Miconia approximata Gamba & Almeda, 2014

Gamba, Diana & Almeda, Frank, 2014, Systematics of the Octopleura Clade of Miconia (Melastomataceae: Miconieae) in Tropical America, Phytotaxa 179 (1), pp. 1-174 : 51-57

6. Miconia approximata Gamba & Almeda , nom. nov. Basionym: Henriettella densiflora Standley (1929: 247) . Clidemia densiflora (Standl.) Gleason (1950: 346) . Henriettea densiflora (Standl.) Williams (1963: 565) . Type: PANAMA. Prov. Bocas del Toro: Region of Almirante, Buena Vista Camp, 11 March 1928, Cooper 575 (holotype: F!; isotypes: K-internet image!, NY!, US-internet image!, WIS-internet image!). Nec Miconia densiflora (Gardner) Naudin (1850: 245; M. pusilliflora (de Candolle) Naudin (1850: 171–172)) nec Miconia densiflora Cogniaux (1886a: 22–23 ; M. caudata (Bonpland) de Candolle (1828: 187)) .

Shrub (0.5–)1–3(–3.5) m tall, commonly forming thickets, main stem branching at around 1 m, the branches pendant and sometimes sprawling over adjacent plants, bark brownish. Upper internodes [(1–) 1.5–6.8 cm long] and cauline nodes terete, nodal line absent. Indumentum on branchlets, petioles, primary and secondary leaf veins abaxially, bracts, hypanthia, calyx lobes abaxially, and exterior calyx teeth densely composed of brown elongate slightly roughened trichomes 0.3–0.7 mm long, each trichome clavate and somewhat thickened, sparsely intermixed with, or occasionally completely replaced by a brown understory of dendritic trichomes 0.05–0.15 mm long with short to moderately long thin-walled (flattened) arms. Leaves of each pair somewhat anisophyllous in size, some pairs isophyllous; the short terete petioles 0.3–0.8(–1) cm long, brownish, occasionally pink; larger blades 12–22 × 4.8–9.3 cm, smaller blades (4.5–)5–11.2 × 2–4.5 cm, narrowly elliptic to elliptic or slightly ellipticobovate, the base acute to obtuse, the margin entire to obscurely undulate-serrulate, the apex gradually acuminate to long-acuminate, firm-membranaceous; mature leaves with adaxial surface glabrescent, in young leaves copiously covered with the general dendritic trichomes, the primary, secondary, tertiary and higher order veins glabrous; abaxial surface essentially glabrous except for a few glands coming on the venules, the tertiary veins moderately covered with the general dendritic trichomes, along with the higher order veins, copiously to sparsely beset with resinous sessile to short-stalked glands 0.05 mm long with thin-walled short to elongate heads, frequently intermixed with and occasionally replaced by white or brown-translucent furrowed sessile glands of the same length; 5-nerved, including the tenuous marginals, areolae 0.5–1 mm, adaxially the primary, secondary, tertiary and higher order veins flat, abaxially the primary and secondary veins elevated and terete toward the blade base, flat toward the apex, the tertiary and higher order veins flat. Inflorescences a congested axillary fasciculate glomerule <1cm long, sessile, unbranched, paired or solitary among upper leafy nodes, seeming cauliflorous on defoliated nodes; bracts 1.5–3 × 0.5–1 mm, triangular to oblong, persistent to tardily deciduous in fruit. Flowers 4- merous, sessile. Hypanthia at anthesis 2.4–2.5 × 1.4–1.5(–2.2) mm, free portion of hypanthium 1 mm long, suburceolate to more or less globose, 8-ribbed, pinkish to green, the ribs frequently concealed by the dense general indumentum, ridged on the inner surface, minutely glandular, the glands sessile and rounded, the torus densely glandular with minute sessile to short-stalked rounded glands adaxially. Calyx open in bud and persistent in fruit, green; tube 0.2–0.25(–0.3) mm long, adaxially with the same vestiture as the torus, abaxially with the same indumentum as the hypanthium; lobes 1–1.5 × 1–1.5 mm, rounded-ovate to triangular, the margin entire, the apex rounded-obtuse, adaxially glabrescent, reflexed at anthesis; exterior teeth 0.5–1 mm long, subulate, inserted on the apical half of the calyx lobes and slightly projecting beyond them. Petals 2.5–2.6 × 0.9–1 mm, ovate-oblong to linear-oblong, the margin entire, the apex rounded-obtuse, white to pale-pink, glabrous on both surfaces, reflexed at anthesis. Stamens 8; filaments 1–1.5 × 0.2–0.25 mm, white, glabrous; anther thecae 1.4–1.5 × 0.25–0.33 mm, linear-oblong, more or less truncate to slightly emarginate at the apex, opening by one dorsally inclined pore 0.07–0.1 mm in diameter, white or pale pink, becoming brown with age; connective white, also becoming brown with age, its prolongation and appendage 0.35–0.45(–0.5) mm long, the appendage deltoid-lanceolate, bluntly acute to obtuse at the apex, copiously gland-edged and covered with stalked or subsessile glandular trichomes to 0.15 mm long, denser dorsally and present throughout the connective. Ovary 4-locular, 3/4 inferior, 1.7–1.8 mm long at anthesis, the apical collar (0.3–)0.4–0.5 × 0.25 mm, conic, copiously glandular-puberulent; style 2.5–3.5 mm long, more or less parallel sided (i.e. subterete), white, glabrous or with few glands at the very base; stigma truncate to capitellate. Berries 2–2.5 × 5–6 mm when dry, globose-oblate, bright orange when ripe, the hypanthium indumentum somewhat persistent at maturity. Seeds 0.52–0.57 × 0.31–0.4 mm, pyramidal, yellow-brown; lateral

