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Monograph of the West Indian fern genus Polystichopsis (Dryopteridaceae)

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Abstract

Keys, full synonymy, descriptions, specimens examined, and illustrations are provided for the fern genus Polystichopsis. The genus occurs in the West Indies, with disjunct populations in Nicaragua and Isla Margarita, Venezuela. Seven species and one hybrid are recognized. The genus can be distinguished from other dryopteroid genera by the combination of creeping rhizomes, distichous leaves, lack of scales on the mature leaves, and straight, whitish multicellular hairs on the leaves. Although the genus has creeping rhizomes, it does not climb as do many other closely related dryopteroid genera. A new species, P. puberula, and new hybrid, P. × sanchezii (=P. chaerophylloides × P. pubescens), are described. Lectotypes are designated for Nephrodium pubescens var. breviculum , P. leucochaete, and P. muscosa. A list is provided of 28 names excluded from Polystichopsis, and another list is provided for accepted names and their synonyms that apply in the genus.

Resumen

Claves, sinonimia completa, descripciones, ejemplares examinados e ilustraciones se suministra para el género de helechos Polystichopsis. El mismo existe en las Antillas con poblaciones disyuntas en Nicaragua y la Isla Margarita, Venezuela. Se reconocen siete especies y un híbrido. El género se distingue de otros géneros dryopteroides por la combinación de rizomas reptantes, hojas dísticas, ausencia de escamas en las hojas maduras y pelos blanquecinos en las hojas. Aunque posee rizomas rastreros, no es trepador como muchos otros géneros driopteroides relacionados. Se describe una especie nueva, P. puberula, y un híbrido nuevo, P. × sanchezii (=P. chaerophylloides × P. pubescens). Se designan lectotipos para Nephrodium pubescens var. breviculum , P. leucochaete y P. muscosa. Se brinda una lista de 28 nombres excluidos de Polystichopsis, y otra lista para los nombres y sus sinónimos que se aplican en el género.

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Acknowledgments

We thank the following herbaria for loaning us specimens: B, BH, BM, F, GH, MO, NY, P, S, UC, US, UPRRP. We also thank Judith Garrison Hanks for help taking SEM photomicrographs of the spores. Carlos Sánchez reviewed the manuscript and provided helpful comments about the Cuban species of Polystichopsis and Cuban specimen localities. We thank Haruto Fukuda for drawing Fig. 2, and Klei Sousa for drawing Figs. 5 and 6.

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Correspondence to Jefferson Prado.

Appendices

Appendix I

List of Species

  1. 1.

    Polystichopsis chaerophylloides (Poir.) C. V. Morton

  2. 2.

    Polystichopsis leucochaete (Slosson) J. Prado & R. C. Moran

  3. 3.

    Polystichopsis lurida (Jenman ex Underw. & Maxon) C. V. Morton

  4. 4.

    Polystichopsis muscosa (Vahl) Proctor

  5. 5.

    Polystichopsis puberula J. Prado & R. C. Moran

  6. 6.

    Polystichopsis pubescens (L.) C. V. Morton

  7. 7.

    Polystichopsis sericea (Mett. ex D. C. Eaton) C. Sánchez.

  8. 8.

    Polystichopsis × sanchezii J. Prado & R. C. Moran

Appendix II

List of accepted names and synomyms for Polystichopsis and excluded taxa. Accepted names are in bold, synonyms in italics, and excluded taxa indicated by “ET”. Number in parentheses refers to the species numbers assigned in the above paragraph.

Arachniodes amabilis (Blume) Tindale (ET)

Arachniodes argillicola (Proctor) Proctor (ET)

Arachniodes aristata (G. Forst.) Tindale (ET)

Arachniodes assamica (Kuhn) Ohwi (ET)

Arachniodes bella (C. Chr.) Ching (ET)

Arachniodes cavaleriei (Christ) Ohwi (ET)

Arachniodes chaerophylloides (Poir.) Proctor (1)

Arachniodes chinensis (Rosenst.) Ching (ET)

Arachniodes denticulata (Sw.) Ching (ET)

Arachniodes haniffii (Holttum) Ching (ET)

Arachniodes lurida (Jenman ex Underw. & Maxon) Proctor (3)

Arachniodes miqueliana (Maxim. ex Franch. & Sav.) Ohwi (ET)

Arachniodes mutica (Franch. & Sav.) Ohwi (ET)

Arachniodes nipponica (Rosenst.) Ohwi(ET)

Arachniodes ochropteroides (Baker) Lellinger (ET)

Arachniodes pubescens (L.) O. Kuntze (6)

Arachniodes rigidissima (Hook.) Proctor (ET)

Arachniodes simplicior (Makino) Ohwi (ET)

