Miconia variabilis 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 : 132-139

32. Miconia variabilis Gamba & Almeda , nom. nov. Basionym: Octopleura robusta Triana (1871: 146) . Ossaea robusta (Triana) Cogniaux (1891a: 1065) . Type: COLOMBIA (Nouvelle Grenade). Dept. Nariño (Prov. de Barbacoas): Chutucal, 600 m, 1851–1857, Triana 4035 (holotype: BM-internet image!; isotypes: BR-internet image!, COL-internet image!, K-internet image!, NY!). Nec Miconia robusta Cogniaux (1887b: 270 ; M. tschudyoides Cogniaux (1887b: 327–328)) .

Shrub or small tree (1.2–)1.5–6(–7) m tall with branching from the upper nodes, bark green to brown-rusty. Upper internodes terete to compressed-rounded to rounded-quadrate 1.5–9 cm long, cauline nodes terete, nodal line present but faint and typically concealed by the dense indumentum. Indumentum on branchlets, petioles, blade bases of young leaves and primary veins adaxially, primary and secondary leaf veins abaxially, inflorescence axes,

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bracts and bracteoles abaxially, pedicels, hypanthia, calyx lobes, calyx teeth, and petals abaxially copiously to moderately covered with brownish dendritic trichomes 0.05–0.08 mm long with short axes and few-moderate number of terete arms. Leaves of each pair isophyllous; the semiterete petiole 1–4.5 cm long, canaliculate adaxially; blades (6–)8.5–34.5 × (2.2–) 3.8–15 cm, elliptic to elliptic-ovate, the base broadly acute to obtuse, occasionally somewhat decurrent on petiole, the margin obscurely and distantly undulate-denticulate to denticulate, the apex acuminate, rarely caudate, chartaceous; mature leaves adaxially with surface and primary vein glabrescent, the secondary, tertiary and higher order veins glabrous; abaxial surface glabrous except for a few minute resinous short-stalked glands 0.015 mm long with thin-walled short heads, the secondary, tertiary and higher order veins sparsely beset with resinous glands of the same type and size; 5-plinerved, including the tenuous marginals, innermost pair of secondary veins diverging from the primary vein asymmetrically (occasionally symmetrically) 0.5–2 cm above the blade base, areolae 0.2–0.3 mm, adaxially the primary and secondary veins impressed, the tertiary and higher order veins flat, abaxially the primary and secondary veins elevated and terete, the tertiary and higher order veins slightly raised. Inflorescences a pseudolateral multiflorous dithyrsoid 4–10 cm long, including a compressed-rounded to rounded-quadrate peduncle 0.5–0.8 cm long, divaricately branched from the peduncle apex, borne in the upper leaf axils, the rachis green; bracts and bracteoles 0.4–0.7 × 0.3–0.5 mm, the bracts narrowly triangular to linear-oblong, the bracteoles triangular, green-brownish, glabrous adaxially, glabrescent abaxially, the bracts early deciduous, the bracteoles persistent in fruit. Flowers 5-merous, sessile or subsessile, pedicels 0.2–0.4 mm long when present. Hypanthia at anthesis 1.1–1.5 × 1.1–1.3 mm, free portion of hypanthium 0.8–1(–1.2) mm long, globose to suburceolate, bluntly 10-ribbed, green-whitish, ridged on the inner surface, sparsely beset with resinous glands as well as the torus adaxially. Calyx open in bud and persistent in fruit, light green; tube 0.2–0.3 mm long with the same vestiture as the torus adaxially and as the hypanthium abaxially; lobes 0.2–0.5 × 0.7–0.9 mm, depressed-rounded, the margin entire, the apex obtuse; exterior calyx teeth 0.1–0.3 mm long, minute, tuberculiform, somewhat perpendicular to the calyx lobes, inserted at the apical half of the lobes and typically not projecting beyond them. Petals 2–4.5 × 0.75–1 mm, lanceolate-triangular, the margin entire, the apex bluntly acute, white, glabrous adaxially, the indumentum abaxially present mainly apically and along the costa, spreading to reflexed at anthesis. Stamens 10; filaments 1.5–2 × 0.25 mm, white, drying yellow to light orange, glabrous; anther thecae 0.75–1.1 × 0.3–0.4 mm, oblong to oblong-obovate, emarginate at the apex, opening by two dorsally inclined pores 0.1–0.2 mm in diameter, pale yellow or white; connective pale yellow or white, its prolongation and appendage 0.4–0.75 mm long, the appendage oblong-lanceolate, bluntly acute at the apex, copiously gland-edged with short-stalked glands throughout the connective and its appendage. Ovary (4-)5-locular, 3/4 inferior, 0.65–0.85 mm long at anthesis, the apical collar absent, the apex 0.3–0.45 mm in diameter, conic, sparsely glandular-puberulent to glabrous; style 3.5–4 mm long, moderately tapered distally, white, glabrous; stigma expanded truncate to capitellate. Berries 2.5–3 × 3–4 mm when dry, globose-oblate, green and ripening white or green and changing to yellow, orange and ultimately purple when fully ripe, the hypanthial indumentum persistent at maturity. Seeds 0.26–0.36 × 0.13–0.21 mm, ovoid, angled, brownish; lateral and antiraphal symmetrical planes ovate, the highest point toward the chalazal side; raphal zone suboblong to sublinear, ca. 80% the length of the seed; 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. Chromosome number n=17.

