Research Article |
Corresponding author: Alexandre Salino ( salinobh@gmail.com ) Academic editor: Pavel Stoev
© 2015 Alexandre Salino, Thaís E. Almeida, Alan R. Smith.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Salino A, Almeida TE, Smith AR (2015) New combinations in Neotropical Thelypteridaceae. PhytoKeys 57: 11-50. https://doi.org/10.3897/phytokeys.57.5641
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288 new combinations of Neotropical Thelypteridaceae taxa are proposed in order to recognize monophyletic genera, based on the results of the most recent molecular phylogeny of the family, as well as the morphological uniformity of characters for each genus. The new nomenclatural combinations correspond to 186 Amauropelta taxa, 77 species of Goniopteris, and 25 Steiropteris taxa. A key to all native Neotropical genera of the family is also presented.
Amauropelta , ferns, Goniopteris , Steiropteris , Thelypteris
The Thelypteridaceae, comprising approximately 1,000 species, is one of the largest families of ferns (
Recent molecular phylogenetic studies have recovered two main groups in the family: the phegopteroid and the thelypteroid lineages (
The following native Neotropical genera of Thelypteridaceae are recognized: Amauropelta, Christella (a more precise circumscription still pending), Cyclosorus , Goniopteris ( ) , Meniscium , Stegnogramma , Steiropteris , and Thelypteris , based on the recent molecular phylogenetic data for this family (
Utilizing our most recent results on molecular phylogeny of the family (
Amauropelta, predominantly Neotropical, is most closely related to several Old World genera: Coryphopteris, Metathelypteris, and Parathelypteris. Amauropelta is recognized mainly by the laminar base usually gradually or abruptly reduced with one to many pairs of auriculiform or glanduliform pinnae, and by the veins from adjacent segments usually meeting the blade margins above the sinuses on pinnatifid pinnae.
Our results (
Goniopteris has long been recognized as a natural group (
The narrow circumscription of Cyclosorus used here has already been adopted by some authors (e.g.,
Following
Meniscium, wholly Neotropical, appears to be monophyletic and more closely related to Cyclosoruss.s. (Cyclosorus interruptus), Mesophlebium , and Ampelopteris (
1 | Forked or stellate hairs present at least on the rachises and costae, and/or on the scales at apices of rhizomes, and may be present in other parts of the frond | Goniopteris |
– | All hairs unbranched (hairs fasciculate, i.e., basally clustered, in some species of Amauropelta) | |
2 | Laminae 1-pinnate, rarely simple; pinnae entire or with an serrate or undulate margin, rarely shallowly lobed; veins regularly anatomosing, areoles 3–25-seriate between costae and pinna margins | 3 |
– | Laminae 1-pinnate-pinnatifid, rarely 2-pinnate; pinnae incised 1/3 their width or more; veins meeting the margins above sinuses, or connivent at sinuses, or with 1 or 2 pairs of anastomosing below sinuses | 4 |
3 | Sori usually in a single row between adjacent costules (except Meniscium andreanum, which often has a double row of sori between main lateral veins), oblong or linear, rarely round; sori exindusiate | Meniscium |
– | Sori usually in a double row between adjacent costules, usually round (except G. holodictya, which sometimes has only one row of somewhat elongate sori between costules); sori indusiate (indusial may be small, or hidden in mature sori!) or exindusiate | Goniopteris |
4 | Sporangia setose; sori elongate, exindusiate | 5 |
– | Sporangia not setose, or if setose (in eight species of Amauropelta) then sori round; sori round, oblong, or rarely elongate; sori indusiate or exindusiate | 6 |
5 | Pinnae broadly adnate and decurrent onto rachises in the distal half of laminae; proximal pinnae not cuneate at bases | Stegnogramma (S. pilosa, S. burksiorum) |
– | Pinnae neither adnate nor decurrent onto rachises, sometimes one distal pair adnate to rachises; proximal pinnae short- to long-cuneate at bases | Steiropteris (S. polypodioides, S. villosa) |
6 | Laminae with all veins usually meeting margins above the sinuses | 7 |
– | Laminae with at least some of the proximal veins running to the sinuses or forming an excurrent vein to the sinuses | 9 |
7 | Laminae narrowed proximally with (1-) 2−20 pairs of greatly reduced pinnae, the proximal pinnae usually auriculiform or glanduliform, rarely proximal pinnae not reduced | Amauropelta |
– | Laminae usually not narrowed proximally, or only slightly so, usually lacking greatly reduced pinnae | 8 |
8 | Rhizomes long-creeping; laminae 1-pinnate-pinnatifid to 2-pinnate; veins of sterile laminae 1- or 2-forked, sometimes simple in the fertile laminae | Thelypteris |
– | Stems ascending to erect, rarely short-creeping; laminae 1-pinnate-pinnatifid; veins of sterile and fertile laminae simple.. Steiropteris (subg. Glaphyropteris) | |
9 | Rhizomes long-creeping to 3 m or more, blackish, scaleless or nearly so | Cyclosorus |
– | Rhizomes short-creeping to ascending or erect, light brown, with conspicuous scales, at least at apices | 10 |
10 | Cartilaginous keel (false vein) present below pinna sinuses, keels extending toward costa but not meeting it; aerophores (tuberculate or often threadlike) at pinna bases usually present, absent in a few species; indusia present or absent | Steiropteris (subg. Steiropteris) |
– | Cartilaginous keel absent below pinna sinuses; aerophores at pinna bases absent; indusia present, persistent | Christella |
The names proposed as needing new combinations were obtained from taxonomic and floristic sources such as floras, monographs, or isolated published papers. All names were checked in IPNI.ORG and TROPICOS.ORG. The main floras consulted were ones for Mexico (
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Thelypteris achalensis ( ) , Darwiniana 13: 566. 1964.
