Miconia laxivenula ( Wurdack 1973a: 407–408 ) 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 : 85-90

19. Miconia laxivenula ( Wurdack 1973a: 407–408) Gamba & Almeda , comb. nov. Basionym: Ossaea laxivenula Wurdack. Type: COLOMBIA. Comisaría del Putumayo: Umbría , 0°54’N, 76°10’W, 325 m, October–November 1930, Klug 1816 (holotype: US!; isotypes: F!, MO!, NY!).

Shrub or small tree 1.5–5(–9) m tall, main stem laxly and poorly branched, bark green to green-brownish, smooth. Upper internodes compressed-rounded, (1.9–) 3.2–6.9 cm long, cauline nodes terete, nodal line absent. Indumentum on branchlets, petioles, primary and secondary leaf veins abaxially, inflorescence axes, bracts, bracteoles, pedicels when present, hypanthia, calyx lobes and calyx teeth sparsely composed of caducous amorphous lepidote trichomes <0.1 mm long with only partially fused radii that superficially resemble minute glandular scales. Leaves of each pair commonly isophyllous, occasionally slightly anisophyllous in size; the semiterete petiole (1–) 1.5–3 cm long, moderately canaliculate abaxially; blades 9–29 × 3.4–13.5(–14) cm, elliptic to oblong-elliptic, or elliptic to elliptic-obovate, the base acute, commonly but not exclusively decurrent on the petiole, the margin entire to obscurely crenulate, the apex acute to bluntly short-acuminate, chartaceous; mature leaves adaxially with surface, primary, secondary, tertiary and higher order veins glabrous; abaxial surface occasionally to rarely flushed red-purple, completely glabrous or glabrescent to sparsely and distantly puncticulate, the points superficially black and consisting of resinous short-stalked glands with thin-walled elongate heads, the tertiary and higher order veins glabrous; 5-plinerved, including the tenuous marginals, innermost pair of secondary veins diverging from the primary vein 1–1.5 cm above the base, areolae 2–3 mm, adaxially the primary and secondary veins impressed, the tertiary and higher order veins slightly impressed to flat, abaxially the primary and secondary veins elevated and terete, the tertiary and higher order veins slightly elevated. Inflorescences a pseudolateral multiflorous dithyrsoid 7–11 cm long, including a peduncle <1 cm long or sessile, divaricately and highly branched from the peduncle apex, with somewhat deflexed lower most branches, when sessile bifurcate or trifurcate from the base, each furcation with the same architecture as the pedunculate dithyrsoid, borne in the upper leafy nodes, the rachis red-purplish; bracts and bracteoles 0.5–1 × 0.25 mm, the bracts triangular-concave, the bracteoles subulate, green-reddish, glabrescent on both surfaces, persistent to tardily deciduous in fruit. Flowers 4- (5-) merous sessile or subsessile, the pedicels 0.1–0.2 mm long when present. Hypanthia at anthesis 1.7–2 × 1.2–1.3 mm, free portion of hypanthium 0.6–0.8 mm long, globose to urceolate, becoming dorso-ventrally compressed in fruit, bluntly 8-(10-)ribbed, green-whitish, ridged on the inner surface, sparsely beset with conspicuous brown rounded glands, the torus adaxially minutely glandular. Calyx open in bud and persistent in fruit, green; tube 0.2–0.3(–0.5) mm long, with the same vestiture as the torus adaxially and as the hypanthium abaxially; lobes 0.1 × 0.75–0.85 mm, or if obsolete then just the tube present, depressed-rounded, the margin vaguely and obscurely undulate, the apex obtuse; exterior calyx teeth 0.2 mm long, minutely depressed-rounded, inserted at the base of the calyx lobes or tube, not projecting beyond the lobes but covering their entire dorsal surface. Petals 2.5–3.5 × 0.5–1 mm, lanceolate-triangular, the margin entire, the apex bluntly acute, white to translucent white, glabrous, reflexed at anthesis. Stamens 8; filaments 1.2–2 × 0.25 mm, white-yellowish, glabrous; anther thecae 1–1.25 × 0.3–0.4 mm, oblong-obovate, truncate to emarginate at the apex, opening by two dorsally inclined pores 0.1–0.15 mm in diameter, white-yellowish; connective pale yellow, its prolongation and appendage (0.5–) 0.6–0.8 mm long, the appendage oblong-deltoid or orbicular, obtuse or rounded-truncate at the apex, copiously and conspicuously gland-edged, the glands stalked and apically rounded. Ovary 4-locular, 3/4 inferior, 1.3–1.5(–1.7) mm long at anthesis, the apical collar 0.3–0.4 × 0.17–0.19 mm, conic, conspicuously glandularpuberulent; style 3–3.3 mm long, narrowed distally (i.e. tapering), white, glabrous; stigma expanded truncate to capitellate. Berries 2–2.5 × 3–3.8 mm when dry, globose-oblate, white or orange when fully ripe, the hypanthium indumentum persistent at maturity. Seeds 0.44–0.5 × 0.16–0.19 mm, typically pyramidal, occasionally ovoid and angled, brownish; lateral and antiraphal symmetrical planes triangular, the highest point toward the chalazal side;

