Elaphoglossum bellermannianum (Klotzsch) T.Moore

Vasco, A., 2011, Taxonomic revision of Elaphoglossum subsection Muscosa (Dryopteridaceae), Blumea 56 (2), pp. 165-202 : 171-173

2. Elaphoglossum bellermannianum (Klotzsch) T.Moore — Fig. 1b , 3; Map 3

Elaphoglossum bellermannianum (Klotzsch) T. Moore (1857a) 352.— Acrostichum bellermannianum Klotzsch (1847) 426. — Type: Moritz 259 (lecto, designated here,B; iso HBG n.v.(photo P ex HBG), LE n.v., US), Venezuela, Aragua, Colonia Tovar, [10°23'N, 67°21'W], s.d.

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Elaphoglossum blandum Rosenst.(1913) 476.— Type: Buchtien 3456 (lecto S, designated here;iso B, US), Bolivia, North Yungas , Polo Polo bei Coroico, [16°24'S, 67°47'W], 1100 m, Oct.–Nov. 1912.

Elaphoglossum atropunctatum Mickel (1991) 124. — Type: Smith & Canne 5835 (holo MO), Peru, Pasco,Prov.Oxapampa, 2–4 km N of Mallampampa , 10°32'S, 75°45'W, 2200–2400 m, 22 Jan. 1984.

Elaphoglossum fuliginosum Mickel (1992) 374.— Type: Taylor & Taylor 11339 (holo NY; iso US), Costa Rica, San José, c. 3 km NW of Cascajal near the union of the Cascajal and Bajo Maquina Rivers, [10°04'N, 83°57'W], 30 July 1972.

Plants epiphytic. Rhizome 4– 6 mm wide, compact to short-creeping; rhizome scales 5 –10 mm long, linear-lanceolate, brown, lustrous, subentire, with small teeth, base truncate, apex acuminate. Sterile leaves (6–)10– 30(– 40) cm long, approximate; phyllopodia present, obscured by the petiole-base scales; petiole 5–16 cm long, 2/5–1/2 the length of the sterile leaves; larger petiole scales 2– 5 mm long, scattered to dense, spreading, lanceolate, light brown to dark brown or mottled, darker at the point of attachment, margin lighter, erose to almost entire, base cordate, apex acute; smaller petiole scales 0.3–1.5 mm long, scattered to dense, appressed, oftentimes hidden by the larger ones, oblanceolate to round, from light brown to black or mottled with lighter margin, darker at the point of attachment, erose to ciliate, base cordate, apex acute; lamina 6 –17 by 1.5 –4 cm, oblong to oblanceolate, coriaceous, base broadly cuneate to truncate, apex obtuse; veins c. 1 mm apart, at an angle of 75° with respect to the costa; adaxial laminar scales 0.5–2 mm long, scattered, lanceolate to round, white, occasionally getting dark at the point of attachment, ciliate, base cordate, apex acute; abaxial laminar scales 0.5 –2 mm long, scattered, slightly elevated above the surface of the lamina, lanceolate to round, brown to black or mottled with lighter margin, ciliate, base cordate, apex acute; abaxial costal scales same as the laminar ones, mostly sessile, thus the scale body not elevated, pointing forward to the apex or at right angles; marginal scales same as the laminar ones. Fertile leaves longer than the sterile leaves; petiole 3/4 the length of the fertile leaves; petiole scales same as the ones of the sterile leaves but the larger ones more spreading; lamina (2.5 –)5 –13 by 1.1– 2 cm, narrowly elliptic to elliptic, coriaceous, base broadly cuneate, apex acute to obtuse; adaxial laminar scales 0.5 –1.5 mm long, dense, lanceolate to round, white to light brown, darker at the point of attachment, ciliate, base cordate, apex acute; abaxial costal scales 0.3 –1 mm long, scattered, not elevated, lanceolate to round, brown to black, ciliate, base stalked, apex acute; intersporangial scales c. 1 mm long, scattered, lanceolate, brown to black or mottled with lighter margin, ciliate, base stalked, apex acute. Spores papillate, without ridges.

i 5 mm a g h 1 cm 1 mm e f 0.2 mm d 5 cm c b 2 mm 1 mm

Distribution — Costa Rica, Panama, Dominican Republic, Venezuela, Colombia, Ecuador, Peru, Bolivia; 1000 –3000 m. Epiphytic in montane forests.