Thelypteris aculeata , Fl. Ecuador 18: 15. F. 1A, 1B. 1983.
Thelypteris acunae , Fontqueria 31: 223. f.1. 1991.
Dryopteris aliena , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16: 23. 1937.
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Aspidium argentinum , Bot. Jahrb. Syst. 22: 367. 1896.
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Aspidium atrorubens , Linnaea 36. 112. 1869.
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Dryopteris bonapartii , Repert. Spec. Nov. Regni Veg. 7: 303–304. 1909.
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Nephrodium brachypodum , Timehri 5: 213. 1886.
Thelypteris brachypoda ( ) , Fieldiana, Bot. 28: 10. 1951.
Nephrodium brachypus , Recens. Crypt. Vasc. Quit. 43. 1883.
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Dryopteris brausei , Hedwigia 46: 337, t. 6, f. 11. 1907.
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Thelypteris burkartii , Darwiniana 13: 550, f. 4, t. 2. 1964.
Thelypteris campii , Fl. Ecuador 18: 27. 1983.
Nephrodium canadasii , Recens. Crypt. Vasc. Quit. 48. 1883.
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Nephrodium caucaense , Bot. Jahrb. Syst. 34: 444. 1904.
Thelypetris caucaensis ( ) , J. Wash. Acad. Sci 48: 233. 1958.
Thelypteris chaparensis , Brittonia 60 (1): 51. 2008.
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Dryopteris christensenii , Bull. Herb. Boissier, sér. 2, 7 (4): 263. 1907.
Thelypteris christensenii ( ) , Phytologia 17 (4): 267. 1968.
Nephrodium cinereum , Sert. Fl. Ecuad. II. 26. 1908.
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Thelypteris cocos , Proc. Biol. Soc. Wash. 98 (4): 918. f. 3. 1985.
Thelypteris comptula , Fieldiana, Bot., n.s., 29: 23. 1992.
Nephrodium conforme , Recens. Crypt. Vasc. Quit. 45. 1883.
Thelypteris conformis ( ) , Fl. Ecuador 18: 37. 1983.
Aspidium consanguineum , Mém. Foug. 11 (Hist. Foug. Ant.): 76, t. 20, f. 3). 1866.
Thelypteris consanguinea ( ) , Rhodora 61 (732): 306. 1959.
Thelypteris cooleyi , Rhodora 68: 468. 1966.
Nephrodium corazonense , J. Bot. 15: 163. 1877, as carazanensis”.
Thelypteris corazonensis ( ) , Fl. Ecuador 18: 38. 1983.
Dryopteris cornuta , J. Wash. Acad. Sci. 19 (12): 245. f. 1. 1929.
Thelypteris cornuta ( ) , Bull. Fan Mem. Inst. Biol. 10: 251. 1941.
Thelypteris correllii , Fl. Ecuador 18: 39. 1983.
Dryopteris crassiuscula , Amer. Fern J. 23: 75. 1933.
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Thelypteris ctenitoides , Fieldiana, Bot., n.s., 29: 37. 1992.
Thelypteris decrescens , Amer. Fern J. 71(2): 57. 1981.
Asplenium decurtatum Fil. sp. 94. 1841.
Dryopteris deflectens , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 25, t. 3, f. 9-11. 1937.
Thelypteris delasotae , Proc. Biol. Soc. Wash. 98(4): 919. f. 5. 1985.
Thelypteris demissa , Fieldiana, Bot., n.s., 29: 21. 1992.
Thelypteris denudata , Willdenowia 35: 159. 2005.
Thelypteris dodsonii , Fl. Ecuador 18: 41. 1983.
Thelypteris dudleyi , Fieldiana, Bot., n.s., 29: 33. 1992.
Nephrodium elegantulum , Crypt. Vasc. Quit. 243. 1893.
Thelypteris elegantula ( ) , J. Wash. Acad. Sci. 48: 233. 1958.
Thelypteris enigmatica , Fieldiana, Bot., n.s., 29: 16. 1992.
Aspidium eriosorus , Crypt. Vasc. Brésil, 2. 73. T. 101. 1873.
Thelypteris eriosorus ( ) Ponce (as T. eriosora), Novon 8: 275. 1998.
Polypodium euchlorum , Recens. Crypt. Vasc. Quit. 58. 1883.
Thelypteris euchlora ( ) , Phytologia 17(4): 275. 1968.
Thelypteris euthythrix , Fl. Ecuador 18: 44. 1983.
Thelypteris exuta , Fl. Ecuador 18: 44. 1983.
Thelypteris fasciola , Brittonia 60 (1): 51. 2008.
Thelypteris fayorum , Brittonia 60(1): 53. 2008.
Nephrodium firmum , J. Bot. 17: 260. 1879.
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Aspidium funckii , Ann. Sci. Nat., Bot., sér. 5, 2: 246. 1864.