Additional specimens studied:— BRAZIL. Acre: (Mancio Lima), P.N. da Serra do Divisor, Serra do Moa , Trilha para a cachoeira do Formoso ( Igarapé do Anil ), 13 November 2007, Goldenberg et al. 978 ( CAS, NY) . COLOMBIA. Antioquia: (Amalfi), 8–27 km NE de Amalfi en la via Vetilla-Fraguas, sitios Salazar y Marengo , 6°56’N, 75°4’W, 1150–1450 m, 7 December 1989, Callejas & et al. 9119 ( HUA, MO, NY) . Cauca: ( Gorgona Island ), "St, George", 91 m, 10 October 1924, S.E.R.A. 562 ( US); (Santa Rosa) , 1 km arriba de la desembocadura del Indiyaco en el Caquetá, 600 m, 15 October 1996, Sánchez et al. 3036 ( COL) . Chocó: (Quibdó), En el sitio de La Platina, 60 m, 29 March 1958, Uribe-Uribe 3135 ( MO, US) ; Serranía de Baudó, along road between Las Animas and Río Pato, along valley of Río Animas , 5 km from beginning of road, 5°16’N, 76°41’W, 100 m, 17 April 1983, Croat 56047 ( MO); (Nuquí) , Alto de Buey, Alto de Buey , 1000 m, 1 June 1940, von Sneidern A 12 ( NY); (Quibdó) , Río Cabi , 20 April 1982, Prance 28008 ( NY) ; Corcovada Region, upper Río San Juan, ridge along Yeracüí Valley , 200–275 m, 24 April 1939, Killip 35252 ( NY, US) ; Pan American Highway (under construction) ca 10 km Wof Las Animas, 100 m, 12 January 1979, Gentry & Renteria 24068 ( MO, US); (Nuquí) , Alto de Buey, Alto de Buey , 1000 m, 12 June 1940, von Sneidern A 51 ( US) ; Hoya del Río San Juan , Pequeño cerro

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Phytotaxa 179 (1) © 2014 Magnolia Press 133 al frente de Palestina, 4°10’N, 77°10’W, 30–40 m, 26 March 1979, Forero et al. 4083 ( COL, MO, US); (La Mojarra), Just upriver from Istmina , Trail through secondary vegetation to nearly primary forest on hills, 5°12’N, 76°37’W , 30–60 m, 5 November 1983, Juncosa 1260 ( MO, US); (Nóvita), Llanadas, Ladera Ndel Cerro Torra , Filo Wdel Río Surama , Camino al Alto del Oso , 600–900 m, 22 February 1977, Forero et al. 3086 ( MO); (Bahía Solano), 25 July 1973, Warner & White 51 ( MO); (Quibdó), Carretera Quibdó-Yutó, Río Cabí, 24 March 1984, García-Cossio 25 ( CAS, CHOCO, MO); (Pizarro), km 30–33 de la carretera Pie de Pepé-Puerto Meluk , 16 November 1985, Espina 1830 ( CAS, CHOCO, MO); ( San José del Palmar ), El Tabor , Finca "El Tabor" , 1650 m, 17 January 1983, Franco et al. 1453 ( COL, MO); (Pizarro), Margen izquierdo del Río Pepé, entre Boca de Pepé y Pié de Pepé, 19 November 1985, Espina 2019 ( CHOCO, MO); (Quibdó), Carretera Yuto-Lloró , 80 m, 30 June 1983, Forero et al. 9696 ( COL, MO); Río San Juan, Estribaciones de La Mojarra (ca. 30 min de Istmina en lancha con motor fuera de borda) , 80–120 m, 25 June 1983, Forero et al. 9520 ( COL, MO); Along road between Las Animas and Pato (on Río Pato) 16 km NW of the jct, with main Quibdó-Istmina road, near Las Animas , 5°20’N, 76°42’W , 100 m, 18 April 1983, Croat 56163 ( MO); Medellín-Quibdó road, ca. 3 km Wof Tutunendó, 5°46’N, 76°35’W , 80 m, 7 January 1981, Gentry et al. 30331 ( MO). Nariño: La Planada R., 7 km from Chucunes , 1°5’N, 78°1’W , 1800 m, 22 December 1987, Gentry & Keating 59686 ( MO); (Barbacoas), Altaquer, El Barro , R.N. Río Ñambí , Vertiente occidental andina, margen derecha del Río Ñambí , 1°18’N, 78°8’W , 1325 m, 1 December 1993, Franco et al. 4721 ( CAS). Putumayo: (Mocoa), Quebrada del Río Afán , 570–680 m, 27 December 1940, Cuatrecasas 11336 ( COL, US); 15 km NW of Puerto Asís , 305 m, 6 August 1965, King & Guevara 6236 ( CAS, COL, NY); (Mocoa), El Medio Afán, El Medio Afán , ca 2 horas arriba del Río Afán , 1200–1600 m, 24 January 1990, Ramos