Selection of other specimens examined. COSTA RICA, Alajuela, above Lechería on Volcán Poás, [10°11'N, 84°13'W], 2290 m, 30 July 1932, Stork 3371 (UC, US). Cartago, Irazú West, [9°58'48"N, 83°50'60"W], 2000 m, 8 Sept. 1908, Brade 184 (NY, UC). Guanacaste, entrance to Finca Montecristo, off. rt. 113, 1800 m, 28 July 1983, Moran 3309 (CR, F, MO, NY). Limón, Cordillera de Talamanca,Atlantic slope, valle del Silencio along the Río Terbi, 0.5 –1.5 airline km W of the Costa Rican-Panamanian border, 9°08'N, 82°57'W, 9 Sept. 1984, Davidse et al. 28772 (MO). Puntarenas, Cantón de Coto Brus, Z.P. Las Tablas, cuenca Terraba-Sierpe, orillas de la quebrada Surú, 8°58'30"N, 82°46'15"W, 1960 m, 11 Feb. 1999, Rojas et al. 4899 (CR, INB, NY). San José, Cantón de Desamparados, Cordillera de Talamanca, El Empalme, 9°43'20"N, 83°57'20"W, 2 Oct. 1994, Rojas 1551 (CR, INB, UC). – PANAMA, Chiriquí, vic. of Casita Alta, Volcán de Chiriquí, [8°48'N, 82°32'W], 28 June–2 July 1938, Woodson et al. 823 (MO, NY, US). – DOMINICAN REPUBLIC, Ázua, Cordillera Central, Valle del Yaque, 1550 m, 5 Oct. 1929, Ekman 13682 (S). La Vega, c. 17 km S of Constanza, vía El Convento, on road to San José de Ocoa, at Arroyo La Siberia , 18°50'N, 70°42'W, 1600 m, 24 July 1980, Mejía 7604 (MO, NY). Peravia, Loma Los Palos Mojados, NNW de El Bejucal, en la cabecera del Río El Canal, 18°37.5'N, 70°35'W, 1700 m, 4 Aug. 1982, Zanoni et al. 22314 (JSBD, NY). San Juan, along Bao river, upper Bao valley, headwaters of Bao river, [19°01'N, 70°59'W], 1700 m, 1–7 Oct. 1968, Liogier 12878 (NY). – COLOMBIA, Antioquia, vereda Los Patos, camino hacia el Páramo de Santa Ines, 6°37'07.8"N, 75°40'43.8"W, 2590 m, 30 July 2005, Vasco 561 (HUA, MO, NY). Boyacá, Chiquinquirá, bosque detrás del Cuartel, [5°37'N, 73°49'W], 2675 m, 12 Oct. 1967, Jaramillo-Mejía et al. 3474 (COL). Caldas, Mun. Manizales,Monteleón,Cordillera Central, 2250 m, 24 Mar.1984, De Fraume et al. 105 (HUA). Cauca, 5 km E of Timbío, [2°21'N, 75°41'W], 2050 m, 29 Dec.1944, Grant 10663 ( US). Cundinamarca, carretera Guasca a Gachetá, Km 60–61, 2720 m, 22 Jan.1974, Acosta-Arteaga 103 (COL). Nariño, entre La Victoria, inspección de policía Mun. Apiales , y el Páramo El Consuelo, [0°50'N, 77°37'W], 2900 m, 12 Jan. 1973, Hagemann 1928 (COL, PSO). Risaralda, Mun. Santuario, PNN Tatamá, camino que lleva al Páramo de Tatamá,subiendo hacia Morro Zancudo, 5°07'28.1"N, 76°02'30.6"W, 2643 m, 17 Apr. 2007, Vasco et al. 734 (HUA, NY). Tolima, Mun. Sta. Isabel, vereda de La Yuca, cerca de Las Bodegas, Finca Buenavista, Alto La Esperanza, [4°43'N, 75°06'W], 2700 m, 31 July 1980, Idrobo 10311 (COL). – VENEZUELA, Amazonas, Dpto. Rio Negro, Cerro de la Neblina, CampII 2.5 –3.5 km NE Pico Phelps (Neblina), 0°50'N, 65°58'28"W, 16 Feb.1984, Funk 6313 (VEN). Aragua, prope Colonia Tovar,[10°23'N, 67°22'W], 1854–1855, Fendler 270 (GH, MO, NY, US). Barinas, Quebrada de Barinitas, 1100 m, 28 July 1960, Vareschi 7427 (VEN). Distrito Federal, Cordillera de la Costa, fila del Avila, lado S, directamente debajo de la estación meteorológica, entre la estación del teleférico y el Hotel Humboldt, 10°32.6'N, 66°52.6'W, 2100 m, 19 Aug. 1991, Meier 302 (NY, VEN). Lara, Mun. Morán,vía al Páramo Las Rosas en las inmediaciones de la pica, 9°37'N, 70°05'W, 2400 m, 5 Dec. 1984, Rivero 775 (NY). Mérida, Parque Nacional La Culata, En el camino hacia el Cerro Pan de Azúcar, 8°41'52.4"N, 71°05'42.5’’W, 2400 m, 26 May 2008, Vasco et al. 799 (NY, VEN). Portuguesa, Mun.Sucre, en La Divisoria de La Concepción, 9°18'N, 70°06'W, 23–26 Oct. 1985, Ortega et al. 2782 (NY). Táchira, Quebrada La Lejia, S of Quebrada Agua Azul, along and above stream on steep slopes, 15–16 km SE of Delicias, 7°30'N, 72°24'W, 2300 m, 25 July 1975, Steyermark 118596 (MO, VEN). Trujillo, carretera Boconó-Trujillo, entrada a Burbusay, 9°26'N, 70°17'W, 1600 m, 23 Nov. 1984, Ortega 2290 (MO, NY, UC). – ECUADOR, Pichincha, old road Chiriboga to Sto. Domingo, 14 km from recinto de Las Palmeras, 0°18'S, 78°51'W, 800 m, 13 Feb.2004, Moran et al. 6843 (NY). Loja, trails c. 5 km ENE of San Pedro de Vilcabamba, from El Bosque to Quebrada Romerillos and Banderilla, 4°14'S, 79°10'W, 30 Nov. 1994, Øllgaard 105943 (AAU). Morona-Santiago, small ravine c. 7 km N of Limón, disturbed forest, 2°54'42"S, 78°24'03"W, 960 m, 3 Mar. 2005, Moran et al. 7604 (NY). Napo, Cantón Quijos, Baeza, along road leading to radio tower behind town, 1°28'18"S, 77°53'51"W, 2016 m, 24 Feb. 2005, Moran et al. 7525 (HUA, NY). Zamora-Chinchipe, road Loja-Zamora, Km

