyctiaceae and Aspleniaceae (Wei et al., 2010). The emparentada con Hemidictyiaceae y Aspleniaceae (Wei
genus name is derived from the Greek homalos, even, et al., 2010). El nombre genérico se deriva del griego
smooth + soros. It refers to the evenly spaced sori. 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.

306
Figure 147. Homalsorus pycnocarpon (Small, 1935. Ferns of the Vicinity of New York)

307
LYCOPODIACEAE
Huperzia Bernh.
DESCRIPTION: Plants epiphytic or terrestrial, pedulous, DESCRIPCIÓN: Epífitas o terrestres; plantas péndulas,
erect, or ascending, homophyllous or heterophyllous, erectas o ascendentes, homofilas o heterofilas;
branched equally; roots emerging from the base of ramificadas igualmente; raíces emergiéndose de la
the stems; sporophylls and vegetative leaves similar, or base del tallo; esporofilos y hojas vegetativas similares,
the sporophylls attached at the base (not peltate), o los esporofilos únidas en la base (no peltados),
persistant after sporangial dehiscence; sporangia persistentes tras la dehiscencia del esporangio;
axillary, reniform, isovalvate, short-stalked spores esporangios axilares, reniformes, isovalvados, con un
foveolate-fossulate; x=67, 68. pedículo corto; esporas foveolado-fosuladas; x=67, 68.

SIMILAR GENERA: Lycopodium and Lycopodiella differ GÉNEROS PARECIDOS: Lycopodium y Lycopodiella
by unequally branched stems (usually a main, long creeping difieren por tallos ramificadas desigualmente (usualmente
stem is present), and their sporophylls are peltate and turn un tallo principal largamente rastrero es presente), y sus
yellowish or straw-colored after sporangial dehiscence. esprofilos son peltatdos y amarillentos o pajizos después
de dehiscencia esporangial.

COMMENTS: Huperzia has about 400 species COMENTARIOS: Huperzia tiene casi 400 especies
worldwide. It is most diverse in the Andes, where nearly mundial. Es más diverso en los Andes, donde casi una
half of the species are endemic.The genus occurs in many mitad de las especies son endémicas. El género existe
habitats, but in the Neotropics most species grow in the en muchos hábitats, pero en el Neotrópico la mayoría
páramos. Epiphytism in Huperzia evolved prior to the de las especies ocurren en el páramo. El epifitismo en
break-up of South America and Africa, and in the high Huperzia se evolucionó antes del romper de América del
Andes there as been a reversion to the terrestrial habit Sur y África, y en los Andes altos ha sido una reversión
at least twice (Wikström & Kenrick, 1999). The genus is al habit terrestre a lo menos dos veces (Wikström &
named for Johann Peter Huperz (d. 1816), author of a Kenrick, 1999). El nombre genérico honra Johann Peter
book on fern propagation. Huperz (d. 1816), autor de un libro sobre propagación
de helechos.

LITERATURE: Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Øllgaard,
B. 1989. Index of the Lycopodiaceae. Biologiske Skrifter 34: 1–135. Øllgaard, B. 1992. Neotropical Lycopodiaceae—an overview.
Annals of the Missouri Botanical Garden 79: 687–717. Rolleri, C. H. 1980. Sinopsis de las especies de Lycopodium de la sección
Crassistachys Herter. Revista Museo de La Plata, Sección Botánica. n.s., 13(71): 61–113. Wikstöm, N. & P. Kenrick. 1999.
Phylogeny of epiphytic Huperzia (Lycopodiaceae): paleotropical and neotropical clades corroborated by rbcL sequences. Nordic
Journal of Botany 20: 165–171.

308
Figure 148. A. Huperzia hippuridea. B. H. crassa. C, D. H. pithyoides. E, F. H. reflexa. (Mickel & Smith, 2004).

309
CYATHEACEAE
Hymenophyllopsis Goebel (= Cyathea)
DESCRIPTION: Terrestrial or rupestral; rhizomes DESCRIPCIÓN: Terrestres o rupestres; rizomas
ascending, short-creeping, or erect, scaly; leaves 10–30(– ascendentes, cortamente rastreros, o erectos, escamosos;
40) cm long, monomorphic, fasciculate; petioles terete hojas 10–30(–40) cm de longitud, monomorfas,
or sulcate, brown to black; blades 1-pinnate-pinnatifid fasciculatadas; pecíolos teretes o sulcados, pardos o
to 3-pinnate-pinnatifid, several cells thick between negros; láminas 1-pinnado-pinnatífidas, de varias
the veins, lacking stomata, glabrous to sparsely scaly cellulas de grosor entre las venas, careciendo de
on both surfaces; pinnules or ultimate segments narrow, estomas, glabras en ambas superficies; pínnulas
often linear; veins usually one per ultimate segment; o segmentos últimos estrechos, a menudo lineares;
sori marginal or submarginal; indusium bivalved, nervaduras usualmente una por segmento último; soros
entire, repand or broadly lacerate; paraphyses absent; marginales o submarginales; indusios bivalvados,
spores tetrahedral-globose, trilete, brown; x=? 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 una célula de grosor entre las venas, y esporas verdes.
low magnification the cells of its laminae are evident, Con magnificación baja las células de sus láminas son
whereas 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 especies, todas endémicas al Escudo Guyanés en
adjacent Guyana and Brazil.The species are usually epipetric Venezuela del sur y partes adyacentes de Guyana y Brasil.
on sandstone cliff faces and ledges or terrestrial at the Las especies son usualmente epipétricas sobre farallones
base of boulders from 1100–2700 m. Hymenophyllopsis de piedra arenosa o terrestres en la base de peñas desde
has recently been shown to be nested within Cyathea, 1100–2700 m. Hymenophyllopsis se ha demonstrado
and its species will eventually be transferred to that genus. ser anidado dentro de Cyathea, e sus epecies deben
Hymenophyllum + the Greek suffix, opsis, like. The plants ser transferido eventualmente a ese género para.
resemble Hymenophyllum by their small size, thin laminae, Hymenophyllum + el sufijo griego, opsis, similar. Las plantas
absence of stomata, and marginal sori. 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.

310
Figure 149. Hymenophyllopsis hymenophylloides.

311
 Hymenophyllum J.E. Sm.
DESCRIPTION: Epiphytes; rhizomes usually filiform, DESCRIPCIÓN: Epífitas; rizomas usualmente filiformos,
long-creeping, pubescent; sterile and fertile leaves typically largamente rastreros, pubescentes; hojas estériles y
1-30 cm long, monomorphic; laminae membranaceous, fértiles típicamente 1-30 cm de largo, monomorfas;
usually translucent, 1-cell thick except around the veins, láminas membranáceas, usualmente translucentes,
simple to 3-pinnate, glabrous or pubescent (the hairs una célula de grosor con excepto alrededor de las venas,
simple, forked, or stellate); veins free, anadromous, false simples a 3-pinnadas, glabras o pubescentes (los pelos
veins absent; sori marginal, round to ovate; indusium simples, furcados, o estrellados); venas libres, anádromas;
(often called an “involucre”) consisting of two soros marginales, redondos a ovados; indusio (a
lobes; receptacle usually not exerted beyond the menudo llamado “involucre”) consiste de dos
indusia; spores green, tetrahaedral-globose; x=11, 13, 18, lóbulos; receptáculos usualmente no exertados
21, 22, 28, 36. 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 extensiones raras en las zonas templadas, y consta de
300 species. All the species grow as epiphytes primarily in casi 300 especies.Todas las especies crecen como epífitas
wet, shaded forests. All of its species are epiphytes. principalmente en bosques húmedos y sombreados.
Three well-defined subgenera of Hymenophyllum Todas de sus especies son epífitas.
occur in the Neotropics: Subgen. Hymenophyllum has Tres subgéneros bien definidos de existen en el
ultimate segments conspicuously serrate and eciliate Neotrópico: Subgén. Hymenophyllum tiene segmentos
(or rarely the serrations tipped by short hairs). Subgen. últimos conspicuamente serrados y eciliados (o raras
Leptocionium has ultimate segments entire and pubescent veces con las serraciones capadas por un pelito).
(sometimes densely so). Subgen. Mecodium has ultimate Subgén. Leptocionium tiene segmentos últimos enteros
segments entire and glabrous. y pubescentes (a veces densamente). Subgén. Mecodium
The genus name is derived from the Greek hymen, tiene los últimos segmentos enteros y glabros.
membrane + phyllon, leaf. The leaves are one-cell thick. 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.
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.
2001b. rbcL data reveal two monophyletic groups of filmy ferns (Filicopsida: Hymenophyllaceae). American Journal of Botany 88:
1118–1130.

312
Figure 150. A-B. Hymenophyllum asplenioides. C, D. H. polyanthos. E–G. H. myriocarpum. H, J. H. brevistipes. (Mickel & Smith,
2004)

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TECTARIACEAE
Hypoderris R. Br. ex Hook.
DESCRIPTION: Terrestrial, occasionally on rocks; DESCRIPCIÓN: Terrestres, a veces sobre rocas;
rhizomes long-creeping with two ranks of leaves rizomas largamente rastreros, con dos hileras de
(distichous); petiole bases diverging widely from the hojas (dísticas), bases de los pecíolos divergiéndose
rhizomes (not ascending), with Eupolypods I vasculature ampliamente desde el rizoma (no ascendentes), con
pattern; laminae lobed to 2-pinnate; veins finely reticulate; el patrón vascular de Eupolypods I; láminas lobuladas
sori round, indusiate or not; spores spiny; x=40. a 2-pinnadas; venas densamente reticuladas; soros
redondeados, inusiados o no; esporas espinosas; x=40.

SIMILAR GENERA: Tectaria differs by polystichous GÉNEROS PARECIDOS: Tectaria difiere por rizomas
rhizomes, these almost always erect or decument, and polísticas, estos casi siempre erectos o decumbentes,
non-spiny spores. Triplophyllum differs by laminae deltate, y esporas no espinosas. Triplophyllum difiere por las
basal pinnae enlarged, and non-spiny spores. Bolbitis may laminas deltadas, pinnas basales agrandadas y esporas
have finely reticulate veins but it differs by rhizomes no espinosas. Bolbitis puede tener venas reticuladas pero
with several ranks of leaves, sterile-fertile leaf dimorphy, difiere por rizomas con varias hileras de hojas, dimorfismo
laminae glabrous, and acrostichoid sori. entre las hojas estériles y fertile, laminas glabras y soros
acrostichoides.

COMMENTS: Hypoderris occurs from Nicaragua hacia COMENTARIOS: Hypolepis existe desde Nicaragua
las Antillas, Venezuela y Bolivia. It consists of three species: hacia las Antillas, Venezuela y Bolivia. Consta de tres
H. brauniana (H. Christ) F. G. Wang & Christenhusz, H. species: H. brauniana (H. Christ) F. G.Wang & Christenhusz,
brownii Hook., and H. nicotianifolia (Baker) R. C. Moran, H. brownii Hook. y H. nicotianifolia (Baker) R. C. Moran,
Labiak & J. Prado. It typically occurs on wet, shaded Labiak & J. Prado.Típicamente existe en el piso de bisques
forest floors, from 200–1100 m. The genus is sister to húmedos sombreados, desde 200–1100 m. The genus is
Triplophyllum, which also has creeping rhizomes with sister to Triplophyllum, lo ccual tiene rizomas reptantes
distichous leaves. The indusium of H. brownii is unusual con hojas dísticas. El indusio de H. brownii es inusual
in that it is attached around the circumference of the porque es únido alrededor del perímetro del receptáculo
receptacle below the sporangia. In contrast, H. brauniana abajo de los esporangios. En contraste, H. brauniana tiene
has a normal reniform indusium, and H. nicotianifolia is un indusio normal reniforme y H. nicotianifolia es no
non-indusiate. The latter two species were previously indusiado. Los dos species últimas fueron previamente
classified in Tectaria. The genus name is derived from the clasificadas en Tectaria. El nombre genérico se deriva del
Greek hypo-, below + derris, covering or hide. It refers to griego hypo, abajo + derris, cubierta o piel. Se refiere a los
the indusial located beneath the sporangia. 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 00: 000–000.

314
Figure 151. Hypoderris brownii. (From Moran et al, 2014.)

315
DENNSTAEDTIACEAE
Hypolepis Bernh.
DESCRIPTION: Terrestrial; rhizomes long-creeping, DESCRIPCIÓN: Terrestre; rizomas largamente
pubescent, the hairs dark, stiff, multicellular; petiole rastreros, pubescentes, los pelos oscuros, tiesos,
bases often with horizontal branches (epipetiolar buds), multicelulares; bases de los pecíolos a menudo con ramas
containing an omega-shaped vascular bundle,; laminae horizontales (brotes epipeciolares), contienen un haz
2-pinnate to 5-pinnate, glabrous or pubescent (not scaly), vascular en forma de una omega; láminas 2-pinnadas a
sometimes long and supporting itself on the surrounding 5-pinnadas, glabras o pubescentes (no escamosas), a veces
vegetation; veins free; sori one at the apex of a vein, largas y apoyándose sobre la vegetación circundante;
marginal or submarginal, indusia formed by the reflexed venas libres; soros cada uno en el ápice de una vena,
margin of the lamina, sometimes absent; x=29, 52. marginales o submarginales,; indusios formados por el
margen reflexo de la lámina, a veces ausentes; x=29,
52.

SIMILAR GENERA: Dennstaedtia resembles Hypolepis GÉNEROS PARECIDOS: Dennstaedtia se parece a

but is distinguished by having cup-shaped, marginal sori. Hypolepis pero se distingue por tener soros marginales
Macrothelypteris can be distinguished by its acicular hairs en forma de copa. Macrothelypteris se puede distinguir
and medial (not marginal or submarginal) sori. Other por sus pelos aciculares y soros mediales (no marginales
ferns with large, finely divided laminae can usually be o submarginales). Otros helechos con láminas grandes y
separated by the form of the sori and indusia. divididas se pueden separar usualmente por la forma de
los soros y indusios.

COMMENTS: Hypolepis is pantropical and consists COMENTARIOS: Hypolepis es pantropical y consta

of about 45 species, with about 20 species in the de casi 45 especies, con alrededor de 20 especies en
Neotropics. It generally grows in open habitats such as el Neotrópico. Generalmente crece en habitats abiertos
pastures, roadsides, and forest margins. Some species have tales como pastizales, taludes y bordes de bosques.
indeterminate leaf growth, where the basal portion of Algunas especies tienen crecimiento indeterminado
the lamina matures but the apex remains as a fiddlehead, de la hoja, donde la porción basal de la lámina madura
continuously extending and producing more pairs of pero el ápice se queda como un cayado, continuamente
lateral pinnae. The genus name is derived from the Greek extendiéndose y produciendo más pares de pinas
hypo, below + lepis, scale. It refers to the scale-like false laterales. El nombre genérico se deriva del griego hypo,
indusia. abajo + lepis, escama. Se refiere a los indusíos falsos
como escamas.

LITERATURE: Brownsey, P. J. 1983. Polyploidy and aneuploidy in Hypolepis, and the evolution of the Dennstaedtiales. American
Fern Journal 73: 98–108.

316
Figure 152. A–C. Hypolepis melanochlaena. D–F. H. pulcherrima. G–J. H. diplotricha. K, N. H. eurychlaena. L–M. H. microchlae-
na. (Mickel & Smith, 2004)

317
ISOETACEAE
Isoëtes L.
Stylites Amstutz

DESCRIPTION: Plants generally less than 30 cm tall, DESCRIPCIÓN: Plantas generalmente menos de 30
tufted, evergreen aquatics or ephemeral terrestrials. cm de largo, amacolladas, siempreverdes, acuáticas o
Stems usually globose, fleshy, 2- or 3-lobed. Leaves linear, terrestres efímeras. Tallos usualmente globosos, carnosos,
subulate, terete or nearly so, one-veined, often winged 2- o 3-lobulados. Hojas lineares, subulatas, teretes o casi
toward the base, containing 4 air chambers, the air así, con una vena, con 4 cámeras para aire; esporangios
chambers septate; sporangia sunken in the leaf base, hundidos en la base de la hoja, simples, usualmente
single, usually ovoid, covered to various degrees by ovoides, cubiertos hasta varios grados por un membrano
a hyaline membrane (the velum); spores of two kinds: hialino (el velum); esporeas de dos tipos: los megasporas
megaspores tetrahedral-globose, trilete, often drying dull tetrahédricas-globosas, triletes, blanqueciñas al secarse, y
white; microspores ellipsoid, monolete, often drying tan, microesporas elipsoides, monoletes, leonadas, pardas o
brown or dark; x=11. 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. COMENTARIOS: Isoëtes tiene 200 especies mundial.
It is typically a submerged or emergent aquatic, although Es típicamente submergido o emergente, aunque
some species regularly grow in seasonally dry habitats. unas especies existen regularmente en habitats
In the Neotropics, most species grow in the páramos. estacionalmente secos. En el Neotrópico, la mayoría de
Stylites, which grows in the high Andes of Peru and las especies existen en los páramos. Stylites, lo cual existe
Bolivia, is sometimes distinguished from Isoëtes. Its stems en los Andes altos de Peru y Bolivia, a veces se distingue
are elongate, erect, and often dichotomously branched, de Isoëtes. Sus tallos son alargados, erectos, y a menudo
with roots arising from a single lateral furrow. Its leaves furcados dicotómicamente, con raíces originándose de
are flat, rigid, and bear sporangia nearly superficially un sólo surco lateral. Sus hojas son planas, rígidas y llevan
a few centimeters above the base. It is necessary to sus esporangios casi superficialmente unos centímetros
examine the megaspores with at least 30X to identify arriba de la base. Hay que examinar los megasporas con
the species. In the northern temperate zones there is a lo menos 30X para identificar las especies. En las zonas
much hybridization, but this phenomenon has not been norteñas templadas hay mucha hibridación, pero este
well studied in the tropics. Many species in the genus fenómeno no ha estado bien estudiado en los trópicos.
have CAM photosynthesis, an adaptation to oligotrophic Muchas especies del género tienen fotosíntesis CAM,
conditions. The genus name is derived from the Greek una adaptación a condiciones oligotróficas. El nombre
isos, equal + etes, year.The green leaves persit throughout genérico se deriva del griego isos, igual + etes, año. Las
the year in many species. 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.

318
Figure 153. Isoëtes. A. Habit. B. Leaf base with sporangium, the triangular structure is the ligule. C, D. Megaspores. FIGURA.
Isoëtes. A. Hábito. B. Base de la hoja con esporangio, la estructura triangular es la lígula. C, D. Megasporas. (Mickel & Smith,
2004).

319
PTERIDACEAE
Jamesonia Hook. & Grev.
DESCRIPTION: Terrestrial; rhizomes 2-4 mm wide, DESCRIPCIÓN: Terrestres; rizomas 2-4 mm de ancho,
creeping, pubescent, the hairs thick, bristle-like; leaves rastreros, pubescentes, los tricomas guesos como cerdas;
generally 11-60 x 0.2-1.5 cm, linear, erect, straight, hojas generalmente 11-60 x 0.2-1.5 cm, lineares, erectas,
stiff, 1-pinnate, the apex circinate, indeterminate; rectas, tiesas, 1-pinnadas, pubescentes, el ápice
pinnae generally less than 1 cm long, pairs 80- circinado, indeterminado; pinnas generalmente
200, rounded, sessile or nearly so, often imbricate and menos de 1 cm, pares 80-200, redondeadas, sésiles
coriaceous, the margins usually enrolled; veins free; sori o casi sésiles, generalmente imbricadas y coriáceas, los
elongate along the veins, without indusia, protected by márgenes típicamente enrrollados; venas libres; soros
the reflexed margins of the pinnae; spores tetrahaedral- a lo largo de las venas, sin indusios, protegidos por los
globose, nongreen. x=87. margenes reflexos de las pinnas; esporangios patentes a
lo largo de las nervaduras; esporas tetraédrico-globosas,
no verdes. x=87.

SIMILAR GENERA: Some species of Eriosorus have GÉNEROS PARECIDOS: Algunas especies de Eriosorus
leaves similar to those of Jamesonia, but their pinnae are tienen hojas parecidas a las de Jamesonia, pero sus pinnas
usually pinnatifid with the margins flat or nearly so (not son usualmente pinnatífidas con los márgenes planos o
reflexed), and the apex is not circinate. casi planos (no reflexos), y el ápice no es circinado.

COMMENTS: Jamesonia is completely neotropical and COMENTARIOS: Jamesonia es completamente

has 19 species. It is characteristic of páramos, usually neotropical y consta de 19 especies. Es característico de
between 3000–4000 m. The genus is polyphyletic, having los páramos, usualmente entre 3000–4000 m. El género
had four independent origins from Eriosorus (Sánchez- es polifilético y ha tendio cuatro orígenes independientes
Barracaldo, 2004). These two genera hybridize frequently a partir de Eriosorus (Sánchez-Barracaldo, 2004). Estos dos
when growing together, and the hybrids have aborted géneros, cuando crecen juntos, hibridizan con frecuencia,
spores. Eventually, the species of Jamesonia should be y los híbridos tienen esporas abortivas. Eventualmente,
combined in Eriosorus. The generic name honors William las especies de Jamesonia deben ser combinadas en
Jameson (1796–1873), Scottish botanist who collected Eriosorus. El nombre genérico honra William Jameson
plants in Quito, Ecuador. (1796–1873), botánico escocés quien colectaba plantas
en Quito, Ecuador.

LITERATURE: Sánchez-Baracaldo, P. 2004a. Phylogenetics and biogeography of the neotropical fern genera Jamesonia and
Eriosorus (Pteridaceae). American Journal of Botany 91: 274–284. Tryon, A. F. 1962. A monograph of the fern genus Jamesonia.
Contributions from the Gray Herbarium of Harvard University. 191: 109–197.

320
Figure 154. A–C. Jamesonia alstonii A. Tryon. (Mickel & Beitel, 1988)

321
DRYOPTERIDACEAE
Lastreopsis Ching
DESCRIPTION: Terrestrial; rhizomes short- to long- DESCRIPCIÓN: Terrestres; rizomas breves a largamente
creeping, scaly; leaves monomorphic; petiole with four rastreros, escamosos; hojas monomorfas; pecíolos con 4 o
or more vascular bundles; laminae 2–4-pinnate-pinnatifid, más haces vasculares; láminas 2–4-pinnadas, lanceoladas,
lanceolate, deltate, or pentagonal, the lower pinnae deltadas o pentagonales, las pinnas más inferiores
enlarged basiscopically; scaly buds absent on the agrandadas basiscópicamente; yemas escamosas
distal parts of the laminae; pinnules anadromic or ausentes en las porciones distales de las láminas;
catadromic; rachis with 2 prominent ridges adaxially, pínnulas dispuestas anadrómica o catadrómicamente;
these continuous with the thickened margins of raquis con 2 prominentes crestas adaxiales, éstas
the ultimate segments (Fig. D,E); costae raised (not continuas con los engrosados márgenes foliares de
grooved), hairy adaxially with 0.1-0.3 mm long, multicellular los segmentos terminales (Fig. D,E); costas elevadas
hairs, costae, and costules scaly abaxially, hairy adaxially; (no surcadas), pelosas adaxialmente con pelos 0.1-0.3
glandular hairs often present on the lower surface of the mm, multicelulares; tricomas glandulosos a menudo
lamina, these cylindrical, oblong appressed, bright yellow presentes en el haz de la lámina, cilíndricos, oblongos,
to orange-red; sori round. x=41. 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 indistinguible morfológicamente. Dryopteris y Arachniodes
adaxially glabrous, not finely pubescent. Megalastrum se difieren por sus ejes adaxialmente glabros, no finamente
differs by coarse, whitish, strigose hairs on the axes pubescentes. Megalastrum difiere por pelos toscos,
adaxially, prominent hydathodes, and the basal veinlet blanqueciños, strigosos sobre los ejes adaxialmente,
of the distal pinnules springing from the costa, not the hidatodos prominentes, y la venilla basal de las pinnulas
costule, with the lobe supplied by this veinlet adnate to distales surgiendo de la costa, no de la cóstula, con el
the costa. 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 in las pínnulas. La parte central de los ejes es prominulo (no
most of the Dryopteridaceae. The genus name is derived sulcados) como en la mayoría de las Dryopteridaceae. El
from Lastrea, a fern genus + opsis, like. nombre genérico se deriva 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. In review. Phylogeny
and historical biogeography of the lastreopsid ferns (Dryopteridaceae). American Journal of Botany [submitted 13 February
2014] 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.

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Figure 155. A–D, H: Parapolystichum effusa. F, G: L. exculta. (Mickel & Smith, 2004)

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POLYPODIACEAE
Lellingeria A. R. Smith, R. C. Moran, & L. E. Bishop
DESCRIPTION: Plants epiphytic or epipetic; rhizome DESCRIPCIÓN: Plantas epífitas o epipéticas; rizomas
short- to long-creeping, ascending, or erect, the scales cortamente reptantes, ascendentes o rectos, las escamas
clathrate, usually blackish, glabrous or provided with clatradas, usualmente negruzcas, glabras o provistas de
hyaline marginal setulae, attached across the entire width sétulas marginales hialinas, unidas por el ancho de la base;
of the base; phyllopodia absent; petioles absent or shorter filopodios ausentes; pecíolos ausentes o más cortos que
than the lamina, continuous with (not articulate to) the la lámina, contínuos (no articulados) al rizoma; láminas
rhizome; laminae pinnatifid to 1-pinnate-pinnatifid, lacking de pinnatífida a 1-pinnada-pinnatífida, setas rojizas
reddish setae, or pubescent (at least along the petiole ausentes, o pubescente (al menos a lo largo del pecíolo
and rachis), the hairs usually with a pale basal cell and y raquis), los tricomas usualmente con una célula
forked unequally, the smaller cell oblique, glandular, basal pálida y furcada desigualmente, la célula
the larger cell acicular; hydathodes present; veins más pequeña oblicua, la célula más larga acicular;
simple, free, several per segment; sori round or elliptic, hidatodos presentes; venas simples, libres, varias por
often somewhat sunken, lacking paraphyses; sporangial segmento; soros redondos o elípticos, a menudo levemente
capsules glabrous or (in 3 species) setose; x=37. hundidos, paráfisis ausentes; cápsulas de los esporangios
glabras o (en 3 especies) setosas; x=37.

SIMILAR GENERA: Cochlidium differs by tan, GÉNEROS PARECIDOS: Cochlidium difiere por
nonclathrate scales and lack of forked hairs. Grammitis escamas leonadas, no clatradas y la ausencia de tricomas
differs by entire laminae with black borders. Stenogrammitis furcados. Grammitis difiere por laminas enteras con
differs by narrow leaves (< 5 mm) and only one sorus and bordes negras. Stenogrammitis difiere por hojas estrechas
vein per segment. Other neotropical grammitids genera (< 5 mm) y solo un soro por segmento. Otros géneros
differ by the presence of dark reddish setae on the leaves. gramitidoides neotropicales difieren por la presencia de
setas oscuras rojizas en las hojas.

COMMENTS: Lellingeria contains about 50 species and COMENTARIOS: Lellingeria consta de casi 50 especies
is entirely neotropical. Most of the species are epiphytes y es completamente neotropical. La mayoría de las
in cloud forests, from 1000–4000 m. The genus is especies son epifitas en bosques nublados, desde 1000–
most diverse in the Andes. Lellingeria is one of the few 4000 m. El género es más diverso en los Andes. Lellingeria
grammitids that lack reddish setae on the leaves. Its hairs es uno de los pocos helechos grammitidoides que faltan
are distinctive, being often forked unequally—a character setas rojizas en las hojas. Sus pelos son distintivos, siendo
it shares with Stenogrammitis (see illustration of that a menudo furcados desigualmente—una característica
genus). The genus is named for David B. Lellinger (1937– que comparta con Stenogrammitis (vea la ilustración de
), American pteridologist who worked at the Smithsonian ese género). El género se nombra para David B. Lellinger
Institution. (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. 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.

324
Figure 156. 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. (Courtesy of Paulo Labiak, 2009)

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POLYPODIACEAE
Leucotrichum Labiak
DESCRIPTION: Plants epiphytic, rarely epipetric; roots DESCRIPCIÓN: Plantas epífitas, raras veces epipetricas;
not proliferous; rhizomes erect, radial, scaly, the scales raíces no prolíferas; rizomas erectos, radiales, escamosos,
clathrate, dark brown to blackish, basifixed, margins ciliate; las escamas clatradas, pardo oscuros a negruzcos;
petioles absent to ca. 1 cm long, pubescent, the hairs up basifijadas, márgenes ciliados; pecíolos ausentes hasta
to 2 mm long, whitish, simple or 1–furcate; laminae erect 1 cm de largo, pubescentes, los pelos hasta 2 mm de
to slightly arcuate, determinate, linear to linear-oblong, longitud, blanquecinos, simples o 1-furcados; láminas
pinnatifid to pinnatisect, with reduced basal segments, erectas a levemente arcuadas, determinadas, lineares
the lowermost decurrent and forming short and broad a linear-oblongas, pinnatífidas a pinnatisectas, con
wings, apices obtuse, subconforme to the lateral segments; segmentos basales reducidos, los más basales decurrentes
rachis with dark sclerenchyma, the sclerenchyma not y formando alas cortas y anchas, ápices obtusos,
visible on the abaxial side, often obscured by the laminar subconformes a los segmentos laterales; ráquises con
tissue adaxially, setose and pubescent, the setae whitish, esclerenquima oscura no visible abaxialmente, a menudo
up to 3 mm long, the hairs simple to 1-furcate; segments oculta por el tejido laminar adaxialmente, setosos y
1–3 × 0.5–1.5 mm, entire, symmetric or asymmetric at pubescentes, las setas blanquecinas, hasta 3 mm de
base, setose and pubescent, the setae whitish, ca. 1–3 longitud, los pelos simples o 1-furcados; segmentos 1–3
mm long, the hairs 0.2–1.0 mm long, 1(2)-furcate; veins × 0.5–1.5 mm, enteros, simétricos o asimétricos en la
1-furcate, obscured by the laminar tissue; hydathodes base, setosos y pubescentes, las setas blanquecinas, ca.
present, lacking whitish deposits; sori one per segment, 1–3 mm de longitud, los pelos 0.2–1.0 mm de longitud,
inframedial, rounded to elongate, superficial, surrounded 1(2)-furcados; venas 1-furcadas, ocultas por el tejido
by the hairs of the laminae; sporangial capsules glabrous or laminar; hydatodos presentes, faltando de depósitos
ciliate; spores trilete, green. x=?. 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 GÉNEROS PARECIDOS: Alansmia difiere por las
rhizome scales, indeterminate leaf growth, pinnate veins, escamas de los rizomas no clatradas, crecimiento
and branched setae. Lellingeria differs by the absence of indeterminado de las hojas, venas pinnadas y setas
setae and whitish hairs, and the presence of pinnate veins. ramificadas. Lellingeria difiere por la ausencia de setas
Stenogrammitis differs by eciliate rhizome scales, absence y pelos blanquecinos, y la presencia de venas pinnadas.
of whitish hairs and setae on the leaves, and one vein per Stenogrammitis difiere por escamas del rizoma no ciliadas,
segment. 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.

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Figure 157. 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. pseudomitchel-
lae (Mickel 3033, NY). Habit. N–P. L. mitchellae (Ekman 14884, K). N. Habit. O. Rhizome scale. P. Sporangium. (Courtesy of Paulo Labiak, 2010)

327
LINDSAEACEAE
Lindsaea Dryander ex J.E. Sm.
DESCRIPTION: Terrestrial, rarely rupestral or epiphytic; DESCRIPCIÓN: Terrestres, ráramente rupestres o
rhizome short- to long-creeping, protoselic, scaly, the scales epífitas; rizoma corto o largamente rastrero, protostélico,
lanceolate to narrow and bristle-like; laminae anadromic, escamoso, las escamas lanceoladas a angostas y como
1-pinnate to 2(-4)-pinnate, when 2-pinnate the apex una cerda; lámina anádroma, 1-pinnada a 2(-4)-pinnada,
(in the neotropics) a conform terminal pinna; ultimate cuando 2-pinnada, el ápice (en el neotrópico) consiste
segments usually dimidiate, with the costule along de una pinna terminal conforme; últimos segmentos
the basiscopic margin, the margins entire; rachises and usualmente dimidiados, con la costilla media a lo largo
costae grooved or sharply angled adaxially, usually del margen basiscópico, los margenes enteros; ráquises
greenish or stramineous; veins free or anastomosing; sori y costas surcados o angulados adaxialmente,
following the margin of the segments, the indusia usualmente verdosos o pajizos; nervaduras libres o
opening toward the margin; annulus of 7-15 cells; anastomosadas; soros siguiendo el margen de los
spores monolete or trilete, 16 or 32 per sporangium; x = segmentos, los indusios abriéndose hacia el margen;
34, 41, 47, 50. 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 COMENTARIOS: Lindsaea consta de casi 50 especies
is pantropical. About 50 species occur in the Neotropics, y es pantropical. Alrededor de 50 especies existen en
mostly in the Guayanas. Lindsaea has an unusual type of el Neotrópico, mayormente en las Guayanas. Lindsaea
protostele because the phloem is surrounded by xylem. It tiene un protostelo raro porque el floema está rodeado
is closely related to Odontosoria. The genus name honors de xilema. Es cercanamente relacionado a Odontosoria.
John Lindsay, a British physician who lived in Jamaica during El nombre genérico honora a John Lindsay, un médico
the late 1700s and made observations on the germination británico quien vivó en Jamaica en las postrimerías de los
of fern spores and formation of gametophytes. 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.

328
Figure 158. Top: Lindsaea portoricensis. G–K: L. quadrangularis subsp. antillensis. L–O: L. lancea. P-R: L. arcuata. S–U: L. stricta.

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PTERIDACEAE
Llavea Lag.
DESCRIPTION: Terrestrial or rupestral; rhizomes short- DESCRIPCIÓN: Terrestres o rupícolas; rizoma erecto
creeping or erect, the apex with lanceolate, stramineous a cortamente rastrero, el ápice con escamas lanceoladas,
to tan scales, sometimes with a black central portion pajizas, leonadas, a veces con una porción central negra
(especially on petiolar scales); leaves hemidimorphic, (especialmente en las escamas del pecíolo); hojas
with the apical pinnae fertile and contracted; laminae hemidimorfas, con las pinnas apicales fértiles y
2–3-pinnate; pinnules oblong to elliptic; veins free; indusia contraídas; pecíolos teretes a subcuadrangulars; láminas
formed by the revolute margin of the pinnule, with short 2–3-pinnadas; pínnulas oblongas a elípticas; nervaduras
glandular hairs on the inner surface; sporangia along the libres; indusios formado por el margen del segmento
veins but confluent at maturity; spores trilete, tetrahedral; reflexo, con pelitos glandulares en la superficie interior;
x=29. 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 COMENTARIOS: Llavea occurs en Texas, México y
Guatemala where it grows on rocky wooded slopes from Guatemala donde crece en taludes rocosos bosqueados
1200–2500 m. It contains only one species: L. cordifolia desde 1200–2500 m. Contiene una sola especie: L. cordifolia
Lag., distinctive by its hemidimorphic leaves. Within the Lag., distinctiva por sus hojas hemidimorfas.Dentro de
Pteridaceae, Llavea forms a clade with Conigramme (Asian) las Pteridaceae, Llavea forma un clado con Coniogamme
and Cryptogramma. The genus name honors Pablo de la (Asiático) Cryptogramma. El nombre genérico honora a
Llave (1773–1833), a traveler in Mexico. 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]

330
Figure 159. A-D. Llavea cordifolia Lag. (Mickel & Smith, 2004).

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LOMARIOPSIDACEAE
Lomariopsis Fée
DESCRIPTION: Hempipiphytes; rhizomes long- DESCRIPCIÓN: Hemiepífitas; rizomas largamente
creeping, in cross section with a broad ventral meristele, rastreros, en sección transversal con un haz vascular
roots borne only on the ventral surfaces; sterile and ancho, raíces se nacen solo en las superficies ventrales;
fertile leaves strongly dimorphic; petioles not pecíolos no articulados al rizoma, con 4-8 haces vasculares;
articulate to the rhizome, with 4-8 vascular bundles; hojas estériles y fértiles fuertemente dimorfas; láminas
laminae 1-pinnate (simple in juveniles of some species), 1-pinnadas (o simples en los juveniles de algunas
with a conform apical pinna; pinnae entire or nearly especies), con una pinna apical conforme; pinnas
so, articulate to the rachis; veins free, simple or with one enteras o casi enteras, articuladas al raquis; nervaduras
fork, parallel; fertile pinnae narrower than the sterile; sori libres, simples o una vez furcadas, paralelas; pinnas
acrostichoid; spores monolete, green; x=41. fértiles más angostas que las estériles; soros acrosticoides;
esporas monoletes, verdes; x=41.

SIMILAR GENERA: Polybotrya differs by pinnatifid leaf GÉNEROS PARECIDOS: Polybotrya difiere por ápices
apices (in all but 1 species) and pinnate groups of veins. de las hojas pinnatífidos (con excepción de una especie)
Also, most species of Polybotrya have decompound leaves. y venas en grupos pinnados. También, la mayoría de
Lomagramma and the climbing species of Bolbitis differ las especies de Polybotrya tienen hojas decompuestas.
by having anastomosing veins. Juvenile plants of certain Lommagramma y las especies trepadoras de Bolbitis
species with simple leaves (the Lomariopsis japurensis difieren por venas anastomosadas. Plantas juveniles
group) resemble Elaphoglossum but can be distinguished de ciertas especies con hojas simples se asemejan
by their cartilaginous-thickened margins. a Elaphoglossum pero se pueden distinguir por sus
márgenes engrosados cartilaginosos.

COMMENTS: Lomariopsis is a genus of wet forests. It COMENTARIOS: Lomariopsis es un género de

is pantropical with about 45 species worldwide and 15 bosques húmedos. Es pantropical con casi 45 especies
in the Neotropics. The sporophytes start growth on the mundialmente y 15 en el Neotrópico. Los esporofitos
base of trunks and climb after establishing contact by long empiezan desarrollo en la base de los troncos y suben
thick roots with the soil. The climbing rhizomes adhere después de establecer contacto por raíces largas y
to the substrate by short lateral roots produced on the gruesas con el suelo. Los rizomas trepadores adhieren
ventral surface. In transverse section the rhizome has a al sustrato por raíces cortos laterales producidas en la
characteristic cat’s paw pattern to the vascular bundles. superficie ventral. En sección transversal el rizoma tiene
The ventral bundle, which bears all the roots, is notched. un patrón característico a las haces vasculares como pata
The articulations of the pinnae can be seen with a de gato. La haz ventral, que da lugar a todas las raíces,
handlens as faint dark lines at juncture with the rachis.The tiene una escotadura. Las articulaciones de las pinnas se
articulations seem non-functional because the pinnae do pueden ver con una lupa como líneas oscuras en la unión
not fall away with age or upon drying. Some evidence, con el raquis. La articulaciones aparecen no funcionales
however, indicates they might function in response to porque las pinnas no se desprenden con edad o al secar.
damage. The genus name is derived from Lomaria, a fern No obstante hay evidence que funcionen cuando la
genus name + opsis, a Greek suffix meaning “like.” pinna se daña. El nombre genérico se deriva de Lomaria,
un nombre de un género de helecho + oasis, un sufijo
griego que significa similitud.

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.

332
Figure 160. Lomariopsis prieuriana. (©R. C. Moran, 2009).

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LONCHITIDACEAE
Lonchitis L.
DESCRIPTION: Terrestrial or occasionally epipetric; DESCRIPCIÓN: Terrestres u ocasionalmente rupícolas;
rhizomes 5–15 mm wide, short-creeping, hairy; leaves rizoma 5–15 mm de ancho, cortamente rastrero, peloso;
0.5–2.5 m, monomorphic; petiole bases with 2 hojas 0.5–2.5 m, monomorfas; bases de los pecíolos
vascular bundles; laminae 2- to 3-pinnate-pinnatifid, con 2 haces vasculares; láminas 2- a 3-pinnado-
slightly succulent, softly hairy, the hairs numerous, pinnatífidas, suculentas, suavemente pelosas, los
septate; pinnules catadromic throughout, but sometimes pelos numerosos, septados; pínnulas catádromas en
anadromic on the basal pinnae; rachises grooved adaxially, toda la pinna o anádromas en la base y catádromas
the groove hairy; costae prominent adaxially; veins free, distalmente; raquis sulcado adaxialmente, el surco peloso;
ending before the margins in clavate hydathodes; sori costas prominentes adaxialmente; nervaduras libres,
marginal, borne along a connecting vein parallel to the terminando antes de los márgenes en ápices claviformes;
margin and connecting (1)2 or 3 vein tips; indusia formed soros marginales, dispuestos en una comisura vascular
by the revolute scarious margin; spores trilete; x=50. 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 COMENTARIOS: Lonchitis consta de dos especies:
L. in the Neotropics and L. occidentalis Baker in Africa- L. hirsuta L. en el Neotrópo y L. occidentale Baker en
Madagascar. Lonchitis hirsuta typically grows on wet, África-Madagasscar. Lonchitis hirsuta existe en típicamente
shaded forest floors from 0–1200 m. The Old World en los pisos mojados sombreados de los bosques de
species differs by net veins along the midribs, but this 1–1200 m. La especie del Viejo Mundo difiere por venas
characteristic is not constant. The petioles contain anastomosadas por las costas, pero esta característica
raphides, probably of calcium oxalate. These are easily no es constante. Los pecíolos contienen rafidios,
viewed in free-hand sections of the petioles. The genus probablemente de oxalato de calcio. Estos se observen
name is derived from the Greek lonche, lance + itis, a fácilmente en secciones transversales del pecíolo. El
suffix indicating a close connection with. An ancient fern nombre genérico se deriva de la Griega lonche, lanza +
name used by Dioscorides. 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.

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Figure 161. A–C. Lonchitis hirsuta. (Mickel & Smith, 2004)

335
DICKSONIACEAE
Lophosoria C. Presl
DESCRIPTION: Terrestrial; stems massive, erect DESCRIPCIÓN: Terrestres; tallos masivos, erectos o
or decumbent, rarely arborescent (up to 1 m tall), decumbentes, raras veces arborescentes (hasta 1 m de
stoloniferous, densely pubescent, the hairs 2-4 cm altura), estoloníferos, densamente pubescentes, pelos
long, golden to dark brown, wooly; leaves 2-5 m long, 2-4 cm de largo, dorados a pardo oscuros, lanosos; hojas
monomorphous; petiole base densely pubescent like 2-5 m de largo, monomorfas; base del pecíolo densamente
the stem; laminae 3-pinnate to 4-pinnate, whitish pubescente como el tallo; láminas 3-pinnadas a
beneath, triangular or ovate; axes pubescent beneath; 4-pinnadas, blanquecinas en el envés, triangulares u
veins free; sori superficial (not marginal), without ovadas; ejes pubescentes en el envés; nervadura libre;
indusia, with long tangled paraphyses; the receptacle soros superficiales (no marginales), sin indusios, con
flat (not elevated); spores tetrahedral, with prominent paráfises largos, enmarañados; receptáculo plano (no
equatorial ridge; x = 65. 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 neotrópical y existe principalmente desde 2000-3600
three species: L. contracta (Hieron.) A. R. Sm. (Ecuador m. Contiene dos especies: L. contracta (Hieron.) A. R.
and Peru), L. quadripinnata (J. F. Gmelin) C. Chr., and L. Sm. (Ecuador y Perú), L. quadripinnata (J. F. Gmelin) C.
quesadae A. Rojas (Costa Rica). Only L. quadripinnata is Chr. y L. quesadae A. Rojas (Costa Rica). Solamente L.
widespread.The genus is often conspicuous because of quadripinnata es muy difundida. El género a menudo
its large size and white undersurface of the leaves. All es conspicua a causa de su tamaño grande y el envés
species of the genus tend to form colonies by stolons de la lámina blanquecino. Todos las especies del género
(Cox & Tomlinson, 1985). Molecular data show Lophosoria tienden formar colonias por estolones (Cox & Tomlinson,
is sister to Dicksonia (Korall et al. 2006, 2007). Lophosoria 1985). Datos moleculares muestran Lophosoria hermana
is derived from the Greek lophos, mane or tuft + soros. It a Dicksonia (Korall et al. 2006, 2007). Lophosoria se deriva
refers to the hairs between the sporangia. 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.

