Miconia neomicrantha Judd & Skean (1991: 62)

Gamba, Diana & Almeda, Frank, 2014, Systematics of the Octopleura Clade of Miconia (Melastomataceae: Miconieae) in Tropical America, Phytotaxa 179 (1), pp. 1-174 : 97-106

22. Miconia neomicrantha Judd & Skean (1991: 62) . Basionym: Melastoma micranthum Swartz (1788: 71) . Octopleura micrantha (Sw.) Grisebach (1860b: 260) . Ossaea micrantha (Sw.) Macfadyen (1850: 49–50) . Type: JAMAICA. Swartz s.n. (holotype: S-internet image!; isotype: LD-internet image!). Nec Miconia micrantha Cogniaux (1896: 12) nec M. micrantha Pilger (1905: 173 ; M. witii Ule (1915: 367)) nec M. micrantha Pittier (1947: 27 ; M. tabayensis Wurdack (1971: 359)) .

Sagraea neurocarpa Naudin (1852: 94) . Type: not found among digital images online but surely at P. In the protologue, Naudin cited Goudot s.n. and Bonpland s.n. from Colombia and Swartz s.n. from Jamaica.

Ossaea caudata Cogniaux (1891a: 1066) . Type : ECUADOR. Prope Quito , September, Jameson 390 (holotype: BR-internet image!; isotypes: E-internet image!, BM-internet image!).

Ossaea tetragona Cogniaux (1891b: 265) . Type: COSTA RICA. Chemin de Carrillo , versant Atlantique, 300 m, 25 November 1890, Biolley 3148 (holotype: BR-internet image!; isotypes: BR-2 sheets-images!).

Octopleura rubescens Triana (1871: 146) . Ossaea rubescens (Triana) Cogniaux (1891a: 1067) . Type: COLOMBIA. Prov. Barbacoas: Arrastradero, 10 m, April 1853, Triana 6258/73 (holotype: BM-internet image!; isotypes: BR-internet image!, COL-internet image!).

Shrub or small tree 1–7 m tall with erect, open branching, bark green to light brown. Upper internodes roundedquadrate to quadrate 1–4.2(–6.6) cm long, cauline nodes terete, nodal line frequently present as a ridge, occasionally not very evident or absent. Indumentum on branchlets, petioles, adaxial surface of young leaves, surface, primary, secondary, tertiary and higher order veins abaxially, inflorescence axes, bracts and bracteoles abaxially, pedicels, hypanthia, calyx lobes, calyx teeth, and petals abaxially copiously to sparsely covered with with hyaline or brownish stellate lepidote trichomes 0.15–0.25 mm long with only partially fused radii, occasionally completely replaced by 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 thin petiole (0.5–) 1–5.5 cm, narrowly sulcate adaxially; blades 6.5–20 × 5–8 cm, elliptic to elliptic-ovate, the base acute to narrowly obtuse, occasionally rounded, the margin entire to obscurely and distantly undulate-serrulate, the apex gradually short- to longacuminate to aristate-caudate, chartaceous; mature leaves adaxially glabrescent with the primary, secondary, tertiary and higher order veins glabrous; abaxial surface along with the primary, secondary, tertiary and higher order veins glabrous with age; 5-nerved or 5-plinerved, including the tenuous marginals, innermost pair of secondary veins diverging from the primary vein symmetrically or asymmetrically 0.3–1.5 cm above the base, with a membrane-like structure or with trichomes like the general indumentum in the spaces where the innermost pair of secondary veins diverge from the primary (mite-domatia?), areolae 0.3–0.4 mm, adaxially the primary, secondary, tertiary and higher order veins flat, abaxially raised and terete. Inflorescences a pseudolateral dithyrsoid 3–6 cm long, including a rounded-quadrate to quadrate peduncle 0.75–1.25 cm long or sessile, divaricately and highly branched from the peduncle apex or the base, borne on the upper foliar nodes, the rachis green; bracts 0.2–0.4 × 0.1–0.3 mm, narrowly triangular to oblong-linear, pale green, adaxially glabrous, tardily deciduous at anthesis; bracteoles 0.3–0.5 × 0.2–0.3 mm, narrowly triangular, pale green, abaxially glabrous, deciduous to subpersistent in fruit. Flowers 4-(5-) merous, sessile. Hypanthia at anthesis 1.6–1.7 × 1.3–1.4(–1.6) mm, free portion of hypanthium 0.5–0.6(–0.8) mm long, globose to urceolate, bluntly 8-ribbed, green-yellowish, ridged on the inner surface, sparsely beset with sessile glands, the torus sparsely glandular and moderately scaly adaxially. Calyx open in bud and persistent in fruit, green; tube 0.3–0.4 mm long, with the same vestiture as the torus adaxially and as the hypanthium abaxially; lobes 0.2–0.4 × 1–1.3 mm, depressed-rounded, erect to flange-like, the margin vaguely undulate, the apex obtuse; exterior calyx teeth 0.15–0.4 mm long, deltoid, merely thickened and inconspicuous,