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Phytotaxa 179 (1) © 2014 Magnolia Press 85 raphal zone suboblong to sublinear, ca. 90% the length of the seed, somewhat ventrally expanded toward the micropyle or from the chalazal side to the micropyle; appendage absent but a small protuberance present; individual cells elongate, anticlinal boundaries channeled, undulate, with Ω- and U-type patterns; periclinal walls convex, low-domed to nearly flat, microrelief verrucose to somewhat striate.

Additional specimens studied:— COLOMBIA. Nariño: (Tumaco), Chajal , 13 June 1951, Romero- Castañeda 2726 ( NY) . Putumayo: 15 km NW of Puerto Asís , 305 m, 6 August 1965, King & Guevara 6205 ( COL, F, NY, US) ; 15 km NW of Puerto Asís , 305 m, 6 August 1965, King & Guevara 6234 ( COL, F, NY, US) ; Selva higrófila del Río San Miguel , entre las quebradas de Sipenae y de Churruyaco, 400 m, 11 December 1940, Cuatrecasas 10945 ( COL, F, US) . COSTA RICA. Alajuela: (Upala), Dos Ríos, 5 km al Sde Brasilia, margen derecha del Río Pizote , 10°55’N, 85°20’W, 500 m, 29 October 1987, Herrera 995 ( CAS, CR, MO); (Upala) , Bijagua, El Pilón, Cabeceras del Río Celeste, 10°49’N, 84°27’W, 700 m, 14 November 1987, Herrera 1265 ( CAS, CR, MO) ; R.B. Monteverde, Río Peñas Blancas , 10°19’N, 84°43’W, 850 m, 6 August 1988, Bello 253 ( CAS, CR, MO) . Guanacaste: P.N. Guanacaste, Estación Pitilla y Sendero El Mismo, 10°59’26"N, 85°25’40"W, 700 m, 15 June 1991, Ríos 380 ( MO) . Heredia: Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Q. El Salto, at S, Boundary , 100 m, 30 August 1980, Hammel 9634 ( CAS) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Along Loop Trail , between Central Trail and Q. El Saltito, 100 m, 15 July 1980, Grayum 2979 ( CAS) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Central Trail , about 2800 m S, 100 m, 1 August 1980, Hammel 9408 ( CAS) ; P.N. Braulio Carrillo, Puesto El Ceibo, Gently sloping area just above the steep rim of the Río Peje gorge on the w side, along side path that goes from guard house to water supply intake, about 15 minutes uphill, Transect 500–1, 10°19’18"N, 84°4’44"W, 520 m, 9 December 1992, Boyle 1415 ( CAS, CR) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , SW Trail, 1400–1600 m line, 100 m, 7 March 1981, Folsom 9262 ( CAS, F) ; 9 km Eof San Ramon , 10°20’N, 84°3’W, 425 m, 1 December 1986, Loiselle 211 ( MO) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , 2600 NS survey cut, Eof 1050 to 850 EW m line, Camino Central , new property, 100 m, 7 October 1982, McDowell 396 ( CAS, MO) ; Cerros Sardinal , ca. 2–2.5 km Nof Chilamate de Sarapiquí, 10°28’N, 84°4’W, 80–160 m, 21 January 1986, Grayum et al. 6158 ( MO); (Sarapiquí) , La Virgen, P.N. Braulio Carrillo, Puesto La Ceiba, 500 m Ssendero El Ceibo, 10°20’N, 84°5’W, 500 m, 21 November 1988, Ballestero 21 ( CR, MO) ; Finca La Selva, Confluence of Río Puerto Viejo and Río Sarapiquí , to Puerto Viejo de Sarapiquí , 10°25’N, 84°0’W, 80 m, 16 October 1979, Denslow 79-30 ( CAS); (Sarapiquí) , Horquetas, Rara Avis, Cerca del Plástico , 10°18’N, 84°2’W, 550–600 m, 24 December 1994, Knapstein 21 ( CAS, CR) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Long spot along Q. El Saltito, back of Loop Trail , 950 m E, 100 m, 9 September 1980, Hammel 9723 ( CAS) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Central Trail , Río Salto at 2500 m, 100 m, 10 February 1981, Folsom 8875 ( CAS) ; Estación Carrillo de 700 a 450 m de la Fila al Cañón del R, Sucio , 12 November 1983, Chacón & Herrera 1678 ( CAS, CR, MO) . Limón: (Pococí), P.N. Braulio Carrillo, Cuenca del Sarapiquí, Estación Quebrada González , a lo largo del sendero Las Palmas , 10°9’50"N, 83°56’20"W, 500 m, 20 January 1998, Rodríguez et al. 2865 ( 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. 1527 ( INB, MO) . Puntarenas: (Golfito), R.F. Golfo Dulce, Península de Osa, Cerro Rincón, Puerto Jiménez , Cabeceras de los Ríos Tigre y Rincón , 8°30’50"N, 83°28’20"W, 700 m, 7 May 1993, Aguilar 1870 ( CAS, INB, MO) ; P.N. Corcovado, Cerro Brujo, Quebrada Vaquedano , 8°38’N, 83°35’W, 400 m, 22 January 1991, Castro 224 ( CR, MO) . ECUADOR. Carchi: Nside of Río Mira, across from Lita, Steep N-facing slope directly across from ( S of) community of Baboso , on Sside of Río Baboso , Transect 750–2, 0°53’N, 78°27’W, 750 m, 11 August 1994, Boyle & Boyle 3562 ( MO); (Tulcán) , R. Étnica Awá, Comunidad de Gualpi Medio , 1°1’N, 78°16’W, 900 m, 21 May 1992, Quelal et al. 539 ( MO); (Tulcán) , R. Indígena Awá, Comunidad El Baboso, 8 km al Nde Lita , 0°50’N, 78°20’W, 800 m, 15 October 1991, Rubio et al. 2135 ( MO, QCNE, US); (Tulcán) , R. Indígena Awá, Parroquia Tobar Donoso, Sector Sabalera, Bosque primario NE de casa comunal, 1°0’N, 78°24’W, 50–100 m, 19 June 1992, Tipaz et al. 1445 ( MO, QCNE) ; Prominent hillcrest directly Nof Lita, on Nside of Río Mira and just to Eof Río Baboso , On steep W-facing slope, Transect 750–1, 0°53’N, 78°27’W, 760 m, 1 July 1994, Boyle & Boyle 3277 ( MO); (Tulcán) , Chical , R. Étnica Awá-Camumbí, 0°53’N, 78°16’W, 1700–1900 m, 20 July 1991, Quelal et al. 143 ( MO, QCNE) . El Oro: Hacienda