336
Figure 162. A–C. Lophosoria quadripinnata. (Mickel & Smith, 2006)

337
LOXOGRAMMACEAE
Loxogramme (Blume) C. Presl
DESCRIPTION: Epiphytic or saxicolous; rhizomes DESCRIPCIÓN: Epífitas o saxícoloas; rizomas corto
short- to long-creeping; roots forming a spongy mass, a largamente rastreros; raíces formando una masa
with dense, opaque, dull-brown hairs; rhizome scales esponjosa, con pelitos densos, opacos, pardos; escamas
clathrate, concolorous, entire, united at the base (not del rizoma clatradas, concoloras, enteras, unidas en la
peltate); plants no sclerenchymatous except for the roots; base (no peltadas); plantas no esclerenquimatosas excepto
sterile and fertile leaves monomorphous or (rarely) las raíces; hojas estériles y fértiles monomorfas o (raras
dimorphous; filopodia present or absent; laminae simple, veces) dimorfas; filopodios presentes o ausentes; láminas
entire, typically linear or narrowly elliptic, glabrous except simples, enteros, típicamente lineares o estrechamente
for minute glandular or clavate hairs; veins anastomosing elípticas, glabras excepto por diminutos pelitos
with many, few or no included free veinlets; hydathodes claviformes a glandulares; nervaduras anastomosadas,
absent; sori usually elongate, oblique or rarely con muchos, pocos o ningún nérvulo libre incluido;
parallel to the costa; indusia absent; paraphyses filiform hidátodos ausentes; soros generalmente alargados,
or absent; spores green, trilete or monolete. x=35, 36. oblicuos o raramente paralelos a la costa; indusios
ausentes; parafisos filiformes o ausentes; esporas verdes,
triletes o monoletes. x=35, 36.

SIMILAR GENERA: Polytaenium differs by non-green GÉNEROS PARECIDOS: Polytaenium difiere por
spores and more than one soral line between the costa esporas no-verdes y más de una línea soral entre la
and margin. Pleopeltis differs by peltate scales on the costa y margen. Pleopeltis difiere por escamas peltadas
blades and (usually) among the sporangia. Microgramma en las láminas y (usualmente) entre los esporangios.
differs yellow spores and (at least in some species) scales Microgramma difiere por esporas verdes y (a lo menos
on the laminae. en algunas especies) escamas en las láminas.

COMMENTS: Loxogramme consists of 33 species, all COMENTARIOS: Loxogramme consta de 33 especies,

of which are paleotropical except L. mexicana (Fée) C. todas paleotropicales excepto L. mexicana (Fée) C. Chr.,
Chr., which occurs from Mexico to Panama. This sole que existe desde México hasta Panamá. Esta única
neotropical species grows in montane forests from especie neotropical crece en bosques montanos de 600–
600–2400 m, usually as a low-trunk epiphyte or on 2400 m, usualmente como epífito en la parte inferior de
mossy rocks along streams. Under dry conditions the los troncos o sobre rocas musgosas por ríochuelos. Bajo
plants are flaccid and limp, but they revive rapidly when condiciones secas las plantas están flácidas y marchitadas,
wetted. Schneider et al. (2004) found that Loxogramme pero recuperan rápidamente cuando mojado. Schneider
forms a clade with the paleotropical genera Anarthropteris et al. (2004) encontraron que Loxogramme forma un
(perhaps not distinct from Loxogramme) and Dictymia. clado con los géneros paleotropicales Anarthropteirs (tal
This clade, recognized here as the Loxogrammaceae, is vez no distincto de Loxogramme) y Dictymia. Este clado,
sister to the Polypodiaceae. Moran and Smith (2001) reconocido aquí como Loxogrammaceae, es hermana a
postulated on the basis of morphology that Loxogramme la Polypodiaceae. Moran y Smith (2001) postularon en
mexicana and L. abyssinica (Baker) M. G. Price were sister la basis de morfología que Loxogramme mexicana and L.
species. This was supported by molecular data of Janssen abyssinica (Baker) M. G. Price fueron especies hermanas.
and Schneider (2007). The genus name is derived from Esto fue apoyado por los datos moleculares de Janssen y
the Greek loxos, oblique + gramme, line, referring to the Schneider (2007). El nombre del género es derivado del
sori that are oblique to the costa. griego loxos, oblicuo + gramme, línea, referiéndose a los
soros que son oblicuos a la 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.

338
Figure 163. Loxogramme mexicana. A. Habit. B. Rhizome. C. Rhizome scale. D. Blade detail. E. Cross section of lamina.
(Mickel & Smith, 2004).

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LOXOMATACEAE
Loxsomopsis H. Christ
DESCRIPTION: Terrestrial; rhizomes creeping, DESCRIPCIÓN: Terrestres; rizomas reptantes,
branched, covered with abundant, rigid, dark ramificados, revestidos con raíces abundantes cerdas
brown, shiny bristles; leaves monomorphic, spaced pardo oscuro, rígidas, lustrosas; hojas monomorfas,
3-10 cm apart; petioles bases with 2 vascular bundles; espaciadas a lo largo del rizoma 3-10.3 cm; pecíolos con
laminae 2-pinnate-pinnatifid to nearly 3-pinnate; 2 haces vasculares en la base; láminas 2-pinnado-
petiole dark to atropurpureious, sparsely bristly toward pinnatífidas a casi 3-pinnadas; pecíolo ligeramente
the base; laminae lanceolate to oblong subdeltate, sulcado, pardo oscuro a atropurpúreo, con cerdas hacia
glabrous above, glabrous to hairy below; veins free, la parte basal; lámina lanceolada a oblonga subdeltada,
extending all the way to the margin; pinnae asymmentical; glabra en el haz, glabra a pelosa en el envés; nervaduras
sori marginal, paraphysate; indusia narrowly cup-shaped libres, bifurcadas, que se extienden a los márgenes;
or uceolate, entire, the receptacle exert; sporangia with pinnas asimétricas; soros marginales, parafisados;
an oblique annulus not interrupted at the stalk; spores indusio estrechamente ciatiforme a urceolado, enteros,
trilete. x=46. el receptáculo exerto; esporangios con anillo oblicuo no
interrumpido por el pedículo; esporas triletes. x=46.

SIMILAR GENERA: Trichomanes differs by GÉNEROS PARECIDOS: Trichomanes difiere por

membranaceous laminae and non-paraphysate sori. laminas membranáceas y soros parfisados. Dennsteadtia
Dennsteadtia usually differs by epipetiolar buds and a difiere usualmente por brotes epipeciolares e un indusio
shallower cup-shaped indusium. Saccoloma has a compact, de forma de copa menos profundo. Saccoloma tiene un
decumbent or erect rhizome. rizoma compacto o erecto.

COMMENTS: Loxomopsis is completely neotropical, COMENTARIOS: Loxsomopsis es completamente

from Costa Rica to Bolivia at 1500–3000 m. It consists of neotropical, desde Costa Rica hasta Bolivia, de 1500–
a single species: L. pearcei (Baker) Maxon. The plants are 3000 m. Consta de una sola especie: L. pearcei (Baker)
terrestrial and occur in humid, semi-open places, especially Maxon. Las plantas son terrestres y existen un lugares
on slopes and roadsides in cloud forests. Loxomopsis is húmedos semiabiertos, especialmente por los taludes y
a member of the tree fern clade, Cyatheales. Only one lados de caminos en bosques nublados. Loxomopsis es
other genus occurs in the family: Loxsoma, endemic to miembro del clado de los helechos arborescentes, las
North Island, New Zealand. The genus name is derived Cyatheales. Solamente otro género existe en la familia:
from the Greek Loxsoma, the name of another fern genus, Loxoma, endémico a la Isla del Norte, Nueva Zelandia.
+ -opsis, like. El nombre del género se deriva del griego Loxsoma, el
nombre de otro género de helecho, + -opsis, parecido.

LITERATURE: Lehnert, M., M. Monnich, T. Pleines, A. Schmidt-Lebuhn & M. Kessler. 2001. The relictual fern genus
Loxsomopsis. American Fern Journal 91: 13–24.

340
Figure 164. Loxomopsis pearcei (© Robbin Moran, 2007).

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POLYPODIACEAE
Luisma M. T. Murillo & A. R. Sm.
DESCRIPTION: Epiphytic; rhizomes radially symmetrical, DESCRIPCIÓN: Epífitas; rizomas simétricos radialmente,
erect or suberect, scaly, the scales 2.2 X 0.3 mm, clathrate, erectos o suberectos, escamosos, las escamas 2.2 X 0.3
ciliate, lanceolate, dark gray, shiny, peltate; petioles 1.6– mm, clatradas, ciliatas, lanceolatas, grisiacas, lustrosas,
2.5 cm long, terete, with 2 vascular bundles in cross peltatas; pecíolos 1.6–2.5 cm de largo, teretes, con dos
section; leaves 9–20 0.3–0.4 cm, simple, entire, narrowly haces vasculares en sección transversal; hojas 9–20 X
elliptic, glabrous on both surfaces, with green borders 0.3–0.4 cm, simples, enteras, angostamente elípticas,
(not blackish); hydathodes prominent; veins free, forked; glabras en ambas superficies, con bordes verdes (no
sori ca. 1 mm wide, round, adjacent to (touching) the negruzcos); hidátodos prominentes; nervaduras libres,
midrib; paraphyses absent; spores green, trilete; x=? furcadas; soros ca. 1 mm de ancho, redondos, adyacente
(tocando) a la costa; paráfisis ausentes; esporas verdes,
triletes; x=?

SIMILAR GENERA: Campyloneurum differs by GÉNEROS PARECIDOS: Campyloneurum difiere por

anastomosing veins. Grammitis differs by dark laminar venas anastomosadas. Grammitis difiere por un margen
margins and entire rhizome scales. Cochlidium differs by laminar oscuro y escamas del rizoma enteras. Cochlidium
non-clathrate, entire rhizome scales. difiere por escamas del rizoma no clatradas, enteras.

COMMENTS: Luisma consists of one species, L. COMENTARIOS: Luisma consta de una sola especie,
bivascularis M. T. Murillo & A. R. Sm., endemic to the L. bivascularis M. T. Murillo & A. R. Sm., endémica al
department of Risaralda, Colombia. It grows in wet forests departmento de Risaralda, Colombia. Crece e bosques
at 1550 m. It was claimed that Luisma was the only genus húmedos alrededor de 1550 m. Se alejó que Luisma fue
of grammitids with two vascular bundles in the petiole, el único género grammitidoide con dos haces vasculares
but this character is more common among grammitids en el pecíolo, pero esta condición es más común entre
ferns than previously believed. Like Lellingeria, it has dark estos helechos que creido previamente (M. Sundue, pers.
clathrate rhizome scales, hydathodes, and lacks reddish com.). Como Lellingeria, tiene escamas del rizoma oscuras
setae on the leaves. It probably represents a simple- clatradas, hidatodos y falta de setas rojizos en las hojas.
leaved species of Lellingeria, but molecular are needed to Probablemente representa una especie de Lellingeria con
test this idea. The genus is named in honor of Luis María hojas simples, pero se necesitan datos moleculares para
Murillo, pioneer Colombian entomologist. probar esta idea. El género se nombra en honor de Luis
María Murillo, pionero entomólogo colombiano.

LITERATURE: Murillo, M. T. & A. R. Smith. 2003. Luisma, a new genus of Grammitidaceae (Pteridophyta) from Colombia.
Novon 13: 313–317.

342
Figure 165. Luisma bivascularis (from Murillo & Smith, 2003).

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LYCOPODIACEAE
Lycopodiella Holub
DESCRIPTION: Terrestrial or rarely epiphytic or DESCRIPCIÓN: Terrestres o raramente epífitas
rupestral; stems of two kinds: 1) horizontal and prostrate o rupícolas; tallos de dos tipos: 1) horizontales y
or arching, non-strobiliferous, and 2), erect and postrados o arqueados, no estrobilíferos, y 2) erectos
strobilifeorus, these produced in a single row from y strobolíferos, estos producidos en una sola hilera
the dorsal side of the horizontal stems; branchlets del lado dorsal de los tallos horizontales; ramitas
isophyilous or anisophyllous; sporophylls turning yellowish isofilas o anisofilas; esporofilas volviéndose amarillentazas
after sporangial dehiscence, subpeltate, with a medial, después de dehiscencia esporangial, subpeltadas, con
basiscopic wing (sect. Lycopodiella y sect. Carolinianae) un ala media basiscópica (sect. Lycopodiella y sect.
or with the bases of the sporophylls coalescent and Carolinianae) o con las bases de esporofilas coalescentes
almost enclosing the sporangia (sect. Campylostachys); casi envolviendo a los esporangios (sect. Campylostachys);
sporangia at the base of the sporophyll or axillary (sect. esporangios sobre la base de la esporofila o axilares (sect.
Lycopodiella), distinctly anisovalvate or equally valvate Lycopodiella), marcadamente anisovalvados o isovalvados
(sect. Carolinianae); epidermal cells of the sporangium (sect. Carolinianae); células del epidermis del esporangio
with thin straight, non-lignified walls, with nodular or con paredes delgadas, rectas, no lignificadas, con
semi-ringlike thickenings; spores rugose; gametophytes engrosamientos nodulosos o semianulares, lignificados;
green, superficial; x=35, 39, 78, 104. esporas rugosas; gametofitos verdes, epigeos; x=35, 39,
78, 104.

SIMILAR GENERA: Huperzia differs by equally forked GÉNEROS PARECIDOS: Huperzia difiere por tallos
stems, sporophylls that remain green after sporangial furcados igualmente y esporofilos que permanezcan
dehiscence, and subterranean gametophytes. Lycopodium verdes después de dehiscencia esporangial y gametofitos
differs by producing two lateral rows of branches from subterráneos. Lycopodium difiere por producir una
the dorso-lateral surface of the creeping stem (versus hilera de tallos erectos de la superficie dorso-lateral
from the dorsal side in Lycopodiella), and subterranean del tallo rastrero principal (versus desde el lado dorsal
gametophytes. Its strobili are generally stalked, not sessile. en Lycopodiella) y por gametofitos subterráneos. Sus
estróbilos usualmente son pedicelados, no sésiles.

COMMENTS: Lycopodiella is cosmopolitan and has COMENTARIOS: Lycopodiella es cosmopolita y consta

about 40 species, mostly neotropical. Most of the de casi 40 especies, principalmente neotropicales. La
neotropical species were illustrated and described by mayoría de las especies neotropicales fueron ilustradas
Øllgaard (1988). The genus occurs in a wide variety of y descritas por Øllgaard (1988). El género existe un
habitats but tends to be especially common in open una variedad amplia de ambientes pero tiende ser
disturbed places such as road banks, exposed slopes, and especialmente común en taludes, laderas expuestas, y
open sandy soils. Lycopodiella cernua (L.) Pic. Serm. is a suelos arenosos abiertos. Lycopodiella cernua (L.) Pic. Serm.
common widespread species throughout the tropics of es común y difundida en el trópico por todo mundo. Sus
the world. Its erect branching systems are treelike, with sistemas erectas de ramificación son como árboles, con
nodding sessile strobili at the branchlet tips. The genus estróbilos péndulos en los ápices de las ramitas. A veces
is sometimes further divided into smaller monophyletic el género se divide en unos más pequeños monofiléticos:
ones: Palhinaea and Pseudolycopodiella. The genus name is Palhinaea y Pseudolycopodiella. El nombre genérico es el
the diminutive of Lycopodium. diminutivo de Lycoodium.

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. Biol. Skr. 34:
1–135.

344
Figure 166. Lycopodiella alopecuroides, high-altitude form (Holm-Nielsen et al. 4839, AAU); A1, vegetative leaf; A2-3, sporo-
phylls. B, Lycopodiella alopecuroides, mid-altitude form (Asplund 19353, S); B1, portion of erect branch; B2, sporophyll. C, Lycop-
odiella riofrioi, composite: C1 (Holm-Nielsen et al. 4103, AAU), basal branchings; C2-4 (Harling & Andersson 12811, AAU); C2,
erect branch system; C3, branchlet leaf; C4, sporophyll. . (from Øllgaard (1988); courtesy of Benjamin Øllgaard).

345
LYCOPODIACEAE
Lycopodium L.
DESCRIPTION: Terrestrial; horizontal stems branched DESCRIPCIÓN: Terrestres; tallos horizontales
unequally, producing two dorso-lateral rows of branch ramificados desigualmente, produciendo dos hileras
systems, these branched repeatedly and generally dorso-laterales de sistemas de ramitas ramificadas
determinate, ascending or erect; strolibi present, repetidamente y generalmente determinadas,
erect, simple or bifurcate, sessile on simple stalks or ascendentes a erectos; estróbilos presentes, erectos,
bifurcate; sporophylls and vegetative leaves similar, or the simples o bifurcados, sésiles o sobre pedúnculos simples
sporophylls gradually to abruptly reduced, peltate or with o bifurcados; esporofilos se tornan pajizos despues
a thin basal wing decurrent on the stalk; straw-colored de dehiscencia esporangial, peltados, o subpeltados
after sporangial dehiscence; sporangia united to the con una delgada ala basal decurrente en el pedicelo;
base of the sporophyll; spores reticulate. x=23, 31, 34, c. esporangios unidos a la base del esporofilo, reniformes,
90. con un pedúnculo corto y grueso; esporas reticuladas;
x=23, 31, 34, c. 90.

SIMILAR GENERA: Huperzia differs by equally forked GÉNEROS PARECIDOS: Huperzia difiere por tallos
stems and sporophylls that are basifixed (non-peltate) furcados igualmente y esporofilos que son basificados (no
and remain green after sporangial dehiscence. Lycopodiella peltados) y permanezcan verdes después de dehiscencia
differs by producing a single row of lateral stems from the esporangial. Lycopodiella difiere por producir una hilera
dorsal surface of the main stem. Its strobili tend to be de tallos subordinados de la superficie dorsal del tallo
sessile, not stalked. rastrero principal. Sus estróbilos tienden ser sésiles, no
pedicelados.

COMMENTS: Lycopodium is pantropical and contains COMENTARIOS: Lycopodium existe en una amplia
about 40 species worldwide. It occurs in a wide variety of variedad de hábitats, pero se halla principalmente arriba
habitats, but is mainly above 1000 m. Five species occur in de 1000 m. Cinco especies existen en el Neotrópico, todo
the Neotropics, all of which are illustrated and described de las cuales están ilustradas y describidas en Øllgaard
in Øllgaard (1988). Lycopodium clavatum L. is common (1988). Lycopodium clavatum L. es común y difundida en
and widespread in the neotropics and is the only species el neotrópico, y es la única especie con ápices filiformes
with a hyaline filiform leaf apices (Fig. E). Two species are hialinas de las hojas (Fig. E). Dos especies son distinctas
distinct by their flattened branchlets: L. thyoides Willd. por sus ramitas achatadas: L. thyoides Willd. (Fig. A–C) y L.
(Fig. A–C) and L. jussiaei Poir. The genus name is derived jussiaei Poir. El nombre genérico se deriva del griego lycos,
from the Greek lycos, wolf + pous, foot. The branch tips lobo + pous, pie. Los ápices de las ramas se asemejan a
resemble a wolf ’s paw. un pato de un lobo.

LITERATURE: Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Øllgaard, B.
1989. Index of the Lycopodiaceae. Biologiske Skrifter 34: 1–135. 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. Wilce, J. H. 1965. Section Complanata of the genus Lycopodium. Beihefte zur Nova Hedwigia 19: i–ix, 1–233.

346
Figure 167. A–C. Lycopodium thyoides. D, E. L. clavatum. (Mickel & Smith, 2004).

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LYGODIACEAE
Lygodium Sw.
DESCRIPTION: Terrestrials; rhizome creeping, DESCRIPCIÓN: Terrestres; rizomas reptantes,
branched, pubescent; leaves vine-like, of indeterminate ramificados, pubescentes; hojas de crecimiento
growth, climbing by means of a twinning rachis; indefinido, trepa por medio de un raquis que
pinnae alternate, short-stalked, the stalk forked with enrosca como un bejuco; pinnas alternas, cortamente
an arrested bud in the axil; veins free or areolate; pecioluladas, peciolulo furcado con un brote latente
sporangia borne singly on the vein tips (not clustered en la axila; nervaduras libres o areoladas; esporangios
in sori), covered by a flap of lamina tissue, arranged in naciendo individualmente en los ápices de las
2 rows on narrow marginal projections of the lamina; nervaduras (no agrupados en soros), cubiertos por una
annulus apical; x=29, 30. 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. alrededor de 25 especies, de las cual solo una es templada
palmatum, of the northeastern United States). Four species (L. palmatum, del noreste de los E.E.U.U.). Cuatro especies
occur in the Neotropics. Two of these are common and existen en el Neotrópico. Dos de ellas son comunes y muy
widespread, especially in seasonally dry forests: Lygodium difundidas, especialmente en bosques secos estacionales:
venustum Sw. and L. volubile Sw. Lygodium venustum Sw. y L. volubile Sw.
Lygodium is distinctive by its twining rachises. The Lygodium es distinctivo por sus ráquises enredadores.
young leaves have reduced pinna and elongated rachises Las hojas jóvenes tienen pinnas reducidas y ráquises
that circumnutate widely until they find a support. muy alargados que hacen circumnutaciones hasta que
After a support is found, then larger, expanded pinnae encuentran un apoyo. Después, producen pinnas más
are produced. The short bifurcated pinna stalk is also grandes, expandidas. El peciolulo corto y bifurcado es
distinctive but easy to overlook, as is the small bud in its distinctivo también pero fácil pasar por alto, tanto como el
axil. brote pequeño en su axila. Se ubica en las Schizaeales, con
The genus is placed in the Schizaeales along with Lygodiaceae y Anemiaceae. Las tres familias comparten la
Lygodiaceae and Anemiaceae. The three families share sinapomorfía un anillo apical completo.
the synapomorphy of a complete apical annulus. El nombre genérico se deriva del griego lygodes, flexible,
The genus name is derived from the Greek lygodes, y se refiere al raquis largo y flexible.
flexible, and refers to the long, flexible rachis.

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.

348
Figure 168. Upper left: Lygodium volubile. G: L. heterodoxum. H, J: L. venustum. (Mickel & Smith, 2004)

349
THELYPTERIDACEAE
Macrothelypteris (H. Itô) Ching
Thelypteris sect. Macrothelypteris H. Itô
DESCRIPTION: Terrestrial; petioles with 2 vascular DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
bundles; lamina 2-pinnate-pinnatifid or more vasculares; lámina 2-pinnada-pinnatífida o más
divided; upper surface of the costae not grooved; pinnae dividida; superficie superior de las costas no sulcada;
with adnate pinnules, the proximal pinnae not reduced, pinnas con pínnulas adnatas, las pinnas proximales
or only slightly so; aerophores absent; veins free, ending no reducidas o poco reducidas; aeróforos ausentes;
before the margin in a clavate apex (seen on the upper nervaduras libres, terminando antes del margen en un
surface) indumentum of needle-shaped hairs, some ápice claviforme (vistas en el haz); indumento de pelos
of which are sepatate; sori round; indusium round- aciculares, algunos tricomas frecuentemente septados;
reniform, rarely absent; sporangia generally with capitate soros redondos; indusios rotundos-reniformes, raramente
glands near the annulus; x=31. exindusiados; esporangios generalmente con glándulas
capitadas cerca del anillo; x=31.

SIMILAR GENERA: Macrothelypteris torresiana has GÉNEROS PARECIDOS: Macrothelypteris torresiana es

the most highly divided laminae of any Thelypteridaceae la especie con hojas más divididas de Thelypteridaceae en
in the Neotropics. Because of its leaf size and shape, it el Neotrópico. A causa del tamaño y forma de sus hojas,
might be mistaken for dryopteroid genera such as Ctenitis, puede ser confundido con otros géneros driopteroides
Megalastrum, y Lastreopsis, but it differs by having needle- como Ctenitis, Megalastrum y Lastreopsis, pero difiere de
shaped hairs and two vascular bundles in the petiole. ellos por tener tricomas aciculares y dos haces vasculares
en el pecíolo.

COMMENTS: All the ten species of Macrothelypteris are COMENTARIOS: Todos las diez especies de
native to the Paleotropics. Only M. torresiana (Gaudich.) Macrothelypteris son nativas en los Paleotrópicos. Sólo
Ching is naturalized in the Neotropics and subtropics.This M. torresiana (Gaudich.) Ching está naturalizada en los
species has spread rapidly and widely and is now one of Neotrópicos y subtropicos. Esta especie ha difundida
the most common weedy species. (It exists in the United rapidamente y ampliamente y ahora es una de las especies
States as far north as southern Illinois.) It generally grows mas comunes y agresivos. (Existe en los Estados Unidos
in disturbed habitats.The genus is related to Phegopteris hasta Illinois del sur.) Generalmente existe en ambientes
(north temperate zones) and Pseudophegopteris abiertos perturbados. El género se relaciona con
(paleotropical). The genus name is derived from the Phegopteris (de las zonas templadas) y Pseudophegopteris
Greek macros, grande + Thelypteris. The species of the (paleotropical). El nombre génerico se deriva del griego
genus are relatively large compared to other thelypteroid makros, grande + Thelypteris. Las especies del género
ferns. son relativamente grandes cuando comparados a otros
helechos thelypteroides.

LITERATURE: Holttum, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris and
Macrothelypteris. Blumea 17: 5–32. 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.

350
Figure 169. Macrothelypteris torresiana (Tryon & Stolze, 1992).

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MARATTIACEAE
Marattia Sw.
DESCRIPTION: Terrestrial, fleshy; stems massive, erect; DESCRIPCIÓN: Terrestres, carnosas; tallos masivos,
sterile and fertile leaves monomorphic, with several erectos; hojas estériles y fértiles monomorfas, con
borne at a time; petiole bases with 2 rounded stipules, varias nacidas a la vez; bases de los pecíolos con 2
without nodes, scaly, the scales narrow, often filiform; estípulas redondeadas, sin nudos, escamosas, las escamas
lamina triangular, 2-5-pinnate, the apex pinnatifid; angostas, a menudo filifomes; láminas triangulares,
pinnae attached to the rachis by swollen nodes, 2-5-pinnadas, el ápice pinnatífido; pinnas unidas al raquis
opposite or nearly so; axes of the distal segments usually por nudos hinchados, opuestas o casi así; ejes de los
winged; veins free; synangia sessile, splitting longitudinally segmentos distales usualmente alados; nervaduras libres;
to reveal two faces each with a row of elongate pores; sinángios sésiles, partiéndose longitudialmente revelar
paraphyses absent or subtending the synangia in a nest- dos caras cada una con una hilera de poros elongados;
like ring; spores monolete; x=40. parafisos ausentes o subtendiendo los sinángios en un
anillo nidiforme; esporas monoletes; x=40.

SIMILAR GENERA: Eupodium differs by stalked GÉNEROS PARECIDOS: Eupodium difiere por los
synangia, the presence of awns along the veins adaxially, snángios peciolados, la presencia de aristas por las venas
and leaves typically one per plant. Danaea differs by adaxialmente y típicamente hojas una por planta. Danaea
1-pinnate leaves, dimorphy between the sterile and fertile difiere por láminas 1-pinnadas, dimorfismo foliar entre
leaves, and sporangia opening by circular pores. las hojas estériles y fértiles y esporangios abriéndose por
poros circulares.

COMMENTS: Marattia contains 7 species and occurs COMENTARIOS: Marattia consta de 7 especies y
from southern Mexico to Panama, the Caribbean, existe desde el sur de México hasta Panamá, el Caribeo,
southern Brazil, and Hawaii. The species, all of which sur de Brazil y Hawaii. Las especies, todos de las cuales
inhabit wet forests, are: Marattia alata Sw. (Jamaica and existen en bosques húmedos, son: Marattia alata Sw.
Cuba); M. cicutifolia Kaulf. (southern Brazil), M. douglasii (Jamaica y Cuba), M. cicutifolia Kaulf. (sur de Brasil), M.
(Hawaii); M. excavata Underw. (from Mexico to Panama), douglasii (Hawai), M. excavata Underw. (México a
M. laxa Kunze (from Mexico to Panama), M. interposita H. Panamá), M. laxa Kunze (México a Panamá); M. interposita
Christ (from Guatemala to Panama), and M. weinmannifolia H. Christ (Guatemala a Panamá) y M. weinmannifolia (
(southern Mexico and Guatemala). The genus name sur de México y Guatemala). El nombre genérico honra
honors Giovanni Francesco Maratti (1723–1777), a Giovanni Francesco Maratti (1723–1777), un abato
Benedictine abbot and later professor and head of the Benedictino and más tarde un profesor y jefe del jardín
botanical garden at Rome. botánico en Roma.

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.

352
Figure 170. A–B: Marattia excavata. C–F: M. weinmanniifolia. G–J: M. laxa. (Mickel & Smith, 2004)

353
MARSILEACEAE
Marsilea L.
DESCRIPTION: Aquatic or semiaquatic; rhizomes DESCRIPCIÓN: Acuáticos o semiacuáticos; rizomas
long-creeping, branched, rooting at the nodes and (in largamente rastreros, ramificados, enraizandos en los
many species) the internodes; leaves 1-pinnate, with 4 nudos y (en muchas especies) además en los entrenudos;
wedge-shaped pinnae clustered at the apex of a hojas 1-pinnadas, con 4 pinnas cuneiformes agrupadas
long petiole; floating leaves with petioles longer than in en el ápice de un largo pecíolo; hojas flotantes con
terrestrial plants, lax or flexuous, the largest pinnae often pecíolos más largos que en plantas terrestres, laxos o
with red bands on the lower surface (hidropotonia); flexuosos, las pinnas más grandes a menudo con bandas
fertile leaves identical in appearance to the sterile ones, rojas en el envés (hidropotonia); hojas fértiles idénticas
with short-stalked sporocarps along the petiole en apariencia a las hojas estériles, con esporocarpos
base or towards its base; mega- and microsporangia in cortamente pediculados a lo largo del pecíolo o
the same sporocarps; x=20. 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 COMENTARIOS: Marsilea es cosmopólito y cuenta con
about 45–70 species, most of which occur in Africa and 45–70 especies, la mayoría de las cuales existen en África y
Australia. It occurs in wet or seasonally wet habitats, Australia. Occure en hábitats mojadas o estacionalmente
and is frequently found around the margins of ponds así y se encuentra con frequencia alrededor de los
and marshes. The sporocarps represent conduplicate, márgenes de chacras y pantanos. Los esporocarpos
indurated pinnae fused along their margins to form a representan pinnas conduplicadas, endurecidas y
closed structure. Phylogenetically, the genus belongs fusionadas por los márgines para forma una estructura
to the clade of heterosporous water ferns, which also cerrada. Filogeneticamente, el género pertenece al clado
includes Azolla, Salivia, and Regnellidium. The leaves of de los helechos acuáticos heterospóricos, que incluye
Marsilea exhibit sleep movements, or nyctinasty. During también Azolla, Salivia y Regnellidium. Las hojas de Marsilea
the day the pinnae are oriented horizontally. Each is exhiben movimientos de dormir, o nictinasty. Durante el
equipped with pulvinus at the base. Toward nightfall, the día las pinnas están orientadas horizontalmente. Cada
two distal pinnae rise up, twist half around and come into una tiene un pulvínulo en la base. Al atardecer las dos
contact with one another. Afterwards, they are embraced pinnas distales se levantan y se torcen media vía y llegan
by the basal pinnae, so that the four pinnae with their a ser en contacto con cada una. Después están abrazadas
lower surfaces turned outward form a vertical packet. por las pinnas basales, de manera que las cuatro pinnas
Marsilea is the only fern that exhibits nyctinasty (Darwin, con sus superficies abaxiales tornadas a fuera forman un
1896).Marsilea is named for Count Luigi Ferndinando paquete vertical. Marsilea es el único helecho que exhibe
Marsigli (1656–1730), Italian botanist at Bologna. 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.

354
Figure 171. 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)

355
ONOCLEACEAE
Matteuccia Todaro
DESCRIPTION:Terrestrial; rhizomes erect, stoloniferous, DESCRIPCIÓN: Terrestres; rizomas erectos,
scaly; sterile and fertile leaves dimorphic; sterile leaves estoloníferos, escamosos; hojas estériles y fértiles
50–130 x 15–25 cm; petioles 1/5–1/10 the length of the dimorfas; hojas estériles 50–130 x 15–25 cm; pecíolos
lamina, with 2 vascular bundles; sterile blades 1-pinnate- 1/5–1/10 la longitud de las láminas, con 2 haces vasculares;
pinnatifid, oblanceolate, gradually tapered toward láminas estériles 1-pinnado-pinnatífidas, oblanceoladas,
the base, apices pinnatifid; sterile pinnae sessile, 20–60 gradualmente angostadas hacia la base, ápices pinnatífidos;
pairs, truncate; veins free; fertile leaves 2-pinnate, 20–40 pinnas estériles séssiles, 20–60 pares, truncadas; venas
cm long, erect, green but turning brown and indurate libres; hojas fértiles 2-pinnadas, 20–40 cm de largo,
at maturity; fertile pinnae strongly ascending; pinnules erectas, verdes pero tornándose pardas y induratas con
enrolled and forming a globose structure enclosing each maduréz; pínulas fértiles enrolladas y formando una
sorus; receptacle prominent; paraphyses absent; indusia estructura globosa encapsulando cada soro; receptáculo
present as a thin hyaline scale; spores bilateral, greenish; prominente; paraphyses ausentes; indusios presentes
x=40. como una escama delgada hialina para cada soro; esporas
bilaterales, verdosas; x=40.

SIMILAR GENERA: Onocleopsis, of southern Mexico GÉNEROS PARECIDOS: Onocleopsis, del sur de
and Guatemala, differs by net veins and fertile leaves México y Guatemala, difiere por rizomas nervación en
2-3-pinnate. Thelypteris subg. Amauropelta differs by forma de red y hojas fértiles 2-3-pinnadas. Thelypteris
aerophores present (at the pinna bases), acicular hairs, subg. Amauropelta difiere por aeróforos presentes (en las
and monomorphic sterile and fertile leaves. bases de las pinnas), pelos aciculares y hojas estériles y
fértiles monomórfas.

COMMENTS: Matteuccia consists of 3 species native COMENTARIOS: Matteuccia consta de 3 especies

to north temperate regions. Two are Asian, and one is nativas a zonas templadas norteñas. Dos son Asiáticos,
North American (M. struthiopteris). They mainly occur on y una es Norteamerican (M. struthiopteris). Existen
shaded forest floors and tend to form colonies by action principalmente en los sotobosques sombreados y
of the stolons. Onoclea stores starch in the thick, wide, tienden formar colonias por acción de sus estolones.
petiole bases that persist after the above part of the leaf Onoclea alménese almidón en las bases gruesos y anchos
has senesced. The fertile leaves persist throughout the de los pecíolos que persistan después de lo demás de la
winter and release their spores the following late winter hoja ha caída. Las hojas fértiles persistan por el invierno y
or early spring. Onoclea is sister to a clade consisting of libran sus esporas en el invierno siguiente o la primavera
Onocleopsis and Matteuccia (Gastony & Ungerer, 1997). temprano. Onoclea es hermana a un clado formado por
These genera, plus the Asian Pentarhizidium (2 species) Onocleopsis y Matteuccia (Gastony & Ungerer, 1997).
constitute the Onocleaceae. The genus name honors Estos géneros, y el asiático Pentarhizidium (2 especies)
Carolo Matteucci (1800–1863), physicist at the University constituyen las Onocleaceae. El nombre genérico honora
of Florence, Italy. Carolo Matteucci (1800–1863), físico en la Universidad
de Florencia, Italia.

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.

356
Figure 172. Matteuccia struthiopteris. Note scale-like leaves on stolons in B. (©Robbin Moran, 2009)

357
DRYOPTERIDACEAE
Maxonia C. Chr.
DESCRIPTION: Lianas or hemiepiphytes; rhizomes DESCRIPCIÓN: Llianas o hemiepífitas; rizomas
creeping, scaly; sterile and fertile leaves dimorphic; reptantes, escamosos; hojas estériles y fértiles
petioles with 6 or more vascular bundles; laminae 2- to dimorfas; pecíolos con 6 o más haces vasculares; láminas
3-pinnate-pinnatifid, glabrous or nearly so, anadromous, 2- a 3pinnado-pinnatífidas, glabras o casi así, anádromas,
chartaceous to coriaceous, the apex pinnatifid; longest cartácea a coriáceas; ráquises y costas sulcados
segments on the acroscopic side of the pinnae; rachis adaxialmente, los surcos más o menos confluentes de un
and costae grooved adaxially, the grooves more or less eje al siguiente, glabros por dentro; costas abaxialmente
continuous with those of the lower order, glabrous within con tricomas uniseriados, rojizos oparduscos, adpresos;
the grooves; veins free; sori round; indusium round, nervaduras libres; soros redondeados; indusios
appearing peltate but actually attached at a narrow sinus; redondeados, apareciéndose peltados pero en realidad
spores monolete; x=41. unidos a un seno estrecho; esporas monoletes; x=41.

SIMILAR GENERA: Arachniodes differs by GÉNEROS PARECIDOS: Arachniodes difiere por

monomorphic sterile and fertile leaves. Cyclodium usually hojas estériles y fértiles monomorfas. Cyclodium difiere
differs by 1-pinnate laminae and anastomosing veins. usualmente por laminas 1-pinnadas y venas anastomosadas.
Polybotrya differs by vascular bundles of the rhizome Polybotrya difiere por los haces vasculares del rizomas
each surrounded by a sheath of black sclerenchyma and cada uno redeado por una vaina de esclerenquima negra
acrostichoid, non-indusiate sori. Lastreopsis differs by y soros acrosticoides no indusiados. Lastreopsis difiere
terrestrial habit and costae puberulent within the grooves por habito terrestre y costa puberulentas dentro de los
adaxially. Rumohra differs by monomorphic sterile and surcos adaxiales. Rumohra difiere por hojas estériles y
fertile leaves and peltate undusia. fértiles monomórfas y indusios peltados.

COMMENTS: Maxonia is entirely neotropical, and its COMENTARIOS: Maxonia es completamente

sole species, M. apiifolia (Sw.) C. Chr., grows in wet forests neotropical, y su única especie, M. apiifolia (Sw.) C. Chr.,
from 0–1200 m. It forms a clade with other dryopteroid existe en bosques húmedos desde 0–1200 m. Forma
genera that are lianas or hemiepiphytes (Arachniodes, un clado con otros géneros dryopteroides que son
Cyclodium, Olfersia, Polybotrya). Two varieties have been lianas o hemiepífitos (Arachniodes, Cyclodium, Olfersia,
recognized: var. dualis (Guatemala to western Ecuador) Polybotrya). Dos variedades ha sido reconocidos: var. dualis
and var. apiifolia (Cuba and Jamaica). They differ only by (Guatemala a Ecuador occidental) y la var. apiifolia (Cuba
the rhizome scales: those of var. dualis are entire, whereas y Jamaica). Difieren solamente en las escamas del rizoma:
those of var. apiifolia are erose to denticulate. The genus en var. dualis son enteras, mientras que en la var. apiifolia
name honors William Ralph Maxon (1877–1948), son erosas a denticuladas. El nombre genérico honora
pteridologist at the United States National Herbarium, a William Ralph Maxon (1877–1948), pteridólogo en el
Washington, D.C. Herbario Nacional de los Estados Unidos., 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.

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Figure 173. Maxonia apiifolia. (Drawn by S. Graedel; ©Robbin Moran, 2009)

359
DRYOPTERIDACEAE
Megalastrum Holttum
DESCRIPTION: Terrestrial; petioles with 4 or more DESCRIPCIÓN: Terrestres; pecíolos con 4 o más
vascular bundles, scaly especially toward the base; lamina haces vasculares, escamosos especialmente hacia la
2–4 pinnate, catadromic above the basal pinnae, the base; lámina 2–4-pinnada, catádroma por encima de las
laminar tissue without cylindrical glands; basal pinnae pinnas basales, el tejido laminar sin glándulas cilíndricas;
enlarged basiscopically; rachises, pinna rachises, and pinnas basales alargadas basiscópicamente; ráquises,
costules adaxially with strigose, multicelular hairs; basal ráquises de las pinnas y cóstulas adaxialmente con pelos
vein of the distal pinnules springing from the costa, strigosos multicelulares; nérvulo basal basiscópico de
not the costule; veins ending in hydathodes behind las pínnulas distales surgiendo de la costa, no de la
the margin; sori round; indusium absent or rarely present; cóstula; nervaduras terminando en un hidatodo detrás
spores monolete, reniforme; x=41. 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.

360
Figure 174. A, B. Megalastrum lunense. C–E. M. atrogriseum. F, G. M. subincisum. H–J. M. pulverulentum. (Mickel & Smith,
2004)

361
POLYPODIACEAE
Melpomene A. R. Sm. & R. C. Moran
DESCRIPTION: Epiphytic, saxicolous or terrestrial; DESCRIPCIÓN: Epífitas, saxícolas o terrestres;
rhizome scales cordate at base, clathrate, generally escamas del rizoma cordiformes basalmente,
blackish or reddish, entire (never setulose), often with one clatradas, generalmente negruzcas o rojizas, enteras
to several papillose cells at the apex, these often falling (nunca setulosas), a menudo con una a varias células
away and not present; laminae sweet-smelling when papilosas en el ápice, éstas a menudo desprendidas y
dried, pinnatifid, pinnatisect or rarely 1-pinnate basally, no presentes; láminas huelen dulce cuando secas,
linear to elliptic, pubescent and setose, the hairs 0.1–0.2 pinnatífidas, pinnatisectas o raramente 1-pinnadas
mm, branched, 2–8 celled, lax, pale reddish, with most basalmente, lineares a elípticas, pubescentes y setosas,
of the color concentrated in the transverse walls, the los tricomas 0.1–0.2 mm, ramificados, 2–8 células, laxos,
setae 0.3–3 mm, reddish, multicellular, erect to spreading; rojizo pálido, con la mayor parte de color concentrado
hydathodes present; veins free, often not visible; sori en las paredes transversales, las setas 0.3–3 mm, rojizas,
round, discrete, superficial or rarely lightly sunken; spores multicelulares, erectas a patentes; hidátodos presentes;
green, trilete; x=37. 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, GÉNEROS PARECIDOS: Lellingeria difiere por los
unequally branched hairs on the petiole and rachis, the pelitos del pecíolo y raquis desigualmente furcados, la
absence of dark setae, and sunken sori. Also, many species ausencia de setas oscuras y soros hundidos. También,
of Lellingeria have setulose rhizome scales, whereas those muchas especies de Lellingeria tienen escamas del rizoma
of Melpomene are entire. 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: 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.

362
Figure 175. A, B. Melpomene leptostoma. C, D. M. deltata. E, F. M. moniliformis. G, H. M. katasophistes. J, K. M. pilosissima. L,
M. M. anfractuosa [=Terpsichore] (Mickel & Smith, 2004)

363
METAXYACEAE
Metaxya C. Presl
DESCRIPTION: Terrestrial; stems densely pubescent DESCRIPCIÓN: Terrestres; tallos densamente
,not arborescent; leaves up to 2 m long, monomorphous; pubescentes, no arborescentes; hojas de hasta 2 m
lamina 1-pinnate with a terminal pinna; pinnae de largo, monomorfas; lámina 1-pinnada con una
alternate, stalked, entire, serrate distally; veins free, parallel pinna terminal; pinnas alternas, pecioluladas, enteras,
to each other; sori superficial (not marginal), without serradas apicalmente; venas libres, paralelas entre sí; soros
indusia, scattered close to the costa, 1-3 on each vein; superficiales (no marginales), sin indusios, difundidos
receptacle flat (not elevated). x=ca. 94–96. cerca de la costa, 1-3 sobre cada vena; receptáculo plano
(no elevado). 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, the only genus in it own family, COMENTARIOS: Metaxya, único género en su propia
contains two species: M. lanosa and M. rostrata. Both familia, contiene dos especies: M. lanosa y M. rostrata.
are forest species, occurring primarily below 800 m. Existen en bosques principalmente bajo de 800 m. Sus
Its juvenile leaves are unusual by being highly lacerate- hojas juveniles son inusuales por ser profundamente
serrate, quite unlike the mature leaves. Metaxya is unique lacerada-serradas, muy diferente que las hojas maduras.
among ferns because it has more than one sorus per Metaxya es el único género de helechos que tiene más de
vein. It belongs to the Cyatheales, forming a clade with un soro por vena. Pertenece a los Cyatheales, formando
Cibotiaceae, Cyatheaceae y Dicksoniaceae. The genus un clado con Cibotiaceae, Cyatheaceae y Dicksoniaceae.
name is derived from the Greek metaxy, between or in El nombre genérico se deriva del griego metaxy, entre
the middle of. It refers to the sori borne along the veins en medio de. Se refiere a los soros nacidos por las enas
between the costa and margin of the pinnae. entre la costa y el margen de las pinnas.

LITERATURE: 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.