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Phytotaxa 179 (1) © 2014 Magnolia Press 97 inserted on the apical half of the lobes and not projecting beyond them. Petals 3–4(–5) × 0.8–1.5(–1.7) mm, lanceolate, the margin entire, the apex bluntly acute to acuminate, white to pinkish, occasionally with red spots or red distally, glabrous adaxially, glabrescent abaxially, reflexed at anthesis. Stamens 8; filaments 1.3–2 × 0.2–0.23 mm, white to cream, glabrous; anther thecae 1–1.5 × 0.25–0.3 mm, linear-oblong, obtuse to slightly emarginate at the apex, opening by (one–)two dorsally inclined pores 0.1–0.2 mm in diameter, cream to yellow, brown-red with age; connective yellow to orange, its prolongation and appendage 0.4–0.9 mm long, the appendage subulate to oblong, bluntly acute to rounded at the apex, copiously gland-edged, the glands rounded and stalked, the stalks linear to subulate. Ovary 4-locular, 2/3 to completely inferior, 1.3–1.6 mm long at anthesis, the apical collar absent, the apex 0.2–0.3 mm in diameter, conic-truncate, sparsely glandular-puberulent, the trichomes to 0.1 mm long; style 3.5–4.5 mm long, narrowed distally (i.e. tapering), white, glabrous to somewhat glandular; stigma capitellate. Berries 4–5 × 4–5 mm when dry, globose-oblate, green turning white when fully ripe, the hypanthial indumentum persistent at maturity. Seeds 0.32–0.43 × 0.13–0.2 mm, ovoid, angled, brownish; lateral and antiraphal symmetrical planes ovate, the highest point toward the chalazal side; raphal zone ovate, nearly as long as the seed, usually ventrally expanded toward 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, lowdomed to nearly flat, microrelief verrucose. Chromosome number: n=17.

