Elaphoglossum lehmannianum Christ

Vasco, A., 2011, Taxonomic revision of Elaphoglossum subsection Muscosa (Dryopteridaceae), Blumea 56 (2), pp. 165-202 : 185-186

8. Elaphoglossum lehmannianum Christ — Fig. 1h , 9 ; Map 9

Elaphoglossum lehmannianum Christ (1899) 81, f. 36. — Type: Lehmann 3557 (lecto B, designated here;iso BAS n.v., G n.v., US), Colombia, Cauca, wächst an Bäumen in lichten Wäldern um Popayán , 1750 m, s.d.

Elaphoglossum decipiens Hieron. (1905) 554.— Type: Lehmann 5075 (lecto B (sheet 070308),designated here;iso F, US), Colombia, Cauca:in a dense forest close to Poblazon supra urbem Popayán , 2000–2500 m, s.d.

Elaphoglossum yarumalense Hieron. (1905) 556. — Type: Lehmann 7675 (lecto B (sheet 072280), designated here; iso US (barcode 00067395), US (sheet 1067595, drawing of B lectotype with a fragment in pocket)), Colombia, Antioquia, Yarumal, habitat ad arborum truncos silvarum pratis interruptarum prope El Yarumal in parte septentrionali provinciae Antioquia, [5°56'N, 75°35'W], 1600–2200 m, s.d.

Elaphoglossum squamiferum A.Rojas (1996) 21, f. 11. — Type: Rivera 28 (holo INB), Costa Rica, Heredia, Parque Nacional Braulio Carrillo, Volcán Barva , 10°07'20"N, 84°06'00"W, 2600 m, 7 Aug. 1989.

Plants epiphytic or terrestrial. Rhizome 2 –7 mm wide, short-creeping to erect; rhizome scales 9 –12 mm long, linear, orange to dark brown, denticulate, base truncate, apex filiform, sometimes tortuous. Sterile leaves 10– 40(– 60) cm long, approximate; phyllopodia present, obscured by the petiole-base scales; petiole 3 –15(–35) cm long, 1/4–1/2 the length of the sterile leaves; larger petiole scales 3 –7 mm long, scattered to dense, spreading, linear-lanceolate, light brown, erose to laciniate, base cordate (sometimes appearing peltate because of the large auricles), apex acute; smaller petiole scales 0.5 – 2 mm long, scattered to dense, appressed, oftentimes hidden by the larger ones, oblanceolate, light brown, erose to ciliate, base cordate (sometimes appearing peltate because of the large auricles), apex acute; lamina 6 – 25 by (1.5–)2– 4(– 5.5) cm, oblanceolate to linear-lanceolate, coriaceous, base broadly cuneate to rounded, apex obtuse to acute; veins 1–2 mm apart, at an angle of 75– 80° with respect to the costa; adaxial laminar scales 1– 2 mm long, scattered to dense, lanceolate, white to light brown, ciliate, base cordate, apex acute; abaxial laminar scales 0.5 – 2 mm long, dense to absent, not imbricate, slightly elevated above the surface of the lamina, lanceolate, light brown to brown, ciliate, base stalked, apex acute; abaxial costal scales same as the laminar ones, typically pointing at right angles, slightly elevated; marginal scales same as the laminar ones. Fertile leaves longer than the sterile leaves;

185 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 (5.5 –)8– 24(–35) by 0.6 – 2.5 cm, narrowly elliptic, coriaceous, glabrescent with age, base broadly cuneate, apex obtuse; adaxial laminar scales 0.5– 3 mm long, dense, imbricate, lanceolate to round, light brown to brown, darker at the point of attachment, ciliate, base cordate, apex acute; abaxial costal scales 0.5– 3 mm long, dense to scattered, pointing at right angles, lanceolate to round, brown, ciliate, base truncate, apex acute; intersporangial scales 1.5– 4 mm long, scattered to dense, sometimes almost obscuring the sporangia, lanceolate, orange to brown, ciliate, base stalked, apex acute. Spores papillate, without ridges.

Distribution — Costa Rica, Panama, Venezuela, Colombia, Ecuador; 1150–3500 m. Epiphytic or terrestrial, in montane forests and páramos.

