Abstract
The Elaphoglossum ciliatum group was supported by a previous study as monophyletic on the basis of cpDNA sequence data. The present paper provides a monograph of this group. It includes keys, full synonymy, descriptions, specimens examined, index to collectors’ names and numbers, photographs, and distribution maps for all species. Morphologically, these species are characterized by abaxial laminar scales reduced to minute dots, resinous rhizomes, and (except for E. dussii) spiny perispores. These characteristics are not unique to the E. ciliatum group because they are found in other species of Elapholgossum sect. Lepidoglossa. Nine species are here recognized in the E. ciliatum group, with no subspecies or varieties. Six species are neotropical and primarily montane, being largely absent from Amazonia. The remaining three species are endemic to the island of St. Helena in the South Atlantic Ocean. A lectoype is designated for E. calaguala.
Resumen
Se presenta la monografía del grupo de Elaphoglossum ciliatum, un grupo definido como monofilético basado en análisis de secuencias de ADN del cloroplasto. La monografía incluye claves, sinonimia completa, descripciones, especimenes examinados, un índice de colectores y sus números de colección, fotografías y mapas de distribución de todas las especies del grupo. Las especies de este grupo se caracterizan por tener las escamas de la superficie abaxial de la lámina reducidas a puntos muy pequeños, rizomas resinosos y (exceptuando E. dussii) perisporas equinadas. Sin embargo estas características no son únicas al grupo de E. ciliatum, y pueden encontrarse en otras especies de Elapholgossum sect. Lepidoglossa. Se reconocen nueve especies en el grupo de E. ciliatum, sin subespecies ni variedades. Seis especies son neotropicales y principalmente de regiones de montaña. El resto de las especies son endémicas de la isla de St. Helena en el sur del océano Atlántico. Se designa un lectotipo para E. calaguala.
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Acknowledgments
This research was partly funded by a grant from the National Science Foundation to Moran (DEB 0211969). Financial support for the first author was also provided by the Oliver Hazard Perry Fellowship from the New York Botanical Garden. We thank Judith Garrison Hanks for helping take some of the SEM images of the spores, John T. Mickel for discussion about Elaphoglossum and comments on the manuscript, and Josmaily Loriga for sending pictures of Cuban specimens. We are grateful to Lawrence Kelly, Paulo Labiak, and Michael Sundue for useful comments that greatly improved the manuscript. We also thank the personnel of the following herbaria for allowing us to examine material and for loans: AAU, B, BR, BM, CAY, COL, CR, F, GH, HUA, INB, JBSD, K, MICH, MO, P, QCA, QCNE, S, SEL, STR, UC, US, USM, USZ, VEN. This paper is part of the first author’s Ph.D. thesis at the City University of New York; she wants to dedicate it to her family (mother, sister and brother) for their continuous support and love while abroad.
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Appendices
Appendix 1. List of Taxa
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1.
Elaphoglossum bifurcatum (Jacq.) Mickel
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2.
E. burchellii (Baker) C. Chr.
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3.
E. ciliatum (C. Presl) T. Moore
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4.
E. dimorphum (Hook. & Grev.) T. Moore
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5.
E. dussii Underw. ex Maxon
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6.
E. gramineum (Jenm.) Urb.
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7.
E. huacsaro (Ruíz) H. Christ
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8.
E. nervosum (Bory) H. Christ
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9.
E. nigrescens (Hook.) T. Moore ex Diels
Appendix 2. List of Exsiccatae
The number in parentheses refers to the species number assigned in the monograph; the numbers in boldface are types.