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 51 symmetrical plane triangular, the highest point near the central part of the seed, with a foot-like projection at the micropylar end; antiraphal symmetrical plane suboblong; raphal zone suboblong, ca. 60–80% the length of the seed; multicellular sculpture rugose throughout the seed; individual cells elongate, anticlinal boundaries channeled, undulate, with Ω- and U-type patterns; periclinal walls convex, low-domed to nearly flat, microrelief striate. Chromosome number: n=17.

Additional specimens studied:— BELIZE. Middlesex: 61 m, 19 July 1929, Schipp 264 (F, US). COLOMBIA. Chocó: (Bahía Solano), Mecana, Jardín Botánico del Pacífico, Ciudad Mutis, Sendero Jaguar (Trail # 3), 6.26435°N, - 77.37045°W, 120 m, 4 February 2012, Almedaet al. 10459 ( CAS, CHOCO, COL); (Bahía Solano), P.N.N. Ensenada de Utría, Camino entre Punta Diego y Caida Cocalito, 6°21’N, 76°26’W, 0–100 m, 18 April 1990, Espina et al. 3644 ( CHOCO, MO); ca 1 mile NE of Camp Curiche, ca 3 miles Eof Curiche, 21 May 1967, Duke & Idobro 11323 ( US); Nridge of Alto Buey, above Dos Bocas del Río Mutatá, tributary of Río El Valle, ESE of El Valle, 200–500 m, 8 August 1976, Gentry & Fallen 17410 ( MO, US); Hills behind Bahía Solano (Puerto Mutis), 0–250 m, 5 January 1973, Gentry & Forero 7204 ( MO); Carretera Quibdó-Guayabal, 40 m, 24 April 1975, Forero et al. 1154 ( COL, MO); Headwaters Río Salaqui, Hydro Camp 3, 335 m, 21 June 1968, Duke 15774 ( MO); (Bahía Solano), El Valle, Trocha El Valle-Boro Boro, 6°21’N, 76°26’W, 17 April 1989, Espina et al. 2674 ( CHOCO, MO). COSTA RICA. Alajuela: (Upala), Dos Ríos, 5 km al Sde Brasilia, margen derecho del Río Pizote, 10°55’N, 85°20’W, 500 m, 29 October 1987, Herrera 986 ( CAS, CR, F, MO); (San Carlos), Entre Dulce Nombre Norte y Dulce Nombre Sur, 530 m, 23 June 1966, Jiménez 4068 (F); 17–20 km NNW of San Ramón by road on way to San Lorenzo, 4 to 7 km Nof Balsa, 10°13’N, 84°32’W, ca. 750 m, 24 April 1983, Liesner & Judziewicz 14680 ( CAS); R.B. Monteverde, Río Peñas Blancas, Laguna de Poco Sol, 10°21’N, 84°40’W, 700 m, 14 December 1989, Bello 1633 ( CAS, CR); Near Artezalea and Methodist Rural Center, about 8 km NE of Villa Quesada , 550 m, 16 February 1966, Molina et al. 17146 (F, US); (San Carlos), Cuenca del San Carlos, Boca Tapada, Entre Santa Rita y la Curena, 10°41’25"N, 84°11’0"W, 100 m, 24 February 1999, Rodríguez & Ramírez 4492 ( INB, MO). Cartago: Valle Escondido, 700 m, 2 April 1966, Schnell 672 ( CR, F). Heredia: Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, Along Far Loop Trail, about 1300 m line, 100 m, 12 July 1980, Grayum 2966 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, Sboundery-E end, 100 m, 2 May 1981, Folsom 9991 ( CAS, US); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, 2200 Holdridge Trail, 100 m, 18 February 1981, Folsom 9020 ( CAS, F); La Selva, Near Puerto Viejo, Loop Trail, 90 m, 21 May 1972, Opler 813 (F, US); (Sarapiquí), Tirimbina, Istarú Farm, 220 m, 17 January 1970, Lent 1872 (F); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, Sboundary, Eof Central Trail, 100 m, 20 February 1981, Folsom 9047 ( CAS, F); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, in SW quarter of new property, 100 m, 15 May 1982, Hammel 12238 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, In forest on ridge Wof Holdridge Trail 2200 m S, 100 m, 19 May 1982, Hammel 12350 ( CAS); Estación Biologica La Selva of the Organization for Tropical Studies, Rain forest off of the Oriental Trail, 100 m, 27 February 1986, Almeda et al. 5096 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, In forest Eof Far Loop Trail, 1100 m line, 100 m, 22 May 1980, Hammel 8750 ( CAS, F); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, Central Trail, 2600 m, 100 m, 10 February 1981, Folsom 8870 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, In forest on slopes between Wboundary of La Selva proper and trail through SE section of new property at 2700 m S, 100 m, 27 May 1982, Hammel 12553 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, Pasos Perdides, 100 m, 31 March 1983, Chacón 649 ( CAS); P.N. Braulio Carrillo, Estación Penal Magsasay, Sendero Principal, 10°24’10"N, 84°3’30"W, 200 m, 20 October 1990, Umaña & Chacón 448 ( CAS, CR); (Heredia), Sarapiquí, La Virgen, P.N. Braulio Carrillo, 200 m Edel Puesto La Ceiba, sobre la Fila y rivera del Río Peje, 10°20’N, 84°5’W, 400 m, 15 October 1988, Ballestero 9 ( CAS, CR, MO); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, 3600 NS × 1350 EW m line, 100 m, 11 December 1982, McDowell 1104 ( CAS, F); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, 2800 m, Central Trail, along path, 100 m, 29 August 1981, Smith 143 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, Central Trail, 1400 m, 100 m, 6 February 1981, Folsom 8794 ( CAS); (Tirimbina),