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Thelypteris furfuracea , Fieldiana, Bot., n.s., 29: 34. 1992.
Dryopteris furva , J. Wash. Acad. Sci. 34: 24. 1944.
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Phegopteris germaniana , Mém. Foug. 11 (Hist. Foug. Ant.): 55. 1866.
Thelypteris germaniana ( ) , Rhodora 61: 306. 1959.
Dryopteris glandulosolanosa , Dansk Bot. Ark. 9(3): 61. 1937.
Thelypteris glandulosolanosa ( ) , Rhodora 69: 6. 1967.
Dryopteris glutinosa , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 18. 1937.
Thelypteris glutinosa ( ) , Amer. Fern J. 53(2): 66. 1963.
Thelypteris gomeziana , Proc. Biol. Soc. Wash. 98(4): 921. f 6. 1985.
Polypodium gracilentum , Bull. Bot. Dept. Jamaica 4. 129. 1897.
Thelypteris gracilenta ( ) , Amer. Fern J. 71(2): 60. 1981.
Gymnogramma gracilis , Mag. Nat. Hist., n.s, 2: 457. 1838.
Thelypteris gracilis ( ) , Bull. Inst. Jamaica, Sci. Ser. 5: 60. 1953.
Thelypteris grayumii , Ann. Missouri Bot. Gard. 77(1): 122. f 2D-E. 1990.
Thelypteris harrisii , Amer. Fern J. 71: 59. 1981.
Dryopteris hastiloba , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 20. t 4, f 4-5. 1937.
Dryopteris heineri , Repert. Spec. Nov. Regni Veg. 6: 380. 1909.
Thelypteris heineri ( ) , Phytologia 17(4): 282. 1968.
Thelypteris hutchisonii , Fieldiana, Bot., n.s., 29: 29. 1992.
Phegopteris hydrophila , Mém. Foug. 11 (Hist. Foug. Ant.): 56. pl 13, f 3. 1866.
Thelypteris hydrophila ( ) , Rhodora 61: 306. 1959.
Dryopteris illicita , Bull. Soc. Bot. Genève sér. 2, 1(5): 225. 1909.
Thelypteris illicita ( ) , Phytologia 17(4) : 284. 1968.
Thelypteris inabonensis , Amer. Fern J. 75(2): 61. f 3. 1985.
Dryopteris euchlora ( ) var. inaequans , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 150. 1913.
Thelypteris inaequans ( ) , Amer. Fern J. 67(2): 60. 1977.
Thelypteris inaequilateralis , Brittonia 60(1): 53. 2008.
Aspidium insigne , Ann. Sci. Nat., Bot., sér. 5, 2: 247. 1864.
Thelypteris insignis ( ) , Bull. Fan Mem. Inst. Biol. 10: 242. 1941.
Dryopteris intromissa , Kungl. Svenska Vetenskapsakad. Handl., ser. 3., 16(2): 22. t 4, f 9-10. 1937.
Dryopteris ireneae , Sellowia 17: 57. f 4. 1965.
Thelypteris ireneae ( ) , Amer. Fern. J. 74(2): 60. 1984.
Dryopteris jimenezii , Amer. Fern J. 4: 79. 1914.
Thelypteris jimenezii ( ) , Phytologia 17(4): 285. 1968.
Nephrodium juergensii , Festschrift Albert von Bamberg 63. 1905.
Thelypteris juergensii ( ) , Phytologia 17(4): 286. 1968.
Thelypteris jujuyensis , Darwiniana 18: 222. f 3, 8c. 1973.
Phegopteris laevigata , Linnaea 36. 112. 1869.
Thelypteris laevigata ( ) , Rhodora 69: 6. 1967.
Thelypteris leoniae , Fieldiana, Bot., n.s., 29: 18. 1992.
Dryopteris lepidula , Hedwigia 46. 328, t. 4, f. 4. 1907.
Thelypteris lepidula ( ) , J. Wash. Acad. Sci. 48: 233. 1958.
Nephrodium longicaule , J. Bot. 19: 204. 1881.
Thelypteris longicaulis ( ) , Phytologia 17(4): 289. 1968.
Nephrodium longipilosum , Anales Univ. Central Quito Ecuador 22(160): 103. 1908.
Thelypteris longipilosa ( ) , Phytologia 17(4): 290. 1968.
Thelypteris longisora , Ann. Missouri Bot. Gard. 77(1): 123. f 3D-E. 1990.
Thelypteris loreae , Mem. New York Bot. Gard. 88: 629–630. f 303A-E, 2004.
Thelypteris loretensis , Fieldiana, Bot., n.s., 29: 30. 1992.
Thelypteris lumbricoides , Brittonia 60(1): 54. 2008.
Thelypteris macra , Fl. Ecuador 18: 54. 1983.
Thelypteris madidiensis , Brittonia 60(1): 54. 2008.
Dryopteris malangae , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 21. t. 4(6−8). 1937.
Thelypteris malangae ( ) , Amer. Fern J. 53(2): 66. 1963.
Dryopteris melanochlaena , Smithsonian Misc. Collect. 52(3): 384. 1909.
Thelypteris melanochlaena ( ) , Phytologia 17(4): 292. 1968.
Dryopteris mertensioides , Kongel. Danske.Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 328, f. 50. 1907.
Thelypteris mertensioides ( ) , Phytologia 17(4): 292. 1968.