364
Figure 176. Metaxya rostrata (Tryon & Stolze, 1989)

365
DRYOPTERIDACEAE
Mickelia R. C. Moran, Labiak & Sundue
DESCRIPTION: Plants terrestrial (2 species) or DESCRIPCIÓN: Plantas terrestres (2 especies)
terrestrial and scandent; rhizomes short- to long-creeping, o terretres y escandentes; rizomas cortamente a
dorsiventral, the ventral surface root-bearing, the largamente rastreros, la superficie ventral llevando raíces;
dorsal surface leaf-bearing; sterile and fertile leaves hojas estériles y fértiles dimorfas; hojas 1-pinnadas
dimorphic; leaves 1-pinnate or (in A. furcata) with a o (en A. furcata) con un segmento basal libre en las pinnas
single free basal segment on the basal pinnae and thus basales y por eso 2-pinnadas; láminas usualmente con
2-pinnate; laminae usually with a free apical segment un segmento apical libre y parecido a las pinnas
similar to the lateral pinnae; lamina buds absent or (in laterales; brotes foliares ausentes o (en dos especies
the two terrestrial species) present at the pinna bases; terrestres) presentes en las bases de las pinnas;; pinnas
pinnae articulate or continuous with the rachis, entire or articuladas o continuas con el caquis, enteras o distalmente
serrate distally; veins variously anastomsing but never serradas; venas anastomosadas valiosamente pero
free, either areolate throughout without main lateral veins nunca libre, areoladas por todas partes sin venas
and without included veinlets, or with main lateral veins laterales principales y sin venas incluidas, o con venas
connected by slightly arched cross veins, and typically with conectadas por venas cruces levamente arcuadas con
included veinlets in the areoles; sori acrostichoid; x=41. venas incluidas en las areolas; soros acrosticoides; x=41.

SIMILAR GENERA: Bolbitis usually differs by terrestrial GÉNEROS PARECIDOS: Bolbitis usualmente difiere
habit, pinnatifid lamina apices, and buds (if present) por hábito terrestre, ápices de las láminas pinnatifidos, y
located toward the leaf or segment apices. Lomariopsis brotes (si presentes) ubicados hacia el ápice de la hoja o
differs by free veins. Tectaria differs by non-acrostichoid segmentos. Lomariopsis difiere por venas libres. Tectaria
(usually round) sori. difiere por soros no acrosticoides (usualmente redondos).

COMENTARIOS: Mickelia consta de 10 especies,

COMMENTS: Mickelia consists of 10 species, all todas neotropicales. El género se encuentra en bosques
neotropical. The genus grows in wet forests, generally húmedos, generalmente abajo de 1200 m, y todas las
below 1200 m, and all but two species (M. hemiotis and M. especies con excepción de dos (M. hemiotis y M. oligarchica)
oligarchica) are scandent on trunks.The species have been son escandentes sobre troncos. Las especies han sido
classified in Bolbitis, but Moran et al. (in press) showed clasificado principalmente en Bolbitis, pero Moran et al.
that Mickelia is distinct and sister to Elaphoglossum. A (en prensa) mostraron que Mickelia es distinta y hermana
synopsis of the genus, including a key and illustrations a Elaphoglossum. Un sinopsis de las especies, incluyendo
of all species, is given by Moran et al. (in review). Only una clave y ilustraciones, es dado por Moran et al. (en
two species of Mickelia (both terrestrial) have leaf buds, revisión). Solamente dos especies de Mickelia (ambas
and those buds are located in a different position than terrestres) tienen brotes foliares, y estos están ubicados
in Bolbitis: In Mickelia the buds occur at the pinna bases en una posición diferente que en Bolbitis. En Mickelia se
on the acroscopic side of the costae where the costa is ubican en las bases de las pinnas en el lado acroscópico,
joined by the lamina tissue (i.e., the buds are produced mientras que en Bolbitis están asociados con la porción
just outside of the pinna “axil”). In contrast, the buds apical o segmento terminal de las lámina. La arquitectura
in Bolbitis are adaxial and associated with the apical del rizoma de M. guianensis fue estudiado por Gay (1993,
portion or terminal segment of the lamina. The rhizome como Lomagramma). El nombre genérico honora John T.
architecture of M. guianensis was studied by Gay (1993, Mickel, pteridólogo estadounidense (1934– ).
as Lomagramma). The genus name honors John T. Mickel,
American pteridologist (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.

366
Figure 177. Mickelia. A. M. bernoullii. B. M. oligarchica. C. M. pergamentacea. D. M. lindigii (original, ©R. C. Moran, 2010).

367
POLYPODIACEAE
Microgramma C. Presl
DESCRIPTION: Epiphytic or rarely saxicolous; DESCRIPCIÓN: Epífitas o raramente saxícolas;
rhizomes long-creeping, the scales peltate, not rizomas largamente rastreros, las escamas peltadas,
clathrate, lanceolate to acicular; sterile and fertile leaves no clatradas, lanceoladas a aciculares; hojas estériles y
monomorphous or (more commonly) dimorphous with fértiles monomorfas a (más comúnmente) dimorfas con
the fertile leaves longer and narrower than the sterile; las fértiles más angostas y largas que las estériles; láminas
laminae simple, entire, ovate to elliptic or oblong; veins simples, enteras, ovadas a elípticas u oblongas; venas
anastomosing, the areoles with included veinlets; anastomosadas, las aréolas con venillas incluídas;
sori round or rarely oblong, in a single row between soros redondos u raras veces oblongos, en una sola hilera
the rachis and margin; paraphyses present, consisting of entre el raquis y el margen; parafisos presentes y consisten
narrow scales. Spores monolete, yellow; x=37. de escamas angostas. Esporas monoletes, amarillas; x=37.

SIMILAR GENERA: Pleopeltis differs by clathrate GÉNEROS PARECIDOS: Pleopeltis difiere por
rhizome scales and circular peltate scales in and around las escamas del rizoma clatradas y escamas peltadas
the sori, especially when young. Microgramma lacks such circulares adentro y alrededor de los soros, especialmente
scales. Campyloneurum and Niphidium differ by their larger los jóvenes. Microgramma carece de dichas escamas.
size (over 20 cm long) and (usually) by having more than Campyloneurum y Niphidium difieren por su tamaño más
one row of sori between the costa and margin. 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 COMENTARIOS: Microgramma tiene casi 30 especies
the Neotropics and 2 in Africa and islands of the Indian en el Neotrópico y 2 en África e islas del Océano Indico.
Ocean. Most of the species grow in lowland forests from La mayoría de especies crecen en bosques de tierras
0–600 m. The genus forms a clade with Niphidium and bajas desde 0–600 m. El género forma un clado con
Campyloneurum. Many species of Microgramma have Niphidium y Campyloneurum. Muchas especies tienen tallos
slightly flattened stems. This tendency reaches its greatest levemente aplanados. Esta tendencia es más desarrollada
development in M. megalophylla (Desv.) Sota, which may en M. megalophylla (Desv.) Sota, que puede tener tallos
have stems up to 30 mm wide and only 3 mm thick. The hasta 30 mm de ancho y sólo 3 mm de grosor. El nombre
genus name is derived from the Greek mikros, small + genérico se deriva del griego mikros, pequeño + gramme,
gramme, line. The sori are slightly elongated in the type línea. Los soros son levemente alargados en la especie
species. 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.

368
Figure 178. A, B. Microgramma piloselloides. C, D. M. reptans. F, G. M. nitida. H-K. M. tecta. L-O. M. percussa. (Mickel &
Smith, 2004)

369
POLYPODIACEAE
Microgramma subgen. Solanopteris (Copel.) Lellinger
DESCRIPTION: Epiphytic; stems slender, long-creeping, DESCRIPCIÓN: Epífitas; tallos delgados, largamente
bearing tubers 2–4 cm wide, globose, hollow, scaly; stem rastreros, con tubérculos 2–4 cm de ancho, globosos,
scales not clathrate, circular or rarely linear, peltate; sterile huecos, escamosos, las escamas no clatradas, circulares,
and fertile leaves monomorphous or dimorphous, entire peltadas; hojas estériles y fértiles monomorfas a
or pinnatifid, glabrous or sparsely and minutely scaly; dimorfas, enteras o pinnatífidas, glabras o esparcida y
veins anastomosing, the areolae containing free included diminutamente escamosas; nervaduras anastomosadas
veinlets; sori round, elongate or linear; paraphyses con venillas libres incluidas; soros redondos, alargados o
numerous, narrowly clavate, with enlarged apical cells; lineares; parafisos numerosos, estrechamente clavados,
spores echinate; x=37. 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.) neotropical y consta de 4 especies: M. bifrons (Hook.)
Lellinger (Amazon basin of Colombia, Ecuador, and Lellinger (cuenca Amazónica de Colombia, Ecuador,
Peru), M. bismarkii (Rauh) L. León (eastern Andes of y Perú), M. bismarkii (Rauh) L. León (Andes oriental
Peru), M. brunei (Wercklé ex H. Christ) Lellinger (Costa de Perú), M. brunei (Wercklé ex H. Christ) Lellinger
Rica, Panama, western Colombia and Ecuador), and M. (Costa Rica, Panama, lados occidentales de Colombia
tuberosa (Maxon) Lellinger (Andes of S. Ecuador). All and Ecuador) y M. tuberosa (Maxon) Lellinger (Andes
are high canopy epiphytes, often occurring on the outer del sur de Ecuador). Todos son epifitos del alto dosel, a
portions of tree branches. Subg. Solanopteris is the only menudo inhabitando las partes extremas de las ramas de
myrmecophytic fern genus in the Neotropics. Its tubers árboles. Subg. Solanopteris el único género mirmecofítico
have hollow chambers inhabited by fierce biting ants. The de helechos en los neotropicos. Los tubérculos tienen
ants bring into the tubers organic material that eventually cámaras huecas las cuales están habitadas por hormigas
decays and liberates nutrients that are absorbed by the mordazes feroces. Las hormigas llevan en los tubérculos
plant. The subgenus name is derived from Solanum, the materia orgánica que eventualmente se pudre y libera
genus of potato + pteris, fern (Greek). It refers to the ant- nutrientes que están absorbidos por la planta. El nombre
inhabited tubers which resemble potatoes. 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.

370
Figure 179. 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 dorsiven-
trally; note roots appressed to chamber walls. I. Paraphyses of M. bifrons. J. paraphyses of M. brunei. K. Tuber of M. bismarckii

371
POLYPODIACEAE
Moranopteris R. Y. Hirai & J. Prado
DESCRIPTION: Epiphytic, rarely saxicolous; rhizome DESCRIPCIÓN: Epifíticas, rara vez saxícolas; escamas
scales either golden, orangish, yellowish or orangish del rizoma doradas, anaranjadas o amarillas a
brown, often shiny, not clathrate, setulose or not, setulae anaranjado-pardusco, a menudo lustrosas, no
when present hyaline, concoloras or, often, darker than clatradas, setulosas o no, sétulas cuando presentes
the body of the scale; leaves short-petiolate or nearly hialinas, concoloras o, a menudo, más oscuras que el
sessile; petioles often setose; laminae (or pinnae if lamina cuerpo de la escama; hojas cortamente pecioladas
is bipinnatifid) commonly 2–15(–25) × 0.2–0.8(–1.2) o casi séssiles; pecíolo a menudo setoso; láminas
cm, linear, shallowly pinnatifid to bipinnatisect; segments comúnmente 2–15(–25) × 0.2–0.8(–1.2) cm, lineares,
generally 20–80 pares, decurrent, sometimes with an someramente pinnatífidas a bipinnatisectas; segmentos
acroscopic lobe; veins simple or with a single acroscopic generalmente 20–80 pares, decurrentes, a veces con
fertile branch; hydathodes present on the upper surface una joroba acroscópica; nervaduras simples o con sólo una
of the lamina; sori one per segment, round, superficial; ramificación acroscópica fértil; hidatodos presentes en
spores tetrahedral, green; x=37. 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. The species were previously placed in tiene 28 especies, todas neotropicales. Las especies
Micropolypodium Hayata, which is now defined as an Asian fueron previamente ubicadas en Micropolypodium Hayata,
genus of only three species (M. okuboi, M. sikkimense, la cual ahora se define como un género Asiático de tres
and M. pulojense). The other genera of grammitid ferns especies (M. okuboi, M. sikkimense y M. pulojense). Los
rarely have shiny, colored rhizome scales and typically otros géneros de helechos gramitidoides raras veces
have more than two veins per segment. In contrast, tienen escamas del rizoma lustrosas y coloreadas y
Moranopteris has only one or two veins per segment, típicamente tienen más de dos venas por segmento. En
which are represented by hydathodes on the upper contraste, Moranopteris tiene solo una o dos venas por
surface of the leaf). The cells of the rhizome scales segmento, las cuales son representadas por hidatodos en
appear swollen or inflated—another difference from el haz). Las células de las escamas del rizoma aparecen
many grammitid ferns.Moranopteris achilleifolum (Kaulf.) hinchadas o túrgidas—otra diferencia de muchos
R. Y. Hirai & J. Prado M. longisetosum (Hook.) R. Y. Hirai helechos gramitoides. Moranopteris achilleifolum (Kaulf.) R.
& J. Prado were formerly classified in Terpsichore. They Y. Hirai & J. Prado y M. longisetosum (Hook.) R. Y. Hirai & J.
are unusual in Moranopteris by their bipinnatisect laminae. Prado fueron previamente clasificadas en Terpsichore. Son
Also unusual are M. grisebachii (Underw. Ex C. Chr.) R. Y. atípicas en Moranopteris por sus láminas bipinatisectas.
Hirai & J. Prado and M. perpusillum (Maxon) R. Y. Hirai & También inusual son M. grisebachii (Underw. Ex C. Chr.)
J. Prado because, unlike their congeners, they lack setae R. Y. Hirai & J. Prado y M. perpusillum (Maxon) R. Y. Hirai
on the leaves. The genus name honors Robbin Moran, & J. Prado porque, a diferencia de las otras especies en
American pteridologist (1956– ). el género, faltan setas 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.

372
Figure 180. A–C. Moranopteris trichomanoides (Sw.) R. Y. Hirai & J. Prado. D, E. M. basiattenuata (Jenman) R. Y. Hirai & J. Pra-
do. F, G. M. blepharodes (Maxon) R. Y. Hirai & J. Prado. (from Mickel & Smith, 2004)

373
POLYPODIACEAE
Mycopteris Sundue, gen. nov. ined.
DESCRIPTION: Mostly epiphytic; roots inserted DESCRIPTION: Principalmente epifítas; raíces
ventrally; rhizomes dorsiventral, creeping, scaly, often with insertadas ventralmente; rizomas dorsiventralses,
branch buds and phyllopodia; rhizome scales orange to reptantes, escamosos, a menudo con brotes y filopodios;
castaneous, setose, the cells turgid, lustrous; fronds escamas de rizoma anaranjadas a castañas, setosas, las
beset with a black clavate fungus (Acrospermum), células túrjidas, lustrosas; hojas con hongos negros
monomorphic, setose throughout, the setae usually calvados (Acrospermum), monomorfas, setosas por
1–2 mm long, reddish,; petiole with one or two vascular todas partes, las setas usualmente 1–2 mm de largo,
bundles, castaneous, often pubescent, the hairs ca 0.1 mm rojizas; pecíolos con uno o dos haces vasculares, castaños,
long, 3-celled, 1-branched; rachises castaneous; laminae a menudo pubescentes; los pelos ca. 0.1mm de largo,
proximally 1-pinnate, distally pinnatisect, generally oblong, 3-celulares, 1-ramificados; ráquises castaños; laminas
with numerous segments, the margins entire; pinna próximamente 1-pinnadas, distalmente pinatisectas,
costae and veins with dark sclerenchyma, the dark color generalmente oblongas con muchos segmentos, enteras;
of which is easily seen on the abaxial side of the lamina; costas y venas con esclerínquima oscura, esto más visible
hydathodes present, usually cretaceous; sori round; abaxialmente; hidatodos presentes, usualmente
sporangia glabrous; spores green; x=37. cretásicos; sori round; sporangios glabros; esporas
verdes; x=37.

SIMILAR GENERA: Ascogrammitis differs by blackish GÉNEROS PARECIDOS: Ascogrammitis difiere por
clathrate rhizome scales and green veins abaxially. escamas del rizoma negruzcas clatradas y venas verdes
Alansmia, Micropolypodium, and Terpsichore differ by lack abaxialmente. Alansmia, Micropolypodium y Terpsichore
of black clavate fungi and dark veins abaxially. Ceradenia difieren por falta del hongo negro clavado y venas
and Zygophlebia differ by lacking hydathodes. oscuras abaxialmente. Ceradenia y Zygophlebia difieren
por carece de hidatodos.

COMMENTS: Mycopteris is entirely neotropical, COMENTARIOS: Mycopteris es completamente

occurring from Mexico to Bolivia, the Guianas, southeastern neotropical, desde Mexico hasta Bolivia, las Guianas,
Brazil, and the West Indies. It comprises ca. 30 species sureste de Brasil y las Antillas. Consta de ca. 30 especies
and grows in humid montane forests. The genus consists y crece en los bosques húmedos montanos. El género
of two main clades: the first, which includes Mycopteris tiene dos clados principales: el primero, que incluye
taxifolia, has rhizome scales with only apical setae, dark Mycopteris taxifolia, lleva escamas del rizoma con solo
sclerenchyma along the costae and veins, and medial setas apicales, esclerinquima oscura por las costas y
sori. The second clade, which includes M. semihirsuta venas y soros mediales. El segundo clado, que incluye M.
and M. alsopteris, has phyllopodia, lack of receptacular semigirsuta y M. alsopteris, tiene filopodios, ausencia de
setae, supramedial sori, and rachis setae of two distinctly setas, soros suprmediales y setas del caquis de dos largos
different lengths. Mycopteris and Ascogrammitis are the diferentes.Mycopteris y Ascogrammitis son los únicos dos
only two grammitid fern genera associated with a black, géneros de helechos grammitidoides asociados con
clavate fungus (Acrospermum, a bitunicate ascomycete). un hongo negro clavado (Acrospermum, a bitunicate
The fungus can be found on nearly every plant; however, ascomycete). El hongo se encuentra en cada planta, pero
it does not appear to harm the plants. The genus name parece que no daña las plantas. El nombre genérico se
is derived from the Greek myco-, fungus, + pteris, fern, deriva del griego myco-, hongo, + pteris, helecho. Refiere
referring to the relationship between this fern genus and a la relación entre el género de los helechos y el hongo
the fungus Acrospermum. 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. 2013. Mycopteris, a new neotropical genus of grammitid ferns.
Brittonia 65: 000–000.

374
Figure 181. 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 longip-
ilosa 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. (©Robbin Moran, 2010)

375
PTERIDACEAE
Myriopteris Fée
DESCRIPTION: Rupestral or terrestrial; rhizomes DESCRIPTION: Rupestres o terrestres; rizomas
compact to long-creeping, scaly, the scales concolorous compactos a largamente rastreros, escamosos, las
or bicolorous with dark central stripe; petioles dark, scaly escamas concoloras a bicoloras con una raya central;
and/or pubescent; rachises adaxially terete or flattened pecíolos oscuros, escamosos y/o pubescentes; ráquises
or grooved, with inudment like that of the petioles; blades adaxialmente teretes o planos o surcados, con indumento
2- to 4-pinnate (rarely 1-pinnate-pinnatifid), adaxially como lo de los pecíolos; láminas 2- a 4-pinnadas (raras
glabrous or pubescent, abaxially glabrous, pubescent, or veces pinnadas-pinnatífidas), adaxialmente glabras o
scaly; ultimate segments round to oblong-ovate, minute to pubescentes, abaxialmente glabras, pubescentes o
> 1 cm long; veins obscure and not ending in prominent escamosas; segmentos últimos circulares a oblongos-
hydathodes; segment margins usually recurved, with a ovados, diminuos a > 1 cm long; venas oscuras, no
poorly differentiated false indusium; sori usually partly to terminándose en hidatodos prominentes; los márgines
completely covered by the recurved segment margins; de los segmentos usualmente recurvados, con un indusio
sporangia clustered at the vein tips, 64-spored in falso poco diferenciado; soros usualmente parcialmente a
sexual species, 32-spored in apomictic ones; spores completamente cubiertos por los márgenes recurvados
globose-tetrahedral, tan to brown; x=87, 90. de los segmentos; esporangios agrupados en los
ápices de las venas, con 64 esporas en las especies
sexuales y 32 esporeas en las apomícticas; esporas
globosas-tetrahédricas, lenonadas a pardas; x=87, 90.

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: Mycopteris consta de 47 especies.

from southern Canada through the Caribbean and Existe desde el sur de Canadá por el Caribe y América
Central America to southern Chile, with one species Central hasta el sur de Chile, con una especie (M.
endemic to Namibia and South Africa. Mexico is the rawsonii) endémica a Namibia y África del Sur. México es
center of diversity of the genus; 34 of the 47 species can el centro de diversidad del género; 34 de las 47 especies
be found in Mexico, 7 of which are endemic. Until recently, se pueden encontrar en México, 7 de las cuales son
most species of Myriopters were classified in Cheilanthes. endémicas. Hasta recién, la mayoría de las especies de
The genus, however, is more closely related to Pellaea and Myriopteris fueron clasificadas en Cheilanthes. El género,
Argyrochosma than to Cheilanthes. A “microphyllous” leaf no obstante, es más relacionado a Pellaea y Argyrochosma
morphology is common in the genus and presumed to que Cheilanthes. Una morfología foliar “microfilia” es
be an adaptation to arid habitats. It consists of numerous común en el género y presumida ser una adaptación a
rounded ultimate segments.This morphology has evolved ambientes áridos. Consiste de numerosos segmentos
in other cheilanthoid genera. The genus name is derived últimos. Esta morfología se han evolucionado en otros
from the Greek myrio-, many, + pteris, fern, referring to géneros cheilantoides. El nombre genérico se deriva del
the many (myriad) ultimate segments of the type species. 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. Mickel, J. T. 1979. The fern genus Cheilanthes in the continental United States.
Phytologia 41: 431–437.

376
Figure 182. Myriopteris. A M. myriophylla. B–H. M. windhamii. J–O. M. clevelandii. P–V. M. lindheimeri. W–Bb. M. yavapensis.
(from Mickel & Smith, 2004)

377
NEPHROLEPIDIACEAE
Nephrolepis Schott
DESCRIPTION: Terrestrial or epiphytic; stems erect, DESCRIPCIÓN: Terrestres o epifitas; tallos erectos,
usually short, bearing long, slender stolons, scaly, the usualmente cortos, llevando estolones largos, delgados,
scales peltate; sterile and fertile leaves monomorphous; escamosos, las escamas peltadas; hojas estériles y fértiles
laminae 1-pinnate, linear to linear-oblong; pinnae monomorfas; láminas 1-pinnadas, lineares a lineares-
numerous, articulate to the rachis by a circular abscission oblongas; pinnas numerosas, articuladas al raquis por
zone, usually cleanly deciduous; veins free, hydathodous, una zona de abscisión circular, usualmente limpiamente
the hydathodes often white; sori round, terminal on the deciduas; venas libres, con hidatodos, los hidatodos a
first acroscopic vein branch, in a single row between the menudo blancos; soros redondos, terminales en la
costa and margin, medial to submarginal; indusia lunate primera rama acroscópica de una vena, en una sola hilera
to circular, attached by a sinus; spores bilateral, brown; entre la costa y el margen, mediales a supramediales;
x=41. 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.

378
Figure 183. A-C. Nephrolepis undulata. D-F. N. brownii. G-L. N. pectinata. M-O. N. rivularis. (from Mickel and Beitel, 1988)

379
POLYPODIACEAE
Neurodium Fée (=Pleopeltis)
Paltonium C. Presl
DESCRIPTION: Epiphytic; rhizomes short-creeping, DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
scaly, the scales clathrate, black in the center with lighter escamosos, las escamas clatradas, negras en el centro con
margins; sterile and fertile leaves monomorphous, 15- márgenes más claros; hojas estériles y fértiles monomorfas;
50 X 1-3.5 cm; laminae simple, entire, glabrous on both láminas simples, enteras, glabras en ambas caras, gruesas,
surfaces, thick, the base and apex long-attenuate; veins la base y el ápice largamente atenuados; nervación poco
hardly conspicuous, anastomosing; sori submarginal in the conspicua, anastomosada; soros submarinales en la parte
distal part of the lamina; paraphyses abundant, reddish distal de la lámina; parafises abundantes, pardo-rojisos.
brown. Spores monolete, yellow. Esporas monoletes, amarillas.

DIAGNOSIS: Leaves simple, entire, with sori in DIAGNOSIS: Hojas simples, enteras, con soros en líneas
submarginal lines in the distal part of the lamina. submarginales en la parte distal de la lámina.

SIMILAR GENERA: Microgramma and Pleopeltis differ GÉNEROS PARECIDOS: Microgramma y Pleopeltis
by having a row of round, discrete sori between the difieren por tener una hilera de soros redondos, separatos
midrib and margin. Dicranoglossum has forked leaves. entre la costa y el margen. Dicranoglossum tiene hojas
Campyloneurum and Niphidium also have simple, entire furcadas. Campyloneurum y Niphidium, también tienen
leaves but can be distinguished by larger size (over 20 hojas simples, enteras, pero se pueden ser distinguidos
cm long) and several rows of sori between the costa and por su tamaño más grande (más de 20 cm) y varias
margin. hileras de soros entre la costa y el margen.

COMMENTS: Neurodium is contains a single species: N. COMENTARIOS: Neurodium contiene una sola especie:
lanceolatum (L.) Fée. It grows in rainforests below 500 m N. lanceolatum (L.) Fée. Crece en bosques lluviosos bajo
and occurs in Mesoamerica, the Antilles, and northern de 500 m, y ocurre en Mesoamérica, las Antillas, y la parte
South America. The species is nested within Pleopeltisi norteña de América del Sur. La especie está anidado
(Alan R. Smith, pers. comm.); therefore, Neurodium will dentro de Pleopeltis (Alan R. Smith, pers. comm.); por eso,
not continue to be recognized as a genus in the future. Neurodium no se continua ser reconocido como género
en el futuro.

LITERATURE: None.

380
Figure 184. Neurodium lanceolatum. (courtesy of John Mickel)

381
POLYPODIACEAE
Niphidium J. Sm.
DESCRIPTION: Epiphytic, terrestrial, or rupestral; DESCRIPCIÓN: Epífitas, terrestres o rupestres; tallos
stems short- to long-creeping, often covered by a dense cortamente o largamente rastreros, a menudo cubiertos
mat of pubescent roots, scaly, the scales clathrate (at por un denso tapete de raíces pelosas, escamosos, las
least medially); sterile and fertile leaves monomorphous; escamas clatradas (al menos en la mitad); hojas estériles
petioles articulate to the stem, the phyllopodia 0-1 cm y fértiles monomorfas; pecíolos articulados al tallo, los
long, often darker than the petioles; laminae simple, filopódios 0-1 cm de largo, a menudo más oscuras
entire, coriaceous, narrowly elliptic to narrowly que el pecíolo; láminas simples, enteras, coriáceas,
oblanceolate, often glaucous beneath; rachis prominent; angostamente elípticas a anogostamente oblanceoladas,
venation of prominent lateral veins, these connected by a menudo glauca en el envés; raquis prominente;
inconspicuous veinlets that anastomose to form irregular nervadura de venas laterales prominentes, conectadas
areoles containing excurrent and recurrent veinlets; sori por venillas inconspicuas que se anastomosan formando
in one row between the main lateral veins; spores aréolas irregulares conteniendo venillas excurrentes
yellowish or hyaline, reniform; x=37. y recurrentes; soros en una hilera entre las venas
laterales principales; esporas amarillas o hialinas,
reniformes; x=37.

SIMILAR GENERA: Campyloneurum differs by the GÉNEROS PARECIDOS: Campyloneurum difiere

several rows of sori between the main lateral veins. por varias series de soros entre las venas principales
Microgramma and Pleopeltis, two other genera of laterales. Microgramma y Pleopeltis, dos otros géneros de
Polypodiaceae with simple, entire leaves, differ by the Polypodiaceae con hojas simples, enteras, difieren por la
single row of sori between the rachis and margin. única hilera de soros entre el raquis y el margen.

COMMENTS: Niphidium is completely neotropical and COMENTARIOS: Niphidium es completamente

contains about 10 species. It is most closely related to neotropical y consta de casi 10 especies. Niphidium es
Campyloneurum and Microgramma. Several species have más cercanamente relacionado a Campyloneurum y
leaves that form a basket or nest. These leaves trap Microgramma. Varias especies tienen hojas que forman
organic material that eventually decomposes and forms una canasta o nido. Estas hojas atrapan materia orgánica
humus into which the roots grow. Niphidium nidiforme, la cual, eventualmente, se descompone integrándose
a high-canopy epiphyte endemic to Costa Rica, has al suelo dentro de la cual las raíces crecen. Niphidium
rhizomes that creep horizontally around trunks, and nidiforme, un epífito de los doseles y endémico a Costa
leaves arranged in a nest-like manner to catch falling Rica, tiene rizomas que serpean horizontalmente
organic matter. Many species of Niphidium have veins that alrededor de troncos, y hojas arregladas como un nido
end in whitish hydathodes.The genus name is derived de manera que pueden recoger matéria orgánica caida.
from the Greek nipha, snow + eidos, like. The fronds of Muchas especies de Niphidium tienen venas que terminan
the type species bear dense white indument. 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.

LITERATURE: Lellinger, D. B. 1972. A revision of the fern genus Niphidium. American Fern Journal 62: 101–120.

382
Figure 185. Niphidium crassifolium. (Mickel & Smith, 2004)

383
PTERIDACEAE
Notholaena R. Br.
DESCRIPTION: Terrestrial; rhizomes erect to short- DESCRIPCIÓN: Terrestres; rizomas brevemente
creeping, scaly; leaves monomorphic; petioles with 1 rastreros a erectos, escamosos; hojas monomorfas;
vascular bundle, light brown to blackish; laminae 1- or pecíolos con un haz vascular, pardo-amarillentos a
2-pinnate-pinnatifid, coraiceous, with the apex pinnatifid; negruzcos; láminas 1-2-pinnado-pinnatífidas, coriáceas,
pinnae white or yellow farinose on the lower con el ápice pinnatífido; pinnas blancas o amarillas
surface, the pinnules sessile or adnate, often scaly, with farinosas en el envés, las pínnulas sésiles o adnatas,
or without hairs; rachises tan to light brown, glabrous or a menudo escamosas y/o pilosas; raquises pardo-
puerulent or farinose; veins free, with two-celled, gland- amarillentos a negruzcos, glabros a puberulentos o
tipped hairs; veins free, ending in slightly clavate tips; sori farinosos; nervaduras libres, terminando en puntas un
intramarginal, terminal on the vein tips; false indusia tanto claviformes; soros intramarginales, terminales en
absent or weakly recurved; x=30. las nervaduras; indusios falsos ausentes o débilmente
recurvados; x=30.

SIMILAR GENERA: Cheilanthes differs by nonfarinose GÉNEROS PARECIDOS: Cheilanthes difiere por
laminae and strongly developed false indusia. Argyrochosma láminas no farinosas y indusios falsos más fuertemente
differs by stalked or sessile (not adnate) pinnules, which desarollados. Argyrochosma tiene también la lámina
are usually entire, elliptic or oblong unlike those typically farinosa en el envés pero difiere por las pínnulas
of Notholaena. 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, COMENTARIOS: Como tratado aquí en un sentido
Notholaena contains about 30 species and is entirely estricto, Notholaena contiene casi 30 especies e es
neotropical. Its center of diversity is the Chihuahua completamente neotropical. Su centro de diversidad
Desert of northern Mexico. The species occur mainly es el Desierto Chihuahuense de México norteño.
in seasonally dry habitats. Molecular data show that Las especies existen principalmente en ambientes
Notholaena is monophyletic and sister to Cheiloplecton secas estacionales. Datos moleculares muestran que
(Gastony & Rollo, 1998; Rothfels et al., 2008). Some species Notholaena es monofilético y cercanamente relacionado
previously placed in the genus have been transferred to con Cheiloplecton (Gastony & Rollo, 1998; Rothfels et al.,
Cheilanthes or Argyrochosma.The genus name is derived 2008). Algunas especies ubicadas anteriormente en el
from the Greek nothos, false + chlaena, cloak. The blade género han sido trasferidas a Cheilanthes o Argyrochosma.
margin is not reflexed as in Cheilanthes. 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.

384
Figure 186. Notholeana. A–C. N. standleyi. E–C. N. sulphurea. H–K. N. affinis. L–N. N. brachycaulis. (from Mickel & Smith,
2004)

385
LINDSAEACEAE
Odontosoria Fée
DESCRIPTION: Terrestrial; rhizomes short-creeping, DESCRIPCIÓN: Terrestres; rizomas cortamente
scaly; leaves scandent, up to 6 m long, glabrous; petioles rastreros, escamosos; hojas escandentes, hasta 6 m
and rachises spiny; laminae 3- to 5-pinnate, pinnae de largo, glabras; pecíolos y ráquises espinosos; láminas
slightly reflexed; costae and costules flexuose; veins free; 3- a 5-pinnadas; pinnas escasamente reflejas; costas y
sori marginal, solitary, ovate, supplied by 1-3 veins; costillas medias flexuosas; venas libres; soros marginales,
indusium double, united by the base and the sides, the solitarios, ovados, abastecidos por 1-3 nervaduras; indusio
adaxial green like the laminar tissue, the abaxial lighter doble, unido por la base y los lados, el adaxial verde como
colored; spores trilete; x=96. 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. las venas.

COMMENTS: Odontosoria is pantropical and consists of COMENTARIOS: Odontosoria es pantropical y consta

22 species, 12 of which occur in the Neotropics. It usually de 22 especies, 12 de las cuales existen en el Neotrópico.
occurs in open places such as slopes and roadsides. Its Típicamente existe en lugares abiertos, como taludes y
leaves exhibit indeterminate growth (the apex always los lados de los caminos. Sus hojas exhiben crecimiento
unfurling) and use surrounded vegetation for support.This indeterminado (el ápice siempre desplegándose) y usan
is an adaptation of the scandent habit can also be seen la vegetación circundante como apoyo. Esta adaptación
in Blotiella, Histiopteris, certain species of Hypolepis, and del hábito escandente se puede ver también en Blotiella,
two species of Eriosorus (E. flexuosus and E. glaberrimus). Histiopteris, ciertas especies de Hypolepis y dos especies
The genus name is derived from the Greek odous, tooth de Eriosorus (E. flexuosus and E. glaberrimus). El nombre
+ soros, sorus. In the type species, the sori are at the end genérico se deriva del griego odous, diente + soros, soro.
of segment teeth. En la especie tipo, los soros están en los ápices de dientes
de los segmentos.

LITERATURE: Maxon, W. R. 1913. Studies of tropical American ferns. No. 4. Contr. U.S. Natl. Herb. 17: 133–179.

386
Figure 187. A-C. Odontosoria aculeata. D-F. O. scandens. (Courtesy of Bobbi Angell)

387
OLEANDRACEAE
Oleandra Cav.
DESCRIPTION: Epiphytic, hemiepiphytic(?) o terrestrial; DESCRIPCIÓN: Epífitas, hemiepífitas(¿), o terrestres;
rhizomes long-creeping, densely scaly, the scales peltate, rizomas largamente rastreros, densamente
non-clathrate; sterile and fertile leaves monomorphous; escamosos, las escamas peltadas, no clatradas; hojas
petioles articulate to phyllopodia; laminae simple, estériles y fértiles monomorfas; pecíolos articulados a
entire, the margins slightly cartilaginous; veins free, filopodios; láminas simples y enteras, los márgenes
long-parallel; sori round, often borne close to the rachis, ligeramente cartilaginosos; venas libres, largamente
indusiate, the indusia reniform, peltate or attached at paralelas; soros redondos, a menudo nacen cerca del
a narrow sinus; spores monolete, bilateral, non-green, raquis, indusiados, los indusios reniformes, peltados
perispores winged, cristate, or spiny; x=41. o unidos en un seno estrecho; esporas monoletes,
bilaterales, no verdes; perisporios alados, cristados o
espinosos; x=41.

SIMILAR GENERA: Elaphoglossum differs by GÉNEROS PARECIDOS: Elaphoglossum difiere

having non-peltate rhizome scales and acrostichoid por tener escamas del rizoma no peltadas y soros
sori. Polypodiaceae with simple, entire leaves (e.g., acrosticoides. Polypodiaceae con hojas simples y enteras
Campyloneuron, Microgramma) differ by net veins. (e.g., Campyloneuron, Microgramma) difieren por venas en
forma de red.

COMMENTS: Oleandra is pantropical with about 8 COMENTARIOS: Oleandra es pantropical con casi 8
species in the Neotropics and 30 in the Paleotropics. It especies en el Neotrópico y 30 en el Paleotrópico. Existe
grows in wet forests. In the Neotropics it often forms en de bosques húmedos. En el Neotrópico a menudo
dense colonies in the canopy. The articulation between forma colonias densas en el dosel. La articulación
the petiole and phyllopodium is slightly swollen. The entre el pecíolo y filopodio es un poco hinchada. Las
articulations are functional, with the leaves falling away articulaciones son funcionales, con las hojas cayéndose
cleanly. Some species with appressed rhizome scales often limpiamente. Algunas especies con escamas del rizoma
have a shrubby habit.Their leaves can be clustered in loose apresas a menudo tienen un hábito arbustivo. Sus hojas
pseudowhorls along the rhizome, indicating rhythmic pueden ser agrupadas por el rizoma en pseudoverticelas
growth. Presumably the leaf primordia are initiated flojas, indicando crecimiento rítmico. Presumablemente
regularly around the apical meristem, but differential las primordias foliares están producidas regularmente
internode elongation causes the pseudowhorls. The alrededor del meristemo apical, pero elongación
species with spreading rhizome scales are apparently diferencial de los entrenudos causan los pseudoverticilos.
hemiepiphytes, starting on trunk bases and climbing into Las especies con escamas del rizoma divergentes son
the canopy. This needs confirmation. The genus name aparentemente hemiepífitas, empezando en las bases
is derived from an allusion of the leaves to those of de troncos y trepando hasta el dosel. Esto se necesita
Oleander (Nerium, Apocynaceae). 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. 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.

388
Figure 188. Oleandra articulata. (Courtesy of Bobbi Angell)

389
DRYOPTERIDACEAE
Olfersia Raddi
DESCRIPTION: Plants terrestrial or low-climbing; DESCRIPCIÓN: Plantas terrestres o bajo-reptantes;
stem short-creeping, in cross-section with the meristeles tallo corto-reptante, en sección transversal con las
arranged irregularly and each one not surrounded by a meristelas arregladas irregularmente y cada uno sin vaina
dark sclerenchymatous shealth; sterile and fertile leaves de esclerénquima oscuro; hojas estériles y fértiles
strongly dimorphic; sterile lamina 1-pinnate with a fuertamente dimórficas; lámina estéril con una
conform terminal pinna; pinnae ovate to lanceolate, pinna terminal conforme; pinnas ovadas a lanceoladas,
excavate on the basiscopic side, entire; veins forking near excavadas en el lado basiscópico, enteras; venas furcadas
or at the base, parallel, connected by a submarginal cerca de la base, paralelas, conectadas por una vena
vein (this sometimes difficult to see in dried material); submarginal (ésta a veces difícil de ver en ejemplares
rachis and costae not or only shallowly grooved, glabrous; secos); raquis y costas sin surcos adaxialmente, glabros;
fertile leaves 2-pinnate, rarely 1-pinnate; sori nonindusiate; hojas fértiles 2-pinnadas, raramente 1-pinnadas; soros sin
spores monolete, echinulate; x=41. indusios; esporas monoletes, equinulatas. x=41.

SIMILAR GENERA: Bolbitis differs by having (usually) GÉNEROS PARECIDOS: Bolbitis difiere por tener
reticulate veins and an elongate ventral meristele (seen venas reticuladas y una meristela ventral elongada (visto
when the rhizome is cut in cross section). Polybotrya differs cuando el rizoma se corta en sección transversal).
by venation (various patterns but never long-parallel like Polybotrya difiere por la venación (varios patrones pero
Olfersia), pinnatifid apex of the lamina, pubescent adaxial nunca es largo-paralela como Olfersia), el ápice pinnatífido
grooves of the rachis, and dark sclerenchymatous sheaths de la lámina, el surco adaxial del raquis pubescente, y el
surrounding the meristeles. patrón del tallo en sección transversal.

COMMENTS: Olfersia consists of three species restricted COMENTARIOS: Olfersia consiste de tres especies
to the neotropics: O. alata Sánchez (eastern Cuba) and restringidas al neotrópico: O. alata Sánchez (Cuba oriental)
O. cervina (L.) Kunze (S. Mexico to Panama, the Antilles, y O. cervina (L.) Kunze (S. México a Panamá, las Antillas,
and French Guiana to Bolivia), and O. corcovadensis Kaulf. Guyana Francesa a Bolivia) y O. corcovadensis Kaulf. ex
ex Raddi (se. Brazil). The fertile leaves are taller and Raddi (se. Brazil). Las hojas fértiles son más altas y rectas
more erect than the sterile ones and soon wither after que las estériles y se marchitan apenas después de que
the spores are shed. The connecting veins of the sterile las esporas son lanzadas. La vena colectora de las pinnas
pinnae is absolutely diagnostic for the genus. estériles es absolutamente diagnóstica para el género.
The generic name honors Ignaz Franz Werner Maria El nombre génerico honora Ignaz Franz Werner Maria
von Olfers (1793–1871), a German naturalist, historian, von Olfers (1793–1871), una naturalista, historiador y
and diplomat who collected in Brazil ca. 1816. diplomático alemán quien colectó en Brasil 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

390
Figure 189. 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)

391
ONOCLEACEAE
Onoclea L.
DESCRIPTION: Terrestrial; rhizomes creeping, scaly, DESCRIPCIÓN: Terrestres; rizomas reptantes,
stolons absent; sterile and fertile leaves dimorphic; escamosos estolones ausentes; hojas estériles y fértiles
sterile leaves 15-50 x 15-35 cm; petioles with 2 vascular dimorfas; hojas estériles 15-50 x 15-35 cm; pecíolos
bundles; sterile blades 1-pinnate, widest at the base con 2 haces vasculares; láminas estériles 1-pinnadas, más
or nearly so, apices pinnatifid; sterile pinnae sessile and anchas en hacia la base o casi así, ápices pinnatífidos;
becoming adnate distally, 5-11 pairs, opposite toward pinnas estériles séssiles y llegando a ser adnatas
the base of the blade, entire to shallowly lobed, bases distalmente, 5-11 pares, opuestas en la base de la lámina,
cuneate, apices acuminate, margins serrulate; veins bases cuneadas, ápices acuminados, márgines serrulados;
netted, without included veinlets; fertile leaves venas en forma de red, sin venillas incluidas; hojas
2-pinnate, 30-50 cm long, erect, green but turning brown fértiles 2-pinnadas, 30-50 cm de largo, erectas, verdes
and indurate at maturity; fertile pinnae 5-11 pairs, strongly pero tornándose pardas y induratas con maduréz; pínulas
ascending; pinnules enrolled and forming a globose fértiles enrolladas y formando una estructura globosa
structure enclosing each sorus; receptacle prominent; encapsulando cada soro; receptáculo prominente;
paraphyses absent; indusia absent or present as a thin paraphyses ausentes; indusios ausentes o presentes
hyaline scale; spores bilateral, greenish; x=37. como una escama delgada hialina para cada soro; esporas
bilaterales, verdosas; x=37

SIMILAR GENERA: Onocleopsis, of southern Mexico GÉNEROS PARECIDOS: Onocleopsis, del sur de
and Guatemala, differs by erect rhizomes, sterile blades México y Guatemala, difiere por rizomas erectos,
reduced toward the base, sterile pinnae truncate or láminas estériles pinatífidas reducidas hacia la base,
subcordate, and fertile leaves 2-3-pinnate. Woodwardia pinnas estériles truncadas o subcordatas, y hojas fértiles
aerolatea, with which this species often grows in North 2-3-pinnadas. Woodwardia aerolatea, que crece con esta
America, differs by serrate pinnae that are alternate especie a menudo en América del Norte, difiere por
throughout. pinnas serratas que son alternas por todo la lámina.

COMMENTS: Onoclea is composed of a single species, COMENTARIOS: Onoclea consta de una sola especie,
O. sensibilis L. It occurs in temperate regions of eastern O. sensibilis L. Ocurre en zonas templadas de América del
North America and eastern Asia, usually in wet habitats Norte oriental y Asia oriental, usualmente en ambientes
such as swamps and marshes, in either sun or shade. It húmedos como pantanos y humedales, en el sol o
often forms dense colonies from its creeping rhizome. sombra. A menudo forma colonias densas por su rizoma
Onoclea stores starch in the thick, wide, petiole bases that reptante. Onoclea alménese almidón en las bases gruesos
persist after the above part of the leaf has senesced. The y anchos de los pecíolos que persistan después de lo
fertile leaves persist throughout the winter and release demás de la hoja ha caída. Las hojas fértiles persistan por
their spores the following late winter or early spring. el invierno y libran sus esporas en el invierno siguiente
Onoclea is sister to a clade consisting of Onocleopsis and o la primavera temprano. Onoclea es hermana a un
Matteuccia (Gastony & Ungerer, 1997). These genera, clado formado por Onocleopsis y Matteuccia (Gastony &
plus the Asian Pentarhizidium (2 species) constitute the Ungerer, 1997). Estos géneros, y el asiático Pentarhizidium
Onocleaceae.The genus name is derived from the Greek, (2 especies) constituyen las Onocleaceae. El nombre
onos, vessel, + kleiein, to close. The fertile pinnules roll up génerico se deriva del griego onos, vasija, + kleiein, cerrar.
into bead-like segments enclosing the sori. Las pinulas fértiles se enrollan en segmentos redondeados
envolviendo de los soros.