52 Phytotaxa 179 (1) © 2014 Magnolia Press

GAMBA & ALMEDA

213 m, 29 May 1971, Proctor 32122 (F, GH, MO); La Selva, Near Puerto Viejo, Loop Trail, Shrub K 10, 90 m, 16 April 1972, Opler 703 (F); La Selva de Sarapiquí , 500 m, 31 October 1965, Schnell 330 ( CR, F, US); Río Sarapiquí above Cariblanco, 15 September 1965, Schnell 99 ( CR, F); La Selva de Sarapiquí , 200 m, 4 February 1966, Schnell 415 ( CR, US); Nof Puerto Viejo , 12 km to ferry, over ferry, 6 km along road , 100 m, 3 February 1983, Garwood et al. 990 ( MO); Sof Puerto Viejo , 2 km Sof Magsasay Penal Colony, Wof the road , 200 m, 5 February 1983, Garwood et al. 1117 ( MO); (Sarapiquí), No protegida, Cuenca del Sarapiquí, Finca Aracuak, Río Frío , Senderos Pilón y Oropendula , 10°13’0"N , - 85°55’0.0001"W , 200–300 m, 1 August 2002, Kriebel 709 ( INB, MO); (Sarapiquí), OET La Selva, Sendero SSA , 950 m, 2 June 2004, Aguilar 8751 ( MO); (Sarapiquí), P.N. Braulio Carrillo, Cuenca del Sarapiquí, Estación El Ceibo, camino al Río Peje , 10°19’37.697"N , - 84°4’39.669"W , 500–546 m, 12 March 2003, González 3076 ( INB, MO). Limón: (Limón), El Progreso, Cordillera de Talamanca. entre Cerro Muchilla y Cerro Avioneta, Fila Matama, siguiendo la fila y los flancos, 9°47’40"N, 83°6’30"W , 850 m, 8 April 1989, Herrera & Madrigal 2555 ( INB, MO); 7 km SW of Bribrí , 100–250 m, 4 May 1983, Gómez et al. 20432 ( CAS, CR); 7 km SW of Bribrí , 100–250 m, 4 May 1983, Gómez et al. 20432 ( CAS, CR); P.N. Braulio Carrillo, Puesto Quebrada González, desde los alrededores de la casa hacia el bosque junto a la Quebrada González, 10°9’N, 85°56’W , 500–600 m, 18 May 1988, Umaña & Chavarría 244 ( CAS, CR); Cerro Coronel, Eof Laguna Danto, 10°41’N, 83°38’W , 20–170 m, 16 January 1986, Stevens 23655 ( CAS, MO); R.B. Hitoy Cerere, From Río Cerere to Cerro Bobókara , 100–600 m, 26 February 1991, Almeda et al. 6824 ( CAS); (Pococí), R.N.F.S. Barra del Colorado, Llanura de Tortuguero, Sardinas , 10°38’38"N, 83°44’10"W , 15–20 m, 12 December 1992, Araya 176 ( CAS, CR); P.N. Tortuguero, Estación Cuatro Esquinas , 400 m al Ede la casa-estación, 10°31’N, 83°30’W, 2 m, 28 November 1987, Robles 1378 ( CAS, CR, MO); Lomas de Sierpe, 5 km NE de La Aurora, Guápiles, Límite S P.N, Tortuguero , junto al Río Sierpe , 10°22’N, 83°31’W , 30 m, 5 December 1988, Robles 2215 ( CAS, CR, F, MO); (Tortuguero), Suretka , 200 m from RECOPE oil drilling platform, 9°34’N, 82°56’W, 27 June 1984, Barringer et al. 3573 ( CAS, CR, F); Hacienda Tapezco-Hacienda La Suerte , 29 air km Wof Tortuguero, 10°30’N, 83°47’W , 40 m, 16 August 1979, Davidson & Donahue 8401 ( CAS); P.N. Tortuguero, Estación Agua Fría, 6 km al SE (aprox), sobre los Cerros Azules , 10°27’N, 83°34’W , 70 m, 19 January 1988, Robles 1532 ( CAS, CR, MO); ( Puerto Viejo de Limón ) , 100 m, 24 March 1966, Schnell 603 ( CR, F, US); (Pococí), P.N. Braulio