Arachniodes simplicior (Makino) Ohwi var. major (Tagawa) Ohwi (ET)

Arachniodes sinomiqueliana (Ching) Ohwi (ET)

Arachniodes speciosa (D. Don) Ching (ET)

Arachniodes sporadosora (Kunze) Nakaike (ET)

Arachniodes standishii (T. Moore) Ohwi (ET)

Arachniodes yasu-inouei Sa. Kurata (ET)

Aspidium muscosum (Vahl) Sw. (4)

Aspidium pubescens (L.) Sw. (6)

Dryopteris chaerophylloides (Poir.) C. Chr. (1)

Dryopteris chaerophylloides (Poir.) C. Chr. var. sericea (Mett. ex D. C. Eaton) C. Chr. (7)

Dryopteris hasseltii (Blume) C. Chr. (ET)

Dryopteris leucochaete Slosson (2)

Dryopteris lurida f. leucochaete (Slosson) C. Chr. (2)

Dryopteris lurida Jenman ex Underw. & Maxon (3)

Dryopteris lurida var. sericea (Mett.) C. Chr. (7)

Dryopteris maximowiczii (Baker) Kuntze (ET)

Dryopteris pubescens (L.) Kuntze (6)

Dryopteris pubescens var. haitiensis C. Chr. (6)

Dryopteris pubescens var. muscosa (Vahl) C. Chr. (4)

Dryopteris wardii (Baker) Kuntze (ET)

Lastreopsis amplissima (C. Presl) Tindale (ET)

Lastreopsis chaerophylloides (Poir.) Tindale (1)

Lastreopsis lurida (Jenman ex Underw. & Maxon) Tindale (3)

Lastreopsis pubescens (L.) Tindale (6)

Nephrodium pubescens (L.) Desv. (6)

Nephrodium pubescens (L.) Desv. var. breviculum Jenman (6)

Phegopteris portoricensis (Spreng.) Fée (1)

Phegopteris sericea Mett. ex D. C. Eaton (7)

Phegopteris villosa Fée (4)

Polypodium chaerophylloides Poir. (1)

Polypodium muscosum Vahl (4)

Polypodium portoricense Spreng. (ET)

Polypodium pubescens L. (6)

Polystichopsis amabilis (Blume) Tagawa (ET)

Polystichopsis amplissima (C. Presl) Abbiatti (ET)

Polystichopsis amplissima (C. Presl) Abbiatti var. phaeochlamys (Fée) Abbiatti (ET)

Polystichopsis argillicola (Proctor) Proctor (ET)

Polystichopsis aristata (G. Forst.) Holttum (ET)

Polystichopsis assamica (Kuhn) Tagawa (ET)

Polystichopsis bella (C. Chr.) Tardieu (ET)

Polystichopsis cavaleriei (Christ) Tagawa (ET)

Polystichopsis chaerophylloides (Poir.) C. V. Morton (1)

Polystichopsis chinensis (Rosenst.) Holttum (ET)

Polystichopsis cubensis (Kuhn) Lellinger (ET)

Polystichopsis haniffii (Holttum) Holttum (ET)

Polystichopsis hasseltii (Blume.) Holttum (ET)

Polystichopsis holttumii (Tagawa) Seriz (ET)

Polystichopsis leucochaete (Slosson) J. Prado & R. C. Moran (2)

Polystichopsis lurida (Jenman ex Underw. & Maxon) C. V. Morton (3)

Polystichopsis lurida f. leucochaete (Slosson) Proctor (2)

Polystichopsis lurida (Jenman ex Underw. & Maxon.) C. V. Morton var. sericea (Mett. ex D.C. Eaton) Duek, Adansonia 11: 724. 1971. (7)

Polystichopsis macrostegia (Hook.) Nakaike (ET)

Polystichopsis maximowiczii (Baker) Tagawa (ET)

Polystichopsis miqueliana (Maxim. ex Franch. & Sav.) Tagawa (ET)

Polystichopsis muscosa (Vahl) Proctor (4)

Polystichopsis mutica (Franch. & Sav.) Tagawa (ET)

Polystichopsis nipponica (Rosenst.) Tagawa (ET)

Polystichopsis ochropteroides (Baker) C. V. Morton (ET)

Polystichopsis pseudoaristata (Tagawa) Tagawa (ET)

Polystichopsis puberula J. Prado & R. C. Moran (5)

Polystichopsis pubescens (L.) C. V. Morton (6)

Polystichopsis pubescens var. haitiensis (C. Chr.) Proctor (6)

Polystichopsis puncticulata (Alderw.) Holttum (ET)