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Daucay, 600 m, 12 October 1993, Cornejo 470 ( MO). Esmeraldas : ( San Lorenzo), R. Indígena Awá , Parroquia Ricaurte , Comunidad Balsareño, Río Palabí, 1°9’N, 78°31’W , 100 m, 15 April 1991, Rubio & Quelal 1310 ( MO, US); ( San Lorenzo), Comunidad Awá Guadalito, 1°16’N, 78°45’W , 125 m, 6 June 1996, Cerón et al. 31347 ( MO, QAP); ( Lita ), Río Lita and tributaries (affluent of Río Mira), 110 km NW of Ibarra, 4.5 km WNW of Lita, 0°52’N, 78°29’W , 600 m, 8 May 1987, Daly & Acevedo 5154 (F, MO, NY, US); ( Eloy Alfaro ), R.E. Cotacachi-Cayapas, Parroquia Luis Vargas Torres , Río Santiago, Estero Angostura, 0°49’S, 78°45’W , 250 m, 8 December 1993, Tirado et al. 793 ( MO, QCNE); Environs of Lita , on the Ibarra-San Lorenzo R.N ., 550–650 m, 11 June 1978, Madison et al. 5206 (F, QCA, US); Cayapas-Cotacachi area , San Miguel and Santiago river, 0°75’N, 78°50’W , 100 m, 1 July 1977, Rodriguez-Carrasquero 5587 ( MO, US); Along road between main Lita-San Lorenzo Hwy, and village of la Ceiba, 2.1 km Wof main hwy, 1°6’38"N, 78°41’33"W , 141 m, 16 July 2000, Croat et al. 84068 ( MO); ( Eloy Alfaro ), Selva Alegre , Estero Gasparito y Estero Felipe, 0°56’N, 78°50’W , 100 m, 21 January 1993, Álvarez et al. 805 ( MO); ( Eloy Alfaro ), Selva Alegre , Estero Gasparito y Estero Felipe, 0°56’N, 78°50’W , 100 m, 21 January 1993, Álvarez et al. 793 ( MO, QCNE); ( San Lorenzo), Finca La Chiquita, 1°13’N, 78°49’W , 80 m, 8 July 1988, Palacios 2666 ( MO); ( San Lorenzo), Carretera Lita-Alto Tambo-La Punta, 1°0’N, 78°35’W , 400 m, 6 February 1991, Gudiño & Moran 1289 ( MO, QCNE); ( San Lorenzo), San Francisco, Recinto Durango , Loma del Rey , Camino al Valle de la Virgen, 2 km al SE de la carretera Lita-San Lorenzo, 1°2’N, 78°37’W , 350 m, 18 October 1999, Valenzuela & Freire 513 ( MO); Zapallo Grande , a mixed black and Cayapa Amerindian community along Río Cayapa, 0°48’N, 78°54’W , 200 m, 11 October 1983, Barfod et al. 48173 ( MO); ( San Lorenzo), Alto Tambo, A 15 km al Wde Lita, 0°50’N, 78°32’W , 400 m, 9 September 1990, Rubio & Quelal 625 ( MO, QCNE); ( Eloy Alfaro ), R.E. Cotacachi-Cayapas , Río San Miguel, Charco Vicente, 0°43’N, 78°53’W , 200 m, 20 September 1993, Tirado et al. 289 ( CAS, MO, QCNE); ( San Lorenzo), R. Étnica Awá, Parroquia Mataje, Centro Mataje, 1°8’N, 78°33’W , 200 m, 21 September 1992, Aulestia et al. 345 ( MO, QCNE); Further along trail to Río Mataje (beginning at point where collecting ended previous day), Awa encampment from Río Palavi encampment, 1°7’N, 78°37’W , 200–230 m, 11 February 1988, Hoover et al. 3929 ( MO); ( Eloy Alfaro ), R.E. Cotacachi-Cayapas , Río Bravo Grando, margen derecha, a 1/ 2 km del Centro Chachi de Corriente Grande, 0°40’N, 78°57’W , 280 m, 3 October 1992, Álvarez & Herrera 709 ( MO, QCNE); Trail from Awá