Carrillo, Cuenca del Sarapiquí, Estación Quebrada González, Sendero Las Palmas , 10°9’48"N, 83°56’20"W , 500 m, 11 September 1996, Rodríguez et al. 1508 (F, INB, MO); (Pococí), P.N. Braulio Carrillo, Cuenca del Sarapiquí, Estación Quebrada González, Sendero Las Palmas , 10°9’48"N, 83°56’20"W , 500 m, 11 September 1996, Rodríguez et al. 1506 ( INB, MO); (Talamanca), San Miguel, Cuenca del Sixaola, Bosque sometidos a explotación de madera entre San Miguel y Gandoca. siguiendo el curso de quebrada innominada (aparentemente Mile Creek ), 9°34’30"N, 82°40’0"W , 30–100 m, 21 January 1997, González et al. 1708 (F, INB); (Barra de Colorado), 0 m, 2 March 1966, Schnell 550 ( CR, F, US); (Pococí), P.N. Braulio Carrillo, Cuenca del Sarapiquí , Estación Quebrada González , Sendero Las Palmas , 10°9’50"N, 83°56’30"W , 550 m, 3 April 1997, Rodríguez et al. 2099 ( INB, MO); (Matina), Z.P. Pacuare, Cuenca del Matina, Sendero Topoyiyo , 9°59’25"N, 83°26’30"W , 300 m, 23 June 1999, Mora-Castro 375 ( INB, MO). Puntarenas: (Golfito), P.N. Corcovado, Valle del Coto Colorado, Alrededores de la Estación Esquinas, 8°46’0"N, 83°15’0"W , 100 m, 19 April 1993, Aguilar 1718 ( CAS, CR, MO); (Golfito), R.N.V.S. Golfito, 3–6 km from Zona Franca near Golfito along the road to La Esquina, 8°38’45"N, 83°10’44"W , 100–120 m, 30 June 1994, Kress & Runk 94-4651 ( US); Eastern Osa Peninsula, More than 500 m from edge of primary moist tropical forest, 8.69539°’N, -83.58246°’W, 25 July 2001, Mayfield 911-1647-1487 ( MO); Osa Peninsula, Along road from Panamerican Hwy at Piedras Blancas to Rincón , 3.7 mi Wof Panamerican Hwy, 8°46’N, 83°18’W , 90–105 m, 16 September 1987, Croat 67666 ( CAS); Marenco Biological Station, On the NW coast of the Osa Peninsula, Unnamed trail leading due Sof the Rainforest trail Tacotal Trail , 60–120 m, 14 March 1986, Almeda et al. 5580 ( CAS); Marenco Biological Station, On the NW coast of the Osa Peninsula, NE sector of the Sendero Camino Publico , 60–120 m, 13 March 1986, Almedaet al. 5554 ( CAS); (Golfito),, P.N. Corcovado, Península de Osa, Manglar del Río Corcovado, Estación Sirena, Playa Llorona , 8°31’35"N, 83°39’10"W , 0 m, 15 May 1995, Churchill a 174 ( CAS, CR); (Sirena), Los Patos, P.N. Corcovado, Sirena, Los Patos, Trail to Río Pavon , 8°28’N, 83°35’W , 1–50 m, 26 June 1989, Kernan 1184 ( CAS, CR, MO); (Golfito), P.N. Corcovado, Península de Osa, Estación Los Patos , alrededores, 8°34’0"N, 83°31’0"W , 200 m, 1 September 1993, Aguilar 2143 ( CAS, CR, MO); (Rincón de Osa), Osa Peninsula, Along the Camino de Altura, 2 to 5 miles Wof Rincón de Osa, Osa Peninsula, Trail to airfield from Mile 4 at about 500 ft, 122–305 m, 9 August 1967, Raven 21670 (F, MO); (Rincón de Osa), Osa Peninsula, Region to W of airstrip , 40–100 m, 21 July 1974, Utley & Utley 1099 ( CR, F); (Rincón de Osa), Osa

