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A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae)

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Abstract

This paper describes Ascogrammitis, a new genus of grammitid ferns formerly combined under Terpsichore. Seventeen species are recognized, including five new ones: A. clavigera, A. colombiensis, A. loxensis, A. oxapampensis, and A. tungurahuae. New combinations are made for A. alan-smithii, A. anfractuosa, A. angustipes, A. athyrioides, A. clathrata, A. cuencana, A. david-smithii, A. dilatata, A. nana, A. pichinchae, A. pichinchensis, A. stuebelii, and a lectotype is chosen for Polypodium induens. A key is provided to distinguish the species, and illustrations are provided for 12 species. The genus occurs primarily in neotropical cloud forests, with the greatest diversity in the Andes. Ascogrammitis is one of two monophyletic groups of grammitid ferns that are consistently associated with Acrospermum, an epibiotic bitunicate ascomycete that produces black clavate fruiting bodies.

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Acknowledgments

I thank the curators of the following herbaria for making their material available for study: AAU, B, F, GH, CR, GOET, HUA, INB, K, LPB, MICH, MO, NY, OS, P, RB, QCA, QCNE, UC, UPCB, US, and USZ. Support for fieldwork was provided by The Graduate Center at the City University of New York, The Organization for Tropical Studies, and The Torrey Botanical Society. Ya-yi Huang and Hannah Stevens assisted with dot maps. Roy Gereau assisted with the Latin diagnoses. Alejandra Vasco helped edit Spanish locality data. Illustrations were provided by Haruto Fukuda. An early draft of this manuscript was greatly improved by comments from Larry Kelly, Fabian Michelangeli, and Robbin Moran. I also thank João Condack, Alejandro Marin, Hilary Martin, Fernando Matos, Joel Nitta, Wilson Rodríguez, Alexander Rojas, Eric Scheuttpelz, Pete Shear, Alejandra Vasco, and Florian Werner for collaborating in field work. This work was done in partial fulfillment of the requirements for a doctoral degree at the City University of New York.

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Authors and Affiliations

  1. The New York Botanical Garden, 200th St. and Southern Blvd., Bronx, NY, 10458-5126, USA

    Michael A. Sundue

  2. The Graduate Center, City University of New York, 365 5th Ave., New York, NY, 10006, USA

    Michael A. Sundue

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  1. Michael A. Sundue
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Correspondence to Michael A. Sundue.

Appendices

Appendix I

Index to collectors’ names and numbers. Numbers in parentheses refer to the species number below.

List of Species

  • 1. Ascogrammitis alan-smithii (A. Rojas) Sundue

  • 2. A. anfractuosa (Kunze ex Klotzsch) Sundue

  • 3. A. angustipes (Copel.) Sundue

  • 4. A. athyrioides (Hook.) Sundue

  • 5. A. clathrata (Sundue & M. Kessler) Sundue

  • 6. A. clavigera A. R. Sm. ex Sundue

  • 7. A. colombiensis Sundue

  • 8. A. cuencana (Hieron.) Sundue

  • 9. A. david-smithii (Stolze) Sundue

  • 10. A. dilatata (Sundue & M. Kessler) Sundue

  • 11. A. loxensis Sundue

  • 12. A. nana (Sundue & M. Kessler) Sundue

  • 13. A. oxapampensis Sundue

  • 14. A. pichinchae (Sodiro) Sundue

  • 15. A. pichinchensis (Hieron.) Sundue

  • 16. A. stuebelii (Hieron.) Sundue (Add species, change numbering, fix in list of collections)

  • 17. A. tungurahuae Sundue

List of collectors names and numbers

The number in parentheses refers to the species numbers assigned in the taxonomic treatment (List of Taxa, above). Boldface indicates types.

Abbott, 17066 (12)

Acuña, 9991 (2)

Alston, 7442, 7476 (7)

Alverson & Flores, 3101 (1)

Andre, K129 (14)

Antonio, 2578 (1)

Arroyo et al., 62 pp (9); 2741 (9); 2762 (12); 2763 (10)

Asplund, 16106 (15)

Atehortúa, 1124 (2)

Aymard & Luteyn, 2459 (2)

Bach et al., 1051 (10); 1349 (2)

Balslev, 1848 (15); 1934 (8); 2700 (15)

Balslev & Steere, 3267 (15)

Balslev et al., 1629 (15)

Barclay & Juajibioy, 6301 (14)

Barringer & Christenson, 3419 (2)

Barringer et al., 2277 (2)

Beck, 2844 (4); 7600 (10); 14474 (9); 21496 (4); 1424a (10)

Bell, 374 (2)