Thelypteris metteniana , Bull. Fan mem. Inst. Biol. 10: 252. 1941.
Thelypteris micula , Fieldiana, Bot., n.s., 29: 33. 1992.
Thelypteris minima , Brittonia 60(1): 55. 2008.
Thelypteris minutula , Amer. Fern J. 43(4): 173. 1953.
Thelypteris mombachensis , Phytologia 52(3): 155–156. 1982.
Thelypteris mortonii , Amer. Fern J. 63(3): 125. f 11. 1973.
Dryopteris mosenii , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 300. f 27. 1907.
Thelypteris mosenii ( ) , Phytologia 17(4): 294. 1968.
Thelypteris muscicola , Rhodora 63: 33. 1961.
Thelypteris namaphila , Amer. Fern J. 75(2): 56. f 1. 1985.
Dryopteris neglecta , Bol. Mus. Nac. Rio de Janeiro 7(3): 142, t.1, f 3, t. 7. 1931.
Thelypteris neglecta ( ) , Bull. Fan Mem. Inst. Biol. Bot. 10: 253. 1941.
Nephrodium negligens , Bull. Bot. Dept. 3: 21. 1896.
Thelypteris negligens ( ) , Amer. Fern J. 71(2): 58. 1981.
Thelypteris nephelium , Brittonia 60(1): 57. 2008.
Polypodium nitens , Mém. Soc. Linn. Paris 6: 240. 1827.
Thelypteris nitens ( ) , Rhodora 69(777): 7. 1967.
Dryopteris novaeana , Arq. Inst. Biol. Veg. 3: 2, t. 2, 4(4-6), 6(4-5). 1936.
Thelypteris novaena ( ) , Novon 8(3): 275. 1998.
Thelypteris nubicola , Darwiniana 18: 225. pl 4, 8b. 1973.
Thelypteris nubigena , Proc. Calif. Acad. Sci., ser. 4, 40(8): 229. f 7C. 1975.
Thelypteris oaxacana , Amer. Fern J. 63(3): 125. f 7-8. 1973.
Thelypteris ophiorhizoma , Proc. Biol. Soc. Wash. 98(4): 925. f 8. 1985.
Thelypteris oviedoae , Fontqueria 31: 225. f 2. 1991.
Aspidium pachyrhachis , Abh. Senckenberg. Naturf. Ges. 2: 367. 1858.
Thelypteris pachyrhachis ( ) , Bull. Fan Mem. Inst. Biol. 10: 253. 1941.
Thelypteris paleacea , Fl. Ecuador 18: 62. 1983.
Gymnogramma patula , Crypt. vasc. Brésil 1: 59, t. 14, f. 3. 1869.
Polypodium pavonianum , Linnaea 20. 386. 1847.
Thelypteris pavoniana ( ) , Rhodora 69(777): 7. 1967.
Thelypteris pelludia , Brittonia 60(1): 58. 2008.
Thelypteris peradenia , Ann. Missouri Bot. Gard. 77(2): 272, f. 11. 1990.
Dryopteris peruviana , Repert. Spec. Nov. Regni Veg. 7: 298-299. 1909.
Thelypteris peruviana ( ) , Rhodora 69(777): 7. 1967.
Dryopteris phacelothrix , Repert. Spec. Nov. Regni Veg. 11: 56. 1912.
Thelypteris phacelothrix ( ) , Rhodora 69(777): 7. 1967.
Aspidium physematioides , Bot. Jahrb. Syst. 24: 115. 1897.
Thelypteris physematioides ( ) , Amer. Fern J. 43(4): 174. 1953.
Dryopteris piedrensis , Smithsonian Misc. Collect. 52: 372. 1909.
Thelypteris piedrensis ( ) , Amer. Fern J. 53(2): 69. 1963.
Thelypteris pilosissima , Fieldiana, Botany 28: 11. 1951.
Nephrodium pilosohispidum , Spec. Fil. 4: 105. 1862.
Thelypteris pilosohispida ( ) , J. Wash. Acad. Sci. 48(7): 233. 1958.
Dryopteris pleiophylla , Fl. Ilustr. Catarin. 1(Aspid.): 245. f 61. 1979.
Thelypteris pleiophylla ( ) , Darwiniana 33(1-4): 270. 1995.
Dryopteris podotricha , Fl. Ilustr. Catarin. 1(Aspid.): 232. f 57. 1979.
Thelypteris podotricha ( ) , Darwiniana 33(1-4): 270. 1995.
Thelypteris proboscidea , Fieldiana, Bot., n.s., 29: 36. 1992.
Thelypteris proctorii , Proc. Biol. Soc. Wash. 98(4): 925. f 9. 1985.
Thelypteris prolatipedis , Amer. Fern J. 67(2): 60, 1977.
Dryopteris coarctata var. longipes , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 292. 1907.
Dryopteris ptarmiciformis , Repert. Spec. Nov. Regni Veg. 12: 472. 1913.
Thelypteris ptarmiciformis ( ) , Phytologia 17(4): 307. 1968.
Polypodium pteroideum , Linnaea 20. 389. 1847.
Thelypteris pteroidea ( ) , Rhodora 69(777): 8. 1967.
Aspidium pusillum , Ann. Sci. Nat., Bot., sér. 5, 2: 245. 1864.
Thelypteris pusilla ( ) , Bull. Fan Mem. Inst. Biol. 10: 254. 1941.