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 53 Peninsula, Trail from Rincón de Osa to Rancho Quemado, 13 November 1972, Kennedy 1952 (F, MO); Top of”Ridge Road", Near Rincón de Osa, 13 November 1972, Kennedy 1971 (MO); 4–6 km from Golfito on road to TV tower and tele-communications center, 400 m, 14 May 1976, Utley & Utley 4873 (F); (Osa), No protegida, Ballena, Playa Hermosa, Dominical, 9°11’30"N, -83°46’23.0001", 0–100 m, 28 June 2002, Kriebel 347 (INB, MO); (Golfito), Península de Osa, Estero guerra de Sierpe, 8°46’0"N, 83°35’10"W, 200 m, 5 June 1995, Rodríguez 768 (INB, MO); (Osa), Rancho Quemado, 8°43’10"N, 83°34’50"W, 150 m, 13 January 1991, Lobo 48 (INB, MO); (Sirena), P.N. Corcovado, Los Patos Forest, 8°27–30’N, 83°33–38’W, 150–200 m, 18 February 1988, Kernan 183 (CR, MO); (Golfito), P.N. Piedras Blancas, Serranías de Golfito, Cerros frente a la Estación, 8°41’10"N, - 83°13’20"W, 200 m, 8 June 2000, Acosta et al. 1609 (INB, MO). San José: Z.P. La Cangreja, Santa Rosa de Puriscal, Bosque primario en las márgenes del Río Negro, SE de la Fila de La Cangreja, 9°42’50"N, 84°23’30"W, 400 m, 8 May 1993, Morales 1451 (CAS, CR); (Vasquez de Coronado), P.N. Braulio Carrillo, Along Hwy San José to Siquerres Hwy, along trail to Río Sucio, site of the Old Carillo Station, 10°9’50"N, 83°57’10"W, 600–700 m, 30 August 1996, Croat 78780 (CAS); (Dota), Zona protectora Cerro Nara, Faldas, 9°29’40"N, 84°0’50"W, 800–900 m, 4 December 1997, Estrada et al. 1400 (CR, F, MO); (Tarrazú), Faldas del cerro Nara, ca Esquipulas, límite Quepos (Puntarenas) y Tarrazú, 9°29’N, 84°3’W, 350–400 m, 11 July 1987, Gómez-Laurito et al. 11590 (CR, F); (Mastatal de Puriscal), 400 m, 4 December 1986, Zamora & Jiménez. 1314 (CR, MO); P.N. Carrillo, Bosque cerca del Río Sucio, Estación Carrillo, 25 April 1984, Sánchez et al. 497 (MO). ECUADOR. Napo-Pastaza: (Mera), 1100 m, 25 November 1955, Asplund 18620 (NY, US). Pastaza: Alongroadto Río Anzu, 17.1 km Nof Mera, 3.5 km Nof Río Anzu, trail W into montains, 1°23’26"S, 78°3’19"W, 1238–1400 m, 6 May 2003, Croat et al. 88712 (CAS). GUATEMALA. Alta Vera Paz: Near The Finca Sepacuite, 23 April 1902, Cook & Griggs 747 (US). Izabal: (Puerto Mendez), Puerto Mendez, 18 km on Poquela Road, 5 June 1970, Contreras 9959 (CAS, MO, UTD); (Puerto Mendez), Puerto Mendez, 7 km on Toquela Road, 8 September 1970, Contreras 10226 (CAS, GH, MO); Montaña del Mico, Between Virginia and Lago Izabal, 50 m, 5 April 1940, Steyermark 38865 (NY); Puerto Mendez, On Toquela River Road, 5 km from the village, 6 September 1969, Contreras 9081 (CAS, GH, MO); Along trail beginning from mile 33.23 between Darthmouth and Morales towards Lago Izabal, Montaña del Mico, 35–150 m, 7 April 1940, Steyermark 39042 (F); Cerro San Gil, Along Río Frío, 75 m, 17 December 1941, Steyermark 39946 (F); (Cadenas), On Toquela River Road, 5 km from the village, 6 September 1969, Contreras 9077 (GH, MO, US). HONDURAS. Atlántida: Southern boundary of Lancetilla Valley, On ridge separating Lancetilla watershed from the watershed to the S, around San Francisco, 15°41’30"N, 87°28’0"W, 380–420 m, 8 November 1988, MacDougal et al. 3377 (CAS, MO); Lancetilla mountain, 100 m, 8 April 1970, Molina & Molina 25610 (F, NY, US); Lancetilla Valley, Near Tela, 20–600 m, 6 December 1927, Standley 52936 (F); Lancetilla Valley, near Tela, 20–600 m, 6 December 1927, Standley 56796 (F); Lancetilla Valley, near Tela, 20–600 m, 6 December 1927, Standley 53107 (F). Gracias a Dios: Camp Tiro, 2 mi NW of Bulebar on third northern branch of Quebrada Tiro, tributary of Río Plátano, Along walk 1 km of camp, 15°43’N, 84°50’W, ca. 61 m, 22 March 1981, Saunders 1087 (CAS, F, UNM); Between Rancho Chico and Cockscomb, Monkey River, 31 March 1943, Gentle 4373 (CAS, UTD). NICARAGUA. Atlántico Norte: (Bonanza), R. de Bosawas, Comunidad de Musawas, SW de Musawas, 14°6’59"N, 84°43’49"W, 50–200 m, 20 September 2003, Coronado & Gurdian 278 (CAS). Bluefields: 1.4 km Nof base camp, Base Camp 3.6 km SE Cerro San Isidro, Río Kama, Río Escondido, 12°05–15’N, 83–84°45–20’W, 0–65 m, 6 March 1966, Proctor et al. 27001 (CAS, US). Matagalpa: (Río Blanco), R.N. Cerro Musún, 12°58’N, 85°13’W, 500–1400 m, 15 July 2000, Rueda & Caballero 14171 (CAS, MO). Río San Juan: (El Castillo), R. Indio-Maíz, a lo largo del caño el Pavon, a 3 km de su desembocadura en el Río Bartola, 11°1’N, 84°16’W, 31 December 1996, Rueda et al. 5136 (CAS, MO); Río Pigibaye, 18 February 1995, Rueda et al. 3197 (CAS); (El Castillo), R. Indio-Maíz, Cerro El Diablo, 11°1’N, 84°13’W, 100–200 m, 7 December 1998, Rueda et al. 9550 (CAS, MO); (El Castillo), R. Indio-Maíz, 3 km al Nde la desembocadura del Caño Chontaleño , 11°5’N, 84°15’W, 24 February 1997, Rueda et al. 6308 (MO). Zelaya: Caño Zamora on Río Rama, 11°57’N, 84°16’W, 10 m, 16 May 1978, Stevens 8828 (CAS); Caño El Hormiguero, On E slope of El Hormiguero, 13°46’N, 84°59’W, 750–800 m, 17 March 1980, Pipoly 6107 (CAS); Caño Costa Riquita, ca. 1.8 km SW of Colonia Naciones Unidas, above (S of) road between Colonia Nueva León and Colonia Naciones Unidas, 11°43’N, 84°18’W, 150–180 m, 6 November 1977, Stevens 5080 (CAS); (Siuna), R. Bosawas, Cerro Saslaya, 13°47’N, 84°59’W, 300–400 m, 10 April 1999, Rueda et al. 10567 (CAS, MO); (Nueva Guinea), R. Indio-Maíz, 11°25’N, 84°13’W, 200–300 m, 5 January 1999, Rueda et al. 9829 (CAS, MO); Costado SW de Cerro El Hormiguero, 13°44’10"N, 84°59’50"W, 900–1000 m, 18 April 1979, Grijalva 463 (MO); Along road to Colonia Yolawa, Colonia La Esperanza of Nueva