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.

392
Figure 190. Onoclea sensibilis L. (©Robbin Moran, 2009)

393
ONOCLEACEAE
Onocleopsis F. Ballard
DESCRIPTION: Terrestrial; rhizomes erect, stout, often DESCRIPCIÓN: Terrestres; rizomas erectos, gruesos,
stoloniferous from bases; rhizome scales concolorous, a menudo estolóniferos desde las bases; escamas del
brown, lustrous; sterile and fertile leaves dimorphic; rizoma concoloras, pardas, lustrosas; hojas estériles y
sterile leaves 50-200 x 25-35 cm; sterile blades fértiles dimorfas; hojas estériles 50-200 x 25-35 cm;
1-pinnate, reduced toward the base, apices pinnatifid; láminas estériles 1-pinnadas, reducidas hacia la
sterile pinnae sessile, 17-28 pairs, bases truncate or base, ápices pinnatífidos; pinnas estériles séssiles, 17-28
subcordate, apices acuminate, margins broadly serrate; pares, bases truncadas o subcordatas, ápices acuminados,
blades glabrous adaxially, short-pilose on veins and costae márgines anchamente serrados; láminas glabras en el haz,
abaxially; veins netted and without included veinlets; cortamente pilosas por las venas y costas en el envés;
fertile leaves 2-3-pinnate, 56-120 cm long, with laminar venas en forma de red y sin venillas incluidas; hojas
tissue nearly absent; fertile pinnules rolled into 3 or 4 fértiles 2-3-pinnadas, 56-120 cm de largo, con tejido
globular divisions, brown at maturity, ultimate segments laminar casi ausente; pínulas fértiles enrolladas en 3 o
with 2 sori; indusia absent or present as a thin hyaline 4 divisiones globulares, pardo en madurez, segmentos
scale for each sorus; spores bilateral; x=? 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 GÉNEROS PARECIDOS: Onoclea difiere por rizomas
rhizomes and pinnatifid, triangular blades (widest at base). reptantes y láminas pinatífidas triangulares (más anchas
Matteuccia differs by free veins and 1-pinnate-pinnatifid en la base). Matteuccia difiere por venas libres y hojas
sterile fronds reduced basally to vestigial pinnae. Onoclea estériles 1-pinado-pinatífido y reducidos hacia la base a
and Matteuccia are north-temperate genera, whereas pinas vestigiales. Onoclea y Matteuccia son de las zonas
Onocleopsis is tropical. templadas norteñas, mientras que Onocleopsis es tropical.

COMMENTS: Consisting of only one species, Onocleopsis COMENTARIOS: Consistiéndose de una sola especie,
hintonii F. Ballard, occurring from central Mexico to Onocleopsis hintonii F. Ballard, ocurriéndose México
Guatemala, usually along streams in forests from 1500– central hasta Guatemala de 1500–2500 m, usualmente
2500 m. Onocleopsis is the sister genus to Matteuccia por riachuelos en bosques. Onocleopsis es el género
(Gastony & Ungerer, 1997). Both share erect rhizome that hermana a Matteuccia (Gastony & Ungerer, 1997). Los
are stoloniferous at the base, basally reduced laminae, and dos comparten rizomas erectos que son estoloníferos
the loss of gametophytic trichomes. The genus name is en la base, láminas reducidas basalmente, y la perdida de
derived from Onoclea + the Greek suffix –opsis, like. 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.

394
Figure 191. Onocleopsis hintonii F. Ballard. (Mickel & Smith, 2004).

395
OPHIOGLOSSACEAE
Ophioglossum L.
Cheiroglossa C. Presl
DESCRIPTION: Terrestrial, rarely epiphytic, fleshy, DESCRIPCIÓN: Terrestres, raras veces epífitas,
completely lacking sclerenchyma; roots in most species carnosas, completamente sin esclerénquima; raíces en
proliferous and forming clones, lacking root hairs; stems la mayoría de las especies prolíferas y formando clones,
erect, subterranean; leaves erect, conduplicate in bud sin pelos sobre las raíces; tallos rectos, subterráneos;
(not circinate); sterile lamina lacking a costa, simple, hojas rectas, conduplicadas en la yema (no circinadas);
entire, most species cordate to oblanceolate, often láminas estériles sin costa, simples, enteras, cordadas
mucronate; veins areolate, usually with free included a oblanceoladas (raramente lineales), a menudo
veilets; fertile spikes borne at the base of the sterile blade, mucronadas; venas areoladas, usualmente con venillas
with two lateral rows of sporangia, usually apiculate; incluídas libres; espigas fértiles nacidas en la base de la
sporangia opening by a transverse slit; x=30. lámina estéril, con dos hileras laterales de esporangios,
generalmente apiculadas; esporangios se abren por una
dehiscencia transversal; x=30.

SIMILAR GENERA: Anemia (Schizaeaceae) differs by GÉNEROS PARECIDOS: Anemia (Schizaeaceae) difiere
two fertile spikes and pinnately divided leaves. por dos espigas fértiles y hojas divididas pinnátamente.
\
COMMENTS: Ophioglossum is cosmopolitan and COMENTARIOS: Ophioglossum típicamente existe en
consists of about 25 species. It typically occurs in open hábitats abiertos a semi-abiertos, desde el nivel del mar
or semi-open habitats, from sea level up to páramos. hasta páramos. Las plantas a menudo forman colonias
The plants often form extensive colonies by proliferous grandes por sus raíces prolíferas, pero a causa del tamaño
roots, but because of the plants’ small size (usually less pequeño de las plantas (usualmente menos de 15 cm de
than 15 cm tall), their colonies are easily overlooked. The largo), sus colonias son fácilmente pasado por alto. La
lamina represents a rachis expanded by intercalary (not lámina representa un raquis expandido por crecimiento
marginal) growth, and therefore it lacks a costa. This is intercalario (no marginal), y por eso no presenta una costa.
similar to what occurs in Plantago. Ophioglossum palmatum Esto es similar a lo que pasa en Plantago. Ophioglossum
L. is a pendent epiphyte with palmately lobed leaves. It palmatum L. es una epífita colgante con hojas lobuladas
often grows on palm trunks in the humus-filled axils of palmatamente. A menudo se encuentra en los troncos
old persistent leaf bases. It is sometimes recognized in de palmas en las axilas llenadas con humus de las bases
a separate genus, Cheiroglossa. Ophioglossum reticulatum persistentes de viejas hojas. A veces se ubica en un género
has the highest chromosome number of any living thing: aparte, Cheiroglossa. Ophioglossum reticulatum tiene el
2n=1262. The genus name is derived from the Greek número cromosómico más alto de cualquier ser viviente:
ophis, snake + glossa, tongue. The fertile spike resembles 2n=1262. El nombre genérico se deriva del griego ophis,
a snake’s tongue. serpiente + glossa, lengua. La espiga fértil se asemeja a
una lengua de serpiente.

LITERATURE: Clausen, R. T. 1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19(2): 1–177.

396
Figure 192. A. Ophioglossum. engelmannii Prantl B. O. palmatum L. (pendent epiphyte; epífito colgante). C. O. reticulatum L. D.
O. nudicaule L.f. (note proliferous root; nótase raíz prolifero). E. O. crotalophoroides Walt.

397
OSMUNDACEAE
Osmunda L.
DESCRIPTION: Terrestrial; stems erect, surrounded DESCRIPCIÓN: Terrestres; tallos erectos, rodeados
by numerous overlapping leaf bases and a dense mat of por numerosas bases de pecíolos traslapadas y una
fibrous roots; sterile and fertile leaves hemidimorphic; densa capa de raíces fibrosas; hojas estériles y fértiles
petioles in cross section with a single C-shaped hemidimórficas; pecíolos en sección cruz con un
vascular bundle, the bases flattened and winged; laminae haz vascular en forma de C, las bases achatadas
1-pinnate-pinnatifid to 2-pinnate, pubescent when young, y aladas; láminas 1-pinnado-pinnatífidas a 2-pinnadas,
glabrescent, scales absent; veins free; sporangia globose, pubescentes cuando jóvenes, a glabrescentes, escamas
borne in clusters, the annulus consisting of a single, slightly ausentes; nervaduras libres; esporangios globosos, nacen
thickened lateral patch; spores green; x=22. 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. principalmente templadas. Solamente una especie se
regalis L., the type of the genus. It typically grows in encuentra en el Neotrópico: Osmunda regalis L., el tipo
wet soils, in sun or shade. The young leaves are often del género. Típicamente crece en suelos mojados, en
reddish or purplish. The Osmundaceae are sister to sol o sombra. Las hojas jóvenes a menudo son rojizas
all other leptosporangiates ferns. It has a long fossil o purpuradas. Las Osmundaceae son hermana a todos
record extending to the Permian. The fossils are easy to los otros helechos leptoesporangiados. Tiene un registro
recognize as belonging to the family because they bear fósil largo extendiéndose al Permian. Los fósiles son
the same stem and petiole anatomy of the present-day fáciles reconocer como pertenecer a la familia porque
species (Miller, 1967, 1970). Osmunda has long included llevan la anatomía distinta de los tallos y pecíolos de
what is now called Osmundastrum cinnamomeum (which las especies actuales (Miller, 1967, 1970). Por mucho
see). To maintain this species in Osmunda would require tiempo Osmunda ha incluido lo que se llama ahora
lumping two Australasian genera (Leptopteris and Todea) Osmundastrum cinnamomeum (veáse). Para mantener
in Osmunda to preserve its monophyly. The derivation esta especie en Osmunda requeriría subsumiendo dos
of the name Osmunda is uncertain. Perhaps named for géneros Australasiaticos (Leptopteris y Todea) en Osmunda
Osmunder, Saxon god of war. 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.

398
Figure 193. A-C. Osmunda regalis. D, E. Osmundastrum cinnamomeum. (Mickel & Smith, 2004.)

399
OSMUNDACEAE
Osmundastrum C. Presl
DESCRIPTION: Terrestrial; stems erect, surrounded DESCRIPCIÓN: Terrestres; tallos erectos, rodeados
by numerous overlapping leaf bases and a dense mat of por numerosas bases de pecíolos traslapadas y una
fibrous roots; sterile and fertile leaves holodimorphic; densa capa de raíces fibrosas; hojas estériles y fértiles
petioles in cross section with a single C-shaped holodimórficas; pecíolos en sección cruz con un
vascular bundle, the bases flattened and winged; sólo haz vascular en forma de C, las bases achatadas
laminae 1-pinnate-pinnatifid, densely pubescent when y aladas; láminas 1-pinnado-pinnatífidas, pubescentes
young, glabrescent, the hairs cinnamon-colored, scales cuando jóvenes, a glabrascentes, los pelos de color
absent; veins free; sporangia globose, borne in clusters, canelo, escamas ausentes; nervaduras libres; esporangios
the annulus consisting of a single, slightly thickened lateral globosos, nacen en grupos, el anillo consiste de un sólo
patch; spores green; x=22. parche lateral ligeramente engrosado; esporas verdes;
x=22.

SIMILAR GENERA: Osmunda differs by hemidimorphic GÉNEROS PARECIDOS: Osmunda difiere por hojas
leaves (i.e., the dimorphic fertile pinnae are restricted hemidimórficas (i.e., the pinnas fértiles dimorfitas están
to only a portion of the leaf, which is otherwise green restingidas a solamente una parte de la hoja, que es en
and photosynthetic). Thelypteris differs by two vascsular otras partes verdes y fotosintéticas). Thelypteris difiere
bundles in the petiole bases. por dos haces vasculares en las bases de los pecíolos.

COMMENTS: Osmundastrum includes only one extant COMENTARIOS: Osmundastrum incluye solo una
species: O. cinnamomea (a fossil species is also known). It especie actual: O. cinnamomea (una especie fósil se
occurs in eastern North America, Central America, South conoce también). Existe en América del Norte oriental,
America, and Asia. Besides holodimorphic sterile and América Central, América del Sur y Asia.Además de
fertile leaves, Osmundastrum differs from Osmunda by two hojas estériles y fértiles holodimórficas, Osmundastrum
anatomical characteristics (Hewitson, 1961; Miller, 1971). difiere de Osmunda por dos características anatómicas
One pertains to the number of clusters of thick-walled (Hewitson, 1961; Miller, 1971). Una pertenezca al número
cells forming the sclerenchyma ring around the vascular de grupos de células con paredes engrosadas formando
strand in the petiole base, the other is the presence of el anillo de esclerénquima alrededor de la haz vascular en
a second endodermis in the stele (one endodermis is la base del pecíolo, la otra es la presencia de un epidermis
located between the inner cortex and the pericycle, and segundo en la estela (un endodermis se ubica entre la
the second is located between the xylem cylinder and the cortex interior y el perciclo, y la segunda se ubica entre
pith).The genus name is derived from Osmunda + astrum, el cilindro de xilema y la médula). El nombre genérico se
a Greek diminutive suffix. 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.

400
Figure 194. 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. (©Robbin Moran, 2009)

401
DRYOPTERIDACEAE
Parapolystichum (Keyserl.) Ching
DESCRIPTION: Terrestrial; rhizomes short- to long- DESCRIPCIÓN: Terrestres; rizomas breves a largamente
creeping, scaly; leaves monomorphic; petiole with four rastreros, escamosos; hojas monomorfas; pecíolos con 4 o
or more vascular bundles; laminae 3–4-pinnate-pinnatifid, más haces vasculares; láminas 3–4-pinnadas, lanceoladas,
lanceolate, deltate, or pentagonal, the lower pinnae deltadas o pentagonales, las pinnas más inferiores
enlarged basiscopically; scaly buds present on the agrandadas basiscópicamente; yemas escamosas en las
distal parts of the rachis of some species; pinnules porciones distales del raquis en algunas especies;
anadromic or catadromic; rachis with 2 prominent pínnulas dispuestas anadrómica o catadrómicamente;
ridges adaxially, these continuous with the thickened raquis con 2 prominentes crestas adaxiales, éstas
margins of the ultimate segments (Fig. D,E); costae continuas con los engrosados márgenes foliares de
raised (not grooved), hairy adaxially with 0.1–0.3 mm los segmentos terminales (Fig. D,E); costas elevadas
long, multicellular hairs, costae, and costules scaly abaxially, (no surcadas), pelosas adaxialmente con pelos 0.1–0.3
hairy adaxially; glandular hairs often present on the lower mm, multicelulares; tricomas glandulosos a menudo
surface of the lamina, these cylindrical, oblong appressed, presentes en el haz de la lámina, cilíndricos, oblongos,
bright yellow to orange-red; sori round. x=41. adpresos, amarillo brillante a anaranjado-rojizo; soros
redondos. x=41.

SIMILAR GENERA: Lastreopsis is indistinguishable GÉNEROS PARECIDOS: Lastreopsis es indistinguible

morphologically. Dryopteris and Rumohra differ by axes morfológicamente. Dryopteris y Rumohra se difieren
adaxially glabrous. Megalastrum differs by coarse, whitish, por sus ejes adaxialmente glabros. Megalastrum difiere
strigose hairs on the axes adaxially, prominent hydathodes, por pelos toscos, blanqueciños, strigosos sobre los ejes
and the basal veinlet of the distal pinnules springing from adaxialmente, hidatodos prominentes, y la venilla basal de
the costa, not the costule, with the lobe supplied by this las pinnulas distales surgiendo de la costa, no de la cóstula,
veinlet adnate to the costa. con el lóbulo suministrado por esta venilla adnato a la
costa.

COMMENTS: Parapolystichum contains 27 species COMENTARIOS: Parapolystichum consta de 27

distributed in the Neotropics (4 spp.), Africa (6 spp.), especies distribuidas en el Neotrópico (4 spp.), África
Madagascar (9 spp.), and Australia and New Zealand (6 spp.), Madagascar (9 spp.), and Australia and Neuva
(8 spp.). The species within each of these geographic Zealanda (8 spp.). En cada area geográfica las especies
areas form clades. Formerly, the species were treated forman un clado. Anteriormente, las especies fueron
in Lastreopsis, but molecular phylogenetic studies show tratadas en Lastreopsis, pero los studios moleculares
that Parapolystichum is distinct. No known morphological filogenéticos demuestran que Parapolystichum es distincto.
character distinguish the two genera. Several species of No hay caracteres morfológicos que separan los dos
Parapolystichum, however, have buds on their laminae géneros. Varias especies de Parapolystichum tienen brotes
distally—a character absent in Lastreopsis. The rachis- en sus láminas distalmente—un carácter ausente en
costa architecture of Lastreopsis and Parapolystichum is Lastreopsis.La arquitectura raquis-costa de Lastreopsis y
unique and unmistakable once learned. It is characterized Parapolystichum es único y no se puede confundir después
by the shape of the axes on the upper surface of the de haber comprendido. Se caracteriza por la forma de
lamina (Fig. D,E). The rachis has two ridges and is densely los ejes en el haz de la lámina (Fig. D,E). El raquis tiene
puberulent between them. The ridges are continuous dos crestas y es densamente puberulento entre ellas. Las
with the decurrent margins of the pinnules. The central crestas son continuas con los márgenes decurrentes de
part of the axes is raised (not grooved) as in most of the las pínnulas. La parte central de los ejes es prominulo (no
Dryopteridaceae. The genus name is derived from the sulcados) como en la mayoría de las Dryopteridaceae.
Greek para, beside + Polystichum. El nombre genérico se deriva del greigo para, al lado +
Polystichum.
LITERATURE: Labiak, P. H., M. Sundue, G. Rouhan, J. Garrison Hanks, J. T. Mickel & R. C. Moran. In review. Phylogeny
and historical biogeography of the lastreopsid ferns (Dryopteridaceae). American Journal of Botany [submitted 13 February
2014] 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.

402
Figure 195. A–D, H: Parapolystichum effusum. F, G: P. excultum. (Mickel & Smith, 2004)

403
POLYPODIACEAE
Pecluma M. G. Price
DESCRIPTION: Epiphytic, rarely terrestrial or rupestral; DESCRIPCIÓN: Epífitas, raras veces terrestres o
proliferous roots often present; rhizomes short-creeping, rupestres; raíces prolíferas a menudo presentes; rizomas
not branched, scaly, the scales basifixed (not peltate), cortamente rastreros, no ramificadas, escamosos, las
not clathrate, concolorous, glabrous or pubescent escamas basificadas (no peltadas), no clatradas,
with dark hairs basally (these resembling rhizoids and concoloras, glabras o pubescentes en de la base con
sometimes obscuring the scales); petioles articulate to low pelitos oscuros (estos se parecen a rizoides y a veces
(1–2 mm) phyllopodia, terete except for narrow lateral oscuren las escamas); pecíolos oscuros, no surcados
ridge decurrent from the lamina base; laminae pectinate, adaxialmente, articulados a filopódios cortos (1–2
with 30 or more segments, usually reduced toward mm),; láminas pectinadas, con 30 o más pares de
the base, the apex pinnatifid; segments adnate, entire or segmentos, usualmente reducidas en la base, el ápice
nearly so; rachises puberulent adaxially, not grooved, pinnatífido; segmentos adnatos, enteros o casi enteros;
abaxially glabrous or scaly and/or pubescent; veins free, ráquises puberulentos adaxialmente, no surcados,
occasionally anastomosing; sori round, non-indusiate; abaxialmente glabros o escamosos y/o pubescentes;
paraphyses present; sporangial capsules glabrous or venas libres, ocasionalmente anastomosadas; parafisos
setulose; spores yellow when fresh, monolete; x=37. presentes; soros redondos, no indusiados; cápsulas
esporangiales glabras o sétulas; esporas amarillas cuando
frescas, monoletes; x=37.

SIMILAR GENERA: Polypodium differs by petioles GÉNEROS PARECIDOS: Polypodium difiere por
and rachises adaxially glabrous and grooved, glabrous pecíolos y ráquises surcados y glabros adaxialmente,
rhizome scales, and non-setulose sporangial capsules. The pecíolos usulmente estramíneos (no oscuros) y cápsulas
“Polypodium dulce group” differs by light-colored petioles, esporangiales no setulosas. El “grupo Polypodium dulce”
fewer pinnae pairs, laminae widest at base, and petioles difiere por pecíolos más claros, menos pares de pinnas,
with several vascular bundles. The Terpsichore taxifolia láminas más anchas en la base, y pecíolos con varios
group differs by long (0.5–2 mm long) hairs on the haces vasculares. El grupo de Terpsichore taxifolia difiere
petioles, and green trilete spores. 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: 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.

404
Figure 196. 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. (Mickel & Smith, 2004)

405
PTERIDACEAE
Pellaea Link
DESCRIPTION: Terrestrial or rupestral; rhizomes DESCRIPCIÓN: Terrestres or rupestres; rizomas
creeping or erect, scaly, the scales usually bicolorous; rastreros o erectos, escamosos, las escamas usualmente
leaves 1–4 pinnate, monomorphic or almost dimorphic, bicoloras; hojas 1–4-pinnadas, monomorfas o casi
coriaceous; petiole with a single vascular bundle; pinnules dimorfas, coriáceas; pecíolos con una haz vascular;
stalked, glabrous or slightly pubescent; petioles and pínnulas pecioluladas, glabras o poco pubescentes;
rachises tan to black, terete, glabrous or hairy; laminar pecíolos y ráquises leonados a negros, teretes, glabros
tissue bluish green, coriaceous, glabrous; veins free, a pubescentes; tejido laminar oscuro azuloso verde,
ending in clavate hydathodes; sori intramarginal on the glabro; venas libres, terminando en hidatodos claviformes;
vein tips; indusium formed by the recurved margin of the soros intramarginales en los puntos de las venas; indusios
pinnule; spores trilete or monolete; x=29. 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 verde-
green). The two are sometimes difficult to distinguish. azulosas). Los dos son a veces difíciles distinguir.

`
COMMENTS: Pellaea contains about 35 species most COMENTARIOS: Pellaea consta de casi 35 especies
commonly found in deserts or seasonally dry habitats, se encuentran más comúnmente en hábitats desérticos
usually in or among rocks. The genus has centers of o secos estacionalmente, usualmente en o alrededor
diversity in the southwestern United States, Mexico, de las rocas. El género tiene centros de diversidad en el
and Africa. Many species are apogamous (Tryon, 1968). suroeste de los Estados Unidos, México y África. Muchas
The neotropical species form a monophyletic group especies son apógamas (Tryon, 1968). Las especies
related to Astrolepis and Argyrochosma. It are not closely neotropicales forman un grupo monofilético relacionado
related African species in sect. Holochlaena, which itself a Astrolepis y Argyrochosma. No es cercanamente
is biphyletic. The genus name is derived from the Greek relacionado a las especies Africanas en sección
pellos, dusky. It refers to the bluish gray lamina tissue. Holochlaena, la cual es bifilético. El nombre genérico
se deriva del griego pellos, obscuro. Se refiere al tejido
laminar azuloso gríseas oscuros.

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.

406
Figure 197. A–C. Pellaea ribae. D–F. P. ternifolia subsp. ternifolia. G, H. P. ternifolia subsp. arizonica. J, K: P. villosa. L–M. P. terni-
folia subsp. brandegeei. O, P. P. wrightiana. (Mickel & Smith, 2004)

407
DRYOPTERIDACEAE
Phanerophlebia C. Presl
DESCRIPTION: Terrestrial or rupestral; rhizomes DESCRIPCIÓN: Terrestres o rupícolas; rizoma
creeping or erect, scaly; leaves monomorphic; petioles with cortamente rastrero o erecto, escamoso; hojas
more than three vascular bundles; laminae 1-pinnate monomorfas; pecíolos con más tres haces vasculares;
(rarely simple) with a terminal pinna that resembles láminas 1-pinnadas (raramente simples) con una
the lateral ones; rachises and costae grooved adaxially, pinna termnal que asemeja a las laterales; raquises
the grooves more or less confluent at their junctures; y costas sulcados adaxialmente, los surcos más o
pinnae spinulose-serrulate, at least distally; veins menos confluentes en las uniones; pinnas espinuloso-
2–5-branched, free or with 1–3 series of marginal serruladas, al menos distalmente; nervaduras
anastomoses; sori round, in (1–)2–4 series between the 2–5-ramificadas, libres o con 1–3 series de anastomosis
costa and margin; indusium absent or present, if present marginales; soros redondos, en (1–)2–4 series entre la
then peltate, persistent or shrunken and falling at maturity, costa y el margen; indusio ausente o presente, si presente
the margins erose; spores monolete; x=41. entonces peltado, persistente o encogido y caedizo al
madurar, los márgenes erosos; esporas monoletes; x=41.

SIMILAR GENERA: Polystichum differs by having an GÉNEROS PARECIDOS: Polystichum difiere por
auricle on the acroscopic side of the pinna base and by a tener una aurícula en el lado acroscópico de la base de
tapered lamina apex. Nephrolepis is also one pinnate but la pinna y ápices de las láminas gradualmente reducidos.
has a pinnatifid, tapered lamina apex. 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 neotropical y consta de 9 especies (Yatskievych, 1996).
from the southwestern United States to northern South Se extiende desde el suroeste de los Estado Unidos hasta
America and Hispaniola. All of its species occur in Mexico América del Sur septentrional y Española. Todos existen
except for P. haitiensis C. Chr., from Española y Honduras. en México con excepción de P. haitiensis C. Chr., de
The species generally grow in wet forests above 1000 Española y Honduras. Las especies existen generalmente
m, often in rocky habitats. Within dryopteroid ferns, en bosques húmedos arriba de 1000 m, a menudo en
Phanerophlebia is sister to a clade consisting of Cyrtomium ambientes rocosos. Entre los helechos dryopteroides,
(Asiatic) and Polystichum. This clade is characterized by Phanerophlebia es hermana a Cyrtomium (Asiático) y
spinulose leaf margins and peltate indusia. The genus Polystichum. Este clado se caracteriza por márgenes de
name is derived from the Greek phaneros, evident + las hojas espinulosos e indusios peltados. El nombre
phelps, veins. The veins are easily visible. 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.

408
Figure 198. A–E. Phanerophlebia juglandifolia. F–L. P. pumila. M–P. P. macrosora. Q, R. P. nobilis. S, T. P. remotispora. (Mickel &
Beitel, 1988)

409
THELYPTERIDACEAE
Phegopteris (C. Presl) Fée
DESCRIPTION: Terrestrial; rhizomes 1-4 mm wide, DESCRIPCIÓN: Terrestres; rizomas 1-4 mm de ancho,
long-creeping, pubescent; petioles with two vascular largamente rastreros, pubescentes; pecíolos con 2
bundles; blades 2-3-pinnatifid at the base, apex gradually haces vasculares; láminas 2-3-pinatifidas en la base, ápice
reduced; suprabasal pinnae adnate to rachis, gradualmente reducido; pinnas suprabasales adnatas al
connected by a wing along the rachis; rachises and raquis, conectadas por una ala a lo largo del raquis;
costae not grooved adaxially, scaly and hairy abaxially; ráquises y costas no surcados adaxialmente; brotes
buds absent; veins free, reaching the margin; indument ausentes; venas libres, alcanzando el margen; indumento
abaxially of unbranched, acicular hairs, also with minute abaxialmente de pelos aciculares no surcados, también
(ca. 0.1 mm), yellowish, capitate-glandular hairs; sori round con pelos capitados-glandulares amarillentosos diminutos
to oblong, supramedial; indusia absent; sporangial capsules (ca. 0.1 mm); soros redondos a oblongos, supramediales;
often bearing a capitate gland or an acicular hair (or both); indusíos ausentes; cápsulas esporangiales a menudo
x=30. llevando una glándula capitada o un pelo acicular (o
ambos); x=30.

SIMILAR GENERA: Other thelypteroid genera differ GÉNEROS PARECIDOS: Los otros géneros
by stalked to sessile (not adnate) suprabasal pinnae, and thelypteroides difieren por pinnas suprabasales
rachises grooved adaxially. Onoclea differs by areolate pecioluladas o sésiles (no adnatas) y ráquises surcados
veins. adaxialmente. Onoclea difiere por venas areoladas.

COMMENTS: Phegopteris consists of only three COMENTARIOS: Phegopteris consta de solo tres
species: two in temperate and boreal North America (P. especies: dos en América del Norte templada y boreal
connectilis and P. hexagonoptera) and one in temperate (P. connectilis and P. hexagonoptera) y una en Asia oriental
and subtropical eastern Asia (P. decursive-pinnata). templada y subtropical (P. decursive-pinnata).Todos crecen
All grow mainly on shaded forest floors. Phegopteris, principalmente en sotobosque sombreado. Phegopteris,
Pseudophyegopteris, and Macrothelypteris form a clade Pseudophyegopteris y Macrothelypteris forman un clado
sister to the rest of the Thelypteridaceae (Smith & hermana al resto de las Thelypteridaceae (Smith &
Cranfill, 2002).The clade is defined by broadly ungrooved Cranfill, 2002). El clado se define por raquises y costas
rachises and costae, adnate pinnae and/or pinnules, veins no surcados, pinnas y/o pínnulas anchamente adnatas,
ending before the margin, and minute capitate-glandular veinas terminando antes del margen y pelitos capitados-
hairs on the laminae and sporangial capsules (pers. obs.). glandulares en las láminas y cápsulas esporangiales (obs.
The genus name is derived from the Greek phegos, beech pers.). El nombre genérico se deriva del griego phegos,
tree + pteris, fern. Presumably referring to a fern that árbol de haya + pteris, helecho. Presumiblemente se
grew beneath beech trees. refiere a un helecho que crecía bajo de árboles de haya.

LITERATURE: Holttum, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophyegopteris, and
Macrothelypteris. Blumea 17: 5–32. Smith, A. R. & R. B. Cranfill. 2002. Intrafamilial relationships of the thelypteroid ferns
(Thelypteridaceae). American Fern Journal 92: 131–149.

410
Figure 199. A–L. Phegopteris connectilis. M–R. P. hexagonoptera. (©Robbin Moran, 2009)

411
POLYPODIACEAE
Phlebodium (R. Br.) J. Sm.
DESCRIPTION: Epiphytes, rupestral, or terrestrial; DESCRIPCIÓN: Epífitas, rupícolas o terrestres;
petioles with several vascular bundles; rhizome creeping, pecíolo con varias haces vasculares; rizoma rastrero,
usually white-farinose, scaly, scales concolorous, usually generalmente blanco-farinoso, escamoso, escamas
orangish, not clathrate, denticulate; leaves pinnatisect concoloras, generalmente anaranjadas, no clatradas,
or 1-pinnate, often glaucous beneath, monomorphic, denticuladas; hojas pinnatisectas a 1-pinnatas, a menudo
articulate to the rhizome; pinnae scaly or glabrous glaucas en el envés, monomorfas, articuladas al rizoma;
beneath, the margins thickened and cartilaginous, entire or pinnas escamosas o glabras en el envés, los márgenes
near so; rachises and costae glabrous adaxially; veins engrosados y cartilaginosos, enteros o casi enteros;
areolate, the areoles with or without included veinlets; ráquises y costas glabros adaxialmente; nervaduras
sori round, without paraphyses, typically supplied by areoladas, las aréolas con o sin nérvulos incluidos; soros
two included veinlets, in 1–7 series between the costa redondeados, sin parafisos, típicamente suministrados
and margin; spores monolete, ellipsoid, yellow; x=37. por dos venillas incluidas, en 1–7 series entre la costa y
el margen; esporas monoletes, elipsoides, amarillas; x=37.

SIMILAR GENERA: Pecluma and the Polypodium dulce GÉNEROS PARECIDOS: Pecluma y el grupo de
group differ by pubescent rhizome scales (at least at the Polypodium dulce difieren por escamas del rizoma
point of attachment), rachises and costae puberulent pubescentes (al menos al punto de unión), requises
adaxially, veins free (most species), and sporangial y costas puberulentos adaxialmente, venas libres (la
capsules often setulose. Polypodium differs by free veins. mayoría de las especies) y cápsulas de los esporangios
Serpocaulon differs by clathrate rhizome scales. a menudo setulosas. Polypodium difiere por venas libres.
Serpocaulon difiere por escamas del rizoma clatradas.

COMMENTS: Phlebodium is entirely neotropical and COMENTARIOS: Phlebodium es completamente

typically occurs as an epiphyte in wet forests from 0–2500 neotropical y típicamente es un epífito en bosques
m. It is sister to a clade consisting of Pecluma and the húmedos de 0–2500 m. Phlebodium es grupo hermana a
Polypodium dulce group. Phlebodium consists of four species: un clado que consta de Pecluma y el grupo de Polypodium
P. aureum, P. decumanum, P. inaequale, and P. pseudoaureum. dulce. Phlebodium consta de cuatro especies: P. aureum,
With the exception of P. inaequale, the sori are usually P. inaequale, P. decumanum y P. pseudoaureum. Con
supplied by two included veinlets—a unique character excepción de P. inaequale, los soros están usualmente
in the Polypodiaceae (only one vein supplying the sorus suministrados por dos venillas incluidas—un carácter
is the typical condition). A key to the species is given by único en las Polypodiaceae (solo una venilla sumistrando
Tejero-Díez et al. (2009). Phlebodium aureum is a fertile el soro es la condición típica). Una clave a las especies
allotetraploid of hybrid origin between P. decumanum and se encuentra en Tejero-Díez et al. (2009). Phlebodium
P. pseudoaureum. In Mexico, Guatemala, and Honduras, the aureum es un allotetraploide fértil de origen híbrido entre
rhizomes are broken and sold in markets. It is commonly P. decumanum y P. pseudoaureum. En México, Guatemala
called “calaguala.” It is prepared as a tea, which is believed y Honduras los rizomas de son fragmentados y vendidos
to cure kidney disorders and other maladies. The genus en los mercados. Se llama comúnmente “calaguala.”
name is derived from the Greek phlebodes, full of veins. Se preparan en una infusión que, según se cree, alivia
The veins are conspicuously netted. trastornos renales y otros males. El nombre genérico se
deriva del griego phlebodes, lleno de venas. Las venas son
anastomosadas conspicuosamente.

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.

412
Figure 200. A-B, F. Phlebodium psuedoaureum. D, E: P. decumanum. (Mickel & Smith, 2004)

413
MARSILEACEAE
Pilularia L.
DESCRIPTION: Rooted aquatics; rhizomes usually long- DESCRIPCIÓN: Acuáticas radicadas en lodo;
creeping, slender, sparsely pubescent; leaves 1–10 cm rizomas usualmente largamente rastreros, delgados,
long, filiform, terete, pubescent or glabrate; sporocarps esparcidamente pubescentes; hojas 1–10 cm de
subterranean, borne at the base of the petioles, largo, filiformes, teretes, pubescentes o glabras;
densely pubescent, indurate, globose, brown to black, esporocarpos subterráneos, en la base de los
splitting into 4 valves (sporangial compartments); spores pecíolos, densamente pubescentes, indurados,
of two types (plants heterosporous): large megaspores globosos, pardos a negros, hendiéndose en 4 valvas
and smaller microspores, these trilete, endosporic; x=10. (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 COMENTARIOS: Pilularia consta de 6 especies. Existe
in. North America, Mesoamerica, South America, Europe, en América del Norte, Mesoamérica, Sudamérica,
Australia, New Zealand, Pacific Islands. A single species Europa, Australia, Nueva Zelanda, Islas Pacíficas. Una sola
of Pilularia occurs in the Neotropics: P. americana A. Br. especie de Pilularia existe en el Neotrópico: P. americana
It grows in wet or seasonally wet habitats (shorelines, A. Br. Existe hábitats mojadas o estacionalmente así
ditches, ephemeral ponds) and can be fully submersed (orillas de lagos, zanjas, charcos efímeros), y se puede ser
but produces sporocarps only on emergent plants. The completamente anegado pero produce esporocarpos
plants form dense colonies but are often overlooked solamente en plantas emergidas. Las plantas forman
because of their small, grass-like leaves. Colonies are colonias densas, pero están pasado por alto a menudo
usually discovered around the margins of ponds and a causa de sus hojas pequeñas que se asemejan a
marshes after the water level has receded.The fiddlehead gramíneas. Usualmente se descubren colonias por las
at the tip of the leaf is the best characteristic to distinguish orillas de chacras y pantanos cuando el nivel de agua ha
it from similar plants. Nagalingum et al. (2008) found that retrodicido. El cayado (prefoliación) en el ápice de las
P. americana was sister to P. novae-hollandiae and that the hojas es la característica mejor para distinguirlo de otras
two had low (compared to ferns in general) sequence plantes similares. Nagalingum et al. (2008) hallaron que P.
divergences. In North America, they found eastern Americana was sister to P. novae-hollandiae y que los dos
and western populations of P. americana differed for tuvieron divergencias de secuencias bajas (comparado
the markers studied. The South American P. mandonii A. con otros helechos). En América del Norte, hallaron
Br. was not uncluded in their study. The genus name is que poblaciones orientales y occidentales difieren en
derived from the Latin pilula, little ball, and refers to the los marcadores estudiados. Pilularia mandonii A. Br., una
globose sporocarps. 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. 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.

414
Figure 201. A–C. Pilularia americana A. Br. (Mickel & Smith, 2004)

415
PTERIDACEAE
Pityrogramma Link
DESCRIPTION: Terrestrial; rhizomes decumbent or DESCRIPCIÓN: Terrestres; rizomas decumbentes o
erect, scaly; sterile and fertile leaves monomorphous; erectos, escamosos; hojas estériles y fértiles monomorfas;
petioles often dark and lustrous, grooved on the upper pecíolos a menudo oscuros y lustrosos, surcados en la
surface; laminae pinnatisect to 4-pinnate-pinnatifid, superficie superior; láminas pinnatisectas a 4-pinnado-
farinose beneath, the farina usually white or yellow, pinnatífidas, farinosas abajo, la farina usualmente blancas
scales rare or absent, sometimes pubescent, the apex o amarillas abajo, escamas escasas o ausentes, algunas
pinnatifid, tapered; veins free; sporangia borne along veces pubescentes, el ápice pinnatífido, gradualmente
the veins, often confluent at maturity; indusia absent; reducido; nervaduras libres; esporangios nacen a lo
x=30, 58, 60. 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 por segmentos elípticos y pecíolos teretes (no surcados).
are usually less than 30 cm long and grow in rocky habitats. Las plantas miden usualmente menos que 30 cm de largo
Pentagramma, a genus of the southwestern United States y crecen en habitats rocosos. Pentagramma, un género
and Mexico, differs by its terete petioles, triangular del suroeste de los Estados Unidos y México, difiere
laminae, and pinnae broadly adnate to the rachis. por sus pecíolos teretes, láminas triangulares y pinnas
anchamente adnatas al ráquis.

COMMENTS: Pityrogramma has about 19 species, all COMENTARIOS: Pityrogramma tiene casi 19
neotropical. They typically grow in open or semi-open especies, todos neotropicales. Crecen típicamente en
places where there has been recent disturbance such as lugares abiertos o semi-abiertos donde ha estado una
road banks, landslides, and gardens. The most common perturbancia reciente, como en lados de los caminos,
and widespread species below 1000 m in the Neotropics deslaves, y jardines. La especie más común y difundida
is P. calomelanos (L.) Link. It has become naturalized in the abajo de 1000 m en el Neotrópico es P. calomelanos (L.)
Old World tropics is and spreading rapidly there. Above Link. Ha llegado a ser naturalizada en el Viejo Mundo
1000 m in the Neotropics, it is replaced by P. ebenea y se difunde rápidamente allá. Arriba de 1000 m en el
(L.) Proctor. Hybrids often occur where two or more Neotrópico, está reemplazado por P. ebenea (L.) Proctor.
species of Pityrogramma grow together (Gómez, 1979). A menudo se encuentran híbridos en lugares donde
Pityrogramma trifoliata (L.) R.Tryon is sometimes placed in dos o más especies de Pityrogramma crecen juntas
its own genus, Trismeria, because of its distinctive lamina (Gómez, 1979). A veces Pityrogramma trifoliata (L.) R.
architecture (digitate pinnae and three-dimensional Tryon se ubica en su propio género, Trismeria, debido a
leaves), bicolorous rhizome scales, and spores that lack su arquitectura laminar distinta (pinnas digitadas y hojas
an equatorial ridge. The genus name is derived from en tres dimensiones), escamas del rizoma bicoloras,
the Greek pityron, scurf, bran + gramme, line. The lower y esporas que faltan un reborde ecuatorial. El nombre
surface of the lamina is covered with a farina, and the genérico se deriva del griego pityron, caspilla + gramme,
sporangia are in lines along the veins. 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.

416
Figure 202. Pityrogramma calomelanos (L.) Link (Mickel and Smith, 2004)

417
PLAGIOGYRIACEAE
Plagiogyria (Kunze) Mett.
DESCRIPTION:Terrestrial; rhizomes erect, subterranean DESCRIPCIÓN: Terrestres; rizomas erectos,
or forming a trunk to 1 meter tall, without hairs or subterráneos o formando un tronco hasta 1 metro
scales; sterile and fertile leaves dimorphic; petiole de altura, sin pelos ni escamas; hojas estériles y
bases with 3 vascular bundles, these uniting distally fértiles dimorfas; base de los pecíolos con 3 haces
to form a “U”; laminae lanceolate or elliptic; segments or vasculares, estos uniéndose distalmente en forma
pinnae adnate to the rachis; mostly approximate, almost de U; láminas lanceoladas o elípticas; segmentos o
all spreading, the proximal ones at times reflexed, the pinnas adnatos al raquis, en su mayoría próximos, casi
distal ones at times ascending or falcate; sterile laminae todos patentes, los proximales a veces reflexos, los
pinnatisect, the segments adnate, the margins serrate distales a veces ascendentes o falcados; láminas estériles
or doubly serrate for most of their length; fertile laminae pinnatisectas, los segmentos adnatos, sus márgenes
pinnatisect or pinnate, the segments adnate or the pinnae en su mayoría serrados o 2-serrados; láminas fértiles
narrowly united, their margins entire or irregular, thin, pinnatisectas o pinnadas, los segmentos adnatos o las
frequently involute; x=66. pinnas angostamente unidas, sus márgenes enteros o
irregulares, delgados, a menudo involutos; x=66.

SIMILAR GENERA: Blechnum differs by scaly rhizomes, GÉNEROS PARECIDOS: Blechnum difiere por rizomas
4 or more vascular bundles in the petioles, reddish young escamosos, 4 o más haces vasculares en los pecíolos,
leaves, and sori in two rows along the costae. Other hojas jóvenes rojizas, y los soros en dos líneas en cada
similar genera have pinnae that are sessile but not adnate lado de las costas. Otros géneros parecidos tienen pinnas
to the rachis. 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 de casi 10 especies (Zhang & Nooteboom, 1998).
usually found above 2000 m in open or semi-open Usualmente ocurre arriba de 2000 m en ambientes
habitats. Only one species (P. pectinata (Liebm.) Lellinger) abiertos o semi-abiertos. Se dicen que solo una especie
is said to occur in the New World, but likely more exist. (P. pectinata (Liebm.) Lellinger) existe en el Nuevo Mundo,
Plants in Mesoamerica have short subterranean rhizomes, pero sin duda más existe. Plantas en Mesoamérica tienen
whereas those from Bolivia have thick trunks up to one rizomas cortos subterráneos, mientras que las de Bolivia
meter tall and can easily be mistaken for an arborescent tienen trocos gruesos hasta un metro de altura, y se
species of Blechnum. The genus name is derived from puede confundir fácilmente con una especie arborescente
the Greek plagios, oblique + gyros, ring. It refers to the de Blechnum. El nombre genérico se deriva del griego
oblique annulus. 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.

418
Figure 203. :A–E. Plagiogyria pectinata (Mexico) (Mickel & Smith, 2004)

419
POLYPODIACEAE
Pleopeltis Humb. & Bonpl. ex Willd.
DESCRIPTION: Epiphytic or rupestral; rhizome scales DESCRIPCIÓN: Epífiticas o rupestres; escamas del
clathrate (at least medially; the cell luminae sometimes rizomas clatradas (a lo menos medialmente; las lúminas
obscured), peltate, with or without rhizoides; sterile and de las células a veces oscurridas), peltadas, sin o con
fertile leaves monomorphous or dimorphous; petioles rizoides; hojas estériles y fértiles monomorfas o dimorfas;
terete (not grooved adaxially), articulate to short pecíolos teretes (no surcados adaxialmente), articulados
phyllopodia; laminae simple to 1-pinnate basally, rarely al filopódios cortos; láminas simples a 1-pinnadas
more divided, scaly, the scales circular, peltate scales, basalmente, raras veces más divididas, escamosas, las
these clathrate at least centrally; veins anastomosing, escamas circulares, peltadas, clatradas al menos en
often hard to see because of the thick lamina; sori borne el centro; venas anastomosadas, a menudo difíciles
at the juncture of several veinlets, non-indusiate, round or ver a causa de la lámina gruesa; soros se nacen en las
rarely elongate or linear, in one row between the rachis uniones de varias venillas, sin indusios, redondos raras
(or costa) and margin, young sori typically protected veces alargados o lineares, en una sola hilera entre el
by overlapping, circular, peltate scales; spores yellow, raquis (a costa) y margen, soros jóvenes típicamente
monolete; x=34, 35, 37? protegidos por escamas traslapadas, circulares,
peltadas; esporas bilaterales, amarillas; x=34, 35, 37?