Acebey, 176 (2)
Acevedo, 2976 (5)
Acosta-Arteaga, 73 (7); 165, 324, 359 (2); 493, 779 (7); 800 (2); 1071, 1082 (7); 1205 (2)
Acuña, 20323 (6)
Agostini, 2835 (2); 2863 (7)
Aguinda, 999 (3)
Allart, 374 (2)
Albert de Escobar, 5573, 7400 (7)
Alfaro, 1117 (3)
Alston, 6517, 7419 (7)
Amorim, 5812 (9)
Anderson, 3223 (6); 7523, 8864 (2)
André, 1961 (3)
Angulo, 71 (9); 602 (3)
Arbeláez, 84, 1244 (7)
Archer, 1278 (2)
Arroyo, 2743 (2)
Asplund, 16945, 17135, 17412 (7)
Atehortúa, 1004, 1014 (7)
Atwood, AN196 (9)
Aulestia, 1722, 1957, 2000 (9)
Azofeifa, 108, 109 (9)
Baker, 5537 (3)
Balansa, 390 (2)
Balslev, 1725, 2474, 2476A (7); 10327 (3); 10336 (2); 10402 (3); 84690, 97427 (9)
Barclay, 4459 (7)
Barringer, 2284 (9)
Bass, 172 (3)
Bayon, 98 (2)
Beck, 465 (9)
Beitel, 85268 (9)
Bell, 666 (7)
Bello, 978 (9); 2712, 4501 (3); 4578 (9)
Bischler, 1233, 1318, 2258 (7)
Bittner, 206 (9); 686, 709 (3); 1014, 1227 (9); 1722 (2); 2024, 2354 (3)
Boyle, 4915, 4958 (3)
Brade, 80, 81 (7); 236 (9); 237(7); 528 (3); 7605, 8415 (9); 9759 (7); 9764 (2)
Brandbyge, 30808 (9); 42517 (7)
Breedlove, 22412, 23167 (7)
Brenes, 19274, 20531 (3)
Bridarolli, 4338, 4402 (2)
Bristol, 456 (3)
Britton, 690 (5)
Broadway, 1900 (5)
Buchtien, 8 (9); 2712 (7); 5169 (2); 5175 (9); 5927 (7)
Bües, 2100 (7)
Burch, 4619 (3)
Burchell, 177 (1); 180 (4); 5155 (2)
Burger, 5018 (3); 5097 (9); 5808 (9)
Callejas, 3834, 3891 (7); 10504 (3); 10518 (7)
Camp, 4661 (7)
Campos, 3677 (2)
Carvajal, 453 (9)
Carvalho, 112 (2)
Chavarría, 24 (2); 485 (9)
Chockerong, 116 (6)
Christenhusz & Katzer, 3221 (7); 4011 (5)
Churchill, 4933, 4935 (9)
Clark, 163, 578, 4875, 6441 (3); 8079 (7); 9059 (3)
Clarke, 5453 (9)
Clute, 82 (7)
Colella, 1366 (7); 1601, 1602 (9)
Core, 451 (7)
Cornman, 1354 (3)
Correa, 18A134 (7)
Correll, E419 (7)
Cotton, 905, 929 (9)
Croat, 821, 824 (7); 27272 (9); 35768 (3); 36458 (9); 41016 (7); 49976 (3); 51523 (7); 59961, 70770 (3); 72196 (7); 88085 (3); 92143, 93854 (7)
Crosby, 2564 (3)
Cuatrecasas, 11469 (7); 13938 (3); 26800 (7)
Cumming, 420, 2464 (1)
Cuninghame, 135 (1)
Davidse, 21137 (2); 23226 (3); 23424 (9)
Dawson, 14507, 14574 (2)
de Garganta, 2047 (7)
de la Sota, 5209 (3); 6206 (7)
de Nevers, 5401 (3)
Díaz C., 2171, 3605 (7)
Díaz S., 1666, 3015 (7)
Dodson, 10670, 10674, 10752, 10756, 10757 (7); 10761 (2); 10774, 10802 (3); 10895, 12088 (7); 14673 (3); 15417 (7)
Dorr, 7292, 7994, 8132, 8214 (7); 8623 (2); 9256 (7)
Duncan, 2607 (7)