et al. 2465 ( CAS, CUVC, NY). Risaralda: (Mistrató), Entre los corregimientos de Geguadas y Puerto de Oro , selva de Pisones , 1550 m, 30 March 1992, Fernández et al. 9657 ( US). Valle: Costa del Pacífico, Río Cajambre, Barco , 5–80 m, 21 April 1944, Cuatrecasas 17033 (F, NY, US); La Trojita, Río Calima (región del Chocó ) , 5–50 m, 19 February 1944, Cuatrecasas 16535 (F, NY, US); Hacienda La Florencia, Río Calima , margen derecha, frente a la Base , 10 m, 21 February 1969, Cuatrecasas & Platino 27461 ( US); Alto Yunda, Río Anchicayá , 1000 m, March 1973, Hilty Mr-9 ( ARIZ, US); (Córdoba), Dagua Valley ,, 80–100 m, 6 May 1922, Killip 5124 ( NY, US); Veneral, Costa del Pacífico, Río Yurumanguí , 5–50 m, 28 January 1944, Cuatrecasas 15872 (F, NY); Queremal, Cordillera Occidental, vertiente occidental, Hoya del Río Dagua , lado izquierdo del Río San Juan en la región del Queremal , Quebradito del km 51, 1540 – 1650 m, 25 February 1947, Cuatrecasas 23715 (F, US); (Dagua), Queremal, El Danubio , P.N.N. Farallones , Alto Anchicayá , Road from campamento EPSA, Yatacue to the dam, Cordillera Occidental , vertiente occidental, 3°33.632’N, 76°52.598’W , 625 m, 5 February 2011, Almedaet al. 10242 ( CAS, COL); La Trojita, Río Calima (región del Chocó ), La Trojita , 5–50 m, 19 February 1944, Cuatrecasas 16378 ( NY); Pacific Coast, Río Calima, Quebrada de López , 20–40 m, 23 September 1961, Cuatrecasas & Willard 26041 ( US); Vicinity of bajo Calima, along road from Buenventura to Málaga , km 49, 4°2’N, 77°4’W , 150 m, 17 July 1993, Croat & Bay 75807 ( MO); Bajo Calima, Concesión Pulpapel / Buenaventura , Carretera Canalete PS2, 3°55’N , 77°’W, 100 m, 13 October 1987, Monsalve 1921 ( CUVC, MO); Bajo Calima, Concesión Pulpapel / Buenaventura , Carretera Canalete PS 2, 3°55’N, 77°W , 100 m, 22 October 1987, Monsalve 2023 ( CUVC, MO); Bajo Calima, Dindo area , 3°59’N, 76°58’W , 100 m, 20 July 1984, Gentry & Monsalve 48427 ( MO); Bajo Calima, Concesión Pulpapel , 100 m, 20 January 1988, van der Werff & Monsalve 9664 ( CAS, MO); Bajo Calima, Concesión Pulpapel / Buenaventura , Carretera a Juanchaco N, 3°55’N , 77°’W, 100 m, 21 September 1987, Monsalve 1752 ( CUVC, MO). COSTA RICA. Alajuela: (San Carlos), No protegida, Cuenca del San Carlos , Boca Tapada , Laguna Lagarto Lodge , cerca de la laguna, 10°41’10"N , - 84°10’50.0001"W , 50–100 m, 23 February 2004, Solano & Kriebel 907 ( INB, NY). Cartago: ( Valle Escondido ) , 700 m, 30 March 1966, Schnell 620 ( US); ( Valle Escondido ) , 700 m, 30 March 1966, Schnell 614 ( US); (Oreamuno), P.N. Volcán Irazú, Cordillera Central , San Juan de Chicuá , carretera al Volcán Irazú , 9°57’0"N, 83°51’45"W , 2800 m, 14 October 1993, Ramírez et al. 165 ( CAS, INB). Guanacaste: P.N. Guanacaste, Estación Pitilla , alrededores, 11°2’N, 85°24’W , 600 m, 15 June 1989, II INBio et al. 21 ( CAS, CR, INB). Heredia: Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí, S Boundary, W end , 100 m, 4 May 1981, Folsom 10009 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , At first stream crossing on Central Trail after Loop Trail , 100 m, 15 July 1980, Grayum 2980 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Along Q. Paralela at crossing with 3600 m S line, new property, 100 m, 18 July 1982, Hammel & Trainer 13235 ( CAS); Finca La Selva, Puerto Viejo de Sarapiquí, F, Almedar Loop , 1300 m line, 15 July 1979, Grayum 1830 ( CAS); (Sarapiquí), La Virgen , Golfito , No protegida, Cuenca del Sarapiquí , parcela del señor Miguel Angel Picado , 10°38’40.0636"N ,