encampment on Río Palaví to Awá encampment on Río Mataje, begining about 1.5 km from Río Palaví, 1°7’N, 78°37’S , 200 m, 10 February 1988, Hoover et al. 3872 ( MO); ( Eloy Alfaro ), San Miguel, Río Cayapas , Propiedad del Sr Miguel Chapiro, Parcela permanente 07 y alrededores, 0°45’N, 78°55’W , 130 m, 3 September 1993, Palacios & Tirado 11137 ( MO, QCNE); Cerro de Río Bravo de Cayapas , 0°41’N, 78°56’W , 250 m, 1 September 1980, Holm-Nielsen et al. 25523 ( CAS, MO). Imbabura : (Lita) , 540 m, 28 May 1949, Acosta-Solís 12537 (F); ( Lita ) , 501 m, 14 April 1949, Acosta-Solís 12215 (F); ( Lita ) , 501 m, 28 April 1949, Acosta-Solís 12312 (F). Morona-Santiago: Centro Shuar Yukutais , Wof Pedro Kunkumas’house and Sof school, 2°30’S, 78°8’W , 900 m, 12 March 1990, Bennett et al. 4046 ( NY, US); ( Morona ), Cordillera de Cutucú , Centro Shuar Uunsuants / TransKutuku, 2°32’S, 77°54’W , 650 m, 18 January 2002, Toasa & Paque 8566 ( MO). Napo : Carretera Hollín-Loreto , 3 km al Edel Río Pucuno, camico al caserío El Pacto, 0°42’S, 77°35’W , 1200 m, 15 July 1988, Neillet al. 8518 ( CAS, MO); ( Orellana ), Zona de amortiguamiento del P.N, Sumaco , Río Bigal , Líneas sísmicas de la compañía petrolera Amoco, Bloque 18, 0°29’S, 77°21’W , 400 m, 9 November 1996, Revelo 125 ( MO); ( Archidona ), Sslopes of Volcán Sumaco, 4 km N of Huahua Sumaco community, Near Río Huataraco, 0°43’S, 77°32’W , 1000 m, 13 December 1989, Neillet al. 9160 ( MO, US); ( Loreto ), Al Nde la carretera Campo Alegre , carretera Hollín-Loreto-Coca. Bloque 19, linea sismica 4, Compañía Triton, 0°36’S, 77°19’W , 420 m, 19 January 1996, Vargas & Cerda 548 ( MO); Road to Bermejo oil field, 35 km N of”km 46" of Lago Agrio-Quito road, ridge top, 10 km Wof Lumbaque, 0°10’S, 77°20’W , 950 m, 21 July 1986, Gentry & Miller 54969 ( MO); Carretera Hollín-Loreto , km 25, entre el Río Hollín y el Río Guamaní, 0°43’S, 77°40’W , 1200 m, 17 May 1988, Neill 8417 ( MO); ( Archidona ), Comuna Huahua Sumaco , Faldas al Sdel volcán Sumaco, Carretera Hollín-Loreto, km 50, 0°43’S , 77°345’W, 1000 m, 1 May 1989, Hurtado et al. 2124 ( MO); ( Archidona ), Comunidad Guagua Sumaco , Carretera Hollín-Loreto , km 50, Faldas al S del volcán Sumaco, 0°38’S, 77°27’W , 1000 m, 29 April 1989, Cerón & Hurtado 6686 ( CAS, MO); ( Archidona ), Comuna Huahua Sumaco , Faldas al Sdel volcán Sumaco, Carretera Hollín-Loreto, km 50, 0°44’S, 77°34’W , 1100 m, 3 May 1989, Alvarado 281 ( MO); ( Archidona ), Comuna Huahua Sumaco , Faldas al Sdel volcán Sumaco, Carretera Hollín-Loreto, km 50, 0°43’S, 77°34’W , 1000 m, 29 April 1989, Hurtado et al. 2004 ( MO); Entre el Río Pucuno y el Río Guamaní, Carretera Hollín-Loreto-Coca, 0°46’S, 77°26’W , 1100–1200 m, 11 December 1987, Cerón 2892 ( MO); ( Archidona ), R. de Biósfera Sumaco, Comunidad Mushullacta, Bosque protector, 0°49’39"S ,