Bello & Mendez, 2663 (1)

Bishop, 826 (2); 857 (2); 1952 (14); 1953 (14); 1955 (14); 1956 (14); 1989 (2); 2506 (9)

Boom, 1432 (11)

Boyle et al., 4389 (4)

Breedlove, 34598 (2)

Breedlove & Smith, 32644 (2)

Breedlove & Thorne, 30159 (2)

Britton, sn, sn (2)

Brooke, 6166 (4)

Bryan, 383 (12); 550b (4)

Buchtien, 891 (4); 2754 (4); 2756 (4)

Bues, 1039 (4); 1259 (4); 1391 (4); 1547 (4); 1822 (4); 1945 (2); 2116 (4)

Caballero, 126 (2)

Camp, 4075 (15); E1394 (11)

Campos & Tenorio, 5047a (2)

Cardenas, 1002 (4)

Carhart, sn (2)

Cazalet & Pennington, 5378 (14)

Ceron, 18127 (15)

Christenhusz & Katzer, 4158 (2)

Cook & Gilbert, 1168 (2)

Cornman, 1042, 1241 (2); 1326 (2)

Croat, 23166, 33054, 33898 (1); 48567 (2), 54917 (6), 58706 (11)

Croat et al., 87817a (2)

Cuatrecasas, 22141 (2)

Dassler, 94-7-13-1 (7)

Davidse, 25257 (2)

Davidse & Herrera, 29122 (2)

Davidse et al., 28598 (2)

Davidson, 195 (2)

De Boer, 1161 (10)

Dodge & Thomas, 7889 (2)

Dodson & Embree, 13394 (2)

Dodson et al., 10929 (2)

Dorr et al., 4984 (2); 5010, 8154 (6)

Ekman, 7373, 13585 (2)

Espinoza, 322 (1)

Evans, 2613 (2)

Farr, s.n. (2)

Fay & Fay, 2449 (10); 2639, 2644 (14); 2961, 3033 (10); 3052 (2); 3105, 3113 (15); 3163, 3193, 3227, 3284 (2); 3285 (15); 3286, 3586, 3897 (2); 4286, 4339 (14); 4446 (11)

Fernández, 632 (4)

Fernández et al., 2202 (10); 2511, 4250 (9)

Fisher, 151 (2)

Fosberg, 20861 (14)

Foster, 85-153, 85-2 (2)

Fuentes et al., 11943 (4)

Fuertes, sn (2)

García & Beck, 4079 (4)

García et al., 4466 (2)

Gavilanes & Funk, 837 (14)

Gómez, 2400, 21778, 22228 (2)

Gonzales, 1541, 1552, 1576 (4)

Grant, 9062 (7)

Grayum, 10303 (2)

Grayum & Yatskievych, 6623 (2)

Grubb et al., 1369 (11)

Harris, 7128, s.n. (2)

Hart, s.n. (2)

Haught, 5288 (2)

Hennipman, 8138 (9)

Henri-Stanislas, 1683 (2)

Herrera, 5564 (1)

Herzog, 2190 (4)

Heyde & Lux, 3254 (2)

Hill et al., 17854 (2)

Hodge, 6102, 6156, 6951, 6961 (2)

Holm-Nielsen, 18144 (2)

Holm-Nielsen & Jeppesen, 1376 (8)

Holm-Nielsen et al., 5656 (15); 6493, 17405 (14); 26638 (11); 41786 (14)

Horwatch, 150 (2)

Howard, 39 (2)

Hutchinson & Bennett, 4754 (2)

Hutchinson & Writght, 6988 (4)

Ibisch, 930102 (9)

Ingela, 95-M046 (2)

Jameson, 137, s.n., s.n. (15)

Jaramillo, 3869 (15)

Jaramillo & Cachago, 4664 (15)

Jenman, 84, s.n. (2)

Jiménez M., A., 1341 (2)

Jiménez, I., 337 (9); 528 (15); 531, 543 (4); 554 (12); 894 (9); 954 (10); 954 (15); 1106 (12); 1156 (10); 1156 (15); 1215,1246, 1352 (9); 1396, 1397, 1399, 1443 (2); 1452 (9); 1689 (10); 1692 (4); 1709 (9); 1753, 1774 (4)

Jiménez, I. & Gallegos, 912C (10)

Jiménez, I. & Moguel, 1609 (10)

Jiménez, I. & Quisbert, 2 (2)

Jiménez, I. & Vidaurre, 528 (10)

Jiménez, I. et al., 398 (2)

Johnson, 533 (2)

Jones & Norris, 1113 (2)

Jørgensen & Madsen, 65795 (14)

Juncosa, 1987 (3)