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84°3’25.1215"W, 0–100 m, 10 December 2005, Santamaría & Solano 3596 ( INB, NY); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Central Trail , 180 m line, 100 m , 28 November 1982, McDowell 969 ( CAS, NY, US); (Sarapiquí), No protegida, Cuenca del Sarapiquí , Rara Avis Lodge and R., Atlantic slope of Costa Rica. (area protegida privada) , 10°16’55"N, 84°2’40.5"W, 700 m, 19 June 2001, Boyle et al. 80 ( INB, MO); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , 1520 m line, S Boundary Line, 100 m , 29 October 1982, McDowell 681 ( CAS, NY, US); (Puerto Viejo), La Selva, Loop Trail , 90 m , 16 April 1972, Opler 702 ( US); La Selva, 3 km SE of Puerto Viejo, Loop Trail , 45 m, 13 April 1973, Opler 1721 ( US); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , 180 m line, Central Trail , 100 m , 28 October 1982, McDowell 642 ( CAS); (Sarapiquí), La Virgen, P.N. Braulio Carrillo, 600 m Sdel Puesto La Ceiba , sobre La Fila , 10°20’N, 84°5’W, 550 m, 14 October 1988, Ballestero 5 ( CAS); ( Magsay ), Borde de la Quebrada Canta Rana, 400 m , 17 January 1983, Chacón 108 ( CAS, CR); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Along Q. Saltito at back of loop, 100 m , 13 June 1982, Hammel & Trainer 12857 ( CAS); (Sarapiquí), Rara Avis, ca 15 km al SW de Horquetas , Camino Leví hacia Catarata y Río Atelopus , aguas arriba , 10°17’N, 84°2’W, 400 m, 19 September 1989, Vargas 121 ( CAS, CR, INB); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Central trail 1500 m S, 100 m , 4 May 1982, Hammel 12000 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Along small water course in SW quarter of new property, 100 m , 18 May 1982, Hammel 12311 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Pantano ridge about 1950 m S, 1820 m E, 100 m , 1 September 1981, Smith 157 ( CAS); Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , Central Trail , 2600 m line-W boundary, 3000 m line, 100 m , 24 April 1981, Folsom 9854 ( CAS); (Puerto Viejo de Sarapiquí) Finca La Selva, First forest , Sabalo-Esquina Trail , 9 July 1979, Sperry 863 ( CAS); Estación Biológica La Selva, R. of the Organization for Tropical Studies , Vicinity of the first swamp on the Central Trail , 100 m , 27 February 1986, Almeda & Anderson 5106 ( CAS, CR). Limón: Hacienda Tapezco-Hacienda La Suerte , 29 air km Wof Tortuguero , 10°30’N, 83°47’W, 40 m, 29 August 1979, Davidson & Donahue 8933 ( CAS, US); (Pococí), P.N. Braulio Carrillo, Cuenca del Sarapiquí , Sector Quebrada González , sendero Las Palmas , 10°9’20"N, 83°56’30.0017"W, 600 m, 29 July 2008, Kriebel 5313 ( INB, NY); ( Toro Amarillo ), 300–350 m , 18 July 1964, Ferreyra 15912 ( US); (Limón), Cerro Muchilla, Fila Matama , Cordillera de Talamanca , 9°47’50"N, 83°5’30"W, 850 m, 6 April 1989, Robles & Chacón 2676 ( CAS); (Talamanca), Fila de exploración minera entre Río Sukut y Río Carbri , Muragubishi , 9°22’50"N, 82°56’50"W, 700 m, 14 July 1989, Herrera 3290 ( CAS, CR). Puntarenas: R.N.V.S. Golfito, Ca , 6 km NW of campo aterrizaje de Golfito , 8°41’0"N, 83°13’10"W, 100 m, 28 January 1992, Hammel & Aguilar 18403 ( CAS, CR); Between Golfo Dulce and Río Térraba 30 m , 5 December 1947, Skutch 5309 (F, US); (Golfito), P.N. Corcovado, Península de Osa, Estación Los Patos , alrededores , 8°34’0"N, 83°31’0"W, 200 m, 6 September 1993, Aguilar 2259 ( CAS); Along road from Panamerican Hwy, at Piedras Blancas to Rincón (on Osa Peninsula ), 3.7 mi Wof Panamerican Hwy , 8°46’N, 83°18’W, 90–105 m, 16 September 1987, Croat 67670 ( CAS). ECUADOR. Carchi: ( El Pailón ), ca 45 km below Maldonado along a foot path to Tobar Donoso, 800 m, 26 November 1979, Madison & Besse 7006 ( US); (Tulcán), R. Étnica Awá, Comunidad de Gualpi Medio , 1°1’N, 78°16’W, 900 m, 21 May 1992, Quelal et al. 694 ( CAS, MO, QCNE); (Gualpi Chico), Vicinity of Awá encampment , 0°58’N, 78°16’W, 1330 m, 17 January 1988, Hoover et al. 2699 ( MO); NW side of Río Gualpi Chico , heading Ndown river , 0°58’N, 78°16’W, 1100–1200 m, 25 January 1988, Hoover et al. 3025 ( MO); (Tulcán), R. Indígena Awá, Parroquia Tobar Donoso, Sector Sabalera , NE Casa Comunal , 1°0’N, 78°24’W, 100–650 m, 19 June 1992, Tipaz et al. 1310 ( MO, QCNE); Border area between Prov, Carchi and Esmeraldas, about 20 km past Lita on Road Lita-Alto Tambo , 550 m , 26 June 1991, van der Werff et al. 12040 ( CAS, MO); (Tulcán), R. Étnica Awá, Parroquia El Chical, Sector Gualpí Medio , Río Canumbí , 1°2’N, 78°15’W, 1150 m, 19 February 1993, Grijalva et al. 506 ( MO, QCNE); (Tulcán), R. Indígena Awá, Parroquia El Chical , Sector Gualpí Medio , Sendero a San Marcos al Nde la casa comunal , 1°2’N, 78°16’W, 3200 m, 23 May 1992, Tipaz et al. 1118 ( MO, QCNE); (Tulcán), Chical, R. Étnica Awá- Camumbí , 0°53’N, 78°16’W, 1700–1900 m, 20 July 1991, Quelal et al. 163 ( MO, QCNE); (Tulcán), R. Étnica Awá, Parroquia Chical, Centro Gualpi Medio , 1°2’N, 78°16’W, 900 m, 18 February 1993, Aulestia & Grijalva 1099 ( MO); Border area between Prov, Carchi and Esmeraldas, about 20 km past Lita on Road Lita-Alto Tambo , 550 m , 26 June 1991, van der Werff et al. 12048 ( MO). Esmeraldas: Road Lita-Alto Tambo-San Lorenzo, km 6.9 from Lita , 0°52’24.6"N, 78°29’33.2"W, 720 m, 30 September 2001, Cotton et al. 1790 ( MO, NY, QCA); Along road between Lita and San Lorenzo , 36.6 km Nof Gasolinera San Lorenzo, 12.6 km Nof Río Tulubí , 1.7 km Sof