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Phytotaxa 179 (1) © 2014 Magnolia Press 87 77°33’47"W, 1160 m, 26 February 2003, Cevallos 42 ( CAS) . Napo-Pastaza : (Veracruz), 900 m, 18 February 1956, Asplund 19461 ( NY) . Pastaza : ( Pastaza ), Edge of Plateau on Sside of Río Arajuno, above river canyon, Proposed ARCO oil pipeline route, km 16, 1°20’S, 77°49’W, 1000 m, 7 September 1997, Neill et al. 10953 ( MO, NY, QCNE); (Pastaza) , Pozo petrolero”Corrientes" de UNOCAL, 1°43’S, 76°49’W, 300 m, 1 August 1990, Gudiño 630 ( CAS, MO, QCNE); (Pastaza) , Villano-Pandanuque , Junto al Río Villano, Terraza de Río Villano, 1°30’S, 77°27’W, 340 m, 26 July 1992, Palacios 10361 ( MO, QCNE) . Pichincha : Carretera Quito-Puerto Quito, km 113, 10 km al Nde la carretera principal, Bosque virgen y alrededores de la reserva, 0°5’N, 79°2’W, 800 m, 23 February 1984, Betancourt 121 (F, MO, NY, QCA, US); (Quito) , R.F. ENDESA: “Corporación Forestal Juan Manuel Durini”, Río Silanche , km 113 de la carretera Quito-Pto Quito, faldas occidentales, a 10 km Nde la carretera principal, 0°5"N, 79°2’0"W, 650–700 m, 20 February 1984, Jaramillo 6380 ( NY, QCA) ; R.F. ENDESA: “Corporación Forestal Juan Manuel Durini”, Río Silanche , km 113 de la carretera Quito-Pto Quito, faldas occidentales, a 10 km Nde la carretera principal, 0°5’N, 79°2’0"W, 650–700 m, 9 June 1984, Jaramillo 6627 ( NY, QCA) ; R.F. ENDESA: “Corporación Forestal Juan Manuel Durini”, Carretera Quito-Puerto Quito , km 113, 10 km al Nde la carretera principal, 0°5’N, 79°2’0"W, 800 m, 19 February 1984, Betancourt 102 (F, NY, QCA) .