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24–25, 3°59'S, 79°045'W, 15 Apr.1973, Holm-Nielsen et al. 3519 (AAU,MO, NY, UC). Sucumbios, Km 15–18, carretera Pun-Aguarico, 13 Aug. 1949, Acosta-Solís 13281 (F). – PERU, Amazonas, a few km from Molina Pampa , [6°52'37"S, 78°12'18"W], 14 Mar.1998, Van der Werff et al. 14960 (MO, UC). Cusco, Machupichu, [13°09'S, 72°29'W], Mar. 1936, Soukup 176, 177 (F). Junín, Prov. Tarma,Agua Dulce, [11°25'S, 75°41'W], 1900 m, 9 Mar. 1948, Woytkowski 35433 (UC, US). Pasco, Prov. Oxapampa, Parque Nacional Yanachaga Chemillén, sector San Alberto, 10°32'S, 75°21'W, 2200 m, 20 Jan.2003, Vásquez et al.27857 (MO,USM). San Martín, Rioja-Pomacochas road, below Venceremos, c. 20 km NW of Rioja near Restaurant El Amigo, 5°45'S, 77°38'W, 1600 m, 8 Feb.1984, Gentry 45177 (MO).– BOLIVIA, Cochabamba, Prov. Franz Tamayo, Piedra Blanca, 10.4 km al S de Pata, sobre el camino a Sta. Cruz del Valle Ameno, 14°37'44"S, 68°40'19"W, 1900 m, 12 Nov. 2003, Fuentes et al. 5999 (LPB, MO, NY, UC). Santa Cruz, Prov. Valle Grande, Loma Larga, 4 km a Masicurí, 18°47'S, 63°53'W, 1600 m, 3 June 1996, Kessler et al. 6289 (LPB, NY, UC).