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Phytotaxa 179 (1) © 2014 Magnolia Press 135 El Durango, 1°38’1"N, 78°38’1"W, 204 m, 18 July 2000, Croat et al. 84147 ( MO) ; Parroquia de Concepción , Playa Rica, 105 m, 10 December 1936, Mexía 8428 (F, MO, NY, US) ; Alongroadbetween Litaand San Lorenzo , 36.6 km Nof Gasolinera San Lorenzo, 12.6 km Nof Río Tulubí, 1.7 km Sof El Durango, 1°38’1"N, 78°38’1"W, 204 m, 18 July 2000, Croat et al. 84160 ( MO) ; Sof bridge Sof Lita on Lita-Alto Tambo road, 0°53’19"N, 78°31’43"W, 900 m, 21 September 2003, Penneys et al. 1576 ( CAS, NY, US); (Eloy Alfaro ) , R.E. Cotacachi- Cayapas , Parroquia Luis Vargas Torres , Río Santiago , estero Pote, 0°49’N, 78°45’W, 250 m, 23 October 1993, Tirado et al. 545 ( MO, NY); (San Lorenzo) , R. Indígena Awá, Comunidad "La Unión", Cañón del Río Mira, 10 km al Wde Alto Tambo, 1°2’N, 78°26’W, 250 m, 16 March 1991, Rubio et al. 1240 ( QCNE, US) ; Road San Lorenzo- Alto Tambo-Lita , km 60.5 from San Lorenzo , fringes of the R. Cotacachi-Cayapas, 0°53’47.7"N, 78°32’32.7"W, 879 m, 3 October 2001, Cotton et al. 1822 ( CAS, MO, QCA); (San Lorenzo) , R. Étnica Awá , Parroquia Alto Tambo, Centro de La Unión, Cañón del Río Mira, 0°52’N, 78°26’W, 250 m, 22 March 1993, Aulestia & Aulestia 1340 ( MO, QCNE); (San Lorenzo) , R. Étnica Awá , Parroquia Alto Tambo, Centro de La Unión, Cañón del Río Mira, 0°52’N, 78°26’W, 250 m, 22 March 1993, Aulestia & Aulestia 1276 ( MO, QCNE) . Morona-Santiago : ( Morona ), Alongroadbetween Macas and Puyo, 31 km Nof Macas, 28.5 km Nof bridge over Río Upano, 2°1’S, 77°56’W, 1125 m, 7 March 1992, Croat 72801 ( MO) ; Along Río Palora 2–5 km downstream from Arapicos, 800–900 m, 9 April 1981, Lugo 6056 ( CAS) . Napo : Baeza-Tena Road, between Osayacu and Jondachi, 10–20 km Nof Archidona, 900 m, 5 February 1980, Harling & Andersson 16237 (F, MO, US) ; 8 km río abajo de Puerto Misahualli, por el Río Napo y 1.5 km al S, 1°4’S, 77°36’W, 450 m, 18 May 1985, Palacios et al. 435 (F); Hacienda Cotapino (Concepción), 550 m, 19 February 1968, Harling et al. 7095 (F, US) ; P.N. Sumaco-Napo-Galeras, Estribaciones occidentales de la cordillera Galeras , Bosque maduro a 100 m de la casa flia Mamallacta, tierra firme a orillas del Río Pusuno, 0°56’S, 77°38’W, 950 m, 27 December 2002, Guevara 601 (F, QCA) ; Estación Biológica Jatún-Sacha , Río Napo, 8 km al Ede Misahualli, 1°4’S, 77°36’W, 450 m, 17 February 1988, Cerón 3588 ( CAS, MO, NY) ; Estación Biológica Jatún-Sacha , Río Napo, 8 km al Ede Misahualli, 1°4’S, 77°36’W, 450 m, 8 September 1987, Neill et al. 7816 ( MO, NY); (Loreto) , Faldas del Volcán Sumaco , Al Wde Avila Viejo, Bloque 19, línea sísmica 8, Compañía Tritón, 0°38’S, 77°27’W, 750 m, 14 February 1996, Freire & Cerda 97 ( MO, NY) ; Estación Biológica Jatún-Sacha , Río Napo, 8 km al Ede Misahualli, 1°4’S, 77°36’W, 450 m, 22 February 1988, Palacios 2439 ( MO, NY) ; R.B. Jatún-Sacha , Margen derecha del Río Napo, 8 km abajo de Misahualli, 1°4’S, 77°36’W, 450 m, 13 February 1987, Palacios & Neill 1534 ( MO, US) ; Estación Biológica Jatún-Sacha , 8 km al E del Puerto Misahualli, Por los senderos de la estación, 1°4’S, 78°36’W, 450 m, 25 March 1989, Macías & Bensman 54 ( MO, QCNE) ; R.B. Jatún-Sacha, Río Napo , 8 km abajo de Misahuallí, 1°4’S, 77°36’W, 450 m, 17 January 1987, Cerón 574 ( MO) ; Estación Biológica Jatún-Sacha , 8 km al Edel Puerto Misahualli, Por los senderos de la estación, 1°4’S, 78°36’W, 450 m, 22 March 1991, Macías 98 ( MO, QCNE); (Tena) , Construcción de carretera Campococha-Chontapunta, Variante Selva Viva-Bosque Protector, abscisas 4+500–12+020, 0°55’S, 77°25’W, 300–500 m, 23 August 1997, Núñez & Tapuy 556 ( MO, QCNE) . Napo-Pastaza : Vicinity of Puyo , Eastern foothills of the Andes , 750–1000 m, September 1939, Skutch 4487 (F); Near Puyo , Puyo forest, 17 February 1953, Prescott 885 ( CAS, NY); (Mera) , Forest near Mangayacu, 1100 m, 14 November 1955, Asplund 18432 ( US); (Mera) , 4 March 1940, Lugo 17 ( MO) . Pastaza : Pacayacu on the Río Bobonaza, c, 16 km NW of Sarayacu, 10 August 1979, Lugo 5241 (F, MO) ; Along road from Puyo to Macas , ca 33 km Sof Puyo, 24.9 km Sof Veracruz, 16 km Sof Escuela Fiscal Cotopaxi, 1°38’S, 77°52’W, 900 m, 3 May 1984, Croat 58939 ( CAS, MO) ; 1 km al Sde Mera, vega a la orilla del Río Pastaza , cerca de la boca del Río Aldayacu, 1°28’S, 78°7’W, 6 March 1984, Baker et al. 5553 ( NY); (Puyopungu) , 15 September 1976, Lugo 4823 ( MO, US); (Pastaza) , Between Shell and Mera, 5.3 km NW of center Shell, along gravel road, 1.1 km Nof highway, disturbed virgin forest at the end of boardcover path, Eend of road, 1°27’S, 78°4’W, 1180 m, 3 April 1992, Croat 73456 ( MO) ; Puyo-Arajuno Road , 1–5 km SW Diez de Agosto, 900 m, 4 March 1980, Harling & Andersson 16864 ( CAS, MO) . Pichincha : R.F. ENDESA, ca. 8 km Ede pedro Vicente Maldonado, 0°5’N, 79°1’W, 780 m, 28 February 2004, Moran et al. 6948 ( NY) ; R.F. ENDESA. "Corporación Forestal Juan Manuel Durini", Río Silanche , km 113 de la carretera Quito-Pto, Quito, faldas occidentales, a 10 km al Nde la carretera principal, 0°5’N, 79°2’W, 650–700 m, 16 December 1984, Cueva 5 (F, QCA) ; R.F. ENDESA. "Corporación Forestal Juan Manuel Durini", Carretera Quito-Pto Quito , km 113, 10 km al N de la carretera principal, 0°5’N, 79°2’W, 800 m, 25 February 1984, Betancourt 126 (F, QCA) ; Carretera Quito- Puerto Quito, km 113, 10 km al Nde la carretera principal, 0°5’N, 79°2’W, 800 m, 27 December 1983, Balslev & Balseca 4665 ( NY, QCA, US) . Sucumbíos : Estación Científica Cuyabeno, Colecciones en el cuadrante de sendero línea B, 220 m, 28 March 1992, Jaramillo & Grijalva 14671 ( QCA, US) ; Estación Científica Cuyabeno , en el cuadrante de sendero línea B, 220 m, 30 March 1992, Jaramillo & Grijalva 14703 ( MO, QCA) . Zamora-