Sucumbíos: (Gonzalo Pizarro), Campo Bermejo 6 N, 30 km al NW de Lago Agrio, 0°14’N, 77°13’W, 1050 m, 23 March 1990, Cerón et al. 9339 ( MO, QCNE, US); (Cascales) , Parroquia El Dorado, Cooperativa Los Angeles, Bloque 11 Compañía Santa Fe , Coleccion 3 km entre La Trocal y Los Angeles, 0°0’S, 77°12’W, 250 m, 3 May 1997, Freire et al. 2166 ( MO, QCNE) . NICARAGUA. Río San Juan: (El Castillo), R. Indio-Maíz, Cerro El Diablo, 11°1’N, 84°12’W, 350–609 m, 9 December 1998, Rueda et al. 9644 ( CAS, MO); ( El Castillo) , Comunidad Aguas Zarcas , 2.5 km al Sdel puesto de guardaparques, 11°14’22"N, 84°14’16"W, 350 m, 10 March 2005, Urbina 160 ( CAS) . PANAMA. Bocas del Toro: Vicinity of Fortuna Dam, Near road to Chiriquí Grande , 8°45’N, 82°15’W, 650 m, 10 August 1986, McPherson 9924 ( MO) . Coclé: Near El Vallede Anton, 8°37’N, 80°7’W, 600 m, 26 November 1985, McPherson 7640 ( CAS, MO); ( El Copé ) , Cerca del aserradero, 31 August 1996, Montenegro 1477 ( CAS); (Anton) , La Mesa, 800 m, 17 January 1973, Kennedy et al. 2109 ( MO) ; Alto Calvario Region, Vivinity of old saw mill works, 4.5 mi Nof El Copé, 2.5 mi Nof Escuela Barrigón , 8°38’N, 80°36’W, 580–740 m, 12 September 1987, Croat 67510 ( CAS, MO); ( El Copé ) , On Pacific side 1/2 hour walk from sawmill, 732 m, 16 October 1979, Antonio 2164 ( CAS, MO) ; In mountains near continental divide, 8 km above El Cope (ca. 0.5 km down logging trail going NE from sawmill), 610 m, 10 January 1978, Hammel 816 ( CAS); ( El Copé ) , Sendero desde la casa de los guardaparques hasta la quebrada, 8°40’N, 80°35’W, 7 July 1996, Aranda et al. 2855 ( CAS) ; Forested slopes above El Copé off of the abandoned road leading to the Contienetal Divide , 8°38’N, 80°38’W, 800 m, 25 January 1989, Almedaet al. 6400 ( CAS) ; NE slopes of Cerro Caracoral, Nrim of El Valle, 823–975 m, 12 March 1981, Sytsma 3774 ( CAS) ; 7.2 km from El Valle main road along the Mesa Road , 700 m, 18 December 1977, Folsom et al. 6936 ( CAS, MO, NY) ; In mountains near continental divide, 8 km above El Copé (ca 5 km down logging trail going NE from sawmill), 610 m, 10 January 1978, Hammel sn ( MO) ; Colón: Trail from Alto Pacora to Cerro Brewster , 9°18’N, 79°16’W, 700 m, 18 November 1985, de Nevers et al. 6237 ( CAS, MO) . Veraguas: Valley of Río Dos Bocas on road between Alto Piedra (above Santa Fe ) and Calovebora , 350–400 m, 29 August 1974, Croat 27469 ( MO) ; Alongroadbetween Escuela Agrícola Alto Piedra (above Santa Fe ) and Río Dos Bocas , ca 10 km from the Escuela , 530–620 m, 26 July 1974, Croat 25895 ( MO) ; Valley of Río Dos Bocas alongroad between Escuela Agrícola Alto Piedra and Calovebora , 15.6 km NW of Santa Fe, Along trail to Santa Fe, Eof river, 450–550 m, 31 August 1974, Croat 27628 ( MO) ; Valley of Río Dos Bocas alongroad between Escuela Agrícola Alto Piedra and Calovebora , 15.6 km NW of Santa Fe, Along trail to Santa Fe, Eof river, 450–550 m, 31 August 1974, Croat 27623 ( MO) . PERÚ. Amazonas: (Bagua), Imaza , Comunidad Samaria , Río Cenepa region , Región NE del Marion , 4°51’S, 78°18’W, 300 m, 1 January 1995, Hodges & Gorham 209 ( CAS, MO); (Bagua) , Imaza, Comunidad Aguaruna de Putuim, Arriba (W) Pueblo de Putuim , 4°55’S, 78°19’W, 680 m, 12 June 1996, Rodríguez et al. 962 ( CAS, MO); ( El Cenepa) , Comunidad de Mamayaque, Cerro Sakee-gaig , 4°34’49"S, 78°14’1"W, 600–800 m, 14 February 1997, Rodríguez et al. 1511 ( CAS, F, MO) . Huánuco: (Pachitea), Codo de Pozuzo, Alluvial fan floodplain of Río Pozuzo after it emerges from mountains, Trail to NW behind settlement, 9°40’S, 75°25’W, 450 m, 18 October 1982, Foster 9253 (F, MO, US) . San Martín: Alongroad between Tocache Nuevo , 114.5 km Sof Juanjui, 16 km Sof Río Pulcache and village of San Martín, 7°50’S, 70°40’30"W, 700 m, 8 April 1984, Croat 58071 ( CAS, MO, US) .