Dryopteris raddii , Hedwigia 56: 367. 1915.
Thelypteris raddii ( ) , Darwiniana 33(1-4): 266. 1995.
Thelypteris randallii , Amer. Fern J. 53(2): 69. 1963.
Dryopteris recumbens , Hedwigia 46. 123. 1906.
Thelypteris recumbens ( ) , Phytologia 17: 308. 1968.
Dryopteris reducta , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 18, t. 2, f. 1-3. 1937.
Dryopteris regnelliana , Kongel. Danske.Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 284, f. 12. 1907.
Thelypteris regnelliana ( ) , Darwiniana 33(1-4): 264. 1995.
Nephrodium retrorsum , Recens. Crypt. Vasc. Quit. 51. 1883.
Thelypteris retrorsa ( ) , Fl. Ecuador 18: 71. 1983.
Thelypteris rheophyta , Amer. Fern J. 75(2): 58. f 2. 1985.
Nephrodium rigescens , Crypt. Vasc. Quit. 239. 1893.
Thelypteris rigescens ( ) , Fl. Ecuador 18: 72. 1983.
Aspidium rivularioides , Crypt. Vasc. Brésil 1: 145, t. 50, f. 1. 1869.
Thelypteris rivularioides ( ) , Revista Mus. La Plata, Secc. Bot. 9(36-37): 19. 1958.
Polypodium roraimense , Timehri 5: 214. 1886.
Thelypteris roraimensis ( ) , Phytologia 17(4): 310. 1968.
Dryopteris rosenstockii , Kongel. Danske.Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 304, f. 30. 1907.
Thelypteris rosenstockii ( ) , Rhodora 69(777): 8. 1977.
Thelypteris rosulata , Brittonia 60(1): 60. 2008.
Dryopteris rudiformis , Univ. Calif. Publ. Bot. 19(9): 305. 1941.
Thelypteris rudiformis ( ) , Fl. Ecuador 18: 74. 1983.
Polypodium rufum , Encycl. 5: 532. 1804.
Thelypteris rufa ( ) Fl. Ecuador 18: 77. 1983.
Polypodium ruizianum , Linnaea 20: 385. 1847.
Thelypteris ruiziana ( ) Fieldiana, Bot., n.s. 29: 35. 1992.
Leptogramma rupestris , Linnaea 20. 415. 1847.
Thelypteris rupestris ( ) , Phytologia 17(4): 310. 1968.
Dryopteris rupicola , Repert. Spec. Nov. Regni Veg. 15: 24. 1917.
Thelypteris rupicola ( ) , Bull. Fan Mem. Inst. Biol. 10: 254. 1941.
Phegopteris rustica , Mém. Foug. 11 (Hist. Foug. Ant.): 55, t. 13, f. 1. 1866.
Thelypteris rustica ( ) , Rhodora 61(732): 306. 1959[1960].
Dryopteris sanctae-catharinae , Hedwigia 46: 126. 1906.
Thelypteris sanctae-catharinae ( ) , Darwiniana 33(1-4): 280. 1995.
Polypodium saxicola , Kongl. Vetensk. Acad. Handl. 1817: 59, t. 3, f. 5. 1817.
Thelpyteris saxicola ( ) , Phytologia 17(4): 312. 1968.
Phegopteris scalpturoides , Mém. foug. 11 (Hist. foug. Ant.): 51. 1866.
Thelypteris scalpturoides ( ) , Phytologia 17(4): 312. 1968.
Dryopteris sellensis , Kungl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 24. t. 3(7−8). 1937.
Nephrodium semilunatum , Recens. Crypt. Vasc. Quit. 46. 1883.
Thelyteris semilunata ( ) , Fl. Ecuador 18: 79. 1983.
Dryopteris shaferi , Amer. Fern J. 4: 77. 1914.
Thelypteris shaferi ( ) , Adansonia, sér. 2, 11: 719. 1971 [1972].
Thelypteris soridepressa , Amer. Fern J. 98(4): 199. f 1. 2009.
Thelypteris steyermarkii , Fl. Ecuador 18: 79. 1983.
Gymnogramma stierii , Festschrift Albert von Bamberg 64. 1905.
Thelypteris stierii ( ) , Phytologia 17(4): 316. 1968.
Nephrodium stramineum , Recens. Crypt. Vasc. Quit. 43. 1883.
Thelypteris chimboracensis , Fl. Ecuador 18: 34. 1993.
Thelypteris strigillosa , Proc. Biol. Soc. Wash. 98(4): 927. F 10. 1985.
Dryopteris struthiopteroides , Smithsonian Misc. Collect. 52(3): 388. 1909.
Thelypteris struthiopteroides ( ) , Phytologia 17(4): 316. 1968.
Thelypteris subscandens , Ann. Missouri Bot. Gard. 77(1): 123. f 4. 1990.
Thelypteris subtilis , Fl. Ecuador 18: 80. 1983.
Nephrodium supinum , Crypt. Vasc. Quit. 241. 1893.
Thelypteris supina ( ) , Fl. Ecuador 18: 82. 1983.
Aspidium tablanum , Bull. Herb. Boissier, sér. 2, 5: 727. 1905.
Thelypteris tablana ( ) , Amer. Fern J. 63(3): 127. 1973.
Dryopteris tamandarei , Hedwigia 56. 365. 1915.