SIMILAR GENERA: Other genera of Polypodiacae GÉNEROS PARECIDOS: Otros géneros de las
differ by non-peltate, non-clathrate laminar scales or the Polypodiaceae difieren por escamas laminares no peltadas
complete lack of laminar scales. Polypodium further differs y no clatradas o la falta completa de escamas laminares.
by (usually) free veins and non-clathrate rhizome scales. Polypodium difiere además por (usualmente) venas libres
Microgramma also differs by non-clathrate rhizome scales. 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.

420
Figure 204. Pleopeltis. Note the peltate scales, which distinguish the genus from other Polypodiaceae. Fijase las escamas pelta-
das 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)

421
POLYPODIACEAE
Pleopeltis (Dicranoglossum J. Sm. group)
DESCRIPTION: Epiphytes; roots dense, densely DESCRIPCIÓN: Epífitas; raíces densas, densamente
brown pubescent, forming a spongy mass; rhizomes pubescentes con pelitos pardos, formando una masa
short-creeping, scaly, the scales lanceolate, bicolorous; esponjiosa; rizoma cortamente rastrero, escamoso, las
sterile and fertile leaves monomorphous, bifurcate escamas lanceoladas, bicoloras; hojas monomorfas,
subdichotomously into several segments, sparsely bifurcadas subdicotómicamente en varios
scaly; veins free or (one species) areolate with included segmentos, esparcidamente escamosas; nervaduras
veinlets; sori submarginal, rounded to elongate or libres o (una especie) anastomosadas con nérvulos
linear, partially covered by the revolute segment margins; incluidos; soros submarginales, redondeados, alargados
paraphyses absent; spores bilateral, yellow; x=36. 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 neotropical y contiene 4 especies, todas de las cuales
from 0–1000(–1500) m. The group is monophyletic and son epifitas en bosques de 0–1000(–1500) El grupo es
nested within Pleopeltis. All the species have a dense mass monofilético y anidado dentro de Pleopeltis. Todas tienen
of spongy roots that hold water. Dicranoglossum desvauxii una masa densa esponjiosa de raíces que cargan agua.
is the only species in the genus with anastomosing veins. Dicranoglossum desvauxii es la única especie del género
The name Dicranoglossum is derived from the Greek con venas anastomosadas. El nombre Dicranoglossum
dikranos, two-pronged + glossa, tongue. It refers to the se deriva del griego dikranos, dos proyecciones + glossa,
bifurcations of the laminae into tongue-like lobes. 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.

422
Figure 205. Dicranoglossum desvauxii (Klotzsch) Proctor (from Tryon & Stolze, 1993).

423
DRYOPTERIDACEAE
Polybotrya Willd.
DESCRIPTION: Plants hemiepiphytic or (in 2 species) DESCRIPCIÓN: Plantas hemiepífitas o (en 2 especies)
terrestrial; stem long-creeping in the hemiepiphytic species, terrestres; tallo largamente rastrero en las especies
short-creeping in the terrestrial ones, in cross-section hemiepífitas, cortamente rastrero en las terrestres, en
with 4–10 circularly arranged vascular bundles, sección transversal con 4–10 haces vasculares
each surrounded by a dark sclerenchymatous dispuestos circularmente, cada uno rodeado por
sheath; sterile and fertile leaves strongly dimorphic; una vaina oscura esclerenquimatosa; hojas estériles
laminae 1-pinnate to 4-pinnate, the apex pinnatifid or y fértiles marcadamente dimorfas; láminas 1-pinnadas
rarely (in South American populations of P. polybotryoides) a 4-pinnadas, el ápice pinnatífido o raramente (en
subconform; veins free or (in subgen. Soromanes) poblaciones de América del Sur de P. polybotryoides)
anastomosing; fertile leaves like a skeletonalized version subconformes; nervaduras libres o (en subgén. Soromanes)
of the sterile ones; indusia absent; x=41. 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 Maxonia se parecen a Polybotrya por sus hábitos
sterile-fertile leaf dimorphy. The first two genera can be hemiepifíticos y dimorfismo foliar. Los dos primeros
distinguished by their conform lamina apices; the third géneros pueden distinguirse por sus ápices conformes
genus by its indusiate sori and glabrous adaxial grooves de la lámina; y el tercero por sus soros con indusios y los
of the leaf axes. 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 es completamente neotropical. Es más diverso en los
23 species occur, 12 of which are endemic. Southeastern Andes, donde se encuentran 23 especies, 12 de las cuales
Brazil has also played a role in the diversification of son endémicas. Suroeste de Brasil ha hecho un papel
Polybotrya because it has six species, all endemic. The importante en la diversificación de Polybotrya porque
fertile leaves are ephemeral and held more erect than tiene 6 especies, todas endémicas. Las hojas fértiles son
the sterile leaves. Polybotrya forms a clade with Cyclodium, efímeras y son más rectas que las estériles. Polybotrya
Maxonia, Olfersia, and Polystichopsis. This clade is united forma un clado con Cyclodium, Maxonia, Olfersia y
by creeping rhizomes. Often its species are climbing and Polystichopsis. Este clado se une por rizomas reptantes.
have dimorphic sterile and fertile leaves (or subdimorphic A menudo sus especies son trepadores y tienen hojas
in Cyclodium). The genus name is derived from the Greek estériles y fértiles dimórfas (subdimorfas en Cyclodium).
poly, many + botrys, bunches, clusters. The sori appear as El nombre genérico se deriva del griego poly, muchos
many bunches or clusters on the fertile leaves. + botrys, grupitos. Los soros aparecen como muchos
grupitos en las hojas fértiles.

LITERATURE: 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.

424
Figure 206. 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. (from Moran, 1987)

425
HYMENOPHYLLACEAE
Polyphlebium Copel.
DESCRIPTION: Epiphytes; roots absent or few; DESCRIPCIÓN: Epífitas; raíces absent or pocas;
rhizomes 0.1–0.8 mm wide, long-creeping, rizomas 0.1–0.8 mm de ancho, largamente
frequently branched, densely pubescent, the hairs rastreros, ramificados con frecuencia, densamente
reddish brown; petioles 0.3–5.5 cm long; sterile and pubescentes, los pelos pardo-rojizos; pecíolos 0.3–5.5
fertile monomorphic; sterile laminae 1-cell thick between cm de largo; hojas estériles y fértiles monomorfas; láminas
the veins, 1-pinnate to 4-pinnate, the apices not proliferous; estériles una célula de grosor entre las venas, 1-pinnadas
veins anadromous, free, false veins absent; sori marginal; a 4-pinnadas, los ápices no prolíferos; venas anádromas,
indusia tubular, the mouth usually dilated; receptacles libres, falsas venas ausentes; soros marginales; indusios
exert; spores green, tetrahaedral-globose; x=36. tubulares, la boca usualmente dilata; receptáculos exertos;
esporas verdes, tetrahédricas-globosas; x=36.

SIMILAR GENERA: Crepidomanes differs by black GÉNEROS PARECIDOS: Crepidomanes difiere por los
rhizome hairs and petioles and petiole-wing margins pelos de los rizomas negruzcos y pecíolos y márgenes de
pubescent with hairs like those of the rhizomes. las alas de los pecíolos pubescentes con pelos parecidos
Didymoglossum differs by the presence of false veins in a los del rizoma. Didymoglossum difiere por la presencia
the laminae and smaller leaves (< 6 cm). Vandenboschia de venas falsas en las láminas y hojas más cortas (< 6
differs by thicker rhizomes (1–2 mm), the presence of cm). Vandenboschia difiere por rizomas más gruesos (1–2
robust roots, and usually longer leaves (> 15 cm) mm), la presencia de raíces robustos y usualmente hojas
más largas (> 15 cm).

COMMENTS: Polyphlebium is pantropical and consists COMENTARIOS: Polyphlebium es pantropical y consta

of 15 species that grow in wet forests. Three species de 15 especies que existen en bosques húmedos. Tres
are common and widespread in the Neotropics: P. especies son comunes y muy difundidas en el Neotrópico:
angustatum, P. capillaceum, P. diaphanum, and (if distinct P. angustatum, P. capillaceum, P. diaphanum y (si distinta
from the latter) P. hymenophylloides. The first two species del anterior) P. hymenophylloides. Los rizomas de todos
are almost entirely restricted to tree fern root mantles. las especies ramifican con frecuencia y producen ramas
The rhizomes of all species branch frequently to produce cortas que parecen a raíces. Ebihara et al. (2009)
short branches that resemble roots. Ebihara et al. (2009) reportaron P. borbonicum de América del Sur y Polinesia,
reported P. borbonicum from South America and Polynesia, pero no puedo distinguir esta especie de P. diaphanum.
but I cannot distinguish this species from P. diaphanum. El nombre genérico se deriva del griego poly, mucho, y
The genus name is derived from the Greek poly, many, phleps, venas.
and phleps, veins.

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.

426
Figure 207. . A–C. Polyphlebium hymenophylloides [=P. diaphanum?]. D–E. P. capillaceum. (from Mickel & Smith, 2004.)

427
POLYPODIACEAE
Polypodium L.
DESCRIPTION: Epiphytic, rupestral, or rarely terrestrial; DESCRIPCIÓN: Epífitas, rupestres, o raras veces
rhizomes creeping, branched, bearing two rows of leaves terrestres; rizomas rastreros, ramificados, llevando dos
on the dorsal surface; rhizome scales nonclathrate, hileras de hojas sobre la superficie dorsal; escamas del
concoloras, peltate, glabrous; sterile and fertile leaves rizoma no clatradas, concoloras, peltadas, glabras; hojas
monomorphous; petioles sulcate adaxially, articulate estériles y fértiles monomorfas; pecíolos surcados
to short phyllopodia, falling off cleanly in old age; laminae en el haz, articulados a filopódios cortos, cayéndose
glabrous, pinnatisect to (rarely) 1-pinnate at the base; limpiamente con edad,; láminas glabras, pinnatisectas
veins free or rarely areolate with each areola containing a (raras veces) 1-pinnadas en la base; venas libres
a single excurrent veinlet; sori terminal on the tips of o areoladas con cada aréola conteniéndose una sóla
included veinlets, round or occasionally elliptic, in one row venilla excurrente; soros terminales en los puntos de
between the costa and margin; indusia absent; paraphyses venillas incluidas, redondos o ocasionalmente elípticas, en
often present among the sporangia; spores yellow (when una hilera entre la costa y el margen; indusios ausentes;
fresh), monolete. x=37. paraphyses a menudo presentes entre los esporangios;
esporas amarillas (cuando frescas), monoletes. x=37.

SIMILAR GENERA: Serpocaulon differs by anastomosing GÉNEROS PARECIDOS: Serpocaulon difiere por venas
veins, typically longer-creeping and unbranched, rhizomes, anastomosadas, rizomas típicamente más largamente
laminae 1-pinnate (at least basally), and clathrate rhizome rastreros y no ramificados, láminas 1-pinnadas (a lo menos
scales. Many species have more than one row of sori en la base), y escamas del rizoma no clatradas. Muchas
between the costa and margin. Pleopeltis differs by scaly especies tienen más de una hilera de soros entre la costa
and generally thicker laminae (the veins often hard to y el margen. Pleopeltis difiere por láminas escamosas y
see). Pecluma differs pectinate laminae with usually more generalmente más gruesas (las venas difíciles ver). Pecluma
than 30 pinna pairs, and petioles not grooved adaxially. difiere por láminas pectinadas con usuamente más de
Phlebodium differs by anastomosing veins and sori 30 pares de pinnas y pecíolos no surcados adaxialmente.
supplied by two veins (versus one in Polypodium). Some Phlebodium difiere por venas anastomosadas y los soros
grammitid ferns resemble Polypodium but differ by short, suministrados por dos venas (versus una en Polypodium).
erect or ascsending rhizomes, radially arranged leaves, and Algunos helechos grammitoides se parecen a Polypodium
green trilete spores. pero difieren por rizomas cortos, erectos o ascendentes,
hojas dispuetas radialmente y esporas verdes triletes.

COMMENTS: Polypodium comprises about 15 species COMENTARIOS: Polypodium consta de casi 15 especies
in temperate regions of America, Europe, Asia and Africa, en las zonas templadas de América, Europa, Asia y África,
and another 15 species in Central America. The species y otras 15 especies en América Central. Las especies son
are primarily epiphytes or saxicolous in wet forests up principalmente epífitas o saxícolas en bosques húmedos
to 2400 m. The genus name comes from the Greek, poly, up to 2400 m. El nombre genérico viene del griego, poly,
many, and podion, foot, alluding to the resemblance of the mucho, y podion, pie, aludiéndose a la semejanza de los
many phyllopodia to feet. numerosos filopodios a pies.

LITERATURE: Tejero-Diez, J. D & L. Pacheco. 2004. Taxa nuevos, nomenclatura, redefinición y distribución de las especies
relacionadas con Polypodium colpodes Kunze (Polypodiaceae, Pteridophyta). Acta Botanica 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.

428
Figure 208. 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. (Mickel & Smith, 2004).

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POLYPODIACEAE
Polypodium dulce Group (sensu Moran, 1995)
DESCRIPTION: Epiphytic, rarely terrestrial or rupestral; DESCRIPCIÓN: Epífitas, raras veces terrestres o
proliferous roots absent; rhizomes short-creeping, not rupestres; raíces prolíferas ausentes; rizomas cortamente
branched(?), dorsiventral (i.e., the leaves borne on the rastreros, no ramificadas(?), dorsiventrales (i.e., las hojas
dorsal surface), scaly, the scales pubescent with dark nacen en la superficie dorsal), escamosos, las escamas
hairs (at least at the point of attachment) not clathrate, pubescentes con pelos oscuros (a lo menos al punto
concolorous; petioles dark, not grooved adaxially, de inserción), no clatradas, concoloras; pecíolos oscuros,
articulate to low (1-2 mm) phyllopodia; laminae with no surcados adaxialmente, articulados a filopódios cortos
10-25 segments, usually widest or nearly so at (1-2 mm); láminas con 10-25 pares de segmentos,
the base, the apex pinnatifid or subconform; segments usualmente más anchas o casi así en la base, el ápice
adnate, entire or serrate; rachises puberulent adaxially, pinnatífido o subconforme; segmentos adnatos, enteros
not grooved; veins usualmente free (anastomosing o serrados; ráquises puberulentos adaxialmente, no
in three species); sori round or (less commonly) elliptic, surcados; venas usualmente libres (anastomosadas
non-indusiate; sporangial capsules often setulose; en tres especies); soros redondos o (menos común)
spores yellow, monolete; x=37. elípticos, no indusiados; cápsulas esporangiales a
menudo sétulas; esporas amarillas, monoletes; x=37.

SIMILAR GENERA: Pecluma differs by rhizomes scales GÉNEROS PARECIDOS: Pecluma difiere por escamas
united across with entire width of their base. It generally del rizomas unidas por la anchura de la base. Usualmente
also differs by dark petioles with only one vascular bundle, difiere por pecíolos oscuros con un solo haz vascular, más
more than 30 pinna pairs, and laminae usually reduced de 30 pares de pinnas y láminas usualmente reducidas
toward the base. Polypodium differs by adaxially glabrous hacia la base. Polypodium difiere por pecíolos y ráquises
and grooved petioles and rachises, glabrous rhizome surcados y glabros adaxialmente, pecíolos usualmente
scales, and glabrous sporangial capsules. estamíneos (no oscuros) y cápsulas esporangiales glabras.

COMMENTS: The Polypodium dulce group contains COMENTARIOS: El grupo de Polypodium dulce consta
about 20 species, all neotropical. Its center of diversity is de casi 20 especies, todos neotropicales. Su centro de
Mexico; only two species occur in South America (P. dulce diversidad es México; solamente dos especies existen
and P. ursipes). Although its species are currently placed in en América del Sur (P. dulce y P. ursipes). Aunque sus
Polypodium, the group is sister to Pecluma (Ranker et al., especies están ubicadas actualmente en Polypodium, el
2004), with which it shares the distinctive characters of grupo es hermana a Pecluma (Ranker et al., 2004), con
hairy rhizome scales, rachises ungrooved and puberulent la cual comparte las características distintivas de escamas
adaxially and, free typically veins, and sporangial capsules del rizoma peludas, ráquises puberulentos adaxialmente
often setulose. Eventually the P. dulce group should be y no surcados, venas típicamente libres y cápsulas
placed in a different genus—perhaps even combined esporangiales típicamente setulosas. Eventualmente el
under Pecluma. Keys to most of its species can be found grupo de P. dulce debe ser ubicado en un género distinto,
in Mickel and Smith (2004) and Moran (1995). The group tal vez aún combinado en Pecluma. Claves a la mayoría
is named for P. dulce, its most common and widespread de sus especies se puede encontrar en Mickel y Smith
species. (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.

430
Figure 209. 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).

431
DRYOPTERIDACEAE
Polystichopsis (J. Sm.) C. Chr.
DESCRIPTION: Terrestrial; rhizomes creeping; DESCRIPCIÓN: Terrestres; rizomas rastreros,
leaves monomorphic; petioles with more than three escamosos; hojas monomorfas; pecíolos con tres o más
vascular bundles; laminae deltate or pentagonal, haces vasculares; láminas deltadas o pentagonales, 2-
2- to 3-pinnate-pinnatifid, pilose on the axes or a 3-pinnado-pinnatífidas, pilosas por los ejes o por
throughout, the hairs whitish, septate; basal pinnae todo, los pelos blanquecinos, septados; pinnas basales
prolonged basiscopically, especially the basal basiscopic prolongadas basiscópicamente, especialmente la pínnula
pinnule, anadromic; medial pinnae catadromic, basal basiscópica; pinnas mediales catadrómicas;
stalked; rachis and costae grooved adaxially, the grooves raquis y costas surcados adaxialmente, los surcos
pubescent within, without proliferous buds; veins free; prolíferos adentro, sin yemas prolíferas; nervaduras libres;
sori round; indusia present or absent, orbicular-reniform; soros redondos; indusios presentes o ausentes, orbicular-
spores monolete; x=41. reniformes; esporas monoletes; x=41.

SIMILAR GENERA: Arachniodes differs by anadromic GÉNEROS PARECIDOS: Arachniodes difiere por
medial pinnae. Dryopteris differs by erect or decumbent pinnas mediales anadrómicas. Dryopteris difiere por
rhizomes and non-pilose laminae. Polystichum differs by rizomas erectas o decumbentes y láminas non pilosas.
pinnae or pinnules with an enlarged acroscopic basal Polystichum difiere por las pinnas o pínnulas con un lóbulo
lobe, spinulose teeth on the laminar margins, and peltate acroscópico basal agrandado, dientes espinulosos en los
indusia. Lastreopsis differs by the upper surfaces of the márgenes de las láminas y indusios peltados. Lastreopsis
rachises and costae rounded or flat (not grooved). difiere por el haz de las raquises y costas redondeado o
Cyclodium and Triplophyllum differs by puberulent rachises achatado (no surcado). Cyclodium y Triplophyllum difieren
and costae adaxially and the lack of pilose hairs on the ráchises y costas puberulentos adaxialmente y la falta de
lamina. pelos pilosos en la lámina.

COMMENTS: Polystichopsis consists of about 8 species COMENTARIOS: Polystichopsis consta de casi 8 especies
endemic to the Antilles. It grows at low to middle endémicas a las Antillas. Existe en elevaciones bajas a
elevations (200–1500 m) in wet forests, limestone, and medias (200–1500 m) en bosques húmedos, piedra
clay banks. It is related to Polybotrya, Cyclodium, Maxonia caliza y taludes de arcilla. Es relacionado a Polybotrya,
and Olfersia, all of which have creeping rhizomes. Proctor Cyclodium, Maxonia y Olfersia, todo de las cuales tienen
(1961) gave a synopsis of the species in Polystichopsis, and rizomas reptantes. Proctor (1961) dio una sinopsis de las
key to most of the species in the Antilles can be found in especies en Polystichipsis y una clave a las especies en
Christensen (1920) and Proctor (1985). A reassessment las Antillas se puede encontrar en Christensen (1920)
of the species is badly needed. Proctor (1985) mentioned y Proctor (1985). Se necesita mucho una revisión de las
several hybrids, but these have never been characterized. especies. Proctor (1985) mencionó varios híbridos, pero
Polystichopsis consists of medium-sized ferns that usually esto nunca han sido caracterizados. Polystichopsis consta
fit, without folding the leaf, on a single herbarium sheet. de helechos de medio tamaño que suelen caber, sin
The laminae are thin relative to other dryopteroid ferns, doblar la hoja, en una sola hoja del herbario. Las láminas
and this plus the whitish pilose hairs characterize the son delgadas relativas a otros helechos driopteroides, y
genus.The genus name is derived from Polystichum + the esto más los pelos blanquecinos pilosos se caracterizan
Greek suffix -opsis, likeness, or resemblance. el género. 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. Morton, C. V. 1960.
Observations on cultivated ferns VI. The ferns currently known as Rumohra. American Fern Journal 50: 145–155. 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.

432
Figure 210. A, B. Polystichopsis muscosa (from Proctor, 1977).

433
DRYOPTERIDACEAE
Polystichum Roth.
DESCRIPTION: Terrestrial or epipetric; rhizomes DESCRIPCIÓN: Terrestre o epipétrico; rizomas
usually thick and covered by old petiole bases, erect usualmente gruesos y cubiertos por las bases de los
to decumbent, scaly apically; sterile and fertile leaves pecíolos viejos, erectos a decumbentes, escamosos
monomorphous; petioles scaly, the scales ciliate; laminae apicalmente; hojas estériles y fértiles monomorfas; pecíolos
usually 2-pinnate to 3-pinnate; pinnae with a basal escamosos, las escamas ciliadas; láminas usualmente
acroscopic lobe or elongated pinnules, otherwise 2-pinnadas a 3-pinnadas; pinnas con una lóbulo basal
equilateral; pinnules usually inequilateral, broader on the acroscópica o una pínnula basal alongada, de otra
acroscopic side, the margins usually spinulose; rachis manera equiláteras; pínnulas usualmente inequiláteras,
and costae often conspicuously scaly; veins free (except más anchas en el lado acroscópico, los márgenes
in P. dubium (H. Karst.) Diels); sori round; indusia absent, usualmente espinulosos; ráquis y costas a menudo
or present and peltate; x=41. conspicuamente escamosos; nervadura libre (con
excepción de P. dubium (H. Karst.) Diels); soros redondos;
indusios ausentes, o presentes y peltados; x=41.

SIMILAR GENERA: Arachniodes and Rumohra differ GÉNEROS PARECIDOS: Arachniodes y Rumohra
by creeping rhizomes. Dryopteris usually differs by difieren por rizomas reptantes. Dryopteris usualmente
basiscopically elongate basal pinnae, and indusia attached tiene pinnas basales elongadas basiscópicamente, y
at a sinus. Dryopteris wallichiana (Spreng.) Hyl., which has indúsisos unidos en un sinus. Dryopteris wallichiana
all pinnae equilateral, can be distinguished by its entire (Spreng.) Hyl. tiene pinnas equiláteras, pero puede
(nonspinulose) segments and laterally attached indusia. distinguirse por sus segmentos enteros (no espinulosos)
e indusios unidos lateralmente.

COMMENTS: Polystichum contains about 260 species COMENTARIOS: Polystichum contiene casi 260
worldwide, about 70 of which are neotropical. Nearly especies mundialmente, casi 70 de las cuales son
all grow above 2000 m (the genus is absent below 600 neotropicales. Casi todas ocurren arriba de los 2000 m
m in the Neotropics). Many of the West Indian species (el género está ausente bajo los 600 m en el Neotrópico).
grow on carbonate rocks (Mickel, 1997). Hybridization Muchas de las especies Antillanas crecen sobre rocas
and polyploidy have been prominent in the evolution calcáreas (Mickel, 1997). La hibridación y poliplodia han
of Polystichum, with 83 sterile interspecific hybrids sido prominentes en la evolución de Polystichum, con 83
having been reported. The New World species form a híbridos interspecíficos estériles habían sido reportados.
monophyletic group nested with the Old World species Las especies del Nuevo Mundo forman un grupo
(Little & Barrington, 2003). The name is derived from the monofilético anidado dentro de las especies del Viejo
Greek poly, many + stichos, rows. The sori of the type Mundo (Little & Barrington, 2003). El nombre genérico se
species are in many regular rows on the pinnae. 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. 1985. The morphology and origin of a new Polystichum hybrid from Costa Rica. Systematic
Botany 10: 199–204. Barrington, D. S. 1985. The present evolutionary and taxonomic status of the fern genus Polystichum.
American Fern Journal 75: 22–28. Barrington, D. S. 2011. The fern genus Polystichum (Dryopteridaceae) in Costa Rica. Annals
of the Missouri Botanical Garden 98: 431–446. 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. Mickel, J. T. 1997.
A review of the West Indian species of Polystichum. Pages 119–143. In: R. J. Johns, editor. 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.

434
Figure 211. A–D. Polystichum rachichlaena. E. P. schizophyllum. (Mickel & Smith, 2004)

435
PTERIDACEAE
Polytaenium Desv.
DESCRIPTION: Epiphytic; rhizomes short-creeping, DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
dorsiventral; rhizome scales clathrate; leaves 2–50 x dorsiventales; escamas del rizoma clatradas; hojas
4–40 cm, distichous, simple, entire, linear to lanceolate 2–50 x 4–40 cm, distichous, simples, enteros, lineales
or oblanceolate, sessile or nearly so, chartaceous to fleshy, hasta lanceoladas a oblanceoladas, sésiles o casi sésiles,
glabrous, with spicular idoblasts (false veins) adaxially; cartáceas a carnosas, glabras, con idioblastos espiculares
costa present; veins areolate, the areoles 2–6(-8) rows (venas falsas) adaxialmente; costa presente; nervaduras
between the costa and margin, elongate, polygonal, areoladas, las aréolas alargadas, en 2–6(-8) hileras
lacking included veinlets; sori elongate along the veins, entre la costa y margen, poligonales, carentes de
forming a reticulate pattern, superficial or in grooves; nérvulos incluídos; soros alargados a lo largo de las
indusia absent; paraphyses absent; spores trilete; x=60. nervaduras, formando un patrón reticulado, superficiales
o en surcos; indusios ausentes; parafisos ausentes;
esporas triletes; x=60.

SIMILAR GENERA: Anetium differs by scattered GÉNEROS PARECIDOS: Anetium difiere por
sporangia between the veins, and long-creeping rhizomes. esporangios salpicados entre las nervaduras, y rizomas
Scoliosorus differs by the presence of paraphyses, monolete largamente reptantes. Scoliosorus difiere por la presencia
spores, and a costa that is elevated abaxially at the base. de parafisos, esporas monoletes, y una costa elevada
abaxialmente en la base.
COMMENTS: Polytaenium occurs in Central and South
America and the West Indies. It consists of 8-10 species, COMENTARIOS: Polytaenium ocurre en América
most of which are low-trunk epiphytes in the understories Central y Sud América y las Antilles. Consiste de 8-10
of wet forests. Formerly, these species were classified in especies, la mayoria de las cuales son epífitas en las bases
Antrophyum, an Old World genus of about 20 species. de los troncos de bosques húmedos. Anteriormente,
Crane (1997) showed that Antrophyum is sister to a estas especies fueron clasificadas en Antrophyum, un
largely neotropical clade comprising Polytaenium, Vittaria, género con casi 20 especies del Viejo Mundo. Crane
Ananthacorus, and Scoliosorus. As in other vittarioids, false (1997) mostró que Antrophyum es hermana a un clado
veins (cells with thick walls replaced by silica) occur in in principalmente neotropical de Polytaenium, Vittaria,
Polytaenium. They appear as minute, slender, prominulous Ananthacorus y Scoliosorus. Como en otros vittarioides,
streaks between the veins. They are always present on las venas falsas (células con paredes gruesas de sílice)
the adaxial laminar surface, but may also occur abaxially. ocurren en Polytaenium. Parecen como rayos minutos
They do not connect with true veins and do not contain delgados promínulos entre las venas. Se presentan
conducting cells (i.e., xylem and phloem). At least one siempre en el haz de la lámina y a veces en el envés. No
species of Polytaenium (P. feei) harbors ants beneath its se conectan con las verdaderas venas y no tienen células
mat of roots. The waste of the ants contribute much conductoras (i.e., xilema y floema). Al menos una especie
nitrogen to the plant. The genus name is derived from de Polytaenium (P. feei) alberguen hormigas debajo de sus
the Greek poly, many + taenia, ribbon, alluding to the sori. 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. Tryon, R. M. 1964. Taxonomic fern notes. IV. Some
American vittarioid ferns. Rhodora 66: 110–117. Watkins, Jr., J. E., C. L. Cardelús & M. C. Mack. 2008. Ants mediate nitrogen
relations of an epiphytic fern. New Phytologist 180: 5–8.

436
Figure 212. A–C. Polytaenium feei. D, E. P. lineatum. G–J. P. chlorosporum. K–M. P. cajenense. (Mickel & Smith, 2004)

437
DRYOPTERIDACEAE
Pradopteris R. C. Moran & Labiak, gen. nov. ined.
DESCRIPTION: Terrestrial; rhizomes creeping; leaves DESCRIPCIÓN: Terrestres; rizomas rastreros
monomorphic; petioles with more than three vascular escamosos; hojas monomorfas; pecíolo con tres o
bundles; laminae widely deltate or pentagonal, 2-pinnate- más haces vasculares; lámina ampliamente deltada o
pinnatifid to 4-pinnate; pinnules anadromically pentagonal, 2-5-pinnada; pinnas medias prolongadas
arranged, stalked; rachises and costae grooves acroscópicamente, el lado basal basiscópico cuneado;
pubescent at least near the junctures, without proliferous pínnulas dispuestas anadrómicamente, pediculadas;
buds; veins free; sori round; indusia orbicular-reniform, surcos de los ráquises y costas pubescentes, al menos
thick, brown, firm; spores monolete; x=41. 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 GÉNEROS PARECIDOS: Cyclodium difiere por hojas
divided, usually 1- to 2-pinnate. Dryopteris differs by menos divididas, usualmente 1- a 2-pinnadas. Dryopteris
rhizomes erect or ascending, pinnules catadromic, and (in difiere por rizomas erectos o ascendentes, pínnulas
the Neotropics) leaves often capitate-glandular. Polybotya catadrómicas y (en el Neotrópico) hojas a menudo
and Maxonia differ by strong sterile-fertile leaf dimorphy, glanduloso-capitadas. Arachniodes difiere por los surcos
Arachniodes differs by grooves of the rachises and costae de los ráquises y costas completamente glabros adentro.
completely glabrous within. Lastreopsis and Rumohra differ Lastreopsis y Rumohra difieren por una fila marginal
by a thickened marginal ridge of the costa or costules engrosada de la costa o cóstula surcurrente por el margen
surcurrent along the green laminar margin of the segment del tejido laminar verde del segmento del próximo orden
of the next highest order (not adaxially along the midrib más alto (no adaxialmente por la cóstula del segmento).
of the costule).

COMMENTS: Pradopteris is neotropical and consists of COMENTARIOS: Pradopteris es neotropical y consta

only two species: P. macrostegia (Hook.) R. C. Moran & de solo dos species: P. macrostegia (Hook.) R. C. Moran
Labiak, and P. ochropteroides (Baker) R. C. Moran & Labiak. & Labiak y P. ochropteroides (Baker) R. C. Moran & Labiak.
It occurs in wet forests from low to middle elevations. Existe en bosques húmedos desde elevaciones bajas
The first species is terrestrial and occurs primarily on hasta medias. La especie anterior es terrestre e existe
sandy soils. The abaxial surfaces of its rachises and costa principalmente sobre suelos arenosos. Las superficies
are glabrous. In contrast, the second species is epiphytic abaxiales de los ráquises y costas son glabras. En contraste,
and has pubescent rachises and costae abaxially. The la secunda especies es epífito y tiene los ráquises y costas
genus name honors Jefferson Prado (1964–), Brazilian pubescentes abaxialmente. El nombre genérico honora a
pteridologist. 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.
In prep. Phylogeny of polybotryoid ferns (Dryopteridaceae). Systematic Botany.

438
Figure 213. Pradopteris. A–C, F. P. macrostegia. D, E, G. P. ochropteroides. (©R. C. Moran, 2012)

439
PSILOTACEAE
Psilotum Sw.
DESCRIPTION: Epiphytic or rupestral, rarely terrestrial; DESCRIPCIÓN: Epífitas o rupestres, raras veces
rhizomes without roots but bearing short brownish hair- terrestres; rizomas sin raíces pero portando rizoides
like rhizoids; aerial stems green, apparently leafless, parduzcos que se parecen a pelos; tallos aéreos verdes,
unbranched basally, repeatedly dichotomously branched aparentemente sin hojas, en la base no ramificados,
distally; branches flattened or triangular in cross-section; distalmente muchas veces ramificados dicotómicamente;
leaves 1-2 mm long, inconspicuous, scale-like, alternate, ramas aplanadas or triangulares en sección transversal;
widely scattered; synangia sessile, 3-lobed, subtended hojas 1-2 mm de largo, inconspícuas, semejantes a
by a forked appendage, yellowish at maturity; spores escamas, alternas, ampliamente difundidas; sinángios
hyaline, reniform; x=52. sésiles, trilobados, subtendidos por un apéndice furcado,
amarillentos al madurar; esporas hialinas, reniformes;
x=52.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ningunos.

COMMENTS: Only two species of Psilotum occur in the COMENTARIOS: Sólo dos especies de Psilotum
Neotropics: P. complanatum Sw. and P. nudum (L.) Pal. They existen en el Neotrópico: P. complanatum Sw. and P.
can be distinguished by the cross-sections of their distal nudum (L.) Pal. Son distinguibles por la sección cruz de
branchlets. Psilotum complanatum has flattened, 2-ranked sus ramitas dístales. Psilotum complanatum tiene ramitas
branchlets; P. nudum has triangular, 3-ranked branchlets. aplanadas con dos ángulos agudos; P. nudum tiene ramitas
Hybrids between the two species are known from Hawaii triangulares con tres ángulos agudos. Híbridos entre
but have not yet been found in the Neotropics. Psilotum is los dos especies se conocen de Hawaii pero no han
extremely variable in habit and habitat. It can be epiphytic sido encontrados todavía en el Neotrópico. Psilotum es
or terrestrial, pendent or erect. A good habitat to look extremamente variable en habitu . Se puede ser epifito
for Psilotum is in the humus-filled leaf axils of palms. o terrestre, pendiente o erecto. Un buen hábitat buscar
The Psilotaceae is sister to the Ophioglossaceae. Both Psilotum es en las axilas de hojas de palmas las cuales han
have sporangia on the adaxial surface of the leaves and acumulado mucho humus. Las Psilotaceae son hermana a
cylindrical subterranean gametophytes. A polyploid series las Ophioglossaceae. Los dos tienen esporangios nacidos
is known for P. nudum in Australia and New Zealand en la superficie adaxial de hojas y gametofitos cilíndricos
(Barber, 1957), but this has not been investigated in the subterráneos. Una serie de poliploides se conoce para P.
Neotropics. The genus name is derived from the Greek nudum en Australia y Nueva Zelanda (Barber, 1957), pero
psilos, smooth or naked. The stems appear naked, without esto no ha sido investigado en el Neotrópico. El nombre
large leaves. genérico se deriva del griego psilos, desnudo o liso. Los
tallos aparecen desnudos, sin hojas grandes.

LITERATURE: Barber, H. N. 1957. Polypoidy in the Psilotales. Proceedings of the Linnean Society of New South Wales.
82:201–208.

440
Figure 214. Left: Psilotum nudum. Right: P. complanatum. In both species, note the trilocular synangia. En los dos especies, fijase
en los sinángios triloculares. (From Mickel & Smith, 2004)

441
DENNSTAEDTIACEAE
Pteridium Gled. ex Scop.
DESCRIPTION: Terrestrial; rhizomes deeply DESCRIPCIÓN: Terrestres; rizomas profundamente
subterranean, long-creeping, pubescent (not scaly); subterráneos, largamente rastreros, pubescentes (no
sterile and fertile leaves monomorphous; petioles about escamosos); hojas estériles y fértiles monomorfas;
equaling the lamina, at base producing epipetiolar buds; pecíolos casi igualando la lámina, produciendo brotes
laminae 3-pinnate to 4-pinnate, coriaceous, deltate epipeciolares en la base; láminas 3-pinnadas a
or pentagonal, pubescent (not scaly), sterile margins 4-pinnadas, coriáceas, deltadas o pentagonales,
reflexed; veins free; sori marginal, continuous (not pubescentes (no escamosas), márgenes estériles
interrupted) along the segment or lobe, supplied by many reflexos; nervaduras libres; soros marginales, continuos
veins; indusia 2, one formed by the reflexed margin of the (no interrumpidos) a lo largo del segmento o lóbulo,
lamina, the other sometimes absent, but when present suministrado por muchas venas; indusios 2, uno formado
delicate, hyaline, and hidden beneath the sporangia; por el margen reflejo de la lámina, el otro a veces ausente,
paraphyses absent; x=52. 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 GÉNEROS PARECIDOS: Algunas especies de
similarly divided laminae and marginal sori. They can be Pteridium tienen láminas divididas y soros marginales.
distinguished by their scaly stems, paraphyses among the Pueden distinguirse por sus tallos escamosos, parafisos
sporangia, plane (not reflexed) sterile margins, and short entre los esporangios, márgenes estériles planos (no
spines on the upper surface of the axes. reflexos) y espinas cortas en la superficie superior de
los ejes.

COMMENTS: Pteridium is an aggressive weed and often COMENTARIOS: Pteridium es una maleza agresiva y
forms dense populations by means of its subterranean a menudo forma poblaciones densas por medio de sus
rhizomes. It invades pastures where it is eaten by cows, rizomas subterraneos. Invade praderas donde se come
and the carcinogens know to be contained in its leaves por ganado, y los carcinagos en sus hojas se pasan en
may be pass on in the cow’s milk. An unusual characteristic la leche de las vacas.Una característica rara es que los
of Pteridium is that the sterile segments have enrolled segmentos estériles tienen márgenes recurvados como
margins like those of the fertile. In the past, bracken los de los segmentos fértiles. En el pasado, Pteridium ha
has been classified as a single species with 12 varieties sido clasificado como una sóla especie con 12 variedades
worldwide. Nowadays, most pteridologists prefer to mundial. Hoy en día, la mayoría de pteridólogos prefieren
recognize the varieties at the specific level. Four species reconocer las variedades al nivel específico. Cuatro
occur in the Neotropics.Their recognition is supported by especies se encuentran en el Neotrópico. Su recognición
morphology and DNA fingerprinting studies (Thomson se apoya por morfología y estudies de DNA (Thomson
et al., 2008). The genus name is derived from Greek, et al., 2008). El nombre genérico se deriva del griego,
pteridion, a small fern. pteridion, un helcho pequeño.

LITERATURE: 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).

442
Figure 215. A–C. Pteridium aquilinum. D–E. P. aquilinum var. pubescens. F. P. aquilinum var. latiusculum. G, J. P. arachnoideum. L,
M. P. caudatum. (Mickel & Smith, 2004)

443
PTERIDACEAE
Pteris L.
DESCRIPTION: Terrestrial; rhizome creeping to erect, DESCRIPCIÓN: Terrestres; rizoma rastrero a erecto,
scaly; leaves 1-pinnate to 4-pinnate; petioles with escamoso; hojas 1-pinnadas a 4-pinnadas; pecíolos con
a single omega-shaped vascular bundle; laminae un hace vascular en forma de una omega; láminas
glabrous or rarely pilose, usually with a conform or glabras or raras veces pilosas, usualmente con un segmento
subconform terminal segment; basal pinnae entire or, if terminal conforme o subconforme; pinnas basales enteras
divided, usually with the basal basiscopic pinnule enlarged; o, si divididas, usualmente con la pínnula basal basiscópica
axes of the penultimate segments in many species aristate agrandada; ejes de los penúltimos segmentos en muchas
adaxially; veins free or anastomosing and without included especies aristados adaxialmente; nervaduras libres o
veinlets; sori linear on a submarginal connecting vein, anastomosadas y sin nérvulas incluídas; soros lineares
covered by the reflexed margin of the lamina (false sobre una nervadura conectora submarginal,
indusium); paraphyses generally present, numerous; cubiertos por el margen reflexo de la lámina (falso
spores tetrahedral, with an equatorial flange; x=29. indusio); parafisos generalmente presentes, numerosos;
esporas tetrahédricas, con un reborde ecuatorial; x=29.

SIMILAR GENERA: None. GÉNEROS PARECIDOS: Ninguno.

COMMENTS: Pteris has about 250 species worldwide, COMENTARIOS: Pteris tiene casi 250 especies
about 60 of which are neotropical. It is heterogeneous, mundial, de las cuales casi 60 son neotropicales. Es muy
probably polyphyletic, and it is difficult to give heterogéneo, y probablemente polifiletico, y es difícil dar
characteristics that define the genus. Many species of características que definan el género. Muchas especies
Pteris have awns on the adaxial surface of the costules of de Pteris tienen aristas sobre la superficie adaxial de la
the penultimate segments. In these species, the ridge of costilla media de los penúltimos segmentos. En estas
the costule is interrupted at the juncture with the costule especies, la cresta de la costilla media se interumpe en la
of the next higher order and is prolonged into a short unión con la costilla media del orden superior siguiente y
awn. A well-marked group is that of Pteris vittata L., which se prolonga en una arista corta. Un grupo bien marcado
is characterized by 1-pinnate and cuneate laminae, and es él de Pteris vittata L., que se caracteriza por láminas
concolorous rhizome scales. Another such group is that 1-pinnadas y cuneadas, y escamas del rizoma concoloras.
of P. podophylla Sw., with huge pedate laminae. The genus Otro grupo asi es eso de P. podophylla Sw., con láminas
name is derived from Greek pteron, wing or feather. It is grandes pedatas. El nombre genérico se deriva del griego
an ancient Greek name for ferns in general, derived from pteron, ala o pluma. Es un nombre antiguo griego para los
a resemblance of the pinnae to wings or feathers. helechos en general, derivado de una semejanza de las
pinnas a las alas o plumas.

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.

444
Figure 216. A, B. Pteris paucinervata. C, D. P. muricella. E, F. P. pungens. (Mickel & Smith, 2004)

445
PTERIDACEAE
Pterozonium Fée
DESCRIPTION: Terrestrial or rupestral; rhizomes DESCRIPCIÓN: Terrestres o rupícolas; rizomas
short-creeping or ascending, either scaly, bristly, or hairy, cortamente rastreros o ascendentes, escamosos o
the indumenta usually dark brown or dark reddish brown; cerdosos o peludos, el indumento usualmente pardo
leaves monomorphic; laminae simple and entire oscuro o pardo rojizo oscuro; hojas monomorfas;
or 1-pinnate with a terminal pinna, coriaceous, láminas simples y enteras o 1-pinnadas con una
glabrous or nearly so; fronds or pinnae reniform, ovate pinna terminal, coriáceas, glabras o casi así; frondes
or oblong; veins free, parallel for most or all of their length; o pinnas reniformes, ovados o oblongos; nervaduras
rachises and costa grooved adaxially, with narrow lateral libres, paralelas en la mayor parte de su longitud, en
wings on either side; veins free, simple, in some species algunas especies terminando en hidatodos conspicuos;
ending in conspicuous hydathodes; sori elongate along soros alargados por las venas, no indusiados, raras
the veins, nonindusiate, rarely glandular, the glands veces glandulares, las glándulas amarillas a anaranjadas;
yellow to orange; spores tetrahedral-globose, trilete; x=? 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 neotropical y consta de 14 especies. En gran parte es
Guayana Highland of Venezuela but with isolated endémico a las Guayana Venezolana pero con poblaciones
populations in Costa Rica, Colombia, Guyana, Suriname, aislados en Costa Rica, Colombia, Guyana, Surinam, Perú
Peru, and northeastern Brazil. Mainly on sandstone rocks. y noreste de Brasil. Principalmente en rocas de piedra
Pterozonium is the sister genus to Jamesonia, and these arenisca. Pterozonium es el género hermana a Jamesonia
two genera are sister to Pityrogramma. All share non- y los dos son hermana a Pityrogramma. Todos comparten
indusiate sori with sporangia that run along the veins, and soros no indusiatos con esporangios que corren por las
the indumenta of the laminae are jointed hairs in stead of venas, e indumento de lás láminas de pelos articulados
scales. The genus name is derived from the Greek pteris, in lugar de escamas. El nombre genérico se deriva del
fern + zone, belt. The lamina has a broad submarginal griego pteris, helecho + zone, faja. La lámina tiene una faja
confluent band of sori. 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.