Notes — Elaphoglossum bellermannianum can be distinguished by its oblong laminae, bicolorous scales on the petioles and laminae (at least along the costae), fertile leaves twice as long as the sterile, and papillate spores. It differs from E. muscosum and E. lehmannianum by the bicolorous petiole and/or lamina scales. These scales are bicolorous because they can be darker at the point of attachment, or mottled, or black medially with white borders. In contrast, the scales of E. muscosum and E. lehmannianum are typically concolorous, or if bicolorous they are darker only at the point of attachment. Another difference is that the costal scales of the sterile and fertile leaves of E. bellermannianum are mostly sessile, thus the scale body is not elevated above the surface of the lamina; whereas scales of E. muscosum and E. lehmannianum have a base typically short-stalked and thus the scale body is elevated above the surface of the lamina. The three species are epiphytes with compact to short-creeping rhizomes, linear rhizome scales with few short teeth up to 0.2 mm, erose petiole scales, and papillate spores without ridges ( Fig. 1b, h, i ). These characteristics distinguish them from E. engelii , which differs by being terrestrial with short- to long-creeping rhizomes and by having linear rhizome scales with many teeth longer than 0.2 mm, ciliate petiole scales, and spores with ridges and irregular deposits ( Fig. 1d ).

There is considerable variation in the colour of the laminar scales of E. bellermannianum . Some plants have all the scales of the abaxial lamina either completely black or mottled; these were described as either E. atropunctatum (from Peru) or E. fuliginosum (from Costa Rica). Other plants have almost all adaxial laminar scales white and have been named E. blandum (type from Bolivia). Throughout the geographic distribution of E. bellermannianum , however, scale colour varies from white to black. Because no other differences distinguish these plants, they are treated here as a single species, E. bellermannianum .

See E. gardnerianum for a comparison with that species.

Figures

Map 3 Distribution of Elaphoglossum bellermannianum (Klotzsch) T.Moore.

Fig. 1 Spores of Elaphoglossum subsect. Muscosa. a. E. aschersonii; b. E. bellermannianum; c. E. corderoanum; d. E. engelii; e. E. gardnerianum; f. E. glabrescens; g. E. laxisquama; h. E. lehmannianum; i. E. muscosum; j. E. novogranatense; k. E. oculatum; l. E. oreophilum; m. E. quisqueyanum; n. E. ruficomum; o. E. yatesii (a: Moran 7645, NY; b: Meier 302, NY; c: Holm-Nielsen 3260, AAU; d: Sánchez 465, US; e: Wacket 186, UC; f: Buchtien 2725, US; g: Vásquez 28162, NY;h: Ruíz-Terán 976, NY;i: Underwood 3212, NY;j: Vasco 746, NY; k: Hutchison 1629, UC;l: Holm-Nielsen 19071, AAU; m: Zanoni 20917, NY;n: Sagástegui 12091, NY; o: Navarrete 1402, NY). — Scale bars = 10 µm.