Thelypteris tamandarei ( ) , Novon 8(3): 277. 1998.
Thelypteris tapantensis , Proc. Biol. Soc. Wash. 98(4): f 11. 927. 1985.
Aspidium tenerrimum , Crypt. Vasc. Brésil 1: 134. t 43, f 1. 1869.
Thelypteris tenerrima ( ) , Phytologia 17(4): 319. 1968.
Thelypteris trelawniensis , Amer. Fern J. 71(2): 58. 1981.
Thelypteris uncinata , Fl. Ecuador 18: 86. 1983.
Dryopteris vattuonei , Darwiniana 1: 100. 1924.
Thelypteris vattuonei ( ) , Darwiniana 13: 566. 1964.
Thelypteris venturae , Mem. New York Bot. Gard. 88: 638. f 302A-C. 2004.
Thelypteris vernicosa , Proc. Biol. Soc. Wash. 98(4): 929. f 12. 1985.
Thelypteris villana , Rev. Biol. Trop. 17(1): 106. f 1-4. 1970.
Thelypteris yungensis , Brittonia 60(1): 61. 2008.
Thelypteris zurquiana , Proc. Biol. Soc. Wash. 98(4): 929. f 13. 1985.
Thelypteris abdita . Amer. Fern J. 75(2): 63. f 4. 1985.
Polypodium abruptum , Mém. Soc. Linn. Paris 6: 239. 1827.
Thelypteris abrupta ( ) , Rhodora 61: 305. 1960.
Thelypteris alan-smithiana , Phytologia 50(7): 458. 1982.
Thelypteris amazonica , Nordic J. Bot. 29(5): 611. f 1-2. 2011.
Dryopteris ancyriothrix , Repert. Spec. Nov. Regni Veg. 7: 305-306. 1909.
Thelypteris ancyriothrix ( ) , Fl. Ecuador 18: 140. 1983.
Aspidium anopteron , Linnaea 36. 113. 1869.
Thelypteris anoptera ( ) , Phytologia 17(4): 260. 1968.
Thelypteris aureola , Ann. Missouri Bot. Gard. 77(1): 118. f 1D-E. 1990.
Thelypteris beckeriana , A.R.Sm. & Labiak, Brittonia 62(2): 149. f 1. 2010.
Thelypteris berlinii , Novon 16: 431. 2006.
Nephrodium bibrachiatum , Gard. Chron., ser. 3, 15: 230. 1894.
Thelypteris bibrachiata ( ) , Ferns Jamaica 337. 1985.
Dryopteris biformata , Repert. Spec. Nov. Regni Veg. 7: 300-301. 1909.
Thelypteris biformata ( ) , Rhodora 69(777): 5. 1967.
Phegopteris blanda , Mém. Foug. 8: 91. 1857.
Thelypteris blanda ( ) , Phytologia 17(4): 264. 1968.
Thelypteris calypso , Brenesia 8: 98. 1976.
Thelypteris chocoensis , Proc. Biol. Soc. Wash. 98(4): 916. f 1. 1985.
Dryopteris clypeata , Bull. Torrey Bot. Club 66: 52. 1939.
Thelypteris clypeata ( ) , Acta Bot. Neerl. 18: 141. 1969.
Phegopteris cordata , Mém. Foug. 5 (Gen. filic.): 244. 1852.
Thelypteris cordata ( ) , Bull. Inst. Jamaica, Sci. Ser. 5: 59. 1953.
Thelypteris crenata , Proc. Biol. Soc. Wash. 98(4): 919. f 4. 1985.
This needs a new name in Goniopteris because of the prior Goniopteris crenata C. Presl, 1836.
Thelypteris crassipila , Willdenowia 34: 513. 2004.
Thelypteris croatii , Ann. Missouri Bot. Gard. 77(1): 122. f 1A-C. 1990.
Dryopteris curta , Bull. Herb. Boissier, sér. 2, 7(4): 263. 1907.
Thelypteris curta ( ) , Phytologia 17(4): 270. 1968.
Dryopteris dissimulans , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 215. 1913.
Thelypteris dissimulans ( ) , Phytologia 17(4): 273. 1968.
Nephrodium equitans , Bull. Herb. Boissier, sér. 2, 6(2): 163. 1906.
Thelypteris equitans ( ) , Phytologia 17(4): 275. 1968.
Thelypteris erythrothrix , Fieldiana, Bot., n.s., 29: 59. 1992.
Dryopteris gonophora , Lloydia 2: 164. f. 1(1−3). 1939.
Thelypteris gonophora ( ) , Ann. Missouri Bot. Gard. 77(1): 203. 1990.
Thelypteris hildae , Amer. Fern J. 75(2): 67. f 6. 1985.
Thelypteris holodictya , Acta Bot. Neerl. 18: 140. f. 2. 1969.
Thelypteris hondurensis , Phytologia 50(7): 458. 1982.
Dryopteris imitata , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 29, t. 6, f. 1-4. 1937.
Thelypteris imitata ( ) , Moscosoa 3: 46. 1978.
Thelypteris indusiata , Phytotaxa 156(5): 280. f 1-3. 2014.
Nephrodium jamesonii , Sp. fil. 4: 66. 1862.
Thelypteris jamesonii ( ) , Rhodora 69(777): 6. 1967.
Thelypteris jarucoensis , Willdenowia 34: 515. 2004.