Dusén, 735, 2045, 2748, 4045, 7225, 7280 (2)
Duss, 1622, 1623, 4130, 4130, 4131, 4150, 4400, 4607, 4688, 4748 (5)
Dwyer, 4493 (9)
Eberhardt, 166, 300, 421, 423, 442 (2)
Ekman, 13878 (7)
Engler, 62 (2)
Evans, 221 (2)
Ewan, 16589 (7)
Fay, 1580, 1605, 1606 (7); 2193, 2487 (2); 2651A (3); 4273, 4308 (7)
Fendler, 276, 280 (2)
Fernández, 27, 223A (7)
Feuerer, 8750A (2)
Folsom, 3298, 3980 (3)
Fonnegra, 2773 (7); 5614 (2)
Forzza, 1219 (2)
Fosberg, 43355 (9)
Foster, 1348, 1513 (7); 10792 (3)
Franco, 5098 (3)
Garcia-Barriga, 12604 (2)
Gehriger, 327 (7)
Gentry, 19319 (7)
Gil, 9 (3)
Giraldo, 1581 (3)
Glaziou, 952, 2433, 2455, 2800, 2855 (2)
Gómez, 247, 2171 (3); 20831 (9); 20845 (9); 22794 (3)
González, 441 (9)
Grant, 9086 (7); 90–00885 (2)
Grayum, 1958, 2426, 2819, 2929 (9); 6738 (3); 7955, 9079 (9)
Grubb, 1314 (3)
Gutiérrez, 20 (3)
Gutiérrez-H, 246 (7)
Hagemann, 81 (2); 369, 641 (3); 1153, 1192 (7); 1261 (3);1865 (3); 2025 (9)
Hahn, 498 (9)
Hamilton, 901, 908 (7)
Hammel, 2003, 2092 (3)
Handro, 1078 (9)
Harley, 50623 (2)
Harling, 809, 4166 (7)
Harris, 7429 (6)
Hassler, 4612 (2)
Hatschbach, 9801 (2); 25799 (9); 31136, 31460 (2)
Haught, 6133 (2); 3163 (7)
Haughton, 1149 (1)
Henkel, 1287 (9)
Henrich, 211 (9)
Herrera, 1377 (9); 1855, 2532 (3); 3221 (9); 3233a (3); 5108 (9); 5239, 5298, 6202, 7857 (3)
Herzog, 2227 (7)
Hinchley, 179 (6)
Hno. Daniel, 1021, 1215 (7); 3277 (2)
Hodge, 16 (5)
Holm-Nielsen, 1366 (7); 3264 (7); 3517 (2); 16171, 18302, 23572, 29352 (7)
Hoover, 1958, 2060 (3)
Huaylla, 1714 (2)
Hutchison, 4818 (7)
Idrobo, 4146 (7); 10298 (3)
Ingram, 1308, 1448, 1825 (9)
Irwin, 15629, 29118, 32893B (2)
Ivey, 292, 327 (3)
James, 13, 109 (9)
Jaramillo, 5778, 7142, 7145 (7); 12234, 14163, 14319 (3)
Jaramillo-Mejía, 2984 (7)
Jenman, 27, 179 (6)
Jimenez, 1052 (2); 2185, 2220, 2256 (9)
Jorgensen, 61898 (7)
Judd, 4517 (7)
Juncosa, 1018 (7)
Jürgens, 61.1, 61a, 144, 144.1 (2)
Kennedy, 3167 (3)
Kessler, 6239, 6264 (2); 6698, 6993 (7); 7395, 7525, 7536B, 7842 (9); 9092, 9147 (2); 459 (7); 9773 (9); 10506 (2); 10507 (7); 10528 (2); 11325 (9); 11515, 11543 (2); 11784 (7)
Killip, 8850, 15798 (7); 16555 (2); 16732, 17190, 19304, 24411, 24414 (7); 25668 (3); 34641 (7); 34648 (3); 38503 (2)
King, 452 (9)
Kirkbride, 1991, 4268 (3)
Knapp, 2488 (3); 7550 (2)
Kress, 91–3359 (9); 94–5110 (3)
Lægaard, 51207 (9)
Langenheim, 3390 (7)
Lawesson, 43444 (9)
Labiak, 2942 (2); 3638 (9); 4236 (2)