446
Figure 217. A. Pterozonium brevifrons (A. C. Sm.) Lellinger. B. P. reniforme (Mart.) Fée. C, D. P. spectabile Maxon & A. C. Sm.
(©Robbin Moran, 2009)

447
PTERIDACEAE
Radiovittaria (Benedict) E. H. Crane
DESCRIPTION: Epiphytic; rhizomes suberect, radial; DESCRIPCIÓN: Epífitas; rizomas suberectos, radiales;
rhizome scales clathrate; leaves 2–50 × 0.2–1(–1.8) escamas del rizoma clatradas; hojas 2–50 × 0.2–
cm, polystichous, simple, entire, linear to narrowly 1(–1.8) cm, polistichous, simples, enteros, lineales a
elliptic, sessile to stalked, chartaceous to fleshy, glabrous, estrechamente elípticas, sésiles o pecioladas, cartáceas a
with spicular idoblasts (false veins) adaxially; petioles carnosas, glabras, con idioblastos espiculares (venas falsas)
dark, flattened to terete; costa present; veins obscure, adaxialmente; pecíolos oscuros, achatados a teretes; costa
anastomosing to form one row of areoles and a presente; nervaduras obscuras, anastomosándose formar
submarginal vein, included veinlets absent; sori linear, una hilera de areolas y una vena submarginal, venas
submarginal, each one parallel to the margin, superficial incluidas ausentes; soros lineales, submarginales, cada
or sunken in grooves; indusia absent; paraphyses with uno paralela al margen, superficiales o hundidos en surcos;
a funnelform (obconic) apical cell; spores monolete; indusios ausentes; parafisos con una célula apical
gametophytes with single gemmae (not T-shaped); x=? 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, COMENTARIOS: Radiovittaria consta de casi 8 especies,
all neotropical. They occur as mostly as both low-trunk todas neotropicales. Ocurren como epífitos en las bases
epiphytes and in the canopies of wet forests. The species de troncos y en los doseles de bosques húmedos. Estas
of Radiovittaria were previously classified in Vittaria, but especies fueron clasificadas anteriormente en Vittaria,
according to a phylogenetic study by Crane (1997), such pero conforme a un estudio filogenético por Crane
a treatment leaves Vittaria polyphyletic. Radiovittaria is (1997), tal clasificación deja Vittaria polifilético. Radiovittaria
sister to Hecistopteris, and those two genera are sister es hermana a Hecistopteris, y estos dos géneros son
to Haplopteris, an Old World genus of about 20 species. hermanas a Haplopteris, un género del Viejo Mundo con
Vittaria is more distantly related within the vittarioid ferns. casi 20 especies. Vittaria es más distantamente relacionado
Most (all?) of the species in the genus have reddish young dentro de los helechos vittarioides. La mayoría (¿todas?)
leaves, a characteristic that it shares with Adiantum and las especies en el género tienen hojas jóvenes rojizas,
certain other vittarioids. The genus name refers to the una característica que comparta con Adiantum y ciertos
fact that this is the only genus of vittarioids ferns with otros vittarioides. El nombre genérico refiere al hecho
radially arranged (polystichous) leaves. 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.

448
Figure 218. Radiovittaria. A–C. R. stipitata. E–J. R. feei. J shows a paraphyses with an obconic apical cell. (Mickel & Smith, 2004)

449
DRYOPTERIDACEAE
Rumohra Raddi
DESCRIPTION: Terrestrial or rupestral; rhizomes DESCRIPCIÓN: Terrestres o rupícolas; rizomas
creeping, scaly; leaves monomorphic, borne alternately rastreros, escamosos; hojas monomorfas, se nacen
in two dorso-lateral rows on the rhizome; petioles alternamente in dos hileras dorsolaterales en
with three or more vascular bundles, these arranged el rizoma; pecíolos con tres o más haces vasculares,
in a U with the two adaxial ones largest; laminae 2- to estas arregladas en forma de U con los dos adaxiales
3-pinnate-pinnatifid, the apices gradually tapered and lo más grandes; láminas 2- o 3-pinnado-pinnatífidas,
pinnatifid (not imparipinnate); pinnules of the lower los ápices gradualmente reducidas y pinatífidas (no
pinnae arranged anadromically; rachises with two imparipinadas); pínnulas de las pinnas inferiores arregladas
adaxial grooves separated by a raised medial ridge; anadrómicamente; ráquises con dos surcos adaxiales
veins free; sori round; indusia peltate, circular; spores separados por una fila medial elevada; venas libres;
monolete; x=41. soros redondos; indusios peltados, circulares; esporas
monoletes; x=41.

SIMILAR GENERA: Polystichum differs by spinulose GÉNEROS PARECIDOS: Polystichum difiere por los
lamina margins and lacking the raised center in the rachis márgines de las láminas espinulosas y por la falta del centro
groove. Arachniodes differs by indusia attached by a sinus elevado en los surcos del ráquises. Arachniodes difiere por
(not peltate) and lacking the raised center in the rachis inducíos unídos por un sinus (no peltadas) y falta de un
groove. Lastreopsis differs by the upper surfaces of the centro elevado en el surco adaxial del raquis. Lastreopsis
rachises and costae rounded or flat (not grooved) and difiere por el haz de las ráquises y costas redondeado o
puberulent. Polybotrya differs by dimorphic sterile and achatado (no surcado) y puberulentos. Polybotrya difiere
fertile leaves. Saccoloma differs by an omega-shaped por hojas estériles y fértiles dimorfas. Saccoloma difiere
vascular bundle in the base of the petiole, and marginal por una haz vascular en forma de una omega en la base
or submarginal sori. del pecíolo y soros marginales o submarginales.

COMMENTS: Rumohra consists of about 8 species COMENTARIOS: Rumohra consta de casi 8 especies
worldwide, with one (R. adiantiformis (Forst. F.) Ching) mundial, con una (R. adiantiformis (Forst. F.) Ching) de
of circumaustral distribution. Two other species occur distribución circunaustral. Dos otras especies existen en
in South America: R. berteroana (Colla) Duek & Rodrig. América del Sur: R. berteroana (Colla) Duek & Rodrig.
is endemic to the Juan Fernández Islands, and another es endémica a las Islas Juan Fernández, y otra (R. turficola
(R. turficola Senna) is endemic to the southern cone of Senna) al Cono Sur de América del Sur. Las especies
South America. The species grow in a variety of habitats, existen en varios ambientes, incluyendo bosques, lugares
including forests, shrubby places, on rocks, and on sandy abustivos, sobre rocas e en suelos areniscos. Los surcos de
soils.The rachis grooves of Rumohra are distinctive among los ráquises de Rumohra son distinctivos entre helechos
dryopteroid ferns. The adaxial rachis surface has consists dryopteroides. La superficie adaxial del los requises
of two grooves with an elevated ridge between them. constan de dos surcos con una fila elevada entre sí. Los
Other dryopteriod ferns lack the medial elevated ridge. otros helechos dryopteroides faltan la fila elevada medial.
Rumohra is sister to Megalastrum, but the two genera Rumohra es hermano a Megalastrum, pero los dos géneros
do not share any obvious morphological similarities to no comparten ningunas simularidades morfológicas para
distinguish them from other dryopteroid ferns. Because distinguirlos de otros helechos dryopteroides. A causa de
its leaves remain green a long time after being cut, sus hojas permanezcan verde mucho tiempo después de
Rumohra adiantiformis is widely used in the florist trade. ser cortadas, Rumohra adiantiformis se usa ampliamente
The leading producers are Florida and Costa Rica. The para los ramos de flores. Los productores principales son
genus is named for Karl F. von Rumohr (1785-1843), an Florida y Costa Rica. El género se nombra para Karl F. von
art student from Dresden, Germany. Rumohr (1785-1843), un estudiante de arte de Dresden,
Alemania.
LITERATURE: 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.

450
Figure 219. Rumohra adiantiformis. (©Moran, 2007)

451
SACCOLOMATACEAE
Saccoloma Kaulf.
DESCRIPTION: Terrestrial or saxicolous; rhizomes DESCRIPCIÓN: Terrestres o saxícolas; rizomas
decumbent to erect, scaly, surrounded by persistant decumbentes o erectos, escamosos, envueltos por bases
petiole bases; sterile and fertile leaves monomorphous; de los pecíolos persistentes; hojas estériles y fértiles
petioles without buds at the base, with an omega-shaped monomorfas; pecíolos sin brotes epipeciolares, con un
(Ω) vascular bundle; laminae 1-pinnate to 5-pinnate; haz vascular en forma de una omega (W); láminas
rachises and costae grooved on the upper surface, 1-pinnadas a 5-pinnadas; ráquises y costas surcados
the grooves glabrous within; veins free; sori marginal or en el haz, los surcos glabros en su interior; nervaduras
submarginal, each at the apex of a vein, often slightly libres; soros marginales o submarginales, cada uno en
sunken in the laminar tissue and prominulous adaxially; el ápice de una vena, a menudo ligeramente hundidos en
indusia hyaline or scarious, usually obconic with a truncate el tejido laminar y promínulos adaxialmente; indusios
apex, opening toward the lamina margin; spores trilete, hialinos o escariosos, por lo común obcónicos con un
with distinctive pattern of parallel, anastomosed ridges; x ápice truncado, se abren hasta el margen de la lámina;
= ca. 63. esporas triletes, con un patrón distintivo de filas paralelas,
anastomosadas; x = ca. 63.

SIMILAR GENERA: Dennstaedtia differs by creeping, GÉNEROS PARECIDOS: Dennstaedtia se difiere por
pubescent rhizomes, leaves with at least some hairs, and rizomas reptantes pubescentes, hojas con al menos
deflexed cup-shaped sori. Unlike the Dennstaedtiaceae, algunos pelos y soros deflexos en forma de copa. A
Saccoloma lacks epipetiolar buds. Other ferns with highly diferencia de las Dennstaedtiaceae, Saccoloma carece de
divided laminae can be distinguished by the form and brotes epipeciolares. Otros helechos con láminas muy
position of their sori. divididas pueden distinguirse por la forma y posición de
sus soros.

COMMENTS: Saccoloma is the only genus in the COMENTARIOS: Saccoloma es el único género en
Saccolomataceae. It is pantropical and contains about 15 las Saccolomataceae. Es pantropical y consta de casi 15
species, 10 of which are neotropical.Whin the Neotropics, especies, de las cuales 10 existen en el Neotrópico. Todas
the genus occurs from Mexico to southern Brazil. All of las especies existen en bosques húmedos y sombreados.
the neotropical species in the genus have decompounds La especie más común, desde México hasta el sur de
laminae except for S. elegans Kaulf. (southeastern Brazil) Brasil, es S. inaequale (Kunze) Mett. Es muy variable y
and S. chartaceum G. B. Nair ex Cremers & K. U. Kramer probablemente consiste de varias especies no descritos.
(widespread). The genus name comes from the Latin La única especie neotropical con láminas 1-pinnadas
saccus, bag + loma, border. It refers to the marginal sori of es S. elegans Kaulf. El nombre genérico se deriva de la
the type species that are located in sac-like depressions. Latín saccus, bolsa + loma, margen. Se refiere a los soros
marginales de la especie tipo que están ubicados en
depresiones como bolsas.

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.

452
Figure 220. Saccoloma. A, B. S. inaequale. C, D. S. chartaceum. (from Tryon & Stolze, Pterid. Peru, 1989)

453
BLECHNACEAE
Salpichlaena J. Sm.
DESCRIPTION: Terrestrial; rhizomes creeping, scaly; DESCRIPCIÓN: Terrestre; rizomas reptantes,
mature leaves vinelike, climbing by means of a escamosos; hojas maduras como bejucos, trepando
twinning rachis; laminae 2-pinnate; pinnae with a por medio de un raquis que enrosca; laminae pinnate;
conform terminal pinnule; pinnules entire, the margins pinnas con pínnulas conformes terminales; pínnulas
cartilaginous-thickened; veins free, connected to the enteras, los márgenes engrosados cartilaginosamente;
cartilaginous margin; sori liinear, parallel along both nervadura libre, conectada al margen cartilaginoso; soros
sides of the costules; indusia present, often lacerate lineares, paralelos por ambos lados de las cóstulas;
with age, opening toward the costule (not the margin), indusios presentes, al menudo lacerados con la edad,
deciduous; x=40. abriéndose hasta la cóstula (no el margen), decíduo; x=40.

SIMILAR GENERA: Lygodium is the only other GÉNEROS PARECIDOS: Lygodium es el único otro
neotropical fern genus that has leaves climbing by género neotropical de los helechos que tiene hojas
means of a twinning rachis. It can be distinguished from que trepan por medio de un raquis enroscador. Puede
Salpichlaena by its pinnae forked at the base, absence of distinguirse de Salpichlaena por sus pinnas furcadas en la
scales on the axes, and sporangia borne in sorophores base, ausencia de escamas sobre los ejes, y esporangios
along the margin. que nacen en soróforos a lo largo del margen.

COMMENTS: Salpichlaena consists of three species and COMENTARIOS: Salpichlaena consta de tres especies
is entirely neotropical. It and Lygodium are the only fern y es completamente neotropical. Él y Lygodium son los
genera that have a twining rachis—a growth characteristic únicos géneros de los helechos que tienen ráquises
totally lacking in angiosperms. Salpichlaena volubilis (Kaulf.) trepadores—una característica de crecimiento que se
J. Sm. is common and widespread in the Neotropics. In falta completamente en las angiospermas. Salpichlaena
Amazonia, S. hookeriana Alston is also common and can volubilis (Kaulf.) J. Sm. es común y muy difundida en
be distinguished from S. volubilis by its fully contracted el Neotrópico. En la Amazonía, S. hookeriana Alston
fertile pinnules and the frequent presence of buds in the también es común y puede distinguirse de S. volubilis
pinna axils and sometimes those of the pinnules. Three por sus pínnulas fértiles completamente contraidas y
morphological notes: The cartilaginous margins of the por la presencia frecuente de brotes en las axilas de las
pinnules appear as thickened and lighter-colored than the pinnas y algunas veces en las de las pínnulas. Tres notas
laminar tissue. Unlike all other ferns, the sori and indusia morfológicas: Los márgenes cartilaginosos de las pínnulas
fall off as a unit. The juvenile leaves of Salpichlaena do not aparecen engrosados y de color más claro que el tejido
climb. They are important taxonomically because they laminar. Diferente de todos helechos, los soros e indusios
may show differences between the species. The genus caen juntos. Las hojas juveniles de Salpichlaena no trepan.
name is derived from the Greek salpinx, pipe, trumpet + Son importantes taxonómicamente por que pueden
chlaena, cloak. The indusium is tubular. mostrar diferencias entre especies. El nombre genérico
se deriva del griego salpinx, pipa, trompeta + chaleña,
capa. El indusio es tubular.

LITERATURE: None; ninguno.

454
Figure 221. Salpichlaena volubilis (from Tryon & Stolze 1993

455
SALVINIACEAE
Salvinia Micheli in Adans.
DESCRIPTION: Floating aquatics; roots absent; stems DESCRIPCIÓN: Acuáticas flotantes; raíces ausentes;
hairy, thin, branched; leaves in verticels of 3, dimorphous, tallos pelosos, delgados, ramificados; hojas en verticilios
with 2 green floating ones and single, whitish, submerged de 3, dimorfas, con 2 verdes flotantes y una sin clorofila,
one that is finely divided and resembles a root; floating sumergida, finamente dividida y semejante a una raíz;
leaves 0.4–3 cm in diameter, circular, ovate or hojas flotantes 0.4–3 cm de diámetro, redondeadas,
oblong, entire, typically with papillae on the upper ovadas u oblongas, enteras, típicamente con papilas
surface, the papillae when present with 3 or 4 hairs at en el haz, las papilas cuando presentes con 3 o 4 pelos en
their apices, these usually united and darkened at the sus ápices, estos usualmente unidos y oscurecidos en los
apex, forming a cage; veins anastomosing, the areoles ápices, formando una jaula; nervaduras anastomosadas, las
elongate, polygonal, without included veinlets; sori borne aréolas alargadas, poligonales, sin venillas includas; soros
on the submerged leaf; indusia globose, whitish; x=9. 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- consta de 9 especies. Se encuentra en agua estancada o
moving water at low elevations. The submerged leaf con flujo lento en elevaciones bajas. La hoja submergida
resembles a root because it is whitish and finely divided; se parece a una raíz porque es blanquecino y finamente
however, it is actually a leaf because it bears sporangia dividida; no obstante, es en realidad una hoja por que lleva
and its development and anatomy is that of a leaf, not a esporangios y su desarrollo y anatomía es eso de una hoja,
root. Most neotropical species have long papilla on the no de una raíz. La mayoría de las especies neotropicales
upper surface of the floating leaves, and these papillae tienen papilas largas en la superficie superior de las hojas
bear 4 hairs at their apex on top of each. In those species flotantes, y estas papilas llevan en sus ápices 4 pelitos. En
belonging to the S. auriculata complex, these hairs are las especies que pertenecen al complejo de S. auriculata,
united at their tips, and the entire structure, (papilla and estos pelitos se unen en sus ápices, y la estructura entera
its hairs) resembles an egg beater. The only neotropical (papila más pelitos) se asemaja un batidor de huevos.
species with hairs not united at the apices is Salvinia La única especies neotropical con pelos no unidos en
minima Baker. A few species lack papillae on the upper sus ápices es Salvinia minima Baker. Algunas especies
surface of the floating leaves, such as S. sprucei Kuhn. A well carecen de papilas en el haz de las hojas flotantes, tales
known species is Salvinia molesta D. S. Mitch., which is a como S. sprucei Kuhn. Una especie bien conocida es
aggressive weed introduced in Africa, Sri Lanka, Indonesia, Salvinia molesta D. S. Mitch., la cual es una maleza agresiva
Australia, and the southern United States (Moran, 2004). introducida en África, Sri Lanka, Indonesia, Australia, y
The genus name honors Antonio Maria Salvini (1633– el sur de los Estados Unidos (Moran, 2004). El nombre
1729), an Italian professor of Greek who helped Micheli genérico honora Antonio Maria Salvini (1633–1729), un
with his botanical work. 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.

456
Figure 222. A–D. Salvinia minima. E–K. S. auriculata. (Mickel & Smith, 2004).

457
SCHIZAEACEAE
Schizaea Sm.
DESCRIPTION: Terrestrial; stems compact, horizontal DESCRIPCIÓN: Terrestres; rizomas compacto,
or erect, pubescent; leaves simple to several times horizontal o erecto, peludos; hojas simples a varias veces
dichotomous, the ultimate divisiones foliaceous to scarcely dicotómicas, las últimas divisiones foliáceas a escasamente
laminar; petioles glabrous, as long or longer than the laminares; pecíolo glabro, tan largo como la lámina a más
lamina, with an adaxial groove; veins free; fertile segments largo que ella, con un surco adaxial; nervaduras libres;
(sporangiophores) pinnatifid, curved, borne at the segmentos fértiles (esporangióforos) pinnatos,
tips of the laminae; sporangia erect, oblong, with curvados, terminales; esporangios rectos, oblongos,
an apical annulus, in 2 rows; spores monolete, foveolate; con anillo apical, 2 hileras; esporas monoletes, foveolate;
gametophytes epigeal, green, filamentous; x=77, 94, 96, gametofitos epigeales, verdes, filamentosos; x=77, 94, 96,
103. 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. casi 45 especies, de las cuales 8 existen en el Neotrópico.
They typically grow below 600 m on shaded forest floors Típicamente crecen bajo de 600 m en el suelo de
or open sandy areas. The species with well-developed bosques sombreados or áreas arenosas abiertas. Las
laminae are sometimes separated as Lophidium, but it is especies con láminas desarolladas están separadas a
doubtful they represent a monophyletic group. Schizaea veces como Lophidium, pero es dudoso si representan un
elegans (Vahl.) Sw. has a broad, expanded lamina and is grupo monofilético. Schizaea elegans (Vahl.) Sw. tiene una
widespread in the Neotropics. Several species with grass- lámina expandida y está muy difundida en el Neotrópico.
like stems (e.g., S. incurvata Schkuhr) exhibit a clumped habit Varias especies con hojas en forma de gramíneas stems
caused by repeated proliferations from buds at the base (e.g., S. incurvata Schkuhr) exhiben un hábito amacollado
of the petioles.Schizaea has filamentous, gametophytes se debe a las proliferaciones de brotes en la base de
that form a mycorrhizal relationships. Specialized, paired, los pecíolos. Schizaea tiene gametofitos filamentosos
globose cells called rhiziodophores house the fungus. The que forman una relación micotrófica. Un par de células
sister genus Actinostachys has subterranean, cylindrical especializadas, llamados rizoidóforos, hospeden el hongo.
gametophytes. Schizaea is sometimes defined to include El género hermano Actinostachys tiene gametofitos
Actinostachys (which see). The genus name is derived subterráneos cilíndricos. A veces se define Schizaea para
from the Greek schizein, to split. The fan shaped fronds of incluir a Actinostachys (véase).El nombre genérico se
some species are split toward the base. deriva del griego schizein, partir o hender. Las hojas en
forma de un abanico de algunas especies son hendidas
hacia la base.

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. Takeuchi, M. 1960. O gênero Schizaea no Amazonia.
Boletim de Museo Paraense Emilio Goeldi 5: 1–26.

458
Figure 223. Schizaea pusilla, the curly-grass fern. A. Habit of plant, showing leaves of two forms: the sterile ones helically coiled,
the fertile (spore-producing) straight and tall. B. Lower surface of the sterile leaf with two rows of stomata (almost all other ferns
have randomly scattered stomata). C. Fertile blade, folded lengthwise. D. Lower surface of pinna detached from fertile blade,
showing sporangia with apical annuli. (©Robbin Moran, 2009)

459
PTERIDACEAE
Scoliosorus T. Moore
DESCRIPTION: Epiphytic; rhizomes short-creeping DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
solenostelic, short-creeping, dorsiventral; rhizome scales solenostélicos, dorsiventrales; escamas del rizoma
clathrate; leaves 30–70 cm long × 1.5–20 cm wide, clatradas; hojas 15–70 × 1.5–20 cm, simples, enteras,
simple, entire, glabrous, fleshy, distichous, clumped; glabras, carnosas, dísticas, amacolladas; pecíolos cortos
petioles short or absent; costae present or absent; veins a ausentes; costas presentes o ausentes; nervaduras
areolate, with several series of areoles between the areoladas, con varias series de areolas entre la costa y el
costa and margin; sori oblique to the costae; indusia margen; soros oblícuos a la costa; indusios ausentes;
absent; paraphyses with a spherical apical cell; parafisos con una célula apical esférica; ssporas
spores monolete, nongreen; gametophytes with paired monoletes, no verdes; gametofitos con yemas pareadas;
gemmae; x=? x=?

SIMILAR GENERA: Polytaenium differs by trilete spores GÉNEROS PARECIDOS: Polytaenium difiere por
and lack of paraphyses. The rachis of this species wide, esporas triletes y falta de parafisos. El raquis de esta
flattened (not or only slightly raised) and often much especie es ancho, achatado (no o poco levantado) y
lighter in color than the lamina tissue on either side. In a menudo más claro en color que el tejido laminar en
Polytaenium the rachis is more prominent. Ananthacorus, ambos lados. En Polytaenium el rachis es más prominentes.
Radiovittaria, and Vittaria differ by a single, long submarginal Ananthacorus, Radiovittaria y Vittaria difieren por un sólo
sorus. soro largo submarginal.

COMMENTS: Scoliosorus comprises three species, one COMENTARIOS: Scoliosorus consta de tres especies,
of which grows from Mexico to Panama (S. ensiforme una de las cuales existe desde México hasta Panamá
(Hook.) T. Moore). The other two occur in Africa and (S. ensiforme (Hook.) T. Moore). Las dos otras existen
nearby islands of the Indian Ocean. The neotropical en África e islas cercanas del Océano Indio. La especie
species grows as an epiphyte in wet, shaded forests at neotropical crece como epífito en bosques húmedos
middle elevations. The genus name is derived from the sombreados en elevaciones medias. El nombre genérico
Greek scolios, curved, tortuous + soros. The sori are se deriva del griego scolios, curvado, tortuo + soros. Los
linear and more or less sinuous. 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.

460
Figure 224. A–F. Scoliosorus ensiforme. (Mickel & Smith, 2004)

461
POLYPODIACEAE
Serpocaulon A. R. Sm.
DESCRIPTION: Epiphytic; rhizomes long‑creeping, DESCRIPCIÓN: Epífitas; rizomas largamente
but short-creeping in a few species, the phyllopodia rastreros, pero cortamente rastreros en algunas especies,
(0.5–)5–20 times the rhizome width apart, sometimes los filopodios (0.5–)5–20 veces distante la anchura del
glaucous; black sclerenchyma strands absent; vascular rizoma; esclerínquima negra ausente; haces vasculares
bundles sometimes with sclerenchyma sheaths; scales a veces con vainas de esclerínquima; escamas peltadas,
peltate, clathrate (sometimes the luminae are occluded), clatradas (a veces la luminas son ocluidas), las paredes
the side walls dark brown or black, surfaces glabrous laterales pardos o negros, superficies glabras (carece de
(lacking rhizoids). Petioles grooved adaxially (never rizoides); pecíolos surcados adaxialmente (nunca teretes);
terete); laminae pinnatifid to pinnate, rarely simple (S. láminas pinatífidas a pinnadas, raras veces simples (S.
levigatum) or shallowly pinnatifid, monomorphic; lamina levigatum) o someramente pinatífidas, monomorfas; tejido
tissue glabrous or hairy, without scales or with sparse de las láminas glabro o pubescente, sin escamas o con
clathrate scales along the costae and a few on the escamas clatradas esparcidas por las costas y algunas sobre
rachises; veins regularly anastomosing, the areoles los requises; venas anastomosándose regularmente,
with a single, free, included vein (goniophlebioid las areolas con una sóla venilla incluida (nervación
veins). Sori round or nearly so, in 1–10 rows between gonofleboide); soros redondos o casi así, en 1–10 hileras
costae and margins, terminal on a free included vein; entre la costa y margen, terminales en el ápice de
indusia absent; paraphyses absent or present and short, una vena incluida; inducíos ausentes; esporas amarillas,
2--3-celled, glandular, not easily seen; sporangia glabrous; monoletes; x=37.
spores yellowish, monolete; x=37.

SIMILAR GENERA: Polypodium differs by (usually) free GÉNEROS PARECIDOS: Polypodium difiere por
veins, non-clathrate rhizome scales, and single row of sori (usualmente) venas libres, escamas del rizoma no
between the costa and margin. Pleopeltis differs by scaly clatradas y una sola hilera de soros entre la costa y el
laminae. Pecluma differs by free veins, or if anastomosing margen. Pleopeltis difiere por láminas escamosas. Pecluma
(rare) then not in a goniophelbioid pattern, a single row difiere por venas libres, o si raras veces anastomosadas
of sori on each side of the costae, often reduced proximal entonces no en un patrón goniofleboide, un sola hilera
pinnae, non-clathrate, often comose rhizome scales, de soros en cada lado de la costa, a menudo pinnas
short-creeping rhizomes, and pinnae usually more than proximales reducidas, escamas del rizoma no clatradas
30 pairs. y a menudo comosos, rizomas cortamente rastreros y
pinnas usualmente más de 30 pares.

COMMENTS: Serpocaulon contains about 40 species, all COMENTARIOS: Serpocaulon contiene casi 40 especies,
neotropical. About 14 species occur in the Antilles and todos neotropicales. Casi 14 especies existen en las
Central America, and 26 are restricted to South America, Antillas y América Central, y 26 son restingidas al América
10 of which occur in southeastern Brazil. All grow del Sur, 10 de las cuales existen en el sureste de Brasil.
primarily as epiphytes in wet forests from about 0–2500 Todos crecen principalmente como epífitos en bosques
m. The generic name comes from the Latin serpens, snake húmedos de 0–2500 m. El nombre genérico se deriva
+ caulon, stem, referring to the long-creeping rhizome. 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. 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.

462
Figure 225. A–C. Serpocaulon pleurosorum. D–E. S. falcaria. F. S. fraxinifolium. G–J. S. triseriale. L–M. S. dissimile. (Mickel &
Smith, 2004).

463
CYATHEACEAE
Sphaeropteris Bernh.
DESCRIPTION: Terrestrial; stems arborescent, up DESCRIPCIÓN: Terrestres; tallos arborescentes,
to 15 m tall; sterile and fertile leaves 2-3(-5) m long, hasta 15 m de alto; hojas estériles y fértiles 2-3(-5) m
monomorphous; petioles without spines, scaly, the scales de largo, monomorfas; pecíolos sin espinas, escamosos,
1–4 cm long, linear, concolorous, white or pale las escamas 1–4 cm de largo, lineares, concoloras,
brownish, the cells of the borders and center equal blancas o cafés claros, las células de los bordes y del
in shape and orientation, margins dark-denticulate; centro iguales en forma y orientación, los márgenes
laminae 2-pinnate-pinnatisect, the apex gradually tapered, oscuro-denticulados; láminas 2-pinnado-pinnatisectas,
pinnatisect; rachis and costae pubescent adaxially, the el ápice gradualmente reducido, pinnatisecto; raquis y
hairs thick, strigose, multiseptate; veins free; sori round, costas pubescentes adaxialmente, los pelos gruesos,
the receptacle elevated, subglosbose; indusia cup-shaped estrigosos, multiseptados; nervaduras libres; soros
or globose; spores tetrahedral-globose, usually 64 per redondos, el receptáculo elevado, subgloboso; indusios en
sporangium, without three large equatorial pores; x=69. forma de copa o globosos; esporas tetrahédrico-globosas,
64 por esporangio, sin poros grandes ecuatoriales; x=69.

SIMILAR GENERA: All other genera in the family have GÉNEROS PARECIDOS: Todos los otros géneros en
petiolar scales with the cells of the margin differing in la familia tienen escamas del pecíolo con las células de los
shape and orientation from those of the center. Dark- bordes diferentes en forma y orientación a las del centro.
denticulate margins occur only in a few species of Cyathea. Márgenes oscuro-denticulados se encuentran en sólo
Many species of Cyathea have spiny petioles, whereas pocas especies de Cyathea. Muchas especies de Cyathea
Sphaeropteris always have nonspiny petioles. tienen pecíolos espinosos, mientras que Sphaeropteris
siempre tiene pecíolos no espinosos.

COMMENTS: Sphaeropteris is pantropical and contains COMENTARIOS: Sphaeropteris es pantropical y consta

about 110 species. In the neotropics, it has 6 species, de casi 110 especies. En el neotrópico, consta de 6
called the Sphaeropteris horrida group, monographed by especies, llamadas el grupo de Sphaeropteris horrida, que
Tryon (1971).The genus as here defined excludes subgen. fue monografiado por Tryon (1971). El género como
Sclephropteris which is placed in Cyathea. Sphaeropteris is definido aquí excluye subgén. Sclephropteris, lo que está
sister to a clade consisting of Alsophila and Cyathea (Korall ubicado en Cyathea. Sphaeropteris es hermana a un clado
et al., 2006). The genus name is derived from the Greek formado de Alsophila y Cyathea (Korall et al., 2006). El
sphaeros, sphere + pteris, fern. It refers to the spherical nombre genérico se deriva del griego sphaeros, esfera +
indiusia. pteris, helecho. Refiere a los indusios esféricos.

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. Tryon,
R. M. 1973. The American tree ferns allied to Sphaeropteris horrida. Rhodora 73: 1–19.

464
Figure 226. B, C. Sphaeropteris quindiuensis. C. Petiole scale showing conform (size shape) cells. (A. is Cyathea poeppigii; from
Tryon & Stolze, 1989).

465
POLYPODIACEAE
Stenogrammitis Labiak
DESCRIPTION: Rhizomes erect, scaly, the scales DESCRIPCIÓN: Rizomas erectos, escamosos, las
entire, clathrate, entire but usually with a single apical escamas enteras, clatradas, enteras pero usualmente
cilia; petioles short, glarous to slightly hirsute, the hairs con una cilia corta apical; pecíolos cortos, glabros a
simple or 1-furcate; laminae 2–5 mm wide, linear, levemente hirsutos, los pelos simples o 1-furcados;
monomorphic or hemidimorphic, erect or slightly laminas 2–5 mm de ancho, lineares, monomórficas
arcuate, subcoriaceous, pinnatifid to pinnatisect, when or hemidimórficas, erectas o levemente arcuatas,
hemidimorphic the fertile portion less divided than the subcoriaceas, pinnatífidas a pinatissectas, cuando
sterile; rachises obscured by the laminar tissue or with the hemidimorficas, entonces las partes fértiles usualmente
dark sclerenchyma exposed, glabrescent to hirsute, the menos divididas que las estériles; raquises escondidos
hairs hyaline or reddish, simple to 1-furcate; segments to por el tejido laminar o visible con su esclerenchima
5 × 1 mm, short decurrent basiscopically, apices acute to expuesta, glabrescentes a hirsutos, los pelos hialinos o
obtuse, glabrous or hirsute, the hairs simple or 1-furcate; rojizos, simples o 1-furcados; segmentos hasta 5 × 1 mm,
veins simple, hidden in the sterile portion of the cortamente decurrente basiscopicamente, los ápices
laminae but visible and dark in the fertile portions; agudos a obtusos, glabros o hirsutos, los pelos simples o
hydathodes present, but lacking calcareous secretions; 1-furcados; venas simples, escondidas en las partes
sori one per segment, oblong to elliptic, glabrous; estériles de las láminas pero visibles y oscuras
sporangia glabrous; spores trilete, green; x=33. 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 GÉNEROS PARECIDOS: Lellingeria difiere por
(usually > 1 cm), several veins per segment, usually several laminas más anchas (usualmente > 1 cm), venas
sori per segment, and x=37. Leucotrichum differs by ciliate pinnadas, usualmente varios soros por segmento y
rhizome scales, whitish hairs of the leaves, and several x=37. Leucotrichum difiere por escamas de los rizomas
veins per segment. 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.

466
Figure 227. 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)

467
GLEICHENIACEAE
Sticherus C. Presl
DESCRIPTION: Terrestrial; rhizomes long-creeping, DESCRIPCIÓN: Terrestres; rizomas largamente
scaly, protostelic; leaves indeterminate, of periodic growth, rastreros, escamosos, protostélicos; hojas indeterminadas,
once to several times forked pseudodichotomously, rarely de crecimiento periódico, una a varias veces
(in S. simplex (Desv.) Ching) simple, bearing pairs of pseudodicotómicamente furcadas, raras veces (en S.
opposite pinnae, the distalmost pair with a scaly resting simplex (Desv.) Ching) simples, llevando pares opuestas de
bud between them; pinnae forked pseudodichotomously, pinnas, el par más distal con una yema latente escamosa
with a scaly bud between the divisions; penultimate entre sí; pinnas furcadas pseudodicotómicamente, con
segments pectinate; veins 1-forked, free; sori round, not una yema escamosa entre las divisiones; segmentos
indusiate; sporangia 3–5(-6) per sorus, sessile or nearly penúltimos pectinados; venas una vez furcadas, libres;
so, opening by a transverse slit that passes through the soros redondos, no indusiados; esporangios 3–5(-6) por
apex; spores monolete; x=34. 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 GÉNEROS PARECIDOS: Dicranopteris y Gleichenella
from Sticherus by bristly rhizomes, pubescent pinna buds difieren de Sticherus por rizomes cerdosos, yemas de
(not scaly), veins 2–4 times forked, and sporangia 8–15 las pinnas pubescentes (no escamosas), venas 2–4 veces
per sorus. Gleichenella further differs by solenostelic furcadas, y esporangios 8–15 por soro. Gleichenella difiere
rhizomes. 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. casi 90 especies, con alrededor de 40 en América tropical.
Like other genera in the family, it often grows in open Como otros géneros en la familia, crece a menudo en
disturbed habitats such as roadsides. Many pteridologists ambientes abiertos y pertubados tales como taludes.
place Sticherus in Gleichenia. Sticherus is here considered Muchos pteridólogos ubican Sticherus en Gleichenia. Aquí
different enough in laminar division, spores shape, and se considera Sticherus bastante diferente en división
chromosome number (34 vs. 20 or 22) to merit a laminar, forma de esporas, y número cromosómico (34
distinct genus. Gleichenia is restricted to southern Africa vs. 20 o 22) para merecer un género distinto. Gleichenia
and Australasia. The genus name comes from the Greek está restringido al sur de África y Australasia. El nombre
stichos, line or row. The sori are born in rows. 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.

468
Figure 228. Sticherus bifidus. (Mickel & Smith, 2004).

469
DRYOPTERIDACEAE
Stigmatopteris C. Chr.
DESCRIPTION: Plants terrestrial; rhizome short- DESCRIPCIÓN: Plantas terrestres; rizoma corto-
creeping to erect; laminae 1-pinnate to 2-pinnate-pinnatifid, reptante a recto; láminas 1-pinnadas a 2-pinnada-
the apex pinnatifid, the tissue usually membranaceous, pinnatífidas, el ápice pinnatífido, el tejido usualmente
with internal punctate glands, lacking hairs (but often membranáceo, con glándulas punctadas internas, carece
with reduced, uniseriate scales that may resemble hairs); de pelos (pero a menudo con escamas uniseriadas que
pinnae apices long-attenuate, serrate; rachis and costae simulan pelos); ápices de las pinnas largo-atenuadas,
lacking hairs abaxially or (in 2 spp.) with minute capitate- aserradas; raquis y costas carece de pelos o (en 2 spp.)
glandular hairs, scaly, the scales thin, ciliate or papillate, con glandulas diminutas capitadas, escamosos, las escamas
the cilia usually ending in a glandular cell, the adaxial delgadas, ciliadas o papiladas, las cilias terminando
surface grooved, the grooves pubescent within, the hairs en una célula glandular, la superficie adaxial surcada,
ca. 0.1 mm long, 1–3-celled; veins ending before the los surcos pubescentes adentro, los pelos ca. 0.1 mm
margin in a conspicuous hydathode, free or (in 3 spp.) de largo, 1–3-celulares; venas terminando antes del
anastomoming; sori round; indusia absent; x=41. margen en un hidatodo conspicuo, libres o (en 3 spp.)
anastomosantes; soros redondos; indusios ausentes;
x=41.

SIMILAR GENERA: Cyclodium differs by the usually GÉNEROS PARECIDOS: Cyclodium difiere por los
indusiate sori, thicker laminae without internal punctate soros usualmente con indusios, las láminas más gruesas
glands, and entire scales. An important tendency is that y sin glándulas punctadas internas, y escamas enteras.
most species of Cyclodium have only a few scales on the Una tendencia importante es que la mayoría de las
axes abaxially, in contrast to Stigmatopteris which is usually especies de Cyclodium tienen pocas escamas sobre sus
scaly. ejes abaxialmente, en contraste a Stigmatopteris que es
usualmente escamoso.

COMMENTS: Stigmatopteris is entirely neotropical, COMENTARIOS: Stigmatopteris es completamente

occurring from Mexico to southern Brazil. It is primarily neotropical, extendiéndose desde México hasta el sur de
montane, from 500–2000 m, and is absent from most Brasil. Es principalmente de las montañas, de 500–2000
of Amazonia. The genus contains 24 species. The internal m, y es ausente de la mayoría de Amazonía. El género
punctate glands in the lamina when the leaf tissue is consta de 24 especies. Pueden verse mejor las glándulas
backlit and examined with a handlens. In herbarium punteadas internas en contraluz con una lupa. Con la
specimens in oblique light, the glands often appear (under luz oblicua en ejemplares montados del herbario, las
a microscope) as minute raised bumps. The glands often glándulas parecen (con un microscopio) como puntitos
dry blackish after pressing.The genus name is derived from elevados. A menudo se vuelven negruzcas al secarse
the Greek stigmatos, punctate + pteris, fern. It refers to después de ser prensadas. El nombre genérico se deriva
the characteristic internal punctate glands of the laminae. del griego stigmatos, punctata + pteris, helecho. Se refiere
a las glándulas características internas punteadas de las
láminas.

LITERATURE: Moran, R. C. 1991. Monograph of the Neotropical fern genus Stigmatopteris (Dryopteridaceae). Annals of the
Missouri Botanical Garden 78: 857–914.

470
Figure 229. Stigmatopteris. A, B. S. heterophlebia (Baker) R. C. Moran. C. S. nephrodioides, adaxial view, note hathodes. D. Punc-
tate 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. (©Robbin Moran, 2009)

471
POLYPODIACEAE
Synammia C. Presl
DESCRIPTION: Terrestrial, saxicolous, or epiphytic; DESCRIPCIÓN: Terrestres, saxícoloas o epífitas;
rhizomes pruinose, long-creeping; rhizome scales rizomas pruinosas, largamente rastreros; escamas
peltate, 2-3 mm wide, thin, ovate, orangish but del rizoma peltadas, 2-3 mm de ancho, delgadas,
blackened toward point of attachment; sterile and ovadas; hojas estériles y fértiles monomorfas; láminas
fertile leaves monomorphous; laminae 1-pinnate to 1-pinnado a 2-pinnado-pinnatífidas, pubescentes o con
2-pinnate-pinnatifid; pinnae adnate to the rachis, margins escamas densas o esparicas; pinnas adnatas al raquis,
cartilaginous-thickened and minutely notched; veins margines cartilaginosos, engrosados, con pequeñas
inconspicuous, more than once forked, anastomosing to hendiduras; venas inconspicuas, más que una vez surcada,
form a single row of costal areolae; areoles with a single anastomosándose, formando una hilera de areolas
included veinlet; sori round to elongate, borne on the tips costales; areolas con una sola venilla incluida; soros
of the included veinlets; indusia absent; paraphyses dense, alargados, se nacen en los puntos de las venillas; inducíos
filiform; spores yellow, monolete. ausentes; parafisos densos, filiformes; esporas amarillas,
monoletes.

SIMILAR GENERA: Pleopeltis differs by peltate scales on GÉNEROS PARECIDOS: Pleopeltis difiere por
the blades and (usually) among the sporangia. Polypodium escamas peltadas en las láminas y (usualmente) entre los
differs by non-pruinose rhizomes, (often) free veins and esporangios. Polypodium difiere por rizomas no pruinosos,
sporangiasters (capitate hairs) among the sporangia. (a menudo) venas libres y esporangiastros (pelitos
capitados) entre los esporangios.

COMMENTS: Synammia consists of four species in COMENTARIOS: Synammia consta de four especies
temperate South America, in Chile, Argentina, and the en las regiones templadas de América del Sur, en Chile,
Juan Fernandez Islands (Rodríguez, 1995). They grow Argentina, and las Islas Juan Fernández. Principalmente
principally in wet forests. Schneider et al. (2006) found existen en bosques húmedos. Schneider et al. (2006)
that Synammia is strongly supported as monophyletic. It encontraron que Synammia es fuertemente apoyado
is either sister to the rest of the Polypodiaceae or to all como monofilético. Es hermana al resto de Polypodiaceae
other neotropical members of the family. Synammia is a todos las otras especies de la familia. =Synammia is
derived from the Greek syn, together + ammatos, knot derivada de la Griega syn, conjunto + ammatos, knot or
or loop, referring to the veins that form a row of costal loop, referiéndose a las venas que forman una hilera de
areoles. areolas costales.

LITERATURE: Looser, G. 1952. El género Polypodium L. y sus representantes chilenos. Revista Univ. (Santiago) 36: 13–82.
Rodríguez, R. 1995. Polylpodium. 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.

472
Figure 230. A–D. Synammia sp. E–F. S. intermedia ssp. masafuerana. H–J. S. feuillei. (©Robbin Moran, 2014)

473
TECTARIACEAE
Tectaria Cav.
DESCRIPTION: Terrestrial or saxicolous; stems DESCRIPCIÓN: Terrestres o saxícolas; tallos erectos a
erect or long-creeping, scaly; sterile and fertile leaves largamente rastreros, escamosos; hojas estériles y fértiles
monomorphous, rarely dimorphous; laminae simple monomorfas, raras veces dimorfas; láminas simples y
and entire to 3-pinnate-pinnatifid; basal pinnae often enteras a 3-pinnado-pinnatífidas; pinnas basales a
with a prolonged lobe or pinnule on the basal menudo con un prolongado lóbulo o pínnula en el
basiscopic side; rachis and costae rounded or shallowly lado basal basiscópico; raquis y costas redondeados
grooved on the upper surface, the grooves not decurrent, o someramente sulcados en el haz, los surcos no
puberulent on the upper surface, the hairs 0.1-0.2 mm decurrentes, puberulentes en el haz, los tricomas 0.1-
long, reddish, pluricellular; veins reticulate, the areoles 0.2 mm de largo, rojizos, pluricelulares; venas reticuladas,
typically with included veinlets, or free in T. brauniana las aréolas típicamente con venillas incluidas, o
(H. Karst.) C. Chr.; sori round or (in T. athyrioides (Baker) libres en T. brauniana (H. Karst.) C. Chr.; soros redondos
Mett.) irregular or J-shaped; indusia present, rarely absent; o (en T. athyrioides (Baker) Mett.) irregulares o en forma
spores bilateral, brown; x=40. de “J”; indusios presentes, raras veces ausentes; esporas
bilaterales, pardas; x=40.