Kessler, 2926 (4); 6567 (10); 6621 pp (6); Kessler, 6621 pp (10); Kessler, 6682 (15); 6723, 6725 (4); 6801 (15); 7029, 7094, 7146 (2); 7231 (4); 9485 (9); 10622, 11714 (4); 11833, 12040 (10); 12128, 12199 (2); 12394 (12); 13507, 13517 (2); 6567, 6621, 6682, 6801 (10); 9049 (9); 9059 (9); 9193 (10); 9401, 9511 (9); 11813, 11872 (10)

Killip, 5194, 5841 (2)

Killip & Garcia, 33843 (3)

Killip & Hazen, 9488 (14)

King & Bishop, 7695, 7695a (10)

Knapp & Alcorn, 7776 (2)

Knapp et al., 2008 (2)

Knight, 2 (2)

Kral, 71973 (2)

Krömer & Aceby, 1237 (5)

Kuntze, s.n. (12)

Labiak et al., 2856, 2864, 2867, 2871 (10)

Lankester, 902 (2)

Lehmann, 3010 (2); 5728 (8); 7365 (2)

Lehnert, 212 (2); 261 (4); 291, 312, 315, 329 (2); 380, 381 (10); 499 (2); 697 (9); 702 (10); 708 (10); 1035 (2); 1233, 1235 (11); 1577 (15)

Lehnert & Kessler, 1164 (8); 1206 (2)

Lellinger, 786, 1735 (2)

Lellinger & White III, 997 (2); 1311 (1); 1582, 1694 (2)

Lems, s.n. (2)

Lent, 1760 (1)

León, 1898 (4)

León & Young, 4487 (9)

Lewis, 88339 (4)

Liesner, 5082 (1)

Lioger, 12807, 16069 (2)

Lioger & Lioger, 22356 (2)

Lojtnant & Molau, 13133 (14)

Lopez, 188 (2)

Macbride, 4305, 4329 (4)

Macbride & Featherstone, 1862 (4)

Madsen, 85650 (11); 86751 (14)

Marin et al., 132 (2)

Mathews, 1103 (4)

Maxon, 278, 1214, 1324, 1332, 2770, 5291, 5486, 5714, 8232, 10221, 10240, 10567 (2)

Maxon & Harvey, 8142, 8434, 8448, 8487 (2)

Maxon & Killip, 711, 1302, 1077a (2)

Meier & Borjas, 7229 (2)

Mellado & Beccera, 2015, 2066 (13)

Merril et al., 15678 (2)

Mickel, 2165, 2195, 2352, 2664, 5562 (2)

Mickel & Hellwig, 4292 (2)

Mickel & Purdue, 6771 (2)

Mickel et al., 8267, 8270, 8292, 8754, 9394 (2)

Mille, s.n. (14); s.n. (15)

Molina, 10673 (2)

Molina & Molina, 21463 (2)

Molina et al., 16807 (2); 17522, 17728 (1)

Moore, s.n. (2)

Moraga et al., 334 (2)

Morales, 3919 (1)

Moran, 2377 (2); 3158 (1); 5670 (2)

Moran & Rohrbach, 5394 (11)

Moran et al., 6863 (14); 7567 (2)

Mori & Kallunki, 2653 (1)

Moritz, 330 (2)

Morton, 7176, 7182, 7313, 7458, 7515, 7524 (2)

Mt. Roraima Expidition, (2)

Munch, 170 (2)

Narváez, 641 (14)

Navarrete, 373 (14); 908 (2); 1385, 1388, 1428 (14)

Navarrete & Ollgaard, 3036 (14)

Navarro, 905 (2)

Nee, 9976 (2); 10686 (1)

Núñez, 29 (12); 57, 333 (10); 448, 542 (9); 644 (10)

Øllgaard, 99568, 99575, 99581 (2)

Øllgaard & Balslev, 8069 (14); 8267, 8331 (15); 8589 (14)

Øllgaard & Holm-Nielsen, 38755 (14)

Øllgaard & Navarrete, 1641 (11); 2297, 3014 (2)

Øllgaard et al., 38632, 38693 (14); 1187 (15); 2235 (2); 34360 (14); 35598, 35900, 37794 (2); 38137 (14); 90445 (11); 98177 (14); 99638 (11)

Ortega & van der Werff, 2236, 2263 (6)

Ortega et al., 2044 (2)

Ownbey, 2616 (2)

Palacios & Tipaz, 9884 (15); 9898 (14)

Paz, 78 (15)

Penland & Summers, 207 (2)

Pennell, 9260 (8)

Penneys, 469 (1)

Peyton & Peyton, 1331 (10)