Leal, 60 (7)
Lehmann, BT1190 (2)
Lehnert, 1338 (2); 1304, 1423 (3)
Leite, 411, 2758, 3480 (2)
Lellinger, 290 (3); 570 (9); 825, 1276 (3); 1834 (9)
León, 618, 955 (7)
Leonard, 4513 (7)
Lewis, 2382 (7)
Licata, 422 (7)
Liesner, 6407, 6645, 7499 (9); 12995 (7); 14862 (3); 17646, 18597, 23561, 23598, 25826 (9)
Linden, 1922b (6)
Liogier, 13422 (2)
Little, 7826, 7891, 8171, 8568, 8582a (7)
Lloyd, 199A, 336, 351, 357, 753 (5); 4182, 4189, 4250, 4256 (3)
Løjtnant, 11927 (7)
López, 7239 (9)
Luederwaldt, 7844 (2)
Luer, 4941 (2)
Luteyn, 5367, 5453 (7); 6628 (2); 7049, 7255, 7572 (7); 8238 (2); 9731, 9831, 9935, 10620, 10995, 10997 (7); 13161 (3); 13167 (7); 13170 (3)14191 (7)
Maas, 5298 (7); 6858 (9)
Macedo, 3071 (2)
Madison, 2260, 2261, 2550 (7)
Madsen, 7111 (2)
Mägdefrau, 1450 (7)
Maguire, 28489, 33543, 34927 (9)
Martínez, 406 (3); 420 (9)
Marulanda, 511 (7)
Matos, 371 (9)
Maxon, 2274, 2847, 2918, 8611 (6)
McPherson, 8869 (3); 8687, 9655, 15387 (9)
Mejía, 3776 (3)
Melo, 139 (2)
Méndez, 924 (7)
Mexia, 7102, 7188 (3)
Mickel, 2577 (3); 3495 (9); 8258, 8268, 8275, 8392, 8406, 8411, 8449 (7)
Miranda, 170 (9)
Montgomery, 8542 (3)
Mora, 928 (9)
Morales, 1256 (9)
Moran, 4118, 5164 (3); 5179 (2); 5183, 5377 (3); 5836 (9); 5925 (2); 6121, 6243, 6332 (9); 6371 (7); 6711, 6762 (3); 6895 (7); 7418 (3); 7491 (9); 7516 (3) 7627 (9)
Moritz, 122 (2); 315 (7); 316 (2)
Mostacero, 344 (7)
Murillo 132, 737,927 (7); 973 (3); 2112, 2954 (7)
Mutis, 3317 (7)
Navarrete, 1540, 1808 (3)
Nee, 49325 (2); 52309 (7)
Neill, N41, 3249, 3253 (9); 9648 (3)
Nuñez, 711 (2)
Øllgaard, 986 (3); 1293 (7); 1569, 1637 (3); 1891, 2384 (9); 2456 (3); 2476 (9); 2639, 2725, 2892, 3033 (3); 8734, 9361, 9378 (7); 99337 (2); 99968 (7); 99985 (3)
Ortega, 1951, 2260 (7)
Pachón, 60 (7)
Palacio, 66 (7)
Palacios, 22, 4134, 5400, 5697 (3); 6479, 8739 (9); 11414 (3)
Peñuela, 60 (7)
Pipoly, 5128, 10915 (9)
Pittier, 3554 (3); 7404 (2)
Plowman, 940, 4513 (3)
Porter, 4665, 4669, 4673 (3); 4797 (9); 4939b, 4941a, 4942 (3)
Porto, 2113, 3014 (2)
Posada, 282 (2)
Prado, 1099 (7); 1109 (2)
Proctor, 3907 (6); 32242 (9)
Ramírez, 4804 (7); 5349 (2)
Rangel, 5583 (3)
Ranker, 1823 (9)
Regnell, 336 (2)
Reitz, 110, 3972 (2); 5714 (9)
Rieckl, 1823 (9)
Roa, 30 (7)
Rodríguez, 3250 (3); 3272 (2); 3464, 5064 (3); 5226, 5445 (7); 5620 (3)
Rohr, 292, 1059 (2)