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Chinchipe: Shore of Río Pastaza opposite Mera , 1000 m, 6 February 1956, Asplund 19257 ( NY, US) . PANAMA. Bocas del Toro: Region of Almirante, Cricamola Valley , January–March 1928, Proctor 478 ( NY) ; Oleoducto Road, near Continental Divide, Fortuna Dam area , 8°48’N, 82°12’W, 1000 m, 5 February 1984, Churchill et al. 4511 ( CAS) . Chiriquí: (Boquete), Fortuna dam site, 1200 m, 7 February 1985, van der Werff & van Hardeveld 6683 ( CAS) ; Gualaca-Chiriquí Grande 8 mi beyond Los Planes de Hornito, 6.9 mi beyond road turnoff to Caldera , 1.4 mi to Wof Centro de Operaciones, along trail to Río Hornito and meteological station, 8°44’"N, 82°14’30"W, 1010–1130 m, 21 September 1987, Croat 67915 ( CAS); (Gualaca - Chiriquí Grande) , 8 mi beyond Los Planes de Hornito, 6.9 mi beyond road turnoff to Caldera, 1.4 mi to Wof Centro de Operaciones, along trail to Río Hornito and meteological station, 8°44’N, 82°14’30"W, 1010–1130 m, 21 September 1987, Croat 67889 ( CAS) ; Fortuna Dam Area, Slope NW of confluence of Río Hornito and Río Chiriquí , 8°44’N, 82°13’W, 1050–1100 m, 10 November 1980, Sytsma & Stevens 2210 ( CAS) ; Upriver from the main camp, Fortuna dam site, 1200–1400 m, 15 September 1977, Folsom et al. 5501 ( CAS, MO) ; Edwin Fabrega Dam and R, in Fortuna, Quebrada Christina ca 7.4 km Nof the Dam site ( Sitio de Presa offices) enroute to Chiriquí Grande , 1060 m, 5 February 1996, Almeda et al. 7508 ( CAS, PMA); (Fortuna) , Trocha Cordillera Central , cerca de la división continental, 28 October 1997, Montenegro 1808 ( CAS) ; Road to Fortuna Dam site Nof Gualaca. 22.7 mi beyond the bridge over Río Estí, 1400 m, 22 November 1979, Antonio 2777 ( CAS) ; Slope NW of confluence of Río Hornito and Río Chiriquí , 8°44’N, 82°13’W, 1050–1100 m, 11 November 1980, Stevens 18263 ( CAS) . Coclé: Above El Copé and Barrigón , near continental divide, 8°38’N, 80°35’W, 850 m, 25 February 1988, McPherson 12168 ( CAS) ; Woods adjacent to chicken farm, La Mesa, above El Valle, 3 January 1974, Dwyer 11906 ( NY, US) ; Elfin Forest, Cerro Caracoral , 1000 m, 19 January 1968, Duke & Dwyer 15095 ( NY); (El Valle) , 800–1000 m, 28 June 1967, Duke 13157 ( US) ; On slopes of Cerro Pilón near El Valle, 700–900 m, 10 June 1967, Duke 12136 ( NY) ; Cerro Gaital, E slope and ridges leading to the summit with montane rainforest and elfin woodland, 8°40’N, 80°7’W, 1050–1100 m, 24 February 1988, Almedaet al. 5904 ( CAS, PMA) ; New works, 7 km Nof El Cope, 750–800 m, 18 August 1977, Folsom 4956 ( CAS); (Alto Calvario) , Above El Copé, ca 6 km Nof El Copé, Atlantic slope, along trail which leads W off old lumber trail which leads down to Las Ricas, Limón and San Juan , 8 or 9°39’N, 80°36’W, 710–800 m, 22 June 1988, Croat 68717 ( CAS) ; Vicinity of La Mesa, Nof El Valle de Antón, along steep slopes above water reservoirs, ca 1 km Wof road between Finca Manadarinas and Finca Furlong , 8°38’N, 80°9’W, 800–900 m, 12 July 1987, Croat 67168 ( CAS) ; Ridge NW of village of Río Blanco de Norte, between Caño Sucio and Río Blanco de Norte, Señor Dideymo Olivera's finca, 8°44’N, 81°40’W, 350 m, 20 February 1982, Knapp 3674 ( CAS) . Colón: Teck Cominco Petaquilla mining concession, Forested slopes below ridgetop road, 8°49’28"N, 80°39’29"W, 180 m, 19 September 2007, McPherson 19716 ( CAS) . Darién: Cerro Pirre , 762–1372 m, 9 August 1967, Duke & Elias 13838 ( US) ; Trocha Río Frío hacia la cima más allá de Rancho Tuque (3h de camino), Trocha de Rancho Frío hacia Rancho tigre, hacia la cima de Cerro Pirre , 1000–1200 m, 21 September 1989, Aranda et al. 891 ( CAS, PMA) ; Trocha Río Frío hacia la cima más allá de Rancho Tuque (3h de camino), Trocha de Rancho Frío hacia Rancho tigre, hacia la cima de Cerro Pirre , 1000–1200 m, 21 September 1989, Aranda et al. 908 ( CAS, PMA) ; Cerro Pirre, ridgetop near Rancho Plástico , 1200 m, 10 July 1977, Folsom 4247 ( CAS) ; Top of ridge leading to Cerro Pirre, Area near Rancho Plastico , 1200–1400 m, 13 November 1977, Folsom et al. 6278 ( CAS) ; SW ridge leading to Alturas de Nique on the Colombian border, 800–900 m, 28 December 1980, Hartman 12352 ( CAS) ; Middle slopes on W side of Cerro Pirre , 7°56’N, 77°45’W, 800–1050 m, 29 June 1988, Croat 68937 ( CAS) . Panamá: High point of ridges Sof Ipetí, 5–6 hours walk from Chocó Village, Serranía de Maje , 8°45’N, 77°30’W, 650–800 m, 31 March 1982, Knapp et al. 4495 ( CAS) . San Blas: Cerro Habú, trail from Río Sidro , 9°23’N, 78°49’W, 427–610 m, 20 December 1980, Sytsma et al. 2777 ( CAS) . Veraguas: Valley of Río Dos Bocas along road 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 27624 ( NY) ; ca 10 km NW of Santa Fe, on road to Calovebora (Panama Hwy 35), at first branch of Río Santa Maria , 4 August 1975, Mori & Bolten 7640 ( US) ; Beyond Río Tres Brazos, 9 km from Santa Fe , 24 July 1974, Croat 25590 ( MO) ; Beyond Río Tres Brazos, along steep descent 11 km beyond Santa Fe , 650 m, 24 July 1974, Croat 25639 ( MO) ; 2–5 km NW de Santa Fe por el camino a Río Calovebora , 500–700 m, 26 August 1984, Hernández et al. 748 ( CAS) ; Vicinity of Santa Fe, along dirt road between Santa Fe and Río San Luis, past Escuela Circlo Alto de Piedra , ca 5 mi Nof school, 8°33’N, 81°8’W, 670 m, 28 June 1987, Croat 66961 ( CAS) . PERÚ. Cuzco: (Quispicanchi), Camanti, Maniri , 8 km Wde Quincemil, Alo largo de la quebrada Yanamayo , 13°17’S, 70°48’W, 720 m, 16 July 1990, Timaná & Astete 610 ( MO) . Huánuco: (Pachitea), Pucallpa region , W part of the "Sira mountains" and adjacent lowland, c, 24 km SE to c, 26 km ESE of Purto Inca. from the beginning of the mountain rain forest next to the "Campamento Pato Rojo" along the mountain crest to the beginning of the