Thelypteris killipii , Proc. Biol. Soc. Wash. 98(4): 923. f 7. 1985.
Thelypteris leonina , Willdenowia 34: 517. 2004.
Aspidium levyi , Bull. Soc. Bot. France 19: 255. 1872.
Thelypteris levyi ( ) , Contr. U.S. Natl. Herb. 38: 37. 1967.
Polypodium meniscioides , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 5, 1: 211. 1849.
Dryopteris liebmannii , Bull. Torrey Bot. Club 65: 348. 1938.
Thelypteris meniscioides ( ) , Phytologia 17: 292. 1968.
The earliest species ephitet, meniscioides, has been used already in Goniopteris – G. meniscioides , a synonym of Ampelopteris prolifera ( ) Copel., a Paleotropical species.
Thelypteris littoralis , Brittonia 54(4): 332. f 1. 2002.
Dryopteris lugubriformis , Repert. Spec. Nov. Regni Veg. 7: 299. 1909.
Thelypteris lugubriformis ( ) , Rhodora 69(777): 7. 1967.
Thelypteris martinezii , Mem. New York Bot. Gard. 88: 654-655. f 308I. 2004.
Dryopteris nicaraguensis ( ) var. minor , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 252. 1913.
Thelypteris minor ( ) , Phytologia 34(4): 232. 1976.
Thelypteris minutissima , Willdenowia 34: 511. 2004.
Thelypteris montana , Brittonia 54(4): 334. f 2D-F. 2002.
Thelypteris multigemmifera , Brittonia 54(4): 336. f 3. 2002.
Thelypteris munchii , Amer. Fern J. 63(3): 120. f 9-10. 1973.
Phegopteris nicaraguensis , Bull. Soc. Bot. France 19: 252. 1872.
Thelypteris nicaraguensis ( ) , Contr. U.S. Natl. Herb. 38(2): 55. 1967.
Dryopteris nigricans , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 31, t. 7, f. 1-2. 1937.
Thelypteris oroniensis , Amer. Fern J. 68(1): 9. f 1. 1978.
Thelypteris paranaensis , Brittonia 54(4): 337. F 2A-C. 2002.
Aspidium paucijugum , Linnaea 20: 368. 1847.
Thelypteris paucijuga ( ) , Univ. Calif. Publ. Bot. 59: 98. 1971
Nephrodium fendleri var. paucipinnatum , Bot. Gaz. 12: 134. 1887.
Thelypteris paucipinnata ( ) , Phytologia 17(4): 302. 1968.
Aspidium pellitum , Sp. Pl. ed. 4. 5: 242. 1810.
Thelypteris pellita ( ) , Bradea 5(40): 384. 1990.
Nephrodium peripae , Recens. Crypt. Vasc. Quit. 52. 1883.
Thelypteris peripae ( ) , Phytologia 17(4): 303. 1968.
Thelypteris pilonensis , Brittonia 60(1): 58. 2008.
Thelypteris pinnatifida , Fl. Ecuador 18: 110. 1983.
Dryopteris praetermissa , Proc. Biol. Soc. Wash. 57: 20. 1944.
Thelypteris praetermissa ( ) , Phytologia 34(3): 232. 1976.
Thelypteris redunca , Ann. Missouri Bot. Gard. 77(1): 123. f 1F-H. 1990.
Dryopteris resiliens , Field Mus. Publ. Bot. 17: 302. 1938.
Thelypteris resiliens ( ) , Amer. Fern J. 63(3): 121. 1973.
Polypodium retroflexum , Sp. Pl. 1089. 1753.
Thelypteris retroflexa ( ) , Bradea 5(40): 385. 1990.
Thelypteris rhachiflexuosa , Amer. Fern J. 79(3): 122. f 1. 1989.
Dryopteris × rolandii , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 258. 1913.
Thelypteris × rolandii ( ) , Rhodora 69: 8. 1967.
Thelypteris sapechoana , Brittonia 60(1): 60. 2008.
Nephrodium schaffneri , Mém. Foug. 8: 108. 1857.
Thelypteris schaffneri ( ) , Phytologia 17(4): 312. 1968.
Dryopteris schippii , Amer. Fern J. 25: 52. 1935.
Thelypteris schippii ( ) , Phytologia 34(3): 233. 1976.
Thelypteris schomburgkii , Ann. Missouri Bot. Gard. 77(1): 202. f 1. 1990.
Thelypteris schunkei , Fieldiana, Bot., n.s., 29: 63. 1992.
Aspidium semihastatum , Linnaea 9: 91. 1834.
Thelypteris semihastata ( ) , Bull. Fan Mem. Inst. Biol. 10: 254. 1941.
Thelypteris semirii , Novon 10(1): 74. f 1A-e. 2000.
Dryopteris septemjuga , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 32, t. 7, f. 4-5. 1937.
Aspidium skinneri , Icon. Pl. 10: t. 924. 1854.
Thelypteris skinneri ( ) , Phytologia 17(4): 314. 1968
Thelypteris stephanii , Brittonia 60(1): 61. 2008.
Thelypteris stolzeana , Phytologia 34(3): 231. 1976.
Dryopteris subsagittata , Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 28, t. 6, f. 5–6. 1937.
Nephrodium tenebricum , J. Bot. 20: 326. 1882.