Illustration:— Wurdack 1980: 346, fig. 38.

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Common names and documented uses:— Ecuador: “cayapa”, for curing cramps and epilepsy (Barfod et al. 48173, MO!); “chicnul” (Quelal et al. 539, MO!); “engal chignul”, for making traps (Tipaz et al. 1445, MO!); “uvitilla” (Quelal et al. 143, MO!); “payatzi" (Vargas & Cerda 548, MO!). The first common name is used by the Cayapa Amerindian community; the next three common names are used by the Awá ethnic community.

Habitat, distribution and ecology:— Locally common in primary or secondary lowland rain forests and montane forests, growing on flat areas or steep slopes, commonly in the understory and near streams, from southern Nicaragua through southern Central America, to Colombia, Ecuador and Perú ( Fig. 15 ), at 42–1200(–1900) m. In Nicaragua it is only known from the Reserva Natural Indio-Maíz; it occurs throughout Costa Rica and in Panama it is known from central areas of the country. In South America the distribution is disjunct. In Colombia it is only known from the southern departments of Nariño and Putumayo, and it occurs throughout Ecuador to northern Perú.

Manakins, Tanagers, Thrushes, the orange-billed Sparrows ( Arremon aurantiirostris ) and Myadestes melanops have been reported to feed on M. laxivenula berries in a Costa Rican premontane wet forest ( Stiles & Rosselli 1993).

Phenology:— Collected in flower and fruit throughout the year, but only flowering material has been collected in March.

Etymology:— The specific epithet refers to the lax (loose) reticulation on the abaxial surfaces of leaves.

Discussion:— Miconia laxivenula has lax foliar reticulation in which the areoles are conspicuously bigger than in other species of the Octopleura clade (2–3 mm vs. commonly <2 mm) and its leaves are commonly narrowly decurrent on the petiole. In the protologue Wurdack (1973a) recognized M. sessilis as the closest relative of M. laxivenula , which is confirmed on the basis of molecular data showing that M. laxivenula is sister to M. sessilis ( Fig. 1 ). Both have a basal position among some species within the Variabilis subclade, including M. magnifolia

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 89 and allies. Morphologically M. sessilis has similar lax venule reticulation (areoles 3–5 mm) and 4-merous flowers, but its leaf blades are long decurrent at the base (leaves sessile), it has more prominent foliar plinervation (innermost pair of secondaries diverging 3–5 cm vs. 1–1.5 cm above the base), and the petals are densely puberulent abaxially (vs. glabrous). Miconia variabilis is similar in appearance and inflorescence architecture but differs in the much denser foliar venule reticulation (areoles ca. 0.2 mm), densely puberulent vegetative and floral indumentum, and 5-merous flowers. The flowers in M. laxivenula are predominantly 4-merous. One specimen from the province of Coclé in Panama (Hammel s.n., MO-2686219!) has fruits that are uniformly 10-costate (and thus 5-merous flowers), a rare and unusual variant.