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 137 elfin forest after the "Campamento Peligroso", 9°27’S, 74°46’W, 1350 m, 25 March 1988, Wallnöfer 12-25388 ( CAS) . Pasco : ( Oxapampa ), Valle del Palcazu , Cordillera de San Matías , Camino de Loma Linda , 700 m, 22 July 1981, Foster 4521 (F); ( Oxapampa ), Palcazu , Río Alto Iscazacin , Ozuz to Río Pescado, 10°19’S, 75°16’W, 400–500 m, 12 May 1985, Foster & d'Achille 10125B (F); (Oxapampa), Pichis Valley, San Matias Ridge , 10–12 km SW of Puerto Bermúdez, above Santa Rosa de Chivis , trail to Puerto Nuevo, 10°20’S, 75°0’W, 500–900 m, 8 September 1982, Foster 8640 ( MO) . San Martín : ( Mariscal Cáceres ), Tocache Nuevo , Palo Alto, Al Wdel Puente, 600–700 m, 26 November 1972, Schunke 5602 (F, NY, US); (Mariscal Cáceres), Tocache Nuevo, Cerro Sinsín , 15 km Wof Tacache Nuevo, alongroadto Puerto Pizana, 8°9’S, 76°34’W, 550–580 m, 17 December 1981, Plowman & Schunke 11480 (F); (Mariscal Cáceres), Tocache Nuevo , Cerro de Palo Blanco, márgen izquierda del Río Tocache, 600–700 m, 9 July 1974, Schunke 7256 ( MO) .