Polystichopsis × sanchezii J. Prado & R. C. Moran

Polystichopsis sericea (Mett. ex D. C. Eaton) C. Sánchez (7)

Polystichopsis simplicior (Makino) Tagawa (ET)

Polystichopsis simplicior (Makino) Tagawa var. major (Tagawa) Tagawa (ET)

Polystichopsis sinomiqueliana (Ching) Tagawa (ET)

Polystichopsis standishii (T. Moore) Tagawa (ET)

Polystichopsis villosa (Fée) Christenh. (4)

Polystichopsis wardii (Baker) Tardieu (ET)

Rumohra chaerophylloides (Poir.) Ching (1)

Rumohra lurida (Jenman ex Underw. & Maxon) Ching (3)

Rumohra pubescens (L.) Ching (6)

Thelypteris chaerophylloides (Poir.) Proctor (1)

Thelypteris lurida (Jenman ex Underw. & Maxon) Proctor (3)

Thelypteris lurida f. leucochaete (Slosson) Proctor (2)

Thelypteris pubescens (L.) Proctor (6)

Appendix III

Index to collectors’ and numbers. The number in parentheses refers to the species number assigned in the taxonomic treatment (see Appendix I, above). Boldface indicates types.

Abbott, W. L.: 1196a, 1227a, 2018, 2512, 2770 (1)

Acevedo-Rodríguez, P.: 13390 (1)

Acevedo-Rodríguez, P. et al.: 7732, 13454 (1)

Ackerman, J. D.: 2109 (1)

Ackerman, J. D. & Montalvo, A. M.: 1438 (1)

Ackerman, J. D. 3933 (1)

Alain, H. & Liogier, A. H.: 15891, 19566, 31726 (1)

Alain, H. et al.: 28389, 28965, 30984, 31460 (1)

Allard, H. A.: 14728 (1)

Areces, F.: 611 (1)

Areces, F. & Khelani, K.: 115 (1)

Arsène, G.: s.n. (1)

Atwood, J. T.: 3908 (6)

Axelrod, F. & Axelrod, B.: 1908, 2318, 4837, 6705, 8055 (1)

Axelrod, F. & Christenhusz, M. J. M.: 12822 (1)

Axelrod, F. & Ackerman, J. D.: 470 (1)

Axelrod, F. & Pascarella, J. B.: 9903 (1)

Axelrod, F. & Ward, S.: 12987 (1)

Axelrod, F. & Wells, J.: 2182 (1)

Axelrod, F. et al.: 663, 1735, 8895, 9794, 14737 (1)

Balch, A.: s.n. (6)

Barker, E. E.: s.n. (1)

Bartlett, H. H.: 17534 (5)

Belanger, M.: 379 (4)

Bello, D. D.: 37 (1)

Benzon, A.: 613 (1)

Blauner, B. F.: 267 (1)

Blomquist, H. L.: 11854 (1)

Boom, B. M.: 7019 (1)

Boom, B. M. et al.: 7019 (1)

Box, H. E.: 512 (4)

Britton, N. L.: 3912 (1)

Britton, N. L. & Britton, E. G.: 5078 (5)

Britton, N. L. & Cowell, J. F.: 4133, 4190 (1)

Britton, E. G. & Marble, D. W.: 638, 1082 (1)

Britton, N. L. & Percy, W.: 5295 (6)

Britton, N. L. & Shafer, J. A.: 2089, 2149 (1)

Britton, N. L. et al.: 5545, 5618, 6280, 8341, 8427 (1)

Caluff, M. G.: 1256 (6)

Cedeño, J. A.: 526 (1)

Christenhusz, M. J. M. et al.: 2675 (4); 3528, 4223(1)

Clément, F.: 775 pro parte (1), 775 pro parte (7); 880 (6); 882 (7); 990 (6); 1052 (8); 1084 (6); 1092 (1); 1292 (8); 1499, 1539, 1575 (7)

D'Arcy, W. G.: 2079 (1)

Day, E. H.: s.n. (6)

Duss, P.: 1582, 1683, 4052, s.n. (4)

Eggers, E.: 5109, s.n. (1)

Ekman, E. L.: 75 (5); 3217 (1); 3801, 5460, 6838 (6); 6931 (1); 6940 (7); 8593 (1); 10415 (5); 11090, 12938a, 14953 (1)

Engelmann, C.: s.n. (4)

Eyerdam, W. J.: 450 (1)

Gonzales, A.: 470 (6)

Graveson, R.: 2049, 2050 (4)

Hahn, L.: 915 (4)

Harris, W.: 7438 (1); 8710 (2); 8770 (3, see discussion); 11023 (2).