Selection of other specimens examined. COSTA RICA, Cartago, Cantón de Paraíso, investigation area of the CATIE, 9°33'30"N, 83°41'30"W, 2600–2700 m, 20 Dec. 1990, Bittner 180 ( CR, HUA, NY) . Heredia, Cantón de Barva, P. N. Braulio Carillo, cuenca del Tárcoles, estación Barva, entre la estación y Laguna del Barva , 10°07'20"N, 84°06'00"W, 2700–2900 m, 30 May 1997, Rojas 3546 ( CR, INB, NY) . Limón, Cantón de Talamanca, P. N. La Amistad, Tararia, sendero a Cerro Kámuk, entre Cerro Nai y Cerro Dudu, 9°13'39"N, 83°04'23"W, 2750–3100 m, 9 Nov. 1996, Rojas 3227 ( CR, INB, NY) . San José, Cerro de la Muerte, along the old oxcart trail above the interamerican Hwy, 9°33.23'N, 83°44.439'W, 2963 m, 28 Jan. 2008, Sundue et al. 1707 ( HUA, INB, NY) . – PANAMA, Chiriquí, above camp Aguacatal , E slope of Chiriquí Volcano,[8°47'N, 82°30'W], 2400–2500 m, 10–13 Mar.1911, Maxon 5297 (US). – COLOMBIA , Antioquia, Mun. Caramanta, límites con el Mun. de Támesis, sector nascimientos Quebrada El Cóndor , 5°33'27.5"N, 75°40'51.2"W, 2650 m, 16Apr.2005, Rodríguez et al.5449 ( HUA, NY) . Boyacá, Mun. Villa de Leyva , SFF de Iguaque, bosques secundarios de Robles, cerca al centro de visitantes, 5°41'43.8"N, 73°27'08.4"W, 2850 m, 29 Apr. 2007, Vasco et al. 760 ( HUA, NY) . Cauca, in parte septentrionali altiplanitiei popayanensis, 1600–1800 m, Lehmann 5139pp (B-070310, P-00577723, US-1067593, syntypes of E. decipiens ) ; Popayán,[2°26'N, 75°37'W], 1500– 1800 m, s.d., Lehmann 6945 ( B, US, syntype of E. decipiens ) ; forests of Popayán, [2°26'N, 75°37'W], 1700–2300 m, Jan. 1901, Lehmann BT1299 ( GH, NY) . Cesar, Sierra del Perijá, E of Manaure , sabana Rubia, [10°21'N, 72°56'W], 3000–3100 m, 7 Nov. 1959, Cuatrecasas 25092 ( US) . Cundinamarca, Bogotá,D. C. Región del Sumapaz,cerca a Santa Rosa, 4°11'21"N, 74°11'36"W, 3160 m, 19 Feb. 1997, Betancur et al. 7026 ( COL, HUA, NY) . Nariño, Mun. Pasto, Bosque Botana, [1°12'N, 77°16'W], 3100 m, 22 Aug. 1963, Soejarto 480 ( COL, GH) . Norte de Santander, hoya del Río Chitagá en Vega, 2880 m, 28 Nov. 1941, Cuatrecasas 13462 ( COL, US) . Quindío, ‘ Pinares’, above Salento, [4°38'N, 75°34'W], 2600 –2800 m, 2–10 Aug. 1922, Pennell 9252 ( GH, NY, US) . 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. 737 ( HUA, NY) . Santander, along road between Pamplona and Chorro Colorado vía Chiquirá, at Km 128 marker, 7°22'N, 72°38'W, 2580 m, 4 May 1983, Croat 56407 ( JAUM, UC) . Tolima, Libano-Murillo road., c. 18–24 km W of Libano, 4°45'N, 75°10'W, 2450–2770 m, 18 May 1984, Luteyn et al. 10627 A ( NY) . – VENEZUELA, Barinas, Mun. Morán, pica que va desde Buenos Aires al Páramo de Las Rosas , 9°34'N, 70°06'W, 2300–2600 m, 15 Nov. 1984, Van der Werff 7968 ( MO, UC) . Mérida, Páramo de La Negra, [8°16'N, 71°52'W], 2980 m, 13 Feb. 1939, Alston 7000 A ( COL, MO, VEN) . 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. 806, 807, 809, 812 ( NY, VEN) . Trujillo, Mun. Carache, Páramo El Jabón, 9°31'N, 70°06'W, 2900–3000 m, 28 May 1988, Rivero et al. 1802 ( NY, UC, VEN) . – ECUADOR. Azuay, E cordillera, 1–8 km N of the village of Sevilla de Oro, [2°48'S, 78°37'W], 2438–2743 m, 27 July–12 Aug. 1945, Camp 4590 ( F, NY, US) . Carchi, Valle de Maldonado, km 53 on the road Tulcán-Maldonado, steep slopes in cloud forest, 0°50'N, 78°03'W, 3150 m, 17–18 May 1973, Holm-Nielsen 5647 ( AAU, MO, NY, UC) . Cotopaxi, in pas c. m. Corazón, [0°53'S, 78°30'W], 3400 m, Sept. 1892, Sodiro s.n. ( P) . Imbabura, N slope of Volcan Cotacachi, protected forest area above town of Peribuela, 0°22'50.7"N, 78°17.787'W, 2879 m, 30 Nov. 2006, Sundue 1066 ( NY, QCA) . Loja, about 10 km SW of Loja, along road past the University towers La Violeta, [4°02'S, 79°10'W], 2950 m, 26 Apr. 1987, Van der Werff 9055 ( MO, NY, UC) . Morona-Santiago, W de la ciudad del Macas, 2°18'S, 78°07'W, 1160 m, 28 Feb.1986, Baker 6689 ( NY) . Napo, Paso de Guamaní, Río Chalpi, at the bridge on road Papallacta-Baeza , [0°22'S, 78°05'W], 2800 m, 6 May 1967, Sparre 15886 ( S) . Pichincha, Nono- Galea, [0°04'S, 78°33'W], Sept. 1899, Sodiro s.n. ( S, US); Volcán Corazón, [0°32'S, 78°39'W], Oct. 1903, Sodiro s.n. ( B) ; carretera Quito-Sto.Domingo, [0°28'S, 78°43'W], 2500 m, 24 May 1987, Van der Werff 9581 ( MO, NY, UC) . Sucumbios, Km 15–18, carretera Pun-Aguarico, 2800–2900 m, 13 Aug. 1949, Acosta-Solís 13283 ( F) .