Additional specimens studied:— BELIZE. Middlesex: 200 ft, 12 July 1929, Schipp 235 ( US). Toledo: Bladen Nature R., Southern Maya Mountains, trail between Roochire Selipan archeological site and "AC Camp", 16°27’54"N, 88°52’49"W 450 m, 2 June 1997, Davidse 36978 ( CAS, MO); Bladen Nature R., Southern Maya Mountains, W Snake Creek, 16°27’24"N, 89°1’1"W, 500 m, 28 May 1997, Holland & Kid 94 ( CAS, MO); On hillside beyond Pate's Camp, Edwards Road beyond Columbia, 28 November 1950, Gentle 7174 ( US). COLOMBIA. Antioquia: (Caramanta), Hojas Anchas, Cerro Viringo, 9.8 km de Caramanta hacia Supía, Cordillera Occidental, 5°31.8’N, 75° 40.69’W, 2140–2420 m, 16 October 1988, Betancur et al. 1098 ( HUA, MO); (Caldas), La Corrala, Finca "La Zarza", 2240 m, 30 March 1985, Palacio & Vallejo 6 ( NY); (Fredonia), Cerro Bravo, Finca de los Betancur, camino hacia la toma de agua, 1770–1900 m, 7 June 1992, Fonnegra et al. 4432 ( HUA, MO); (Medellín), La Cascada, Carretera a Rionegro vía Santa Helena, 6°14’N, 75°32’W, 1500–1700 m, 11 August 1991, Gómez et al. 58 ( MO); (Dabeiba), 32 km SE de Mutatá en la via Mutatá-Dabeiba, Río Chever, sitio Chever, 280 m, 5 August 1987, Callejas et al. 5065 ( HUA, MO, US); (Salgar), Camono de ascenso a Cerro Plateado, quebrada La Liboriana, 2200–2400 m, 3 November 1985, Franco et al. 2284 ( MO). Caldas: (Manizales), Alrededor de la cañada entre helados La Fuente y tuburio, 11 June 1985, Escobar-Cardona 35 ( US); (Salento), Río Santa Rita, 1600–1800 m, 26 August 1922, Killip & Hazen 10127 ( NY, US); Finca La Oculta, 2020 m, 14 January 1983, de Escobar & Uribe 3640 ( HUA, US); (Manizales), Monteleón, Cordillera Central, 2250 m, 16 February 1985, de Fraume et al. 363 ( US). Chocó: (Bahía Solano), Mecana, On Pacific Coast, Quebrada Resquiata, 6°16’N, 77°21’W, 2 m, 2 January 1984, Juncosa 1640 ( MO, US); Carretera Ansermanuevo-San José del Palmar, Alto del Galápago, Límite con el Valle del Cauca, 2000 m, 19 February 1977, Forero et al. 2910 ( COL, MO, US); Alongroadbetween Pueblo Rico (Risaralda) and Istmina (Chocó), along Quebrada Antón, 15 km Wof Santa Cecilia, 6 km Wof Chocó-Risaralda border, ca 20 km Eof Playa del Oro, 5°20’30"N, 76°13’45"W, 240 m, 22 February 1990, Croat 70914 ( MO); Río Quito, 5 km above junction with Río Atrato, 22 April 1982, Prance 28029 ( NY). Cundinamarca: (Medina), Farallones de Medina, 4°35’09”N, 73°23’06”W, 1000 m, 25 February 1997, Mendoza 2160 ( COL, FMB). Magdalena: (Santa Marta), 200 ft, November 1898, Smith 457 (F, MO, NY, US); (Santa Marta), Sierra Nevada de Santa Marta, Alto Río Buritaca, Alto de Mira, por el camino a la Quebrada Julepia, 11°5’N, 73°48’W, 700–1100 m, 13 July 1989, Madriñán & Barbosa 198 (F, GH, MO, NY); (Santa Marta), Sierra Nevada de Santa Marta, Alto Río Guachaca, Escuela La María (Macho Solo), al margen izquierdo del Río Guachaca, 11°4’N, 73°55’W, 800 m, 11 July 1989, Madriñán & Barbosa 130 (F, GH, MO); (Santa Marta), Sierra Nevada de Santa Marta, Alto Río Buritaca, Alto de Mira, por el camino a la Cascada del Caño Negro , 11°5’N, 73°48’W, 1000–1100 m, 14 July 1989, Madriñán & Barbosa 230 ( GH, MO). Nariño: (Ricaurte), 1300 m, 24 April 1941, von Sneidern A.623 ( US); (Ricaurte), km 200 carretera Tumaco-Pasto, 2000 m, 20 November 1982, de Benavides 4109 ( MO); R.N. La Planada, 7 km above Chucunes (on road between Tuquerres and Ricaurte), along trail to Pialapí, past entrance to La Planada field station, 1°6’N, 77°53’W, 150–200 m, 10 March 1990, Croat 71255 ( MO); Trail to El Hondón, 5–12 km SW of La Planada, 1°4’N, 78°2’W, 1750–1800 m, 6 January 1988, Gentry et al. 60454 ( MO); (Ricaurte), R.N. La Planada, Vertiente occidental, Cordillera Occidental, 1800 m, 22 May 1989, Restrepo 472 ( MO); R.N. La Planada, A 7 km de Chucunes, 1°10’N, 77°58’W, 1800 m, 14 November

