Elaphoglossum engelii (H.Karst.) Christ

Vasco, A., 2011, Taxonomic revision of Elaphoglossum subsection Muscosa (Dryopteridaceae), Blumea 56 (2), pp. 165-202 : 175-179

4. Elaphoglossum engelii (H.Karst.) Christ — Fig. 1d , 5 ; Map 5

Elaphoglossum engelii (H.Karst.) Christ (1899) 81. — Acrostichum engelii H.Karst. (1860) 118, t. 59. — Type: Karsten s.n. (lecto B, designated by Mickel (1991) 131; second-step barcode B20 0070391 excluding the fertile leaf portions in the pocket, designated here; iso W [sheet W0018475, excluding the fertile leaf on the right], LE n.v.), Venezuela (in the original description ‘Colombia’), Merida, cordillera Meridensis, s.d.

Elaphoglossum caulolepia (H.Karst.) Hieron. (1905) 548. — Acrostichum caulolepia H.Karst. (1860) 121, t. 60. — Type: Karsten s.n. (lecto W (sheet W0018476 ),designated here;iso B n.v., LE n.v.), Colombia, Cundinamarca, Bogotá, s.d.

Elaphoglossum atrorubens Mickel (1987) 314, f. 3D, E. — Type: Ruiz-Terán & M. López-Figueiras 1754 (holo NY), Venezuela, Mérida, Distrito Campo Elías, Municipio Pueblo Nuevo, Páramo de Quirorá , alto de las Cruces, [8°20'N, 71°27'W], 3200 m, 23 Apr. 1971.

Elaphoglossum cristatum Ballenth., Dudenh. & M.Kessler (2009) 46–49. — Type: Kessler et al. 6875 (holo GOET n.v.; iso LPB n.v., NY, UC), Bolivia, Cochabamba, Prov. José Carrasco Torrico, 63 km antigua carretera Cochabamba-Villa Tunari , 17°15'S, 65°43'W, 3700 m, 2 July 1999.

Plants terrestrial. Rhizome 4 – 8 mm wide, short to long-creeping; rhizome scales 6–11 mm long, linear-lanceolate, light brown

h

f

to dark brown, sometimes bicolorous, margin lighter, ciliate, with abundant cilia (up to 0.4 mm long), base cordate, apex filiform and tortuous to acute. Sterile leaves 14 –47(– 63) cm long, approximate to up to 1 cm apart; phyllopodia present, sometimes obscured by the petiole-base scales; petiole 6 –25(–41) cm long, 2/5– 3/5 the length of the sterile leaves; larger petiole scales 4 – 6 mm long, dense, spreading, sometimes pointing forward, linear-lanceolate to oblanceolate, light brown to dark brown, or mottled, darker at the point of attachment, margin lighter, ciliate, base cordate to truncate, apex acute; smaller petiole scales 0.5 –1(– 3) 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, ciliate, base cordate, apex acute; lamina 8 – 23 by 1.5 – 4.5 cm, linear-lanceolate to oblanceolate, coriaceous, glabrescent with age, base truncate to broadly cuneate, apex obtuse; veins 0.5–1.5 mm apart, at an angle of 70– 85° with respect to the costa; adaxial laminar scales 1–3 mm long, dense, imbricate, oftentimes pointing downwards, lanceolate, white to light brown, ciliate, base cordate, apex acute; abaxial laminar scales 2 – 3 mm long, dense, imbricate, slightly elevated above the surface of the lamina to patent, lanceolate to oblanceolate, from light brown to black or mottled with lighter margin, darker at the point of attachment, ciliate, base stalked (up to 0.4 mm), apex acute; abaxial costal scales same as the laminar ones, pointing forward to the apex or at right angles, slightly elevated, sometimes darker; marginal scales same as the laminar ones. Fertile leaves longer than the sterile leaves; petiole 1/2– 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 7–22 by 1–2 cm, narrowly elliptic, coriaceous, base truncate to broadly cuneate, apex obtuse; adaxial laminar scales 0.5– 3 mm long, dense, imbricate, lanceolate to round, from light brown to black or mottled with lighter margin, darker at the point of attachment, ciliate, base cordate, apex acute; abaxial costal scales 0.5– 2(–3) mm long, dense, pointing forward to the apex or at right angles, lanceolate to round, from light brown to black or mottled, darker at the point of attachment, ciliate, base cordate, apex acute; intersporangial scales 1–2 mm long, absent to scattered, lanceolate, from light brown to black or mottled with lighter margin, darker at the point of attachment, ciliate, base cordate to short-stalked, apex acute. Spores with irregular deposits and ridges.

Distribution — Mexico, Guatemala, Costa Rica, Venezuela, Colombia, Ecuador, Peru, Bolivia; 1300– 4300 m. Mainly terrestrial growing in montane forest and páramos in exposed, rocky, dry areas.