SIMILAR GENERA: Bolbitis differs by acrostichoid sori, GÉNEROS PARECIDOS: Bolbitis difiere por soros
axes of the lamina glabrous on the upper surface, and an acrosticoides, ejes de la lámina glabros en el haz, y un haz
elongated ventral vascular bundle in the rhizome (seen in vascular alargado ventral en el rizoma (visto en sección
cross-section). transversal).

COMMENTS: Tectaria is pantropical and subtropical COMENTARIOS: Tectaria es pantropical y subtropical

with about 200 species. Approximately 30 species occur y consta de casi 200 especies. Aproximadamente 30
in the New World and urgently needs a monograph especies existen en el Nuevo Mundo y necesitan una
because many taxonomic and nomenclatural problems monografía, ya que muchos problemas taxonómicos y
require solution. Most of the species occur in wet de nomenclatura requieren solución. La mayoría de las
forests from 0–1500 m, but some are characteristic of especies existen en bosque húmedos de 0–1500 m,
drier habitats and occur primarily on limestone. The pero algunas caracterizan ambientes más secas y existan
genus is sister to Triplophyllum.Tectaria contains Tectaria principalmente sobre piedra caliza. El género es hermana
panamensis (Hook.) R. M.Tryon & A. F.Tryon is sometimes a Triplophyllum. Tectaria panamensis (Hook.) R. M. Tryon &
put in its own genus, Dictyoxiphium, because of its simple A. F. Tryon es algunas veces puesto en su propio género,
leaves and submarginal sori. This species hybridizes with Dictyoxiphium, a causa de sus hojas simples y soros
Tectaria incise (see section on hybrid in this manual). The submarginales (vea la sección sobre híbridos en este
genus name is derived from the Latin tectum, roof + aria, manual). El nombre genérico se deriva del latín tectum,
a substantive suffix. It alludes to the roof-like indium in techo + aria, un sufijo sustantivo. Alude al indusio que es
many species. 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.

474
Figure 231. A: Tectaria panamensis. B–D. T. heracleifolia. E–G. T. mexicana. H–L. T. incisa. (Mickel & Smith, 2004)

475
POLYPODIACEAE
Terpsichore A. R. Sm
DESCRIPTION: Epiphytic; rhizomes short-creeping, DESCRIPCIÓN: Epífitas; rizomas cortamente rastreros,
compact; rhizome scales setulose on margins and compactos; escamas del rizoma setulosas por los
often on the surfaces, not clathrate, orange to brown, márgenes y a menudo las superficies, no clatradas,
basifixed (not cordate); petioles ca. half to equaling anaranjadas a pardas, basifijas (no cordadas); pecíolos
the length of the laminae, setose (especially basally), ca. una mitad hasta igual la longitud de las láminas,
the setae 0.5–3 mm long, generally castaneous, numerous, setosos (en especial basalmente), las setas 0.5–3 mm de
spreading, the hairs 0.1–0.2 mm long; laminae determinate, largo, generalmente castañas, numerosas, patentes, los
pinnatisect to 1-pinnate, rarely 1-pinnate-pinnatifid, tricomas 0.1–0.2 mm de largo; láminas pinnatisectas a
without black clavate fungi, setose, the setae simple (not 1-pinnadas, raramente 1-pinnado-pinnatífidas, sin hongos
branched); hydathodes present, without whitish chalk clavados negros, setosas, las setas simples (no ramificadas);
deposits; veins simple, free; sori separate, round, superficial hidatodos presentes, sin puntos blanquecinos calizos;
(i.e., not sunken in the lamina); sporangial capsules nervaduras simples, libres; soros separados, redondos,
setose (except T. atroviridis); spores green, trilete. x=37. superficiales (i.e., no hundidos en la lámina); cápusulas
esporangiales setosos (excepto T. atroviridis) esporas
verdes, triletes. x=37.

SIMILAR GENERA: Alansmia differs by indeterminate GÉNEROS PARECIDOS: Alansmia difiere por hojas
leaves, short petioles, laminar setae often paired, branched indeterminadas, pecíolos cortos, setas de las láminas a
or stellate, and blades tapered toward the base. The menudo pareadas, ramificadas o esteladas y láminas
Terpsichore taxifolia group typically has black costae, black reducidas hacia la base. El grupo de Terpsichore taxifolia
club-shaped fungi on the lower surface of the laminae, tiene costas negras y hongos negros claviformes en el
white hydathodes, and glabrous sporangial capsules. envés de la lámina, hidatodos blanquecinos y cápsulas
Pecluma differs by monolete, yellowish spores and lacking esporangiales glabras. Pecluma difiere por monoletes
setae on the petioles and blades. amarillentosas y la falta de setas en los pecíolos y láminas.

COMMENTS: Terpsichore is entirely neotropical and COMENTARIOS: Terpsichore es completamente

contains 12 species, all of which are epiphytes in wet neotropical y consta de 12 especies, todas epífitas en
shaded forests from low to high elevations. Many of the bosques húmedos sombreados de elevaciones bajas
species have arching or horizontal petioles from which a altas. Muchas especies tienen pecíolos arcuados o
the laminae are abruptly pendent. Although occurring on horizontales desde que las láminas son abruptamente
various substrates, Terpsichore species are often common pendientes. Aun que existiendo en sustratos varios,
on the root mantles of tree fern trunks. The genus is las especies de Terpsichore son a menudo frecuentes
recognized here in the strict sense, with the removal of sobre los manteles de raíces en los troncos helechos
Alansmia and the T. taxifolia group.These groups, originally arborescentes. Se reconoce el género aquí en el sentido
included in Terpsichore (Smith, 1993), have been shown stricto, quitando Alansmia y el grupo de T. taxifolia. Estos
to be distinct lineages more closely related to other grupos, originalmente incluidos en Terpsichore (Smith,
grammitid genera (Ranker et al., 2004). Terpsichore is 1993), han sido mostrado ser líneas evolutivas distinctas
named for the Greek Muse of dance and choral singing. más relacionadas a otros géneros grammitoides (Ranker
The leaves appear to “dance” when the wind blows. 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.

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.

476
Figure 232. A–G. Terpsichore chrysleri. H–L. T. lehmanniana. O, P. T. asplenifolia. (A–L © R. C. Moran, 2010; O, P from Mickel
& Smith, 2004)

477
THELYPTERIDACEAE
Thelypteris subgen. Amauropelta (Kunze) A.R. Sm.
DESCRIPTION: Terrestrial; rhizomes erect to suberect, DESCRIPCIÓN:Terrestres; rizomas erectas a suberectas,
rarely creeping or decumbent; leaves monomorphous; raras veces reptantes o decumbentes; hojas monomorfas;
petioles bases with 2 vascular bundles; laminae las bases de los pecíolos con 2 haces vasculares;
1-pinnate-pinnatifid, reduced toward the base, with láminas 1-pinnato-pinnatífidas, reducidas hacia la
the apex gradually reduced (not conform); aerophores base, con el ápice gradualmente reducido (no conforme);
present or rarely absent; proximal pinnae gradually or aeróforos presentes o raras veces ausentes; pinnas
abruptly reduced, sometimes to vestigial pinnae; veins proximales gradualmente o abruptamente reducidas, a
meeting the margin above the sinus, never united veces a pinnas vestigiales; venas encontrándose con el
below the sinus, usually simple (not forked); indumentum margen por arriba del seno, nunca unido por abajo del
of needle-shaped (rarely hooked) hairs; sori round, rarely seno, simples (no furcadas); pelos en forma de aguja, raras
elongate, indusiate or not; sporangia glabrous, rarely veces ganchudo; soros redondos, raras veces elongados,
setose; spores finely reticulate; x=29. indusiados o no; esporangios glabros, raramente setosos;
esporas finamente reticuladas; x=29.

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

COMMENTS: This is the most common and widespread COMENTARIOS: Este es el subgénero de Thelypteris
subgenus of Thelypteris in the American tropics. It contains más común en los Neotropicos. Contiene alrededor de
about 200 species, only a few of which occur outside 200 especies, sólo unas de las cuales existen afuera de
the Neotropics. Smith (1974) recognized 9 sections los Neotropicos. Smith (1974) reconoció 9 secciones
within the subgenus, providing a key to the sections dentro del subgénero, dando una clave a las secciones y
and a list of their species. The phylogenetic relationships una lista de las especies en cada sección. Las relaciones
among these sections have not been studied. In some filogenéticos entre estas secciones no han sido estudiados.
species the bases of petioles or even entire crosiers are En algunas especies las bases de los pecíolos o aún las
covered with mucilage. This occurs mostly in extremely prefoliaciones enteras están cubiertos con mucílago. Esto
wet habitats. The mucilage is secreted by dense, glandular ocurre a menudo en ambientes extremamente húmedos.
hairs (Hennipman, 1968). Aerophores usually protrude El mucílago está secretado por pelos glandulares densos
beyond the layer of mucilage. (Hennipman, 1968). Aeróforos usualmente se extienden
a fuera la capa de mucílago.

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. 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.

478
Figure 233. Thelypteris (subgen. 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)

479
THELYPTERIDACEAE
Thelypteris subgen. Cyclosorus (Link) C.V. Morton
Cyclosorus Link

DESCRIPTION: Terrestrial; petioles with 2 vascular DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
bundles; lamina 1-pinnate-pinnatifid, with the apex vasculares; lámina 1-pinnado-pinnatífida con un ápice
pinnatifid and confluent or conform and resembling the pinnatífido confluente o abruptamente acortada y
lateral pinnae; proximal pinnae not reduced, or up to 6 con una pinna terminal similar en forma a las pinnas
pairs gradually reduced in a few species; aerophores and laterales; pinnas proximales no reducidas o hasta 6
buds absent; veins meeting the margin just above the pares gradualmente reducidos en unas cuantas especies;
sinus, or, generally, connivent to the base of the sinus, o aeróforos y yemas ausentes; nervaduras llegando al margen
a pair united below the sinus with an excurrent veinlet; en o justo arriba del seno o, generalmente, conniventes a
indumentum of needle-shaped hairs; sori round; indusium la base del seno, o un par unido abajo del seno con una
round-reniform, persistent; sporangial capsules glabrous. nervadura excurrente; indumento de tricomas aciculares;
soros redondos; indusio redondo-reniforme persistente;
cápsulas de los esporangios glabras.

DIAGNOSIS: Petioles with 2 vascular bundles; lamina DIAGNOSIS: Pecíolo con 2 haces vasculares; lámina
1-pinnate-pinnatifid, proximal pinnae the longest or only 1-pinnato-pinnatífida, pinnas proximales lo más largas o
slightly reduced (never vestigial); hairs needle-shaped. poco reducidas (nunca vestigial); venas libres; pelos en
forma de aguja.

SIMILAR GENERA: Some species of subgen. Goniopteris GÉNEROS PARECIDOS: Algunas especies de subgén.
also have 1-pinnate-pinnatifid laminae, but they can be Goniopteris también tienen láminas 1-pinnado-pinnatífidas,
distinguished by the presence of minute, forked hairs. pero se distinguen por la presencia de pelitos furcados.

COMMENTS: This subgenus contains about 70 species. COMENTARIOS: Este contiene alrededor de
It is pantropical. Two species of this subgenus, T. dentata 70 especies. Es pantropical. Dos especies de este
and T. opulenta, are commonly found in the Neotropics subgénero, T. dentata and T. opulenta, se encuentran con
but are naturalized from the Old World. frequencia en el Neotrópico pero son naturalizadas del
Viejo Mundo.

LITERATURE: Smith, A. R. 1971. Systematics of the neotropical species of Thelypteris section Cyclosorus. University of
Calififornia Publications in Botany. 59: 1-143; Smith, A. R. 1983. 14(4). Polypodiaceae—Thelypteridoideae. In: G. Harling & B.
Sparre, Flora of Ecuador, no. 18.

480
Figure 234. Thelypteris (subgen. Cyclosorus) depilata. (from Smith in Tryon & Stolze, 1992)

481
THELYPTERIDACEAE
Thelypteris subgen. Goniopteris (C. Presl) Duek
DESCRIPTION: Terrestrial; petioles with 2 vascular DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
bundles; lamina simple to 1-pinnate-pinnatifid; vasculares; lámina simple a 1- pinnada-pinnatífida;
proximal pinnae not reduced, or only a few slightly pinnas proximales no reducidas, o sólo unas pocas
reduced; distal pinnae gradually shortened with a escasamente reducidas; pinnas distales gradualmente
confluent, pinnatifid apex or with a conform apex acortadas con un ápice pinnatífido confluente o con
resembling the lateral pinnae; aerophores absent; veins una pinna apical similar en forma a las pinnas laterales;
free or regularly united and forming a series of areoles; aeróforos ausentes; nervaduras libres o regularmente
indumentum of minute acicular and often forked anastomosadas y formando una serie de aréolas;
hairs (in some species stellate or anchor-shaped); sori indumento de pelitos aciculares y generalmente
round; indusium round-reniform or absent; sporangia furcados, en algunas especies estrellados o en forma
glabrous or with stellate or furcate hairs on the capsule de ancla; soros redondos, con un indusio reniforme o
or stalk; x=36. 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: Subgenus Goniopteris is completely COMENTARIOS: Subgénero Goniopteris es

neotropical and consists of about 90 species. It generally completamente neotropical y consta de 90 especies.
grows below 800 m in wet forests and is common in Generalmente existe abajo de 800 m en bosques
Amazonia. The most distinctive characteristic of subgenus húmedas y es común en la Amazonía. La característica
Goniopteris is the furcate hairs, but these are small and a más distinctiva del subgénero Goniopteris es los pelitos
handlens is necessary to see them. They are most easily furcados, pero son pequeños y una lupa es necesario
seen on the scales at the apex of the rhizome and in verlos. Estos son más evidentes en las escamas del ápice
the adaxial grooves of the lamina, but also on the lower del rizoma y en los surcos adaxiales, pero a menudo
surface of the axes. The name of the subgenus is derived también en el envés en los ejes. El nombre del subgénero
from the Greek gonia, angle + pteris, fern. It refers to the se deriva del griego gonia, ángulo + pteris, helecho. Refiere
basal veins that converge at sharp angles. 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.

482
Figure 235. A–C. Thelypteris blanda. D–F. T. tetragona. G–J. T. hatchii. L–N. T. paucipinnata. O, P. T. toganetra. Q, R. T. imbrica-
ta. (Mickel & Smith, 2004)

483
THELYPTERIDACEAE
Thelypteris subgen. Meniscium (Schreb.) C.F.
DESCRIPTION: Terrestrial; petioles with 2 vascular DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
bundles; laminae 1-pinnate, rarely entire; pinnae vasculares; láminas 1-pinnatas, raras veces enteras;
entire or serrate, the basal ones the largest or nearly pinnas enteras or serradas, los basales lo más
so; aerophores (found at the pinna base) absent; veins largas o casi así; aeróforos ausentes; nervaduras
netted regularly in pairs, with the transverse veins anastomosadas regularmente en pares, con las
producing excurrent veins that run toward the nervaduras transversas produciendo nervaduras
margin, the areoles 4–25(–35) between the costa and excurrentes, las aréolas en 4–25(–35) hileras entre
margin; hairs needle-shaped; sori on the transverse veins, la costa y el margen; pelos en forma de aguja; soros en
elongate (not round); indusia absent; sporangia glabrous las nervaduras transversas, alargados (no redondeados);
or setose; x=36. indusios ausentes; esporangios glabros o setosos; x=36.

SIMILAR GENERA: Some species of Serpocaulon (e.g., GÉNEROS PARECIDOS: Algunas especies de
S. fraxinifolium and S. triseriale) also have 1-pinnate blades Serpocaulon (e.g., S. fraxinifolium and S. triseriale) también
and netted veins with excurrent veinlets in the areoles. tienen láminas 1-pinnadas y nervaduras anastomosadas
They differ from subgen. Meniscium by having four or con venillas excurrentes en los aréolas. Difieren de
more vascular bundles in the petiole, lack of acicular hairs, subgén. Meniscium por tener 4 o mas haces vasculares en
and round sori. el pecíolo, pelos aciculares ausentes, y soros redondeados.

COMMENTS: Subgenus Meniscium is completely COMENTARIOS: Subgénero Meniscium es

neotropical and consists of 23 species. It occurs mostly completamente neotropical y consta de 23 especies.
below 1000 m in wet forests. It is easy to recognize by Existe principalmente bajo de 1000 m en bosques
its characteristic pattern of venation with straight and húmedos. Es fácil reconocer por el patrón característico
regular rows of veins between the costa and margin. de la nervadura con hileras rectas y regulares entre la
The genus name is derived from the Greek meniskos, costa y el margen. El nombre genérico se deriva del
diminutive of mene, moon. It refers to the shape of the griego meniscos, diminutivo de mene, luna. Refiere a la
sori, which spread across the transverse veins and form a forma de los soros, los cuales se difunden por la venilla
short crescent. transversa y forman un creciente corto.

LITERATURE: Smith, A. R. 1983. 14(4). Polypodiaceae—Thelypteridoideae. In: G. Harling & B. Sparre, Flora of Ecuador, no. 18.

484
Figure 236. Thelypteris subgen. Meniscium. A, B. T. falcata. C, D. T. angustifolia. E–G. T. reticulata. H, J. T. serrata. K–N. T. standleyi
(Mickel & Smith, 2004)

485
THELYPTERIDACEAE
Thelypteris subgen. Stegnogramma (Blume) C. F. Reed
DESCRIPTION: Terrestrial or rupicolous; rhizomes DESCRIPCIÓN: Terrestres o rupícolas; rizomas
short-creeping to erect; petioles with 2 vascular bundles; cortamente rastreros a erectos; pecíolos con 2
laminae 1-pinnate-pinnatifid, proximal pinnae not or only haces vasculares; láminas 1-pinnada-pinnatífidas;
slightly reduced; distal pinnae broadly adnate and pinnas proximales no reducidas o poco reducidas;
often decurrent on the rachis; aerophores absent; basal pinnas distales anchamente adnatas y a menudo
veins ending at the base of the sinus (in our one species) decurrentes al raquis; aeróforos ausentes; nervaduras
or anastomosing to form a complex web (several Old libres, encontrándose con el margen en el seno
World species); indument of acicular hairs; sori elongate, (Mesoamérica) o anastomosándose para formar una
exindusiate; sporangial capsules setose; spores spinulose. red compleja (en algunas especies del Viejo Mundo);
x=36. indumento de tricomas aciculares; soros alargados,
exindusiados; cápsulas esporangiales setosas; esporas
finamente espinulosas; x=36.

SIMILAR GENERA: Subgen. Amauropelta differs by GÉNEROS PARECIDOS: Subgén. Amauropelta difiere
having one or more pairs of reduced proximal pinnae. por tener uno o más pares de pinnas proximales reducidas.
Subgen. Cyclosorus differs by round, indusiate sori. Subgen. Subgén. Cyclosorus difiere por soros redondeados,
Steiropteris differs by having a cartilaginous keel between indusiados. Subgén. Steiropteris difiere por tener una quilla
the base of the sinus and the costa. Subgen. Goniopteris cartilaginosa entre la base del sino y la costa. Subgén.
differs by the presence of minute, forked hairs (use Goniopteris difiere por la presencia de pelitos furcados
handlens). (use lupa).

COMMENTS: Thelypteris subgen. Stegnogramma has COMENTARIOS: Thelypteris subgén. Stegnogramma

about 15 species worldwide, only one of which occurs in consta de casi 15 especies mundial, solo una de las cuales
the Neotropics: T. pilosa (M. Martens & Galeotti) Crawford. existe en el Neotrópico: T. pilosa (M. Martens & Galeotti)
It occurs from northwestern Mexico to Honduras, from Crawford. Existe desde el noroeste de México hasta
1800–2700 m, usually on limestone. In rock houses of Honduras, de 1800–2799 m, usualmente sobre piedra
the southeastern United States (Alabama) occurs T. caliza. En lugares rocosos en el sureste de los Estados
burksiorum J. E. Watkins & D. R. Farrar. It differs subtly from Unidos (Alabama) existe T. burksiorum J. E. Watkins & D.
the Central American species.The name Stegnogramma is R. Farrar. Difiere sutilmente de la especie mesoamericana.
derived from the Greek stegnos, cover + gramme, line. It El nombre Stegnogramma se deriva del griego stegnos,
refers to the short, line-like sori that cover the veins. 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.

486
Figure 237. A–D. Thelypteris pilosa. (Mickel & Smith, 2004)

487
THELYPTERIDACEAE
Thelypteris subgen. Steiropteris (C. Chr.) K. Iwats.
DESCRIPTION: Terrestrial; petioles with 2 vascular DESCRIPCIÓN: Terrestres; pecíolos con 2 haces
bundles; lamina 1-pinnate-pinnatifid or pinnatifid (with a vasculares; lámina 1-pinnado-pinnatífida o pinnatífida (con
few pinnae free in 2 spp.); proximal pinnae the longest or unas cuantas pinnas libres en 2 spp.); pinnas proximales
nearly so; distal pinnae gradually reduced or sometimes las más largas o casi así; pinnas distales gradualmente
with an apical pinna somewhat similar in shape to the reducidas o a veces con una pinna apical algo similar
lateral pinnae; aerophores often strongly developed at the en forma a las pinnas laterales; aeróforos a menudo
pinnae bases; buds absent (except in T. seemannii); veins fuertemente desarrollados en la base de las pinnas; yemas
free, proximal ones of the adjacent segments either joining ausentes (excepto en T. seemannii); nervaduras libres, las
the margin above the sinus base (sect. Glaphyropteris), or proximales de los segmentos adyacentes encontrándose
connivent to the sinus or a cartilaginous keel (false vein) con el margen arriba del seno (sect. Glaphyropteris) o
extending from the base of the sinus (sect. Steiropteris); conniventes en el seno (sect. Steiropteris) con una quilla
indumentum of needle-shaped hairs, sometimes septate; cartilaginosa (vena falsa) que se extiende desde el seno
sori round; indusia present or absent; sporangia glabrous, hacia la costa; indumento de tricomas aciculares, a veces
rarely setose. x=36. septados; soros redondeados; indusios presentes o
ausentes; esporangios glabros, raramente setulosos. x=36.

SIMILAR GENERA: Thelypteris subgen. Amauropelta, GÉNEROS PARECIDOS: Thelypteris subgén.

the most common subgenus in the Neotropics, differs by Amauropelta, el subgénero más común en el Neotrópico,
highly reduced proximal pinnae. Subgenus Cyclosorus can difiere por pinnas proximales altamente reducidas. El
be distinguished by lack of aerophores and sinus keels. subgénero Cyclosorus se distingue por la falta de aeróforos
y quillas desde del sinus.

COMMENTS: Subg. Steiropteris consists of 21 species, all COMENTARIOS: Subg. Steiropteris consta de 21
Neotropical. It is divided into two sections: sect. Steiropteris especies, todas Neotropicales. Tiene dos secciones: la
(5 spp.) and sect. Glaphyropteris (16 spp).The first has the secc. Steiropteris (5 spp.) y secc. Glaphyropteris (16 spp).
sinus keel prominent and raised, veins connivent at the La primera tiene la quilla del sinus prominente y elevada,
sinus or uniting below the sinus, and glands lacking on the venas conniventes en el sinus o unidas abajo del sinus, y
leaf tissue. In contract, sect. Glaphyropteris has the sinus glándulas faltantes en el tejido laminar. En contraste, sect.
keel absent, veins meeting margin above the sinus, and Glaphyropteris tiene la aquilla del seno ausente, venas
glands usually present on the laminar tissue. Steiropteris terminando en el margen arriba de la base del senos y
is often most evident on the upper surface of the lamina. glandulas usualmente presents en el tejido lamina. Los
The aerophores at the pinna bases are usually well aeróforos en las bases de las pinnas usualmente son bien
developed, conical or peg-like, and whitish or yellowish. desarrollados, cónicos or tuberculiformes, y blanquecinos
They usually darken upon drying. It is believed that they o amarillentos. Se tornan oscuros al secar. Se cree que
function in aerating the fiddlehead, which in several funcionan en impregnar con aire la prefoliación, cual es en
species (e.g., T. decussata, T. thomsonii) is covered by a thick varias especies (e.g., T. decussata, T. thomsonii) es cubierta
mucilage through which the aerophores protrude. The con un mucílago desde que los aeróforos proyectan. El
genus name is derived from the Greek steira, keel + pteris, nombre genérico se deriva del griego steira, carina +
fern. It refers to the false vein present in sect. Steiropteris pteris, helecho. Refiere a la vena falsa presente en secc.
that runs from the costa to the base of the sinus. 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.

488
Figure 238. Thelypteris (subgen. Steiropteris) valdepilosa. Note in B the dark aerophore at base of pinna. (from Smith, Pterid.
Peru, 1992)

489
THYRSOPTERIDACEAE
Thyrsopteris Kunze
DESCRIPTION: Terrestrial; stems erect to decumbent, DESCRIPCIÓN:Terrestres; tallos erectos a decumbentes,
to ca. 2 m tall, massive, sometimes producing stolons; hasta 2 m de altura, massivos, a veces produciendo
leaves 2-3.5 m long, partially dimorphic, the fertile ones estolones; hojas 2-2.5 m long, parcialmente dimorfas, las
with strongly modified lower pinnae; petioles with long, fértiles con pinnas inferiors dimorfas; pecíolos con pelos
matted hairs; laminae 3–5-pinnate, broadly ovate, largos y enredados; láminas 3–5-pinnadas, anchamente
tapered to the tip; pinnae and pinnules stalked, ascending; ovadas, reducidas gradualmente hacia el ápice; pinnas y
rachises and costae grooved adaxially; veins free; pínnulas pecioluladas, ascendentes; ráquises y costas
sori terminal on narrow, non-laminate segments, lacking surcados adaxialmente, los surcos decurrentes; venas
paraphyses, often restricted to the proximal portions of libres; soros terminales en segmentos estrechos no
the pinnae; receptacle columnar, clavate; indusia cup- laminados, carente de paráfises, a menudo restingidos a
shaped, firm, entire; spores tetrahedral-globose, trilete. las partes proximales de las pinnas; receptáculo columnar,
x= ca. 76–78. clavado; indusios en forma de una copa, firmes, enteros;
esporas tetraédrico-globosas, triletes. x= ca. 76–78.

SIMILAR GENERA: Culcita has monomorphic fertile GÉNEROS PARECIDOS: Culcita tiene pínulas fértiles
pinnules and clam-shaped indusia. Dicksonia also differs by dimorfas y indusios en forma de una bivalva. También
clam-shaped indusia and by leaves long-tapered toward Dicksonia difiere por indusios en forma de una bivalva y
the base. por hojas largamente reducidas hacia la base.

COMMENTS: The Thyrsopteridaceae consists of a COMENTARIOS: La Thyrsopteridaceae consta de una

single species, Thyrsopteris elegans Kunze, which is of única especie, Thyrsopteris elegans Kunze, la cual es de
uncertain relationship to other tree ferns. It is endemic una relación incierta de otros helechos arborescentes.
to the Juan Fernández Islands where it grows from 500- Es endémica a las Islas Juan Fernández donde existe de
1000 m in woodlands and heaths, often with Dicksonia 500-1000 m en bosques y lugares abiertos abustivos,
berteriana. It is unusual by the sori restricted to only the a menudo con Dicksonia berteriana. Es inusual por
proximal pinnae. Many species related to Thyrsopteris los soros retrigidos únicamente a las pinnas inferiores.
were widespread world-wide during the Mesozoic, but Muchas especies relaciondas a Thyrsopteris estaban muy
nowadays only Thyrsopteris survives. Cretaceous fossils of difundidas mundialmente durante el Mesozóico, pero hoy
Thyrsopteris are known from mainland Chile. The genus día solamente Thyrsopteris sobrevive. Fósiles Cretásicos
name is derived from the Greek thyrsos, bunch + pteris, de Thyrsopteris se conocen de Chile continental. El
fern. The sori are arranged in grape-like clusters. nombre genérico se deriva del griego thyrsos, macolla
+ pteris, helecho. Los soros están agrupados en grupos
como uvas.

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.

490
Figure 239. Thyrsopteris elegans Kunze. (©Robbin Moran, 2004)

491
PTERIDACEAE
Trachypteris H. Christ
DESCRIPTION: Terrestrial or rupestral; rhizomes erect DESCRIPCIÓN: Terrestres o rupestres; rizomas erectos
or decumbent, scaly; leaves monomorphic or dimorphic, o decumbentes, erectos, escamosos; hojas monomorfas o
10–25 cm long; blades densely scaly abaxially, the dimorfas, 10–25 cm; láminas densamente escamosas
scales obscuring the surface, glabrous to sparsely scaly en el envés, las escamas oscuriendo la superficie,
adaxially; veins hard to see because of the thick blades, adaxialmente glabras a esparcidamente escamosas;
anastomosing without included veinlets; sori borne in nervadura difícil ver a causa de las láminas gruesas,
a narrow to broad band near the margin; indusia anastomosada sin venillas incluidas; soros nacen en una
and paraphyses absent; spores globose, trilete, strongly faja estrecha cerca del margen; indusios y parafises
cristate, without an equatorial flange; x=29 or 30. ausentes; esporas globosas, triletes, fuertemente cristadas,
sin un engrosamiento ecuatorial; x=29 o 30.

SIMILAR GENERA: Doryopteris differs by laminae GÉNEROS PARECIDOS: Doryopteris difiere por
glabrous beneath. Some species of Cheilanthes can be las láminas glabras en el envés. Algunas especies de
densely scaly beneath, but these species usually have Cheilanthes puede ser muy escamosas en el envés, pero
finely divided leaves. estas especies usualmente tienen hojas más finamente
divididas.

COMMENTS: Trachypteris consists of 3 species and COMENTARIOS: Trachypteris consta de 3 especies

occurs from Ecuador (including Galapagos Islands) to y existe desde Ecuador (incluyendo los Galápagos) a
Argentina and southern Brazil, and Madagascar. It usually Argentina y sur de Brasil, y Madagascar. Usualmente crece
grows on rocks or among rocks and in dry soils, usually sobre o entre rocas y en suelos secos, usualmente en
in seasonally dry habitats. The leaves have the ability to ambientes estacionales secos. Las hojas tienen la capaz
lose most of their normal moisture content and enroll, de perder su contenido de agua normal y enroscarse,
then after a rain rapidly absorb water and expand. The y luego después de una lluvia absorber agua y expandir.
American species grow from 700-2900 m. The three Los especies americanas existen de 700-2900 m. Las tres
neotropical species are: T. gilliana (Baker) Svenson, T. induta especies neotropicales son: T. gilliana (Baker) Svenson,
(Maxon) R. M. Tryon & A. F. Tryon and T. pinnata (Hook. f.) T. induta (Maxon) R. M. Tryon & A. F. Tryon y T. pinnata
C. Chr. Trachypteris induta having a basal rosette of sterile (Hook. f.) C. Chr. Trachypteris induta tiene una roseta basal
leaves and erect, long-petioled fertile leaves, whereas T. de hojas estériles y hojas fertiles erectas y largamente
pinnata lacks the rosette and has monomorphic leaves. pecioladas, mientras que T. pinnata falta de una roseta
Trachypteris gilliana resembles T. pinnata but differs by basal y tiene hojas monomorfas. Trachypteris gilliana se
reticulate or verrucose spores and trifid fertile leaves. asemeja a T. pinnata pero difiere por esporas reticuladas
Another species, T. drakeana C. Chr., occurs in Madagascar. o verrucosas y hojas fértiles trífidas. Otra especie, T.
It has a basal rosette of sterile leaves, as in T. pinnata, but drakeana C. Chr., existe en Madagascar. Tiene una roseta
each leaf bears apical bud. According to rbcL studies, basal de hojas estériles, como en T. induta, pero cada
Trachypteris is the sister genus to Doryopteris (Gastony & hoja lleva una yema apical. Conforme a estudios de rbcL,
Rollo, 1998). The genus name is derived from the Greek, Trachypteris es el género hermana a Doryopteris (Gastony
trachys, rough + pteris, fern.The scales covering the fronds & Rollo, 1998). El nombre genérico se deriva del griego,
impart a rough texture. trachys, áspero + pteris, helecho. Las escamas cubriendo
las hojas imparten una textura aspera.

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.

492
Figure 240. Trachypteris. A-H. T. induta. J-M. T. pinnata. (©Robbin Moran, 2009)

493
HYMENOPHYLLACEAE
Trichomanes L. subgen. Davalliopsis
DESCRIPTION: Terrestrial; roots many, robust; DESCRIPCIÓN: Terrestres; raíces numerosas,
rhizomes 4-15 mm wide, erect or ascending, pubescent, robustos; rizomas 4-15 mm de ancho, erectos o
the hairs dark, multicellular; sterile and fertile leaves up ascendentes, pubescentes, los pelos oscuros, multicelulares;
to 60 × 35 cm, monomorphic, polystichous; sterile hojas estériles y fértiles hasta 60 × 35 cm, monomorfas,
laminae 3 cells thick between the veins, opaque (not polistícas; láminas estériles 3 células de grosor
translucent), 3- to 4-pinnate, usually iridescent, the entre las venas, no translúcidas, 3- a 4-pinnadas,
apices pinnatifid, not flagellate or proliferous; veins free, usualmente iridiscentes, los ápices pinnatífidos, no
anadromous, false veins absent; sori marginal, oriented flagelados y prolíferos; venas libres, anadrómas, falsas
downward, out of the plane of the lamina; indusia venas ausentes; soros marginales, orientados de abajo,
tubular, the mouth truncate or slightly dilate; receptacles a fuera del plano de la lámina; indusios tubulares,
exert; spores green, tetrahaedral-globose; x=32. las bocas truncadas o levemente dilatas; receptáculos
exertos; esporas verdes, tetrahédricas-globosas; x=32.

SIMILAR GENERA: Abrodictyum differs by laminae non- GÉNEROS PARECIDOS: Abrodictyum difiere por
iridescent and only a few rows of cells on either side of láminas no iridiscentes y solo unas hileras de células por
the veins. Trichomanes subgen. Feea differs by flagellate, los lados de las venas. Trichomanes subgén. Feea difiere
proliferous apices of the sterile leaves and (usually) por ápices de las hojas estériles flagelados y prolíferos y
dimorphic sterile and fertile leaves. Trichomanes subgen. (usualmente) hojas estériles y fértiles dimórfas. Trichomanes
Trichomanes differs by translucent laminae and epiphytic subgén. Trichomanes difiere por láminas translucidas y
habit. Vandenboschia and Trichomanes subgen. Lacostea hábito epifítico. Vandenboschia y Trichomanes subgén.
differ by long-creeping rhizomes that climb up trunks. Lacostea difieren por rizomas largamente rastreros
Polyphlebium differs by epiphytic habit and creeping que trepan los troncos. Polyphlebium difiere por hábito
rhizomes. epifítico, rizomas rastreros.

COMMENTS: Trichomanes subgen. Davalliopsis consists COMENTARIOS: Trichomanes subgén. Davalliopsis

of 2 neotropical species: T. elegans (widespread) and T. consta de 2 especies neotropicales: T. elegans (ampliamente
resinosum (Guayana, s. Venezuela). Both occur terrestrially difundida) y T. resinosum (Guayana, s. Venezuela). Ambos
in wet forests. Their rhizomes are erect and supported son terrestres en bosques húmedos. Sus rizomas son
by roots on all sides. The leaves are unusual in the family erectos y apoyados por raíces en todos lados. Las hojas
by being 3 cells thick between the veins and (if in shade) son atípicas dentro de la familia por ser 3 células de
iridescent. The sori bend downward, out of the plane of grosor entre las venas y (si en la sombra) iridiscentes. Los
the lamina. The chromosome number of x=32, shared soros doblan hacia abajo, a fuera del plano de la lámina.
with other species of the genus, is derived within the El número cromosómico de x=32, compartido con las
family (Ebihara et al., 2007) The name Davalliopsis is otras especies del género, se deriva dentro de la familia
derived from Davallia and the Greek suffix -opsis, like. (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.

494
Figure 241. . A, B. T. elegans. C. D. T. resinosum. (Moran, 2000)

495
HYMENOPHYLLACEAE
Trichomanes subgen. Feea (Bory) Hook.
DESCRIPTION: Terrestrial; roots many, robust, often DESCRIPCIÓN: Terrestres; raíces numerosas,
stilt-like; rhizomes 2-8 mm wide, suberect or ascending, robustos, a menudo como raíces zancudos; rizomas 2-8
unbranched, pubescent, the hairs dark, multicellular; mm de ancho, suberectos a ascendentes, no ramificados,
sterile and fertile leaves up to 30 × 18 cm, dimorphic pubescentes, los pelos oscuros, multicelulares; hojas
(except T. mougeotii), polystichous; sterile laminae 1-cell estériles y fértiles hasta 30 × 18 cm, dimorfas (con
thick between the veins, 1-pinnate, the apices often excepción de T. mougeotii), polistícas; láminas estériles
becoming flagellate and proliferous; veins free, una célula de grosor entre las venas, 1-pinnadas, los
catadromous, false veins absent; sori marginal; indusia ápices a menudo se tornan flagellados y prolíferos;
tubular, the mouth truncate or slightly dilate; receptacles venas libres, catádromas, falsas venas ausentes; soros
exert; spores green, tetrahaedral-globose; x=32. marginales; indusios tubulares, la boca entera, tuncada;
receptáculos exertos; esporas verdes, tetrahédricas-
globosas; x=32.
SIMILAR GENERA: Trichomanes subgen. Trichomanes
usually differs by monomorphic sterile and fertile leaves, GÉNEROS PARECIDOS: Trichomanes subgén.
and sori that tend to be clustered toward the pinna tips. Trichomanes difiere usualmente por hojas estériles y
Trichomanes subgen. Davalliopsis differs by laminae opaque fértiles monomórfas y soros que tienden ser agrupados
(3-cells thick between the veins), usually iridescent, and hacia los ápices de los pinnas. Trichomanes subgén.
not flagellate and proliferous. 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 neotropical y típicamente ocurre en bosques húmedos
rocks. It contains 5 species: T. botryoides, T. diversifrons, T. sobre suelos arcillosos o rocas. Consta de 5 especies: T.
mougeotii, T. osmundoides, and T. trollii. These plants typically botryoides,T. diversifrons,T. mougeotii,T. osmundoides y T. trollii.
have short erect rhizomes supported by prop roots, Estas plantas típicamente tienen rizomas cortas erectas
imparting the aspect of a small tree.The sterile leaf apices apoyadas por raíces zancudos, dando el aspecto de un
may become flagellate and proliferous. If a lamina is arbolito. Los ápices de las hojas estériles pueden tornarse
present on the fertile leaves, it is more than one cell thick. flagelados y prolíferos. Si una lámina está presente en
All but one species (T. mougeotii) have dimorphic sterile las hojas fértile, es más de una célula de grosor. Con la
and fertile leaves. The chromosome number of x=32, excepción de una especie (T. mougeotii), todas tienen hojas
shared with other species of the genus, is derived within estériles y fértiles dimorfas. El número cromosómico de
the family (Ebihara et al., 2007) The genus name honors x=32, compartido con las otras especies del género, se
A.L.A. Fée (1789–1874), a pteridologist from Strasbourg. 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.

496
Figure 242. A–C. Trichomanes diversifrons. D–F. T. osmundoides. G–K. T. botryoides. L, M. T. muegotii. (©Robbin Moran, 2009)

497
HYMENOPHYLLACEAE
Trichomanes subgen. Lacostea (Bosch) C. Chr.
DESCRIPTION: Lianas; rhizomes 0.5-1.5 mm wide, DESCRIPCIÓN: Lianas; rizomas 0.5-1.5 mm de ancho,
long-creeping, climbing, often branched, pubescent, the largamente rastreros, a menudo ramificados, pubescentes,
hairs dark, multicellular; sterile and fertile leaves up to 30 × los pelos oscuros, multicelulares; hojas estériles y fértiles
9 cm, monomorphic, short-petiolate, distichous,; laminae hasta 30 × 9 cm, monomorfas, cortamente pecioladas,
pressed flat against the substrate, 1-cell thick between dísticas; láminas aplastadas contra el sustrato, una
the veins, simple, 1-pinnate or 1-pinnate-bipinnatifid; célula de grosor entre las venas, simple a 1-pinnadas o
pinnae symetrical; veins free, anadromous, false veins -pinnado-bipinnatífidas; venas libres, anádromas, falsas
absent; sori marginal, pedicellate, projecting at right venas ausentes; soros marginales, pedicelladas,
angles to the substrate and lamina; indusia tubular, the llevados a angulos rectos al sustrato y lámina;
mouth truncate or sometimes dilate; receptacles exert; indusios tubulares, las bocas tuncadas o a veces dilatas;
spores green, tetrahaedral-globose; x=32. 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 hábito hemiepifítico o hemiepipétrico y hojas divergentes
away from substrate (not appressed). Trichomanes subgen. del sustrato (no aplastadas). Trichomanes subgén.
Trichomanes differs by catadromous venation and leaves Trichomanes difiere nervación catadrómica y hojas
spreading away from the substrate. Hymenophyllum differs divergentes desde el sustrato. Hymenophyllum difiere por
by two-valved indusia and included receptacles. indusios bivalvados y receptáculos incluidos.

COMMENTS: Trichomanes subgen. Lacostea is COMENTARIOS: Trichomanes subgén. Lacostea es

neotropical and occurs in wet forests below 800 m. It neotropical y existe en bosques húmedos abajo de 800
contains five species: T. ankersii, T. pedicellatum, T. tanaicum, m. Consta de cinco especies: T. ankersii, T. pedicellatum,
T. tuerckheimii, and T. subsessile. According to Dubuisson T. tanaicum, T. tuerckheimii y T. subsessile. Conforme a
et al. (2003), these species are lianas, starting in the soil Dubuisson et al. (2003), estas plantas son lianas, empezando
and secondarily climbing trunks. Their leaves are unusual el suelo y secundariamente subiendo troncos. Sus hojas
by being plastered against the trunk with pedicellate sori son inusuales por ser aplastadas contra el tronco con los
projecting outward. The petioles are short or apparently soros pedicelados proyectándose a fuera. Los pecíolos
absent. The chromosome number of x=32, shared with son cortos a aparentemente ausentes. El número
other species of the genus, is derived within the family cromosómico de x=32, compartido con otras especies
(Ebihara et al., 2007) The name Lacostea honors Cornelius del género, se deriva dentro de la familia (Ebihara et al.,
Marinus van der Sande Lacoste (1815-1887), student of 2007). El nombre Lacostea honora Cornelius Marinus van
East Indian bryophytes. 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.

498
Figure 243. A–D. Trichomanes ankersii. E–K. T. tuerckheimii. L–O. T. tanaica. (©Robbin Moran, 2010)

499
HYMENOPHYLLACEAE
Trichomanes L. subgen. Trichomanes
DESCRIPTION: Terrestrial, rarely epiphytic; roots DESCRIPCIÓN: Terrestres, raras veces epífitas;
many, robust; rhizomes 1-5 mm wide, erect or short- to raíces numerosas, robustos; rizomas 1-5 mm de
long-creeping, pubescent; sterile and fertile leaves up to 70 ancho, erectos o cortamente a largamente reptantes,
× 25 cm, monomorphic or rarely dimorphic, polystichous; pubescentes; hojas estériles y fértiles hasta 70 × 25 cm,
sterile laminae 1-cell thick between the veins, 1-pinnate, monomorfas o raras veces dimorfas, polistícas; láminas
the apices pinnatifid (flagellate and proliferous only in two estériles una célula de grosor entre las venas, 1-pinnadas,
species), pubescent, the hairs with a distinct basal cell; los ápices pinnatífidos (flagelados y prolíferos solo en dos
pinnae symmetric; veins free, catadromous (anadromous especies), pubescentes, los pelos con una célula distinta
in T. scandens and T. anadromum), false veins absent basal; venas libres, catádromas, falsas venas ausentes
(present only in T. pinnatum); sori marginal, immersed (presentes solo en T. pinnatum); soros marginales,
in the laminae or (in two species) pedicellate; indusia inmersas en la lámina o (en dos especies) pediceladas;
tubular, the mouth truncate or slightly dilate; receptacles indusios tubulares, las bocas enteras, truncadas o
exert; spores green, tetrahaedral-globose; x=32. levemente dilatas; receptáculos exertos; esporas verdes,
tetrahédricas-globosas; x=32.

SIMILAR GENERA: Trichomanes subgen. Feea differs GÉNEROS PARECIDOS: Trichomanes subgén. Feea
by flagellate, proliferous apices of the sterile leaves and difiere por ápices de las hojas estériles flagelados y
(usually) dimorphic sterile and fertile leaves. Trichomanes prolíferos y (usualmente) hojas estériles y fértiles dimórfas.
subgen. Davalliopsis differs by laminae opaque (3-cells thick Trichomanes subgén. Davalliopsis difiere por láminas
between the veins) and usually iridescent. Vandenboschia no translucidas (3 células de grosor entre las venas) y
and Trichomanes subgen. Lacostea differ by long-creeping usualmente iridiscentes. Vandenboschia y Trichomanes
rhizomes that climb the base of trunks. Polyphlebium subgén. Lacostea difieren por rizomas largamente
usually differs by epiphytic habit, creeping rhizomes, and rastreros que trepan la base de los troncos. Polyphlebium
more divided leaves. difiere por hábito epifítico, rizomas rastreros y hojas más
divididas.