54 Phytotaxa 179 (1) © 2014 Magnolia Press

GAMBA & ALMEDA

Guinea , inmediatly upriver from bridge over Cano Sardina , 180–200 m, 11 February 1978, Vincelli 144 ( MO) . PANAMA. Bocas del Toro: Between Quebrada Logrom and Cerro Bonyc, near Río Terebe , 91–274 m, 13 April 1968, Kirkbride & Duke 638 ( MO) . Coclé: P.N. Omar Torrijos, Road to La Rica. ca. 0.5–1.5 km beyond pass on Atlantic slope, 8°40.76’N, 80°35.84’W, 700–800 m, 12 February 2005, Penneys & Blanco 1755 ( CAS, US) ; Alto Calvario, Area surrounding Rivera Sawmill , 7 km Nof El Cope, continental divide, 700–850 m, 25 November 1977, Folsom & Collins 6536 ( CAS, MO) ; Alto Calvario, 7 km Nof El Copé de Veraquas, lumber camp at circa 900 m, Continental Divide at 1300 m, Circa Rivera sawmill, 900–1300 m, 11 January 1977, Folsom 1213 ( CAS, MO) ; Near continental divide along lumbering road 8.4 km above El Cope (1 km beyond sawmill), 900 m, 19 January 1978, Hammel 964 ( CAS, MO) ; Alto Calvario, Forest above sawmill, on continetal divide, 5.2 mi above El Cope, 930 m, 6 December 1979, Croat 49187 ( CAS) ; Small patch of forest on continental divide ridge along road to Coclecito , between La Pintada and Cascajal, 488 m, 6 March 1982, Hammel & Kress 11283 ( CAS) . Colón: Santa Rita Ridge, along road, ca. 1 mi from Boyd-Roosevelt hwy, 9 July 1971, Croat 15302 ( MO, NY, US) ; Santa Rita Ridge, along road, ca 1 mi from Boyd-Roosevelt Hwy , 9 July 1971, Croat 15332 (F, MO) ; Río Guanche, ca 2.5 km upriver from bridge on road to Portobelo, Along small quebrada to the Sof the river, 10–100 m, 14 December 1974, Mori & Kallunki 3694 ( MO) ; In forest at Loma La Toba, Off Hwy , 5–8 (82), ca 4 km NE of Pina, 0–20 m, 17 September 1974, Mori & Kallunki 1982 ( MO) ; Along Río Guanche ca 3–5 mi inland, 10–100 m, 3 August 1974, Croat 26160 ( MO) . Comarca de San Blas: El Llano-Carti Road, 19 km from Interamerican Hwy, Ridge down to creek on Atlantic side, 9°19’N, 78°55’W, 130–350 m, 4 September 1984, de Nevers & Herman 3836 ( CAS, MO) ; Cerro Habú, Trail from Río Sidro , 9°23’N, 78°49’W, 427–762 m, 18 December 1980, Sytsma et al. 2663 ( CAS, MO) ; Cerro Habú, Vicinity of peak, 9°23’N, 78°49’W, 762 m, 19 December 1980, Sytsma et al. 2741 ( CAS, MO) . Darién: 10 km NE of Jaqué, slopes of Río Tabuelitas above Birogueirá, Indian village on Río Jaqué below mouth of Río Pavarandó. , 122 m, 30 January 1981, Sytsma & D'Arcy 3302 ( CAS, MO) ; Rancho Frio , 7°58’N, 77°42’W, 600 m, 9 August 1986, McDonagh et al. 597 ( MO); (Mamey) , 6 March 1982, Whitefoord & Eddy 383 ( CAS, MO) ; Serranía de Pirre, along ascent of Serranía de Pirre above Cana Gold Mine between Río Cana and Río Ascucha Ruido, 600–1000 m, 27 July 1970, Croat 37690 ( CAS, MO) ; Serranía del Sapo, Area surrounding Río Chado , 7°40’N, 78°10’W, 350 m, 2 January 1981, Hahn 282 ( CAS, MO) ; 0.5 to 1.5 mi Eof Manene , 21 December 1980, Hartman 12077 ( CAS, MO); (Chipigana) , Canal-Cuasi trail (camp 2), 610 m, 9 March 1940, Terry & Terry 1424 (F, MO) ; Sof El Real, on slopes of Cerro Pirre about 3 miles Eof Dos Bocas of the Río Pirre , 500–1000 m, 26 September 1969, Foster & Kennedy 