Thelypteris tenebrica ( ) , Bull. Inst. Jamaica, Sci. Ser. 5: 65. 1953.
Thelypteris tryoniorum , Fieldiana, Bot., n.s., 29: 62. 1992.
Thelypteris tuxtlensis , Acebey & A.R.Sm., Amer. Fern J. 97(3): 137–139. 2007.
Polypodium urbani , Crypt. Vasc. Quit. 301. 1893.
Thelypteris urbani ( ) , Fl. Ecuador 18: 116. 1983
Thelypteris verecunda , Amer. Fern J. 75(2): 65. f 5. 1985.
Thelypteris yaucoensis , Mem. New York Bot. Gard. 53: 208. 1989.
Glaphyropteris , Crypt. Vasc. Brésil, 2. 40. 1873.
Dryopteris subg. Glaphyropteris ( ) , Biol. Arb. Til. Eug. Warming 80. 1911.
Thelypteris subg. Glaphyropteris ( ) , Alston, J. Wash. Acad. Sci. 48: 234. 1958.
Thelypteris buchtienii , Univ. Calif. Publ. Bot. 76: 10. 1980.
Nephrodium holmei , Ann. Bot. 5: 317. 1891.
Thelypteris clypeolutata ( ) var. holmei ( ) , Univ. Calif. Publ. Bot. 76: 11. 1980.
Dryopteris comosa , J. Wash. Acad. Sci. 28: 528. 1938.
Thelypteris comosa ( ) , Amer. Fern J. 51(1): 38. 1961.
Polypodium decussatum , Species Plantarum 2: 1093. 1753.
Thelypteris decussata ( ) , Bull. Inst. Jamaica, Sci. Ser. 5: 59. 1953.
Dryopteris decussata var. brasiliensis , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 161. 1913.
Thelypteris decussata var. brasiliensis ( ) , Univ. Calif. Publ. Bot. 76: 15. 1980.
Thelypteris decussata var. costaricensis , Univ. Calif. Publ. Bot. 76: 16. 1980.
Dryopteris mapiriensis , Repert. Spec. Nov. Regni Veg. 6: 313. 1909.
Thelypteris decussata var. mapiriensis ( ) , Univ. Calif. Publ. Bot. 76: 16. 1980.
Polypodium velutinum , Recens. Crypt. Vasc. Quit. 59. 1883.
Thelypteris decussata var. velutina ( ) , Univ. Calif. Publ. Bot. 76: 17. 1980.
Aspidium fraseri , Linnaea 36: 109. 1869.
Thelypteris fraseri ( ) , Univ. Calif. Publ. Bot. 76: 32. 1980.
Polypodium brachyodus , Linnaea 9: 48. 1834.
Thelypteris glandulosa var. brachyodus ( ) , Phytologia 34: 233. 1976.
Thelypteris glandulosa ( ) var. longipilosa , Univ. Calif. Publ. Bot. 76: 22. 1980.
Thelypteris hatschbachii , Univ. Calif. Publ. Bot. 76: 22. 1980.
Dryopteris hottensis , Kongl. Svenska Vetenskapsacad. Handl., ser. 3, 16(2): 26. t 2(4-6). 1937.
Thelypteris hottensis ( ) , Univ. Calif. Publ. Bot. 76: 23. 1980.
Nephrodium costale , Syn Fil, 2. 495.1874.
Thelypteris leprieurii ( ) var. costalis ( ) , Univ. Calif. Publ. Bot. 76: 25. 1980.
Thelypteris leprieurii ( ) var. glandifera , Univ. Calif. Publ. Bot. 76: 25. 1980.
Aspidium incanum , Hedwigia 44: 367. 1905.
Thelypteris leprieurii ( ) var. incana ( ) , Univ. Calif. Publ. Bot. 76: 26. 1980.
Thelypteris leprieurii ( ) var. subcostalis , Univ. Calif. Publ. Bot. 76: 26. 1980.
Dryopteris mexiae , Univ. Calif. Publ. Bot., 17: 32. 1932.
Thelypteris mexiae ( ) , Bull. Fan Mem. Inst. Biol. 10: 252. 1941.
Thelypteris parva , Brittonia 60(1): 57. 2008.
Thelypteris pennellii , Univ. Calif. Publ. Bot. 76: 28. 1980.
Dryopteris polyphlebia , Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 161. 1913.
Thelypteris polyphlebia ( ) , Amer. Fern J. 51(1): 38. 1961.
Ceterach polypodioides , Opusc. Sci. 3: 284. 1819.
Thelypteris polypodioides ( ) , Phytologia 7: 305. 1968.
Phegopteris seemannii , in Seem., Bot. Voy. Herald. 228. 1854.
Thelypteris seemannii ( ) , Univ. Calif. Publ. Bot. 76: 30. 1980.
Gymnogramma villosa , Hort. Berol. 2: 51. 1833.
Thelypteris villosa ( ) , Phytologia 7: 323. 1968.
We thank the curators and the staff of the following herbaria: B, BHCB, BM, COL, F, GH, HUA, K, MO, NY, P, PMA, PR, PRC, QCA, QCNE, QPLS, S, UC, and US. Thanks also to CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) for productivity grants and financial support to A. Salino, and to Fapemig (Fundação de Amparo à Pesquisa do Estado de Minas Gerais) for financial support and grants. We thank the two reviewers for helpful comments.