Wurdack (1973a) suspected that O. robusta (Triana) Cogn. fma. glabrata Mgf., which was described as a glabrous population from eastern Ecuador (Pacapaca region, apparently in the province of Pastaza), is the same taxon as M. laxivenula . The holotype chosen by Wurdack was previously determined as O. robusta fma. glabrata Mgf., but it was not cited by Markgraf (1941) in the protologue. The only specimen that Markgraf (1941) based this form on (Schultze-Rhonhof 2717) was not seen; all the Pastaza specimens of M. laxivenula studied have 4- merous flowers and were not glabrous, but covered with the squamate-amorphous indumentum that characterizes this species. Markgraf (1941) did not specify the merosity of the specimen he cited, and no records from that region (locality details are not clear) were seen. Specimens of M. variabilis examined for this study have 5-merous flowers, and those from the province of Pastaza had the normal M. variabilis indumentum. The taxon described by Markgraf (1941) will probably remain unknown based on available evidence. The only specimen cited by Markgraf was probably deposited in Balong with the other Schultze collections. It was destroyed during the Second World War.

The lax foliar reticulation and squamate-amorphous indumentum of M. laxivenula are consistent throughout its geographic range. However, two distinguishable morphological variants have come to light in the course of this study. The type of this species was collected in Colombia (Putumayo) and in accord with South American specimens, has leaves that are elliptic to oblong-elliptic with the base acute and not decurrent along the petiole, or only slightly so. The abaxial leaf surfaces are completely glabrous, and the berries are orange at maturity, which can change to purple-black according to some specimen labels (Acosta-Solís 12215, F!). These populations are known from Colombia (Nariño, Putumayo), Ecuador (Esmeraldas, Napo, Pastaza, and Sucumbíos) and Perú (Huánuco), at 250–1200 m. The second morphological variant includes specimens with leaves that are elliptic to elliptic-obovate. The blade base is acute and conspicuously decurrent on part of the petiole, the abaxial leaf surfaces are black-puncticulate (with minute resinous trichomes), and the berries are white at maturity. Populations with these characteristics have been collected more abundantly than those that conform to type material. These populations are found where the species has been collected in Mesoamerica , and also in Ecuador (Carchi, Cotopaxi, El Oro, Esmeraldas, Pichincha, and Morona-Santiago) and Perú (departments of Amazonas , and San Martín), at 42–975(–1900) m. The floral morphology and measurements of these two morphological variants are consistent, as are the characters mentioned above which supports the view that they belong to the same taxon. The fact that they co-occur in the province of Esmeraldas in Ecuador would prohibit recognizing them as subspecies, since there is no geographic isolation. Field work and population studies are needed to determine how these two variants are interacting where they are sympatric .

Several Ecuadorian specimens have been reported to have the abaxial surface of the leaves red-purple, which might be another kind of morphological variation within this species. This foliar coloration occurs in individuals of both variants described above. The presence of this coloration is not uncommon among tropical plants that live in the dark forest understory, but the function of this anthocyanin pigmentation remains unclear.

Conservation status:— This species would be considered Endangered EN B2ab(iii) based on IUCN criteria ( AOO). However, it occurs in many protected areas throughout its range, which warrants a status of Least Concern LC. Protected in Costa Rica in the Monteverde Biological Reserve (Alajuela) ; in the Guanacaste National Park (Guanacaste); in La Selva Biological Station and the Braulio Carrillo National Park (Heredia, the latter also in Limón); in the Golfo Dulce Forest Reserve and in the Corcovado National Park (Puntarenas) . In Ecuador it is protected in the Awá Indigenous Reserve (Carchi and Esmeraldas) ; in the Cotacachi-Cayapas Ecological Reserve (Esmeraldas) ; and in the ENDESA Forest Reserve (Pichincha) . In Nicaragua it is protected in the Indio-Maíz Reserve (Río San Juan) .

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Figures

FIGURE 1. Bayesian phylogenetic tree of the Octopleura clade, resulting from a combined molecular dataset of nrITS, nrETS, psbkpsbL, and accD-psa1 sequences. Numbers above branches correspond to estimated posterior probabilities. Numbers below branches correspond to estimated bootstrap support values from the Maximum Likelihood phylogenetic tree resulting from the same molecular dataset. The three black bars correspond to the three subclades within the Octopleura clade: Approximata, Quinquenervia, and Variabilis.

FIGURE 15. Distributions of Miconia laxivenula, M. magnifolia, M. neocoronata, M. neomicrantha and M. aff. neomicrantha.