COMMENTS: Trichomanes subgen. Trichomanes is COMENTARIOS: Trichomanes subgén. Trichomanes

neotropical except for T. crenatum of western Africa. es neotropical con excepción de T. crenatum de África
The subgenus contains ca. 30 species that typically are occidental. El subgénero consta de ca. 30 especies que
terrestrial in wet forests. The neotropical species were típicamente ocurre en bosques húmedos. Las especies
monographed by Windisch (1992).. Most have 1-pinnate neotropicales fueron monografiadas por Windisch
leaves with sori borne at the pinna apices, not along (1992). La mayoría tienen hojas 1-pinnatas con soros
the sides. Many have the laminae pubescent by long, llevados en los ápices de las pinnas, no los lados. Muchas
multicellular hairs, these with a distinct basal cell. The especies tienen las láminas pubescentes por pelos
chromosome number of x=32, shared with other species largos multicelulares, estos con una célula basal distinta.
of the genus, is derived within the family (Ebihara et al., El número cromosómico de x=32, compartido con las
2007) The genus name is derived from the Greek thrix, otras especies del género, se deriva dentro de la familia
hair + manes, cup. It alludes to the hair-like receptacle that (Ebihara et al., 2007). El nombre genérico se deriva del
protrudes from the cup-like indusium. griego thrix, pelo + manes, copa. Se refiere a receptáculo
piliforme que sobresale del indusio en forma de copa.

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.

500
Figure 244. A, B. T. pinnatum. C, D. T. polypodioides. E, F. T. lucens. G, H. T. crispum. (Mickel & Smith, 2004)

501
TECTARIACEAE
Triplophyllum Holttum
DESCRIPTION: Terrestrial; rhizomes creeping, scaly; DESCRIPCIÓN: Terrestre; rizomas reptantes,
leaves monomorphous, long-petiolate; vascular bundles in escamosos; hojas monomorfas, largamente pecioladas;
petiole 4 or more; lamina typically tripartite, deltate or haces vasculares en los pecíolos 4 o más; láminas
pentagonal, the basal pinnae typically conspicuously larger típicamente tripartidas, deltadas o pentagonales, las
than the suprabasal ones and wider on the basiscopic pinnas basales típicamente conspicuosamente más grandes
side; rachis and costae not grooved, usually puberulent que las adyacentes y más anchas en el lado basiscópico;
below and usually above with reddish hairs, these 0.1-0.4 raquis y costas no surcados, usualmente puberulentos
mm long, reddish, articulate, below provided with a few en la superficie inferior e usualmente superior con
dark scales; veins free; sori round; indusia present, circular pelos 0.1-0.4 mm de largo, rojizos, articulados, en el lado
to reniform, attached laterally (not peltate); x=41. inferior provisto con pocas escamas oscuras; nervadura
libre; soros redondeados; inducíos presentes, circulares a
reniformes, unidos lateralmente (no peltados); x=41.

SIMILAR GENERA: Tectaria typically differs by GÉNEROS PARECIDOS: Tectaria difiere típicamente
anastomosing veins. Megalastrum and Ctenitis usually differ por venas anastomosadas. Megalastrum y Ctenitis
by erect or decumbent rhizomes and scalier rachises and usualmente difieren por rizomas erectos o decumbentes y
costae. Similar dryopteroid genera, such as Arachniodes, ráquises y costas más escamosos. Géneros dryopteroides
Dryopteris, and Polybotrya, differ by adaxially grooved parecidos, tales como Arachniodes, Dryopteris, y Polybotrya,
rachises and costae. difieren por los raquises y pinnas surcados adaxialmente.

COMMENTS: Triplophyllum occurs in Africa (14 spp.) and COMENTARIOS: Triplophyllum existe en África (14
the Neotropics (9 spp.). A key to the neotropical species spp.) y el Geotrópico (9 spp.). Una clave a las especies
can be found in Prado & Moran (2008).The genus is most neotropicales está presentada en Prado & Moran (2008).
species-rich and abundant in the Guayanas and northern El género es más diverso en las Guayanas y Brasil del
Brazil. It usually grows on wet, shaded forest floors, norte. Existe en sotobosques húmedos y sombreados,
generally from 0–1200 m. The usually tripartite laminae, generalmente de 0–1200 m. Las láminas usualmente
with basal pinnae greatly elongated on the basiscopic tripartitas, con pinas basales alargadas fuertemente en
side, make this an easy genus to recognize. The species el lado basiscópico, contribuyen a la recognición fácil de
are more challenging to identify, being distinguished by este género. Las especies presentan un reto identificar,
indument characteristics that require at least a 30X siendo distinguidas por características del indumento que
dissecting microcope and a strong light source. requieren a lo menos a 30X microscopio de disección y
The genus name is derived from the Latin triplo, three una fuente de luz fuerte. El nombre genérico se deriva
times + the Greek phyllon, leaf. It refers to the tripartite del latín triplo, tres veces + el griego phyllon, hoja. Se
leaf. refiere a la hoja tripartita.

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.

502
Figure 245. Triplophyllum. A. T. chocoense. B. T. dicksonioides, pinnule. C. T. dicksonioides, habit. D. T. funestum, sorus. E. T. funes-
tum, 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).

503
PTERIDACEAE
Tryonia Schuettp., J. Prado & A. T. Chochran
DESCRIPTION: Terrestrial or saxicolous; rhizomes DESCRIPCIÓN: Terrestres o saxícolas; rizomas
creeping, bristly; leaves 6–100 cm; petioles brown basally reptantes, cerdosos; hojas 6–100 cm; pecíolos pardos
and straminous distally, pubescent; laminae linear to basalmente e estramíneous distalmente, pubescentes;
elongate-triangular, 1- to 3-pinnate-pinnatifid, determinate; láminas lineares a alargadas-triangulares, 1- 3-pinnado-
rachises stramineous, straight, terete or sulcate adaxially, pinnatífidas, determinadas; ráquises estramineos,
pubescent; pinnae 0.5–10 × 0.5–5 cm, membranaceous rectos, teretes o surcados adaxialmente, pubescentes;
to herbaceous, densely pubescent on both surfaces, the pinnas 0.5–10 × 0.5–5 cm, membranosas a herbáceas,
hairs glandular or non-glandular; veins free; sporangia densamente pubescentes en ambos lados, los pelos
borne along the veins; spores borwn, trilete, tetrahedral- glandulosos o no; venas libres; esporangios nacidos por
globose; x=?. als venas; esporas pardas, triletes, tetrahédricas-globosas;
x=36.

SIMILAR GENERA: Jamesonia differs castaneous GÉNEROS PARECIDOS: Jamesonia difiere por
rachises. ráquises castaños.

COMMENTS: Tryonia consists of four species of the COMENTARIOS: Tryonia consta de cuatro especies
Atlantic forests of southeastern Brazil, with one species de los bosques Atlánticos del sureste de Brasil, con
occurring in adjacent Uruguay. Generally along streams una especie en Uruguay adyacente. Generalmente por
on damp, shaded sandstones; 600–2300 m. The species riachuelos en piedra arenosa sombreada e húmedo; 600–
of Tryonia were formerly-treated in Eriosorus, a genus 2300 m. Las especies de Tryonia fueron anteriormente
now subsumed under Jamesonia. Tryonia is distinguished clasificadas en Eriosorus, un género subsumido ahora
from Jamesonia only by stramineous rachises. Instead abajo de Jamesonia. Tryonia se distingue de Jamesonia solo
being most closely related to Jamesonia, Tryonia is most por ráquises estramíneos. Tryonia es más cercanamente
closely related to the Old World genera Austrogramme, relacionados a los géneros del Viejo Mundo Austrogramme,
Pterozonium, Syngramma and Taenitis. The four species of Syngramma y Taenitis. Las cuatro species de Tryonia son T.
Tryonia are T. areniticola (Schwartsb. & Labiak) Schuettp., areniticola (Schwartsb. & Labiak) Schuettp., J. Prado & A.
J. Prado & A. T. Cochran, T. myriophylla (Sw.) Schuettp., J. T. Cochran, T. myriophylla (Sw.) Schuettp., J. Prado & A. T.
Prado & A. T. Cochran, T. schwackeana (Christ) Schuettp., Cochran, T. schwackeana (Christ) Schuettp., J. Prado & A.
J. Prado & A. T. Cochran y T. sellowiana (Kuhn) Schuettp., J. T. Cochran y T. sellowiana (Kuhn) Schuettp., J. Prado & A. T.
Prado & A.T. Cochran. Commemorates Alice Faber Tryon Cochran. Conmemora a Alice Faber Tryon (1920–2009),
(1920–2009), American pteridologist who monographed pteridólogo Americano quien hizo una monografía a
Eriosorus and Jamesonia, the genera from which Tryonia Eriosorus y Jamesonia, los generos de los cuales Tryonia
was segregated. fue segregado.

LITERATURE: Cochran, A. T. 2014. Tryonia, a new taenitidoid fern genus segregated from Jamesonia and Eriosorus. 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.

504
Figure 246. Tryonia. Above, A–H. T. myriophylla. Below, A–H. T. sellowianus. (From Tryon, 1970)

505
HYMENOPHYLLACEAE
Vandenboschia Copel.
DESCRIPTION: Hemiepiphytic or hemiepilithic; DESCRIPCIÓN: Hemiepifitos o hemiepilíticos;
roots many and robust; rhizomes 1-2 mm wide, long- raíces numerosas y robustos; rizomas 1-2 mm de ancho,
creeping, irregularly branching, densely pubescent, the hairs largamente rastreros, ramificándose irregularmente,
dark, multicellular; sterile and fertile leaves up to 40 × 20 típicamente densamente pubescentes, los pelos oscuros,
cm, monomorphic, distichous; laminae membranaceous, multicelulares; hojas estériles y fértiles hasta 40 × 20
translucent, 1-cell thick between the veins, 1-pinnate to cm, monomorfas, dísticas; láminas membranáceas,
5-pinnate; veins free, anadromous, false veins absent; sori translucientes, una célula de grosor entre las venas,
marginal; indusium (often called an “involucre”) 1-pinnadas a 5-pinnadas; venas libres, anádromas, falsas
tubular to campanulate, the mouth entire or flaring; venas ausentes; soros marginales; indusio (a menudo
receptacles exert, extending beyond the mouth of the llamado “involucre”) tubular, la boca entera o
indusium; spores green, tetrahaedral-globose; x=36. ensanchada; receptáculos exertos, extendiéndose
fuera de la boca del indusio, filiforme; esporas verdes,
tetrahédricas-globosas; x=36.

SIMILAR GENERA: Hymenophyllum differs by two- GÉNEROS PARECIDOS: Hymenophyllum difiere por
valved indusia and included receptacles. When dried, its indusios bivalvados y receptáculos incluídos. Cuando se
laminae are often pale reddish, whereas those of the secan, sus láminas son a menudo rojizas pálidas, mientras
trichomanoid genera are green. Polyphlebium differs by que las de los géneros trichomanoides son verdes.
epiphytic habit, filiform rhizomes, and lack of conspicuous Polyphlebium difiere por su hábito epifítico, rizomas
roots. Trichomanes subgen. Trichomanes usually differs by filiformes y falta de raíces conspicuas. Trichomanes subgen.
terrestrial habit, erect to short-creeping rhizomes, and Trichomanes difiere usualmente por hábito terrestre,
catadromous venation. 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. rupestre. Among Mundo: V. boschiana, V. collariata, V. radicans, and V. rupestre.
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.

506
Figure 247. A-C. Vandenboschia radicans. D-F. T. collariatum. (Mickel & Smith, 2004)

507
PTERIDACEAE
Vittaria J. E. Sm.
DESCRIPTION: Epiphytic; rhizomes short-creeping, DESCRIPCIÓN: Epífito; rizomas cortamente rastreros,
dorsiventral; rhizome scales clathrate. Leaves distichous, dorsiventrales, escamas del rizoma clatradas; hojas
clumped or slightly remote; petioles absent or nearly so; dísticous, amacolladas o un poco remotas; pecíolos
laminae 14–40 cm long, 1-4 mm wide, simple, entire, ausentes o casi así; láminas 15–40 cm longas, 1–4 mm
elongate-linear, coriaceous to subcoriaceous, glabrous; de ancho, simples, enteras, alargados-lineares, coriáceas
veins obscure, anastomosing, with a single row of areoles a subcoriáceas, glabras; venas obscuras, anastomosadas,
between the rachis and margin, the areoles without con una hilera de aréolas entre el raquis y el margen, las
included veinlets; sori linear, in one row between the aréolas sin venillas incluídas; soros lineares, en una sóla
rachis and margin, parallel to the margin, sunken in a hilera entre el raquis y el margen, paralelas al margen,
groove; paraphyses with a slender or slightly clavate apical hundidos en un surco; parafisos con una célula apical
cell (not obconic); spores trilete or monolete, nongreen; delgada o levemente clavada (pero no obcónica); esporas
gametophytes with paired gemmae; x=60. triletes o monoletes, no verdes; gametofitos con gemas
pareadas; x=60.

SIMILAR GENERA: Radiovittaria differs by dark petioles, GÉNEROS PARECIDOS: Radiovittaria difiere por
usually wider blades (0.2–1.0(–1.8) cm), and paraphyses pecíolos oscuros, usualmente láminas más anchas (0.2–
with an obconic apical cell. Other epiphytic genera with 1.0(–1.8) cm) y parafisos con una célula apical obcónica.
simple, entire leaves (e.g., Campyloneurum, Elaphoglossum) Otros géneros epifíticos con hojas simples y enteras (e.g.,
usually differ by non-linear sori and the presence of hairs Campyloneurum, Elaphoglossum) usualmente difieren por
or scales on the blades. soros no lineares y la presencia de pelos o escamas sobre
la lámina.

COMMENTS: Vittaria comprises 6 species, all neotropical COMENTARIOS: Vittaria consta de 6 especies, todas
except for one that occurs in Africa and offshore islands neotropicales con excepción de una que ocurre en África
in the Indian Ocean. All the species typically grow in wet e islas cercanas en el Océano Indio. Todas las especies
forests as epiphytes on trunks or in the canopy. All species crecen típicamente en bosques húmedos como epífitas
have pendulous leaves. Because of the long, linear leaves sobre troncos o en el dosel. Todas tienen hojas péndulas.
of some species, the genus has the common name of Debido a que las hojas son largas y lineares en algunas
“shoe-string fern.” The genus name is derived from the especies, el género tiene el nombre común de “helecho
Latin vitta, ribbon. The leaves are narrow and ribbon-like. 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.

508
Figure 248. A–C. Vittaria dimorpha. E–F. V. flavicosta. G–O. V. graminifolia. L–O. V. lineata. (Mickel & Smith, 2004)

509
WOODSIACEAE
Woodsia R. Br.
DESCRIPTION: Terrestrial or saxicolous; stems erect or DESCRIPCIÓN: Terrestres o saxícolas; tallos erectos
decumbent, scaly, the scales thin, translucent; sterile and o decumbentes, escamosos, las escamas delgadas,
fertile leaves monomorphous; petioles with 2 vascular translúcidas; hojas estériles y fértiles monomorfas;
bundles; laminae usually 10–35 cm long, 1-pinnate- pecíolos con 2 haces vasculares; láminas usualmente
pinnatifid to 2-pinnate-pinnatifid, scaly (on axes), usually 10–35 cm de largo, 1-pinnado-pinnatífdas a 2-pinnado-
pubescent with articulate hairs and sometimes with pinnatífidas, escamosas (sobre los ejes), usualmente
capitate-glandular hairs; pinnae sessile or nearly so; axes pubescentes con pelos articulados y a veces con pelos
grooved, the grooves decurrent; veins free, the tips slightly capitados-glandulosos; pinnas sésiles o casi sésiles; ejes
thickened (hydathodous); sori round; indusia attached surcados, los surcos decurrentes; nervaduras libres,
around the base of the receptacle, splitting at los ápices un poco engrosados (hidatodos); soros
maturity into several lobes or many hairs that form redondos; indusios unidos alrededor de la base del
a loose cup around the sporangia; x=38, 39, 41. receptáculo, se hende en varios lobulos o muchos
pelitos que forman una copa suelta alrededor de los
esporangios; x=38, 39, 41.

SIMILAR GENERA: Cystopteris differs by the indusium GÉNEROS PARECIDOS: Cystopteris difiere por el
attached on the proximal side of the receptacle only. indusio unido solo al lado proximal del receptáculo.
Cheilanthes differs by one vascular bundle at the base of Cheilanthes difiere por un haz vascular en la base del
the petiole and marginal sori protected by false indusia. pecíolo y soros marginales protegidos por indusios falsos.

COMMENTS: Woodsia contains about 35 species and COMENTARIOS: Woodsia consta de casi 35 especies
is primarily in the temperate zones of North America, y es principalmente en zonas templadas de América del
Europe, and Asia. In Latin America, it is most diverse in Norte, Europa y Asia. En América Latín, es más diverso en
Mexico (8 spp.). Only one species occurs in South America México (8 spp.). Solamente una especie existe en América
and the Antilles (Hispaniola): Woodsia montevidensis del Sur y las Antillas (Hispaniola): Woodsia montevidensis
(Spreng.) Hyl. It usually grows on rocks from 2500–4000 (Spreng.) Hyl. Usualmente crece sobre rocas desde 2500–
m. The indusia of this species are peculiar because they 4000 m. Los indusios de esta especie son peculiares por
often have thickened, whitish cells. The genus name que tienen a menudo células engrosadas blanquecinas.
honors Joseph Woods (1776–1864), British architect and El nombre genérico honora Joseph Woods (1776–1864),
author of A Tourist’s Flora. arquitecto británico y autor de 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.

510
Figure 249. A–E. Woodsia plummerae. F–K. W. cystopteroides. L–O. W. cochisensis. (Mickel & Smith, 2004)

511
BLECHNACEAE
Woodwardia Sm.
Anchistea C. Presl; Lorinseria C. Presl

DESCRIPTION: Terrestrial; rhizomes decumbent DESCRIPCIÓN: Terrestres; rizomas decumbentes

or long-creeping, scaly; sterile and fertile leaves o largamente rastreros, escamosos; hojas estériles
monomorphous or (in Lorinseria) dimorphous; laminae y fértiles monomorfas o (en Lorinseria) dimorfas;
1-pinnate-pinnatifid (in Mesoamerica) to 2-pinnate; láminas 1-pinnado-pinnatífidas (en Mesoamérica)
pinnae pinnatifid, the segments spinulose (in Mesoamerica) a 2-pinnadas; pinnas pinnatífidas, los segmentos
or entire; aerophores absent at the base of the pinnae espinulosos (en Mesoamérica) o enteros; aeróforos
abaxially, and along the petioles; veins areolate, at least ausentes en las bases de las pinnas en el envés y a lo largo
along the costae, without included veins; sori parallel del pecíolo; nervaduras areoladas, al menos a lo largo de
to the costae, one per areole, in most species deeply las costas, sin nérvulos incluidos; soros paralelos a las
immersed; indusia opening toward the costae, not the costas o cóstulas, uno por aréola, en la mayoría de las
margin. Spores monolete, nongreen. x=34, 35. especies profundamente inmersos; indusios abriéndose
hacia la costa, no el margen. Esporas monoletes, no
verdes. x=34, 35..

SIMILAR GENERA: Tectaria differs by included veinlets GÉNEROS PARECIDOS: Tectaira difiere por nérvulos
in the areoles, round sori, and plane (not grooved) upper incluidos en las areolas, soros redondeados, y ráquises
surfaces of the rachises. Thelypteris has two vascular planos (no surcados) en la superficie superior. Thelypteris
bundles in the petiole and (usually) round sori. tiene dos haces vasculares en el pecíolo y (usualmente)
soros redondos.

COMMENTS: Woodwardia is mostly north-temperate COMENTARIOS: Woodwardia es principalmente

and contains about 14 species. In the Neotropics, it en el norte templado y consta de 14 especies. En el
extends from Mexico to Nicaragua in wet forests above Neotrópico se extiende desde México hasta Nicaragua
1500 m. It contains two neotropical species, W. spinulosa en bosques húmedos arriba de 1500 m. Contiene
M. Martens & Galeotti and W. martinezii Maxon ex Weath., dos especies neotropicales, W. spinulosa M. Martens &
and their hybrid, W. X semicordata Mickel & Beitel. The Galeotti y W. martinezii Maxon ex Weath., y su híbrido W.
center of diversity for the genus is in China, where six or X semicordata Mickel & Beitel. El centro de diversificación
more species occur. Woodwardia areolata, a species of the está en China, en donde existen seis o más especies.
eastern United States, is often segregated as Lorinseria Woodwardia areolata, una especies de los Estados Unidos
because of its dimorphic fronds and adnate pinnae. The oriental, es a menudo segregado como Lorinseria a causa
common name for the genus is “chain fern,” alluding to de sus hojas dimorfas y pinnas adnatas. El nombre común
the chain-like row of sori along the costae or costules. para el género es “helecho de cadena,” en alusión a la
The genus name honors Thomas Jenkinson Woodward hilera de soros que asemejan cadenas por las costas y
(1745–1820), British phycologist. cóstulas. El nombre génerico honora a Thomas Jenkinson
Woodward (1745–1820), ficólogo británico.

LITERATURE: 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.

512
Figure 250. A-E. Woodwardia spinulosa. (Mickel & Smith, 2004).

513
POLYPODIACEAE
Zygophlebia L.E. Bishop
DESCRIPTION: Epiphytic; rhizome scales not clathrate, DESCRIPCIÓN: Epífitas; escamas del rizoma no
shiny, cilate or glandular-papillose on the margins; clatradas, lustrosas, ciliadas o glanduloso-papilosas en los
phyllopodia present, swollen; petiole equalling or longer márgenes; filopodios presentes, hinchados; pecíolo tan
than the lamina; laminae pinnatisect, with or without setae, largo como la lámina o más largo; láminas pinnatisectas, con
with simple or branched glandular hairs; hydathodes o sin setas, con tricomas glandulosos simples o bifurcados;
absent; veins 1—2(—3) forked, connivent at the apices hidatodos ausentes; nervaduras 1—2(—3) bifurcadas,
or rarely free; sori separate, round, paraphyses presnt, conniventes en los ápices o raramente libres; soros
consisting of a 1-seriate stalk with 2 or 3 brown glands separados, redondos, parafisos presentes, constituidos
distally; spores green. 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 características importantes de Zygophlebia (filopodios
veins) are often difficult to observe due to the covering y nervaduras areoladas), son a menudo difíciles de
of scales on the rhizome and the thickness of the lamina. observar debido a la cobertura de escamas del rizoma y
The phyllopodia are evident as a swollen base with al grosor de la lámina. Los filopodios son evidentes como
a darker color than the petiole. The change in color is una base hinchada con un color más oscuro que el del
usually abrupt. The absence of hydathodes on the upper pecíolo. El cambio de color es típicamente abrupto. La
surface of the lamina will separate Zygophlebia from the ausencia de hidatodos en el haz separará Zygophlebium
other neotropical genera of Grammitidaceae except de los otros géneros neotropicales de Grammitidaceae
for Ceradenia and Enterosora (which see). The leaves of con excepción de Ceradenia y Enterosora (véase). Las
Zygophlebia have a peculiar habit, but this characteristic hojas de Zygophlebia tienen un hábito peculiar, pero esta
is found in other grammitid ferns: the petiole is nearly característica la poseen otros helechos gramitidáceos por
horizontal, i.e., perpendicular to the branch or trunck on igual: el pecíolo es casi horizontal, i.e., casi perpendicular
which the plant is epiphytic and is sharply geniculate at its a la rama o tronco en el que la planta es epífita y es
apex so that the lamina is pendulous. 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.

514
Figure 251. Zygoplebia werfii. Note in C the anastomosing veins. These are often difficult to see because the laminae are
thick. (from Bishop, 1989)

515
 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, place a small drop of ACH (50% acetocarmine and 50% Hoyer’s solution 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.

516
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 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. 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 (50% aceto-carmine, 50% Hoyer’s solution), it is best to view the material un-
der a phase-contrast microscope because the chromosomes stain much lighter than if straight aceto-carmine is used.
For further information about the use of Hoyer’s solution for mounting spores and leaf fragments, see Anderson, L.
W. 1954. Hoyer’s solution as rapid permanent mounting medium for bryophytes. Bryologist 57: 242–244.

518
Appendix 2 | Principal Fern and Lycophyte 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. & J. Beitel 1988. Pteridophyte flora of Oaxaca, Mexico. Memoirs of the New York Botanical Garden 48: 1–568.
Mickel, J. T. & A. R. Smith. 2004. The pteridophytes of Mexico. Memoirs of the New York Botanical Garden 88: 1–1055.
Mickel, J. T. 1992. Pteridophytes. In: R. McVaugh, editor, Flora Novo-Galiciana 17: 120–455.
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
Francisco.
Stolze, R. G. 1976. Ferns and fern allies of Guatemala, part I. Ophioglossaceae through Cyatheaceae. Fieldiana, Botany 39: i–v
+ 1–130.
Stolze, R. G. 1981. Ferns and fern allies of Guatemala, part II. Polypodiaceae. Fieldiana, Botany new series, 6: i–ix + 1–522.
Stolze, R. G. 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.
Kramer, K. U. 1978. The pteridophytes of Suriname. Uitgaven. Natuurwetenschappelijke Studiekring voor Suriname
Nederlandse Antillen 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.
Rodríguez R., R. 1995. Pteridophyta. Pages 119–309. In: C. Marticorena & R. Rodríguez, Flora de Chile,Vol. I, Pteridophyta—
Gymnospermae. Univ. de Concepción, Chile.
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.
Smith, A. R. 1992. 16. Thelypteridaceae. In: R. M. Tryon & R. G. Stolze, Pteridophyta of Peru. Fieldiana, Botany new series 29: iii
+ 1–80.
Smith, A. R. 1995. Pteridophytes. In: Steyermark et al., general editors. Flora of the Venezuelan Guayana, volume 2. Pages
1–334. Timber 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.
Tryon, R. M. & R. G. Stolze 1989. Pteridophyta of Peru, Part II, 13. Pteridaceae-15. Dennstaedtiaceae. Fieldiana, Botany new
series 22: iii + 1–128.

519
Tryon, R. M. & R. G. Stolze 1991. Pteridophyta of Peru, Part IV, 17. Dryopteridaceae. Fieldiana, Botany new series 27: iii +
1–176.
Tryon, R. M. & R. G. Stolze 1993. Pteridophyta of Peru, Part V, 18. Aspleniaceae—21. Polypodiaceae. Fieldiana, Botany new
series 27: iii + 1–176.
Tryon, R. M. & R. G. Stolze 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

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
Vasculares. 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
Harvard University, Jamaica Plain, Massachusetts.
Proctor, G. R. 1985. Ferns of Jamaica. British Museum (Natural History), London.
Proctor, G. R. 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 Neotropical Pteridophytes

LYCOPHYTES 11 Anemiaceae 24. Saccolomataceae

Lycopodiaceae Anemia Saccoloma
Huperzia Ornithopteris
Lycopodiella 25. Dennstaedtiaceae
Lycopodium 12. Schizaeaceae Blotiella
Actinostachys Dennstaedtia
Selaginellaceae Schizaea Histiopteris
Selaginella Hypolepis
13. Marsileaceae Microlepia
Isoëtaceae Marsilea Paesia
Isoëtes Pilularia Pteridium
Regnellidium
26. Pteridaceae
FERNS 14. Salviniaceae Acrostichum
1. Psilotaceae Azolla Adiantopsis
Psilotum Salvinia Adiantum
Allosorus
2. Ophioglossaceae 15. Thyrsopteridaceae Ananthacorus
Botrychium Thyrsopteris Anetium
Ophioglossum Anogramma
16. Loxsomataceae Antrophyum
3. Equisetaceae Loxsomopsis Argyrochosma
Equisetum Aspidotis
17. Culcitaceae Astrolepis
4. Marattiaceae Culcita Bommeria
Danaea Ceratopteris
Eupodium 18. Plagiogyriaceae Cheilanthes
Marattia Plagiogyria Cheiloplecton
Cryptogramma
5. Osmundaceae 19. Cibotiaceae Doryopteris
Osmunda Cibotium Gaga
Osmundastrum Hecistopteris
20. Cyatheaceae Hemionitis
6. Hymenophyllaceae Alsophila Jamesonia
Hymenophyllum Cyathea Llavea
Trichomanes Gymnosphaera Mildella
Sphaeropteris Myriopteris
7. Gleicheniaceae Nephropteris
Dicranopteris 21. Dicksoniaceae Notholaena
Diplopterygium Dicksonia Pellaea
Gleichenella Lophosoria Pentagramma
Sticherus Pityrogramma
22. Metaxyaceae Polytaenium
8. Dipteridaceae Metaxya Pteris
(Old World) Pterozonium
23. Lindsaeaceae Radiovittaria
9. Matoniaceae Lindsaea Scoliosorus
(Old World) Lonchitis Trachypteris
10. Lygodiaceae Odontosoria Vittaria
Lygodium Sphenomeris

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27. Cystopteridaceae 38. Dryopteridaceae 42. Davalliaceae
Cystopteris Anapausia (Old World)
Gymnocarpium Arachniodes
Atalopteris 43a. Polypodiaceae
28. Aspleniaceae Bolbitis Campyloneurum
Asplenium Ctenitis Loxogramme
Cyclodium Microgramma (incl. Solanopteris)
29. Hemidictyaceae Dryopteris Niphidium
Hemidictyum Elaphoglossum Pecluma
Lastreopsis Phlebodium
30. Diplaziopsidaceae Lomagramma Platycerium
(Asia; temperate E North Amer.) Maxonia Pleopeltis (incl. Dicranoglossum &
Megalastrum Neurodium)
31. Rhachidosoraceae Mickelia Polypodium
(Old World) Olfersia Serpocaulon
Parapolystichum Synnamia
32. Woodsiaceae Phanerophlebia
Woodsia Polybotrya 43b. Grammitid clade
Polystichopsis Alansmia
33. Athyriaceae Polystichum Ascogrammitis
Athyrium Pradoia Ceradenia
Diplazium (incl. Callipteris) Rumohra Cochlidium
Stigmatopteris Enterosora
34. Thelypteridaceae Galactodenia
Macrothelypteris 39. Lomariopsidaceae Grammitis
Phegopteris Cyclopeltis Lellingeria
Thelypteris Dracoglossum Leucotrichum
Lomariopsis Lomaphlebia
35. Blechnaceae Nephrolepis Luisma
Blechnum Melpomene
Salpichlaena 40. Tectariaceae Micropolypodium
Woodwardia Dictyoxiphium Moranopteris
Hypoderris Mycopteris
36. Onocleaceae Tectaria Stenogrammitis
Matteuccia Triplophyllum Terpsichore
Onoclea Zygophlebia
Onocleopsis 41. Oleandraceae
37. Hypodematiaceae Oleandra
Didymochlaena

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Appendix 4 | accepted genera of ferns world-wide
List includes their main synonyms, and the families to which they belong. A more complete list of rarely used syn-
onyms (and accepted generic names) can be found in Christenhusz et al. (2011).

FAMILY & NUMBER OF GENERA Acystopteris (33) Callipteris = Diplazium

Adenoderris = Polystichum Callistopteris (6)
Ophioglossaceae (1) Adenophorus (44) Calochlaena (21)
Psilotaceae (2) Adiantopsis (26) Calymmodon (44)
Equisetaceae (3) Adiantum (26) Camptodium = Tectaria
Marattiaceae (4) Aenigmopteris (41) Campyloneurum (44)
Osmundaceae (5) Afropteris = Pteris Caobangia = Lemmaphyllum
Hymenophyllaceae (6) Aglaomorpha (44) Cardiomanes = Hymenophyllum
Gleicheniaceae (7) Alansmia (44) Cassebeera (26) = Doryopteris
Dipteridaceae (8) Aleuritopteris = Allosorus Cephalomanes (6)
Matoniaceae (9) Allosorus (26) Ceradenia (44)
Lygodiaceae (10) Alsophila (20) Ceratopteris (26)
Anemiaceae (11) Amauropelta = Thelypteris Cerosora (26)
Schizaeaceae (12) Ampelopteris = Cyclosorus Ceterach = Asplenium
Marsileaceae (13) Amphiblestra = Tectaria Ceterachopsis = Asplenium
Salviniaceae (14) Amphineuron = Cyclosorus Cheilanthes (26)
Thyrsopteridaceae (15) Ananthacorus (26) Cheilanthopsis (33)
Loxomataceae (16) Anapausia = Bolbitis Cheiloplecton (26)
Culcitaceae (17) Anarthropteris = Loxogramme Cheiroglossa = Ophioglossum
Plagiogyriaceae (18) Anchistea = Woodwardia Cheiropleuria (8)
Cibotiaceae (19) Anemia (11) Chieniopteris = Woodwardia
Cyatheaceae (20) Anetium (26) Chingia = Cyclosorus
Dicksoniaceae (21) Angiopteris (4) Chlamidogramme = Tectaria
Metaxyaceae (22) Anisocampium (33) Christella = Cyclosorus
Lindsaeaceae (23) Anogramma (26) Christensenia (4)
Saccolomataceae (24) Anopteris = Pteris Christiopteris (44)
Dennstaedtiaceae (25) Antigramma = Asplenium Chrysogrammitis (44)
Pteridaceae (26) Antrophyum (26) Cibotium (19)
Cystopteridaceae (27) Arachniodes (38) Cionidium = Tectaria
Aspleniaceae (28) Araiostegia (43) Cnemidaria = Cyathea
Hemidictyaceae (29) Araiostegiella (43) Cochlidium (44)
Diplaziopsidaceae (30) Archangiopteris = Angiopteris Colysis (44)
Rhachidosoraceae (31) Argyrochosma (26) Coniogramme (26)
Woodsiaceae (32) Arthrobotrya (38) Coptodipteris (25)
Athyriaceae (33) Arthromeris (44) Cornopteris (33)
Thelypteridaceae (34) Arthropteris (41) Coryphopteris = Thelypteris
Blechnaceae (35) Ascogrammitis (44) Cosentinia (26)
Onocleaceae (36) Aspidotis (26) Costaricia = Dennstaedtia
Hypodematiaceae (37) Aspleniopsis = Austrogramme Coveniella = Lastreopsis
Dryopteridaceae (38) Asplenium (28) Crepidomanes (6)
Lomariopsidaceae (39) Astrolepis (26) Crypsinus = Selliguea
Nephrolepidiaceae (40) Atalopteris (38) Cryptogramma (26)
Tectariaceae (41) Ataxipteris (38) Ctenitis (38)
Oleandraceae (42) Athyriopsis = Deparia Ctenitopsis = Tectaria
Davalliaceae (43) Athyrium (33) Ctenopterella (44)
Polypodiaceae (44) Austrogramme (26) Ctenopteris (44)
Azolla (14) Culcita (17)
GENUS & NUMBER OF SPECIES Belvisia = Lepisorus Currania = Gymnocarpium
Blechnum (35) Cyathea (20)
Abrodictyum (6) Blotiella (25) Cyclodium (38)
Acrophorus (38) Bolbitis (38) Cyclogramma = Cyclosorus
Acrorumohra (38) Bommeria (26) Cyclopeltis (39)
Acrosorus (44) Botrychium (1) Cyclosorus (34)
Acrostichum (26) Botrypus = Botrychium Cyrtogonellum (38)
Actiniopteris (26) Brainea (35) Cyrtomidictyum (38)
Actinostachys (12) Calciphilopteris (26) Cyrtomium (38)

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Cystoathyrium (38) Hemitelia = Cyathea Melpomene (44)
Cystodium (23) Heterogonium = Tectaria Meniscium = Cyclosorus
Cystopteris (27) Hewardia = Adiantum Menisorus = Cyclosorus
Danaea (4) Hicriopteris = Dicranopteris Merinthosorus = Aglaomorpha
Dasygrammitis (44) Himalayopteris (44) Mesophlebion = Cyclosorus
Davallia (43) Histiopteris (25) Metathelypteris = Thelypteris
Davallodes (43) Holcochlaena (26) Metaxya (22)
Dendroconche (44) Holodictyum = Asplenium Mickelia (38)
Dennstaedtia (25) Holostachyum = Aglaomorpha Microgonium = Didymoglossum
Deparia (33) Homalosorus (30) Microgramma (44)
Diacalpe (38) Humata = Davallia Microlepia (25)
Dicksonia (21) Hyalotrichopteris = Campyloneurum Micropolypodium (44)
Dicranoglossum = Pleopeltis Hymenasplenium (28) Microsorum (44)
Dicranopteris (7) Hymenocystis = Woodsia? Microstaphyla = Elaphoglossum
Dictymia (44) Hymenoglossum = Hymen-ophyllum Microtrichomanes = Hymenophyllum
Dictyodroma = Diplazium Hymenophyllopsis = Cyathea Mildella (26)
Dictyoxiphium = Tectaria Hymenophyllum (6) Mohria = Anemia
Didymochlaena (37) Hypodematium (37) Monachosorum (25)
Didymoglossum (6) Hypoderris (41) Monogramma (26)
Diellia = Asplenium Hypolepis (25) Monomelangium = Diplazium
Diplaziopsis (30) Isoloma = Lindsaea Moranopteris (44)
Diplazium (33) Jamesonia (26) Mycopteris (44)
Diplopterygium (7) Japanobotrychium = Botrychium Myriopteris (26)
Dipteris (8) Kaulinia (44) Nematopteris (44)
Doodia (35) Kontumia (44) Neocheiropteris (44)
Doryopteris (26) Lastreopsis (38) Neolepisorus (44)
Dracoglossum (39) Lecanopteris (44) Neolindsaea (23)
Drymoglossum = Pyrrosia Lellingeria (44) Nephelea = Alsophila
Drymotaenium = Lepisorus Lemmaphyllum (44) Nephopteris (26)
Drynaria (44) Lepidomicrosorium (44) Nephrolepis (40)
Dryoathyrium = Deparia Lepisorus (44) Neurocallis = Pteris
Dryopolystichum (38) Leptochilus (44) Neurodium = Pleopeltis
Dryopsis (38) Leptolepia (25) Niphidium (44)
Dryopteris (38) Leptopteris (5) Niphobolus = Pyrrosia
Edanyoa = Bolbitis Leptorumohra = Arachniodes Notogrammitis (44)
Egenolfia = Bolbitis Leucostegia (37) Notholaena (26)
Elaphoglossum (38) Leucotrichum (44) Nothoperanema = Dryopteris
Enterosora (44) Lindsaea (23) Ochropteris (26)
Equisetum (3) Lithostegia (38) Odontosoria (23)
Eriosorus = Jamesonia (26) Litobrochia = Pteris Oenotrichia (25)
Eupodium (4) Llavea (26) Oenotrichia p.p (38)
Fadyenia = Tectaria Lomagramma (38) Oleandra (42)
Fourniera = Sphaeropteris Lomaphlebia (44) Oleandropsis (44)
Gaga (26) Lomariopsis (39) Olfersia (38)
Galactodenia (44) Lonchitis (23) Onoclea (36)
Glaphyropteridopsis = Cyclosorus Lophosoria (21) Onocleopsis (36)
Gleichenella (7) Lorinseria = Woodwardia Onychium (26)
Gleichenia (7) Loxogramme (44) Ophioderma = Ophioglossum
Goniophlebium (44) Loxoma (16) Ophioglossum (1)
Goniopteris = Cyclosorus Loxoscaphe = Asplenium Oreogrammitis (44)
Grammitis (44) Loxsomopsis (16) Oreopteris = Thelypteris
Gymnocarpium (27) Luisma (44) Ormoloma = Lindsaea
Gymnogramma = Hemionitis Lunathyrium = Deparia Ormopteris = Pellaea
Gymnopteris = Hemionitis Lygodium (10) Ornithopteris (11)
Gymnogrammitis (44) Macrothelypteris (34) Orthiopteris = Saccoloma
Gymnosphaera (20) Mankyua (1) Osmolindsaea (23)
Haplopteris (26) Marattia (4) Osmunda (5)
Hecistopteris (26) Marsilea (13) Pachypleuria (43)
Helminthostachys (1) Matonia (9) Paesia (25)
Hemidictyum (29) Matteuccia (36) Papuapteris = Polystichum
Hemigramma = Tectaria Maxonia (38) Paraceterach (26)
Hemionanthes (26) Mecodium = Hymenophyllum Paragramma (44)
Hemionitis (26) Megalastrum (38) Paragymnopteris (26)

524
Parahemionitis (26) Pterozonium (26) Trigonospora = Cyclosorus
Parapolystichum (38) Ptilopteris = Monachorosum Triplophyllum (41)
Paraselliguea (44) Ptisana (4) Trismeria = Pityrogramma
Parkeria = Ceratopteris Pycnodoria = Pteris Tryonella (26)
Parathelypteris = Thelypteris Pyrrosia (44) Vaginularia = Monogramma
Pecluma (44) Quercifilix = Tectaria Vandenboschia (6)
Pelazoneuron = Cyclosorus Radiogrammitis (44) Vittaria (26)
Pellaea (26) Radiovittaria (26) Wagneriopteris (44)
Peltapteris = Elaphoglossum Regnellidium (13) Wibellia (43)
Pentagramma (26) Revwattsia = Dryopteris (38) Woodsia (32)
Pentarhizidium (36) Rhachidosorus (28c) Woodwardia (35)
Peranema (38) Rheopteris (26) Xiphopterella (44)
Pessopteris = Niphidium Rosenstockia = Hymenophyllum Xiphopteris = Cochlidium
Phanerophlebia (38) Rumohra (38) Xyropteris (23)
Phanerophlebiopsis = Arachniodes Saccoloma (24) Zygophlebia (44)
Phanerosorus (9) Sadleria (35)
Phegopteris (34) Salpichlaena (35)
Phlebodium (44) Salvinia (14)
Photinopteris = Aglaomorpha Sceptridium = Botrychium
Phyllitis = Asplenium Schaffneria = Asplenium
Phymatopteris (44) Schizaea (12)
Phymatosorus (44) Scleroglossum (26)
Pilularia (13) Scoliosorus (26)
Pityrogramma (26) Scyphularia = Davallia
Plagiogyria (18) Selliguea (44)
Platycerium (44) Serpocaulon (44)
Platygyria = Lepisorus Serpyllopsis = Hymenophyllum
Platyloma = Pellaea Sinephropteris = Asplenium
Platyzoma (26) Sinopteris = Aleuritopteris
Plecosorus = Polystichum Solanopteris = Microgramma
Pleocnemia (41) Sphaerocionium =Hymenophyllum
Pleopeltis (44) Sphaeropteris (20)
Plesioneuron = Cyclosorus Sphaerostephanos = Cyclosorus
Pleuroderris = Tectaria Sphenomeris (23)
Pleurosoriopsis (44) Steenisioblechnum (35)
Pleurosorus = Asplenium Stegnogramma = Cyclosorus
Pneumatopteris = Cyclosorus Steiropteris = Cyclosorus
Podosorus (44) Stenochlaena (35)
Polybotrya (38) Stenogrammitis (44)
Polyphlebium (6) Stenolepia (38)
Polypodioides (44) Sticherus (7)
Polypodiopteris (44) Stigmatopteris (38)
Polypodium (44) Stromatopteris (7)
Polystichopsis (38) Synammia (44)
Polystichum (38) Syngramma (26)
Polytaenium (26) Taenitis (26)
Pradoia (38) Tapeinidium (23)
Pronephrium = Cyclosorus Tectaria (41)
Prosaptia (44) Teratophyllum (38)
Protowoodsia (33) Terpsichore (44)
Psammiosorus (41) Thamnopteris = Asplenium
Pseudocolysis = Pleopeltis Thelypteris (34)
Pseudocyclosorus = Cyclosorus Themelium (44)
Pseudocystopteris = Athyrium Thylacopteris (44)
Pseudodrynaria = Aglaomorpha Thyrsopteris (15)
Pseudophegopteris (34) Thysanosoria (39)
Pseudotectaria = Tectaria Tmesipteris (2)
Psilotum (2) Todea (5)
Psomiocarpa (41) Tomophyllum (44)
Pteridium (25) Trachypteris (26)
Pteridoblechnum (35) Trichipteris = Cyathea
Pteridrys (41) Tricholepidium (44)
Pteris (26) Trichomanes (6)

525
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 253)

Figure 252. 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, lep-
tosporangiates; 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 253.

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 (Figure 254)

Figure 253. Continued from Figure 252. 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 254.

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 (Figure 255)

Figure 254. Continued from Figure 253. 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. Fam-
ilies recognized in the most recent classification of extant ferns (Smith & al., 2006b) are indicated in boxes between trees: Asp, As-
pleniaceae; Ble, Blechnaceae; Ono, Onocleaceae; The, Thelypteridaceae; Woo, Woodsia ceae. Phylogeny continues in Figure 255.

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 (Figure 256)

Figure 255. Continued from Figure 254. Major ciades dis cussed in text are indicated in circles on trees: dc, di morphic climb-
ers; 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 256.

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Figure 256. Continued from Figure 255. 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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