Rojas, 406 (3); 591 (9); 616, 1022 (3); 1025 (9); 1040 (3); 1054, 1295 (9); 1629, 1777 (3); 1802, 1827, 2096 (9); 2107, 2218, 2265, 2345, 2441, 2606 (3); 2716 (7); 2795, 2966 (3); 3029 (9); 3278 (2); 3361 (7); 3367 (9) 3999, 4197 (3); 4230, 4545 (9); 4627 (3); 4663 (7); 4824, 4856 (3); 4871 (9); 5105 (2); 5458 (3); 5492 (7); 6050 (2); 6051, 6310 (3)
Romero, 8 (7)
Romoleroux, 2070 (9)
Rueda, 6207, 10552, 10692, 17043 (9)
Ruiz, 54 (7)
Ruiz-Terán, 6124 (7)
Rusby, 298 (2)
Saldarriaga, 27 (7)
Salino, 1780, 1810, 4635, 5196 (2); 5399 (9); 5527 (2); 6480 (9); 6871, 7454, 9260 (2)
Sánchez, 137, 138 (5); 1665, 6001 (7)
Sandeman, 5841 (9)
Santa, 1078 (7)
Saravia, 3019A (7)
Savatier, 1993 (1)
Schultes, 15285, 15739 (9)
Schunke, 136, A145 (9)
Seidenschnur, 127, 128 (3); 137 (9); 155, 164, 168 (3); 173 (9); 175, 191a (3); 194 (9); 202 (3); 261a (7); 280 (3); 281 (9); 381 (9)
Shafer, 592, 3433, 3482, 3669 (5)
Skog, 5042 (9)
Smith, 1707, 1855 (9); 2103 (3); 5133 (9) 5525 (3)
Sodiro, 22 (7)
Soejarto, 3031 (7)
Solomon, 11748, 14956 (2); 4903 (7)
Spannagel, 10 (2)
Sparre, 16526 (2); 16973, 17389 (7)
Spruce, 5230 (3); 2187 (9)
Standley, 36417, 36494, 37767, 37847, 37963 (3)
Stergios, 4253 (9); 17340, 17341, 17474 (7); 20609 (2);
Steyermark, 144 (9); 56989 (2); 110109 (7); 129728, 130312A (9)
Stimson, 5400 (2)
Stones, 2041 (7)
Stork, 9892 (7)
Tate, 940 (2)
Thompson, 7872 (7)
Torres, 19 (7); 96, 2277 (3)
Torres-Rojas, 181 (9)
Tryon, 5309, 6020 (7)
Tuomisto, 7749 (9)
Ule, 265 (7)
Underwood, 479 (7); 1337, 1388, 1619, 2057, 2120, 2144 (6); 2589 (7)
Valeur, 634 (6)
van Cotthem, 1523, 1588 (2); 1708, 1999 (7)
van der Werff, 5939, 5943 (7); 6649, 6877, 7064 (3); 7670 (7); 8745 (2); 9544 (9); 13428, 15821 (3); 16256 (9); 17612 (3)
Vareschi, 5556, 7167 (7)
Vargas, 2365 (3); 2501 (7)
Vasco, 468, 504 (3); 555, 556 (2); 562, 567, 568, 576 (7); 579 (3)
Vásquez, 28869 (7)
Velásquez, 99 (3)
Vélez, 1306 (3)
Vivas, 1452 (9)
Wacket, 114 (9)
Webster, 13417 (5)
Wercklé, 17102 (9)
Werling, 248 (7)
Werner, 278 (3); 1547 (2)
White, 508 (2)
Wiggins, 10991 (7)
Williams, 15338 (9); 19943, 20323 (3)
Woytkowski, 6740, 6741 (7); 7693 (2)
Zanoni, 19347, 20278 (7)
Zardini, 40813 (2)
Zarucchi, 6212 (7)
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Vasco, A., Moran, R.C. & Rouhan, G. Monograph of the Elaphoglossum ciliatum group (Dryopteridaceae). Brittonia 61, 241–272 (2009). https://doi.org/10.1007/s12228-009-9075-y
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DOI: https://doi.org/10.1007/s12228-009-9075-y