Notes — Elaphoglossum lehmannianum can be distinguish- ed by its epiphytic habit (although it can be terrestrial above 3000 m), short-creeping to erect rhizomes, long, linear, denticulate rhizome scales, erose to laciniate (never ciliate) larger petiole scales, and papillate spores without ridges. It exhibits considerable variation in the density of the abaxial laminar scales. Some plants have their abaxial surfaces completely obscured by the scales, whereas others are subglabrous, with the laminar scales restricted to the margins and costae. The two extremes intergrade completely. Moreover, field observations of the plants in Colombia and Venezuela reveal that scaly and glabrous laminae can be found on the same rhizome (e.g. Vasco 821, NY). The plants with glabrous laminae were described from Costa Rica as E. squamiferum , here considered a synonym.

Elaphoglossum lehmannianum resembles E. muscosum , and the latter name is the one that has been commonly applied to E. lehmannianum in Costa Rica, Panama, and northern South America. The differences between these two species, especially in the scaly specimens of E. lehmannianum , are extremely subtle. Phylogenetic molecular analyses ( Vasco & Moran in prep.), however, recovered these two species in two different clades, and thus they are recognized here as two different entities. Although similar, E. lehmannianum and E. muscosum can be distinguished by scales of the rhizomes and petioles and by geographic distribution. Elaphoglossum lehmannianum has orange to dark brown denticulate rhizome scales with teeth 0.1–0.2 mm long. Elaphoglossum muscosum has reddish orange, entire rhizome scales. Occasionally these scales have teeth, but they are less than 0.1 mm long. The larger petiole scales of E. lehmannianum , although erose like those of E. muscosum , tend to have slightly longer teeth. These teeth are never longer than the scale body, as is the case in E. engelii . As to geographic distribution, E. lehmannianum occurs in Costa Rica, Panama, Venezuela, Colombia, and Ecuador ( Map 9 ). In contrast, E. muscosum occurs from Mexico to El Salvador, and in Cuba, Jamaica, Haiti, Dominican Republic, and Southern Brazil ( Map 10 ). The two species do not overlap geographically. Because of the variation within each species in scale shape, scale density, and lamina shape, it is difficult to find more characters that distinguish them; however, E. lehmannianum has a tendency toward darker rhizome scales, acute leaf apex, and smaller abaxial laminar scales that do not obscure the abaxial surface (especially in specimens of lower elevations). The abaxial laminar scales are stalked in both species, but tend to be longer-stalked, therefore appearing more elevated above the surface, in E. lehmannianum .

Another species with which E. lehmannianum is likely to be confused is E. ruficomum , endemic to Peru and north-western Bolivia. These two species can be distinguished by their rhizome and petiole scales. The rhizome scales of E. lehmannianum are less than 12 mm long, orange to dark brown, and dull; in contrast, those of E. ruficomum are typically 12 –20 mm long, maroon, and lustrous. The petiole scales of E. lehmannianum are erose to laciniate, whereas those of E. ruficomum are ciliate.

Although Lehmann’s main set of specimens is deposited at B, I designated the specimen at US as the lectotype. This specimen is more complete and has a rhizome and fertile leaves all belonging to a single plant. In contrast, the B specimen does not have a rhizome, and the collection consists of one fertile and three sterile leaves detached from each other .

See E. bellermannianum and E. engelii for a comparison with those species.

Figures

Map 9 Distribution of Elaphoglossum lehmannianum Christ.

Map 10 Distribution of Elaphoglossum muscosum (Sw.) 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.

Fig. 9 Elaphoglossum lehmannianum 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 (a, f–i: Rentería 5165; b–d: Rodríguez 5449, all HUA).