1267 ( MO, US) ; Río Balsa , between Río Areti and Nanane, 14 September 1966, Duke 8766 ( MO, US) ; Elfin Forest, Cerro Campamento ( Sof Cerro Pirre ), 20 March 1968, Duke 15647 ( MO) . Panamá: Cerro Jefe, Along trail off Pacora Rd , 750 m, 21 February 1988, Almeda et al. 5845 ( CAS) ; Road past Altos de Pacora, 3–3.5 mi NE of Altos de Pacora, 7.8–8.2 mi above Panamerican Hwy , 11.1–11.6 mi beyond Lago Cerro Azul , 9°15’N, 79°25’W, 700–750 m, 19 June 1988, Croat 68622 ( CAS, MO) ; Along trail to Cerro Brewster from Río Pacora valley , 9°20’N, 79°15’W, 670 m, 19 November 1985, McPherson 7515 ( MO) ; Cerro Jefe, Forest on trail around summit tower, Trail around summit tower, 9°15’4"N, 79°30’4"W, 900 m, 1 March 1987, McPherson 10595 ( MO) ; Cerro Jefe, 1.5 mi down right turnoff 6.7 mi past Goofy Lake , 700 m, 27 December 1980, Sytsma et al. 2866 ( CAS, MO) ; Cerro Pelón, About 23 km from the Interamerican Hwy off of a dirt road toward Cerro Jefe , 675 m, 26 February 1996, Almedaet al. 7692 ( CAS) ; P.N. Chagres, Sendero ”El Cantar", Cerro Azul , 5 December 1991, Carrasquilla et al. 3334 ( CAS, PMA) ; Altos de Pacora , 9°16’N, 79°20’W, ca. 700 m, 26 February 1996, Galdames et al. 2545 ( CAS, F) ; Cerro Jefe , 945 m, 5 January 1972, Dwyer & Gentry 9462 ( MO) . Veraguas: (Isla de Coiba), P.N. Coiba, Subida por Río Escondido , 17NMU1341, 40 m, 11 March 1996, Castroviejo & Velayos 8145MV ( CAS); ( Isla de Coiba ) , Distrito de Montijo, Nde la isla, Sendero Yuma, 7°35’N, 81°43’W, 30 m, 22 February 1995, Galdames et al. 2166 ( CAS, PMA); ( Isla de Coiba ) , Siguiendo la Quebrada del Río Escondido, 1341, 19 November 1994, Espinosa et al. MB535 ( CAS); ( Isla de Coiba ) , Río de la Boa, márgenes del río, 23 November 1994, Espinosa et al. MB649 ( CAS); ( Isla de Coiba ) , Ensenada al Nde Punta Cirilo, playa de las Boyas, 17NMU1139, 25 m, 27 November 1994, J. Cuadras et al. 8069MV ( CAS); ( Isla de Coiba ) , Distrito de Montijo, Playa Rosario, 8 March 1996, Galdames et al. 2580 ( CAS, F, MO, PMA); ( Isla de Coiba ) , Distrito de Montijo, Punta Cirilo, 17 March 1996, Galdames et al. 2819 ( CAS, PMA); ( Isla de Coiba ) , Distrito de Montijo, Playa Rosario, 8 March 1996, Galdames et al. 2575 ( CAS, PMA); ( Isla de Coiba ) , Distrito de Montijo, Río Escondido, corriente arriba hasta la caida de agua, 11 March 1996, Galdames et al. 2686 ( CAS, PMA); ( Isla de Coiba ) , Distrito de Montijo, Desde Playa Hermosa hacia el borde superior de la falla geológica, 18 March 1996, Galdames et al. 2857 ( CAS, PMA); ( Isla de Coiba ) , Playa Rosario, northern tip of Coiba Island , 26 August 1970, Foster 1611 (F,

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 55 MO , US); ( Isla de Coiba ), Penal Colony , 8 July 1962, Dwyer 2401 ( MO, US); ( Isla de Coiba ), 28 July 1962, Dwyer 2330 ( MO). Hills Nof Frijoles , Canal Zone, 19 December 1923, Standley 27554 ( US); Nof Punta Guayabo Grande, NW of Ensenada El Guayabo, 7°24’N, 78°7’W, 0–200 m, 25 January 1982, Knapp & Mallet 3151 ( MO) .

Illustration:— None found.

Common names and documented uses:— Honduras: “sirin” (Standley 52936 and Gentle 4373).