Illustration:— Almeda et al. 2007: 555.

Common names and documented uses:— Ecuador: “chignul” used as pitchforks (Aulestia 1276, 1099, MO!, Quelal et al. 694, CAS!, Tipaz 1310, 1118, MO!); “mora” ( Mexia 8428, CAS!).

Habitat, distribution and ecology:— This species is common to occasional in primary and secondary rainforests, cloud forests, swamp forests, and disturbed sites, growing in the understory, in light gaps or near streams in flat areas, from Costa Rica throughout southern Central America south to Colombia, Ecuador, Perú, and with one record from western Brazil ( Fig. 17 ), at 5–2800(–3200) m. In Costa Rica it grows on the Caribbean slope along all the principal Cordilleras, to the Tortuguero plain, and south to the Golfo Dulce region on the Pacific slope. In Panama it is present throughout the country, both on the Pacific and Atlantic slopes. In Colombia it occurs on the Pacific slope of the Western Cordillera of the Andes, encompassing the Biogeographic Chocó Region, with fewer collections from the Central Cordillera. In Ecuador it is common throughout ranging south to Perú. In this study M. variabilis is reported for the first time in Brazil, where it occurs in the western region of Acre close to the Peruvian border.

Manakins, Tanagers, Thrushes, the orange-billed Sparrow ( Arremon aurantiirostris ) and Myadestes melanops (in a Costa Rican premontane wet forest) have been reported to feed on M. variabilis berries ( Stiles & Rosselli 1993). The White-ruffed Manakin ( Corapipo altera ) has also been reported to feed on M. variabilis fruits ( Boyle 2006).

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

Etymology:— The specific epithet refers to the variable colors of the mature fruits of this species.

Discussion:— Miconia variabilis has a distinctive puberulent indumentum of dendritic trichomes on the petals abaxially, petiolate 5-plinerved leaves, 5-merous flowers, tuberculiform exterior calyx teeth that do not project, and oblong-obovate anthers that are emarginate at the apex. Miconia sessilis has similar vegetative and floral pubescence, and anther shape, but differs in having 4-merous flowers and leaf bases that are conspicuously decurrent on the petioles. The indumentum is also somewhat similar to that in M. bensparrei , M. incerta , and M. palenquensis , which collectively differ in their sessile, amplexicaul leaves. Miconia variabilis is the most basal species within the Variabilis subclade.

This species is fairly consistent in indumentum and floral characters but the pubescence varies in quantity without any apparent correlation with elevation or other characters. The color of mature berries is highly variable in this species as well. There are at least two variants across its geographic and elevational range. The first variant has berries that are consistently white at maturity. These plants which are probably found throughout the species range are more abundant on the Atlantic slope of Costa Rica south to the Pacific slope in Panama (Bocas del Toro down to Darién) with populations also in Colombia, Ecuador and Perú, at 5–2800 m. It is difficult to delimit the exact geographic extent of this variant because collectors do not always record mature berry colors on specimens labels. The other variant includes individuals in which mature berries undergo a change of color through the maturation process. The fruits are initially green or green-white, then turning yellow, orange and in some cases red-purple. Berries with different colors can be seen on the same plant. The geographic distribution of this variant is clearer than the one with white berries. It occurs on the Pacific coast of Costa Rica (Parque Nacional Corcovado), south to Panama in the Atlantic province of San Blás to Darién. In Colombia, Ecuador and Perú this is the predominant variant, at 5–1325(–3200) m. From these distributions it appears that in Central America the two variants are clearly separated, but this is not clear in South America where both variants may be sympatric.

The only record from Brazil was reported to have white berries at maturity. These two variants appear to represent divergent lineages within the same species. It would be of interest to learn whether the two berry color variants are selectively favored by the bird species that feed on them.

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There is one collection (Cuatrecasas 17033, F!, NY!, US!) that probably represents a rare variant from lowland Pacific Colombia (Valle), at 5–80 m. This specimen has smaller leaves than usual (mostly 6 × 2.2 cm), larger exterior calyx teeth (to 0.8 mm), and floral parts that are smaller overall than the norm. This collection is otherwise identical to M. variabilis in the puberulent indumentum and leaf venation, which are consistent diagnostic characters of this species .

As discussed under M. laxivenula , the vegetatively glabrous population from eastern Ecuador described as Ossaea robusta f. glabrata Markgraf , will probably remain unknown until the “Pacapaca” region is more thoroughly explored ( Wurdack 1973a).

The only illustration of this species ( Almeda et al. 2007), is accurate except that the seed looks cochleate and the testa appears verrucose. Cochleate seeds are not present in the tribe Miconieae , and in M. variabilis they are clearly ovoid, with the testa seemingly smooth and glossy, the verrucose microrelief is only visible with a SEM.

Conservation status:— This species would be considered Vulnerable VU B2ab(iii) according to IUCN criteria ( AOO). However, because it occurs in many protected areas, a status of Least Concern LC is justified. It is protected in Brazil in the Serra do Divisor National Park (Acre) . In Colombia it is protected in the Isla Gorgona National Park (Cauca); in La Planada Reserve (Nariño); and in in the Farallones National Park (Valle). In Costa Rica it is protected in the Volcán Irazú National Park (Cartago) ; in the Guanacaste National Park (Guanacaste); in La Selva Biological Station (Heredia) ; in the Braulio Carrillo National Park (Limón) ; in the Golfito Wild Life Reserve and in Corcovado National Park (Puntarenas) . In Ecuador it is protected in the Awá Indigenous Reserve (Carchi and Esmeraldas) ; in the Sumaco-Napo-Galeras National Park and in the Jatún-Sacha Ecological Reserve (Napo) ; in ENDESA Forest Reserve (Pichincha) ; and in the Cuyabeno Scientific Station (Sucumbíos) .

Figures

FIGURE 17. Distributions of Miconia renatoi, M. sessilis, M. spiciformis, and M. variabilis.