Pipoly et al., 17680 (2)

Pittier, 2000 (2)

Pizarro, 9 (4)

Popenoe, 1095 (2)

Portugal, 289 (9)

Proctor, 9704, 10844, 25071 (2)

Questel, 3228 (2)

Ramirez et al., 7794 (2)

Ramos, 7392 (15)

Ramos et al., 6372 (2); 6601 (15)

Rimbach, 22 (17); 37 (15); 113a (14); 127a, 131, 131a (17)

Rivero & Díaz, 1422 (2)

Rodríguez, J., 603, 641, 646 (15)

Rodríguez, J. V., 756, 1945 (2)

Rodríguez, W. D., 5182 (3)

Rodríguez, W. D. et al., 4746, 4755, 5195 (2); 5128 (3)

Rojas, 703 (2); 2430 (1); 3128 (2); 4279 (1)

Rojas et al., 2820 (2)

Ruíz, 58 (2)

Rusby, 382 (2)

Salinas, 2460 (9)

Santiago, 2626 (14)

Schneider, 2400 (6)

Scolnik et al., 19an445 (2)

Seibert, 95, 196, 648, 987 (2)

Silverston-Sopkin et al., 9809, 9934 (2)

Skutch, 3229 (2)

Smith, A. R. & Beliz, 2109 (2)

Smith, A. R. et al., 1569 (2); 1572 (6); 2388, 2800 (2)

Smith, Austin, 85 (2)

Smith, D. N. & Canne, 5837 (9)

Smith, D. N. et al., 2607 (9)

Sodiro, 33, 136, s.n. (15); s.n. (2); s.n. (14); s.n. (17)

Solomon, 7261 (10); 8656 (12); 9112 (9)

Sperling, 5267 (15); 5269 (14)

Sperling & Bleiweiss, 5024 (2); 5165 (17)

Spross, 117 (2)

Spruce, 4642 (2); 5372 (15)

Standley, 4902

Standley, 15653, 35403, 38367, 38566, 38667, 38749 (2)

Standley & Torres, 47752 (2)

Standley & Valerio, 44598, 44610 (1); 48052, 50466 (2)

Steinbach, 9554, 9720 (4)

Steyermark, 30020, 42665, 43610, 53612, 58710, 58731, 75401 (2)

Steyermark & Liesner, 118862 (16)

Steyermark et al., 128039 (2)

Stolze, 1416 (2)

Stolze & Stolze, 1663 (11); 1678 (2)

Stork, 1969 (2)

Stübel, 663 (16)

Sundue, 1125 (11); 1128, 1176 (2)

Sundue & Martin, 1031, 1035, 1050 (2)

Sundue & Schuettpelz, 1073 (11); 1075 (2); 1078, 1084 (11); 1086 (2); 1088 (11); 1091 (8)

Sundue & Vasco, 1208 (3); 1232 (2); 1237 (3); 1306, 1312, 1316, 1331, 1334, 1335 (7)

Sundue & Werner, 1164 (11)

Sundue et al., 782, 785 (9); 866 (12); 1070 (15); 1689 (2)

Tate, 504 (2)

Tejero-Díaz, 4073 (2)

Tillet, 673-231 (2)

Tonduz, 12197 (2)

Torres, 66, 69a (2)

Tryon & Tryon, 5310 (2); 6148 (14)

Ulloa, 92952 (15)

Underwood, 540, 773, 1273, 1369, 2346, 2448, 2451, 2489, 3157 (2)

Valenzuela et al., 1749 (9)

van der Werff & Palacios, 10622 (14)

van der Werff et al., 8451, 8452 (13); 8799, 8822 (6); 15007 (13); 15535; 16676, 16700 (11); 16773 (2); 17706, 18537 (13); 18599, 18622 (9); 19568 (11); 19569 (2)

Vargas & Narvaez, 3531 (2)

Vasco & Sundue, 650, 702 (3)

Vasquez et al., 28446 (4); 28474 (13); 30123 (9)

Weatherwax, 325 (15)

Webster & Addison, 27573 (2)

Webster et al., 29317, 30299 (2)

Werner, 734 (2)

Williams & Molina, 10287, 10387, 11142, 12790, 13688, 18869, 40732 (2)

Wilson, 2816 (14)

Windish, 2363 (2)

Woodson Jr. et al., 829 (2)

Woytowski, 7840 (2)

Wurdack, 703 (4)

Yunker et al., 6197 (2)

Zanoni et al., 20416, 24947, 37502 (2)

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Sundue, M.A. A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae). Brittonia 62, 357–399 (2010). https://doi.org/10.1007/s12228-009-9108-6

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