Heller, A. A.: 883, 4557 (1)

Hespenheide, H. A. et al.: 889b (3)

Hess, W. E.: 7014 (1)

Hieram, B.: 4682 (1)

Hioram, J.: 909 (1)

Hioram, L. F.: 119 [Rosenstock Filices Portoricensis no. 20] (1)

Hodge, W. H. & Howard, R. A.: 4540 (6); 4445 (8)

Holdridge, L. R.: 692 (1); 2031 (6)

Howard, R. A. & Proctor, G. R.: 15053, 15165 (3)

Jack, J. G.: 7098, 7114 (6)

Jenman, G.: s.n. (3); s.n. (6); s.n. (8)

Johnston, J. R.: 195 (4); 231 (1)

L'Herminier, M.: 31, 121 (4)

Laubengayer, A.: s.n. (1)

León, B.: 10393 (7)

León, B. & Clément, F.: 4051 (1)

León, B. & Roca, ?: 7859 (6)

Leonard, E. C.: 4435, 4710, 7918 (6)

Leonard, E. C. & Leonard, G.: 14151 (6)

Linden, J.: 1875 (6)

Liogier, A. H.: 37103, 37132 (1); 14682 (6)

Liogier, A. H. & Oquendo, J.: 72 (1)

Liogier, A. H. & Liogier, P.: 26223, 26234 (6)

Maxon, W. R. & Killip, E. P.: 455 (3); 1352 (6)

Maxon, W. R.: 1465 (6); 1957, 2260, 2311 (3); 2860 (2); 8530, 8531, 8543, 9088, 9103, 9154, 10318 (6); 10447, 10466 (3)

Mejía, M. et al.: 796 (1)

Mickel, J. T. et al.: 8647, 9027 (6)

Miller, J. S. & Sherman, C. D.: 6413 (1)

Moore, A.: 118, s.n. (1)

Morton, C. V.: 4157 (6); 9537 (7); 10390, 10829 (6)

Morton, C. V. & Acuña, J.: 3451 (1)

Orcutt, C. R.: 3794, 5913 (6)

Otero, J. I.: 343 (1)

Parkhust, V. P.: 36 (6)

Percy, W.: 188, 246 (1)

Perkins, A. E.: s.n. (1)

Philipson, W. R.: 973 (6)

Pollard, C. L. & Palmer, W.: 148 (1)

Portugal, A. & Collignon, G.: 46 (1)

Portugal, A. & Fernández, H.: 43, 45 (1)

Proctor, G. R.: 3186, 4070 (6); 20384, 21652 (4); 28263 (6); 28365, 28643 (1); 35654 (3); 39337 (1)

Proctor, G. R. & Pinto, B.: 40497 (1)

Read, R. W.: 1929 (2)

Ryan, J.: 25 (4)

Saldaña, J.: 1316 (1)

Sánchez, C. & Liogier, A. H.: 80, 218 (1)

Sargent, F. H.: 187, 3151 (1)

Scamman, E.: 8118 (1)

Schwannecke, ?: s.n. (1)

Shafer, J. A.: 3170, 3224, 8507 (1); 8600 (8); 8807 (1)

Shaw, S. W. et al.: 484 (1)

Sintenis, P.: 435, 437, 2013, 2112, 6537 (1)

Stehlé, H.: 3282, 4446, 5200 (4)

Stevens, G. L.: 5978 (1)

Stimson, W. R. & Yantin, P.: 1630 (1)

Sundue, M. et al.: 2051, 2093 (1)

Taylor, C. M.: 6730, 7450 (1)

Torres, L. K. et al.: 18 (1)

Trejo, J. C. et al.: 2937 (1)

Underwood, L. M.: 1187, 1593 (6); 1825 (3); 2018, 2073, 2479, 2685, 2714 (6)

Valeur, E. J.: 343 (6); 1016 (1)

Vincent, M. A. et al.: 15366 (1)

Wagner, W. H.: s.n. (1)

Walker, T. T.: T2225 (3)

Watt, D. M.: 80 (1); 139 (6); 180, 435 (1)

Webster, R. M. & Goldberg, ?: 236 (6)

Williams, M.: 4 (1)

Wood, C. E. & Atchison, E.: 7477 (8)

Woodbury, R.: s.n (1)

Wright, C.: 815 (6); 851, 1000 (1); 1054 (7)

Yuncker, T. G.: 18234 (3)

Zanoni, T. & Jiménez, F.: 44459 (1)

Zanoni, T. et al.: 12856, 18968x, 21224, 24984a (1); 29688 (6); 35438, 38144 (1)

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Prado, J., Moran, R.C. Monograph of the West Indian fern genus Polystichopsis (Dryopteridaceae). Brittonia 68, 1–24 (2016). https://doi.org/10.1007/s12228-015-9387-z

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