Selection of other specimens examined. MEXICO, Mexico, cerca de Amealco, camino a Amecameca Tlamacas , vertiente NW del Popocatépetl, [18°38'N, 99°55'W], 3100 m, 9 Oct. 1966, Rzedowski 23279 ( NY) . Morelos, Zempoala , [19°03'N, 99°18'W], Dec. 1936, Lyonnet 1413 ( UC, US) ; Lagunas de Zempoala , [19°03'N, 99°18'W], Dec. 1946, Sánchez 465 ( US) . – GUATEMALA, Huehuetenango , Sierra Chuchumatanes between Kms 136 to 150 to San Juan Ixcoy, [15°30'N, 91°30'W], 3000–3500 m, 12–23 Jan. 1966, Molina et al. 16539 ( F, NY, US) . Quezaltenango , NW-facing forested Abies slopes of barranco of Volcán Zunil, [14°44'N, 91°27'W], 3000 m, 22 Jan. 1940, Steyermark 34911 ( F, US) . – COSTA RICA, Limón , Cordillera de Talamanca,SW foot of Cerro Kámuk, 9°16'N, 83°02'30"W, 3200–3350 m, 24 Mar. 1984, Davidse et al. 25952 ( CR, MO, NY, UC) . – COLOMBIA, Antioquia , Mun. Urrao , Inspeccion Jaiperá, vereda El Chuscal, Páramos de Frontino, Sitios Pico de Aguila-La Laguna, 6°27'N, 76°46'W, 3590–3750 m, 6 Apr. 1989, Callejas et al. 7672 ( HUA, NY) . Arauca , Sierra Nevada del Cocuy, Quebrada El Playón, Hoya S. Luis 1.5 km al NNE de la finca El Playón, [6°23'N, 72°16'W], 3540 m, 13 Mar. 1973, Cleef 9122 ( COL, NY) . Boyacá , Mun.Chisacá-San Pedro de Iguaque,SFF de Iguaque,sendero subiendo a la laguna de Iguaque, 5°41'03.2"N, 73°26'22.3"W, 3536 m, 30 Apr. 2007, Vasco et al. 765 ( HUA, NY) . Caldas , PNN los Nevados, Alto del Cisne, 4°51'17.1"N, 75°22'03.5"W, 4127 m, 7 Aug. 2006, Vasco 591 ( HUA, MO, NY) . Cauca , Páramo de Puracé, [2°15'N, 76°25'W], 3400 m, 22 Sept. 1967, Hagemann 572 ( COL, PSO) . Cundinamarca , Páramo de Sumapaz, páramo y bosque