Habitat, distribution and ecology:— This is a locally common species in the understory of primary rainforests where it also colonizes disturbed sites throughout Central America from Belize and Guatemala (except El Salvador), to southern Panama, and extending to Colombia and Ecuador ( Fig. 12 ), at 0–1400(–1759) m. In the TROPICOS database there are two Ecuadorian specimens that were not examined in this study. One is from Pichincha (Øllgaard 37630, AAU), and the other from Esmeraldas (Holm-Nielsen 25472, AAU). Miconia approximata is one of the most widespread species in the Octopleura clade. It appears to be especially common in Central America.

White-ruffed Manakins ( Corapipo altera ) have been reported to feed on M. approximata berries in a wet forest of northeastern Costa Rica ( Boyle 2010). Other fruit-eating birds of the tropical forest understory (at Estación Biológica La Selva, Costa Rica) have been reported to consume M. approximata berries, including Chlorothraupis carmioli , Corapipo leucorrhoa , Euphonia gouldi , Hylocichla mustelina , Mionectes oleaginous , and Pipra mentalis ( Loiselle & Blake 1999, 2000).

Phenology:— Collected in flower and fruit throughout the year.

Etymology:— The specific epithet refers to the close proximity of the flowers in the congested glomerules that characterize this species.

Discussion:— This species is readily distinguished by its brownish thickened-clavate elongate trichomes mixed with a furfuraceous indumentum of dendritic trichomes, short and pubescent petioles, narrowly elliptic to elliptic leaves, and sessile congested fasciculate glomerules. Its closest relatives include species that share the

56 Phytotaxa 179 (1) © 2014 Magnolia Press

GAMBA & ALMEDA

congested fasciculate glomerules ( M. chocoensis , M. quadridomius , and M. veraguensis ) and also have elongate roughened trichomes (but different sizes and colors). Wurdack (1981) provides a detailed summary of similarities and differences between M. approximata and M. quadridomius (which he treated as Clidemia densiflora and C. cuatrecasasii respectively). Miconia veraguensis , a new species from Veraguas, differs primarily in berry dimensions which are bigger and modal differences in leaf shape.

Miconia evanescens , which is also closely related and somewhat similar to M. approximata , has been confused in the past with this species, probably due to the poorly developed inflorescences in both, especially in Colombian material, where M. approximata only occurs in Chocó, and M. evanescens in Nariño. Although both species present abaxial foliar venules densely resinous-glandular, in M. evanescens the indumentum is whitish (vs. brownish), the hypanthium is green-whitish (vs. pinkisk green), and the ovary is completely superior (free from the hypanthium vs. 3/4 inferior).

Quite a few populations from Costa Rica (Limón, Heredia / San José), Panama (Bocas del Toro, Coclé, Darién, Panama), Colombia (Chocó) and Ecuador (Pastaza), lack the claviform elongate slightly roughened trichomes, the vestiture being only composed of a dense to moderate brown furfuraceous indumentum of dendritic trichomes with short to moderately long thin-walled (flattened) arms. These pubescence variants do not appear to correlate with geography or elevation since these populations occur throughout the elevational range of the species and appear to be sympatric with typical forms of M. approximata . In all other vegetative and floral features M. approximata is quite uniform.

Aclosely related taxon, and possibly distinct, was recently collected from the department of Chocó in Colombia (Almeda et al. 10459, CAS!, COL). It has narrower elliptic leaves, fewer flowers per node, and abaxial foliar venules densely covered with white furrowed sessile glands that are also sparsely present on the hypanthium and exterior calyx teeth. Superficially it is easily confused with M. approximata , but the white furrowed glands in the latter are fewer, although common not consistently present, and restricted to the leaves abaxially. This specimen only has immature fruits, making floral comparisons impossible. It is probable that the indumentum differences, specifically the location of these white furrowed glands (vegetatively or/and on hypanthia/calyx) is taxonomically significant, but it is premature to describe this entity based on the one imcomplete collection. Based on molecular data these two species are close to one another.

Conservation status:— Vulnerable VU B2ab(iii), based on IUCN criteria ( AOO). However, this species occurs in many protected areas, warranting a status of Least Concern LC. Miconia approximata is protected in Colombia in the Ensenada de Utría National Park. In Costa Rica in the Monteverde Biological Reserve (Alajuela and Cartago) ; in La Selva Biological Reserve and the Braulio Carrillo National Park (Cartago, the latter also in Limón and San José); in the Hitoy Cerere Biological Reserve , Barra del Colorado Wild Life Refuge , the Pacuare Protected Zone , and the Tortuguero Nationl Park (Limón) ; in the Corcovado National Park, the Golfito Wild Life Refuge, the Piedras Blancas National Park, the Marenco Biological Station (Puntarenas) ; and in La Cangreja the Cerro Nara Protected Zones (San José) . In Nicaragua it is protected in the Indio-Maíz Reserve (Río San Juan) . In Panama it is protected in the Omar Torrijos National Park (Coclé) ; in the Chagres National Park ( Panama); and in the Coiba National Park (Veraguas) .

Figures

FIGURE 12. Distributions of Miconia anchicayensis and M. chocoensis, M. approximata, M. atropurpurea, M. aurantiaca and M. aff. aurantiaca.