177 alto-andino cerca de Lagunitas al S de San Juan,Alto de Chorreras; 2.5 km al N de la laguna Gobernador, [3°45'N, 74°25'W], 3780 m, 27 Jan. 1937, Cleef 8359 ( COL, MO, NY) . Magdalena, Sierra Nevada de Santa Marta , S of Cerro Icachui, Mamancanaca-Camberimena,[10°45'N, 73°58'W], 3900 m, 30 May 1905, Waston 10384 ( CR) . Meta, Páramo de Sumapaz , hoya de la quebrada Sitiales, [4°05'N, 74°12'W], 3550 m, 26 Jan. 1972, Cleef 1043 ( COL) . Nariño, Cumbal, Laguna La Bolsa , páramo, [0°54'N, 77°47'W], 3400 m, 21 Jan. 1973, Hagemann 1956 ( COL, PSO) . Norte de Santander, Páramo de Romeral , 3800–4200 m, 30 Jan. 1927, Killip 18672 ( NY, US) ; Páramo de San Turban , near Vetas, [7°12'N, 72°59'W], 3950–4160 m, 17 Jan.1927, Killip 17571 ( NY, US) . Tolima, Mun.Murillo,cerca del Nevado del Ruiz, carretera a Murillo, 4°56'N, 75°17.5'W, 3900 m, 9 May 1991, Churchill 17881 ( COL, HUA, NY) . – VENEZUELA, Mérida, entre Timote y Pico de Aguila , 3000–3500 m, 24 Nov.1984, Ortega 2339 ( MO, NY, UC) . Táchira: carretera Bailadores-La Grita,vía Páramo La Negra , 8°15'20"N, 71°52'46"W, 3000 m, 29 May 2008, Vasco et al. 808 ( NY, VEN) . Trujillo, Mun. Boconó, Laguna Larga,vía Laguna Las Parias to Laguna Eco.Páramo Motumbo, Monumento Natural Teta de Niquitao-Guirigay ,[8°56'N, 70°30'W], 3400–3600 m, 15 Sept. 2003, Stergios et al. 20424 ( NY, UC, US) . – ECUADOR, Azuay, 16 km NE of Azogues along road to Pindilíg, 2°40'S, 78°50'W, 3340 m, 19 Jan. 1985, Luteyn 11137 ( HUA, NY, UC) . Carchi, Páramo El ángel, in the pass on road El ángel-Tulcán, 0°41'N, 78°54'W, 3450 m, 15 May 1973, Holm-Nielsen et al. 5504 ( AAU, F, MO, NY, UC) . Chimborazo, Río Chimbo , 3400 m, Sept. 1905, Rimbach Rosenst. Filices ecuadoresnes, No. 7 ( B, P, S, US) . Cotopaxi, Parque Nacional Cotopaxi, 3–6 km S de la Estación Ferrocarril, 0°40'S, 78°30'W, s.d., Balslev 2655 ( AAU, GH, NY) . Imbabura, Lago San Marcos,Cayambe, 3413 m, 29 Nov.1961, Cazalet 5408 ( NY, UC, US) . Loja, Páramos de Saraguro , 10 km S of Saraguro, [3°36'S, 79°13'W], 3050 m, 2 Jan. 1979, Luteyn et al. 6663 ( NY) . Napo, Parque Nacional Llanganates, vía Salcedo-Tena, Km 45–55, cabecera del Río Anatenorio , Siete Vueltas, 0°59'S, 78°20'W, 3600 m, 17 Sept.1998, Vargas et al. 2586 ( MO, NY, QCNE, UC) . Pichincha, above Hacienda Monjas, Concepción, [0°28'S, 78°35'W], 3962 m, 29 Mar. 1951, Bell 91 ( S) ; in monte Pichincha, [0°10'S, 78°35'W], 3400 m, 1917, Mille s.n. ( GH, NY) . Tungurahua, Parroquia San José de Poaló , Loma Potrerillos, 1°05'S, 78°20'W, 3200 m, 4 Apr. 1987, Cerón 1122 ( MO, UC, US) . – PERU, Amazonas, Prov. Chachapoyas, Jalca Calla-Calla, [6°28'S, 77°43'W], 3450 m, 29 July 1991, Mostacero et al. 2634 ( MO) . Ancash, Prov.Huari, Huascarán National Park ,quebrada Pachachaca, a lateral valley of quebrada Rurichinchay, 9°23'S, 77°17'W, 3700–3860 m, 12 June 1986, Smith et al. 12543 ( MO) . Junín, Prov. Satipo, Cordillera Vilcabamba, Río Ene slope,near summit of divide, 11°39'36"S, 73°40'02"W, 3350–3400 m, 8 June 1997, Boyle et al.4262 ( NY) . Lambayeque, Prov. Ferreñafe, Laguna 0.5 mm Tembladera , distrito Incahuasi, [6°09'S, 79°18'W], 3100 m, 11 Sept. 1985, Sagástegui et al. 12780 ( GH, MO, NY, UC) . Pasco, Prov. Oxapampa, Dist. Huancabamba, sector Santa Barbara, 10°20'55"S, 75°39'08"W, 3379 m, 27 Jan.2005, Mellado 2658 ( MO) .– BOLIVIA, Cochabamba, Prov.Carrasco, Km 84 antigua carretera Cochabamba-Villa Tunari, 17°15'S, 65°43'W, 3750 m, 30 June 1996, Kessler et al. 6833 ( LPB, NY, UC) . La Paz, Prov. Murillo, bajando de la cumbre 16, 7 km hacia Unduavi, pasando Pongo, [16°19'S, 67°57'W], 3700 m, 28 Apr. 1991, Beck 18754 ( HUA, LPB, NY) .

Notes — Elaphoglossum engelii and E. muscosum are two names commonly applied to species of subsect. Muscosa in the Andes. What has been called E. muscosum in the Andes is actually E. lehmannianum (see notes under that species for a discussion on the application of this name). The Andean species E. lehmannianum and E. engelii are often confused in herbaria, but they are easy to differentiate. Elaphoglossum engelii is terrestrial with short- to long-creeping rhizomes, linear rhizome scales with many teeth longer than 0.2 mm, ciliate petiole scales, and spores with ridges and irregular deposits ( Fig. 1d ). In contrast, E. lehmannianum is typically epiphytic with compact to short-creeping rhizomes, linear rhizome scales with few short teeth, erose petiole scales, and papillate spores without ridges ( Fig. 1h ).

In E. engelii , density of the laminar scales varies, and there is a slight geographic component to this variation. Typically, the scales of both surfaces of the lamina are imbricate, although the adaxial ones tend to fall off when the leaves get older. Specimens at the southern and northern limits of the distribution ( Mexico, Guatemala, and Peru) sometimes have abaxial laminar scales more scattered and not necessarily obscuring the surfaces of the lamina.

Some specimens of E. engelii from Ecuador (e.g. Clemants 2142, NY) have unusually black rhizome scales. These might be confused with E. yatesii , a species restricted to Ecuador and Colombia. Elaphoglossum yatesii differs by thinner and longer creeping rhizomes, petiole scales that generally point forward, and papillate spores with ridges ( Fig. 1o ).

The type material from B and W has some fertile-frond fragments not attached to the sterile parts. These fragments do not belong to E. engelii or any other species of subsect. Muscosa .

Figures

Map 5 Distribution of Elaphoglossum engelii (H.Karst.) Christ.

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.

Fig. 5 Elaphoglossum engelii (H.Karst.) Christ. a. Habit; b. rhizome scale; c. larger petiole scale; d. smaller petiole scale; e. abaxial laminar surface; f. abaxial laminar scale; g. adaxial laminar surface; h. adaxial laminar scale; i. fertile lamina (all: Rentería 5387, HUA).