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1987, de Benavides 8871 ( MO, PSO) ; R.N. La Planada, Isla de los Osos, 1°10’N, 77°58’W, 1500–1800 m, 1 May 1988, de Benavides 9746 ( MO, PSO); ( Ricaurte ) , Trayecto San Isidro-La Planada , 1°10’N, 77°58’W, 1500–1800 m, 13 February 1988, de Benavides 9246 ( MO, PSO) ; La Planada, Trail to El Hondón, 6–12 km SW of La Planada, 1°4’N, 78°2’W, 1750–1800 m, 5 January 1988, Gentry et al. 60386 ( MO, US) . Norte de Santander: Cordillera Oriental, Región del Sarare , Hoya del Río Margua , cabeceras del Río Negro , ladera Nentre El Amparo y La Mesa, 1400–1700 m, 7 November 1941, Cuatrecasas 12874 ( COL, US) . Putumayo: (Mocoa), San Antonio , Alto Campucana , Finca La Mariposa , Vertiente amazónica de Colombia , 1°12’N, 76°38’W, 1350–1420 m, 20–1 April- May 1994, Fernández 10756 ( COL, US) . Quindio: (Pijao), Arenales , Bosque ubicado en la intersección de las carreteras hacia los Juanes , margen izquierda, 1850 m, 19 November 1987, Arbeláez et al. 2388 ( HUA, US) . Risaralda: (Pereira), La Florida, S.F.F. Otún Quimbaya, 4°44’17"N, 75°34’1"W, 1900 m, 22–28 February 2004, Alzate et al. 2108 (F, HUCO); (Pereira) , Bajada de "La Pastora" a "El Cedral", 17 October 1989, Rangel et al. 5961 ( MO); (Pereira) , Bajada de "La Pastora" a "El Cedral", 17 October 1989, Rangel et al. 5970 ( MO); (Pereira) , El Cedral, Trocha a El Silento, 20 km de Pereira, 2140 m, 16 October 1989, Rangel et al. 5894 ( MO) . Tolima: (Santa Isabel), Vereda Guaimaral , 4° 35’58.7”N, 74°49’44”W, 2100 m, 9 May 2007, Pava & Esquivel 40 ( COL, TOLI); (Toche) , 2300 m, 20 May 1942, von Sneidern 3085 ( US) . Valle: (Cali), Cordillera Occidental , vertiente oriental, Hoya del Río Cali , vertiente derecha: Quebradahonda , arriba de La Glorieta, camino a Miralindo , 2100–2250 m, 31–1 October-November 1944, Cuatrecasas 18434 (F, NY, US); ( La Cumbre) , Bitaco, Chicoral, Detrás de Dapa , en los nacimientos del Río Bitaco , vertiente del Pacífico , bosque arriba del cultivo de Té, 2000 m, 14 January 1990, Ramos 2412 ( CAS, CUVC); ( Cali ) , La Elvira, Finca Los Sauces , km 4 on the road to Dapa (km 18 on the road from Cali to Buenaventura ), near Quebrada Aguas Claras , Cordillera Occidental , vertiente oriental, 3°32.424’N, 76°36.107’W, 1890 m, 2 February 2011, Almedaet al. 10223 ( CAS, COL, CUVC); (Dagua) , Queremal, P.N.N. Farallones, Along the road from vereda El Danubio to Campamento EPSA ( Yatacue ), Cordillera Occidental , vertiente occidental, 3°35.277’N, 76°52.352’W, 580 m, 7 February 2011, Almedaet al. 10284 ( CAS, COL) ; La Laguna, Cordillera Occidental, vertiente occidental, Hoya del Río Sanjuniquín , lado izquierdo, 1250–1400 m, 10–20 December 1943, Cuatrecasas 15482 (F, NY); ( Cali ) , Bosque de San Antonio , Wof Cali, near television tower, 1950–2050 m, 15 July 1984, Gentry et al. 48168 ( MO); (Sevilla) , Vía Sevilla-Barragán, a 28 km de Sevilla, finca Alsanieves , 2400 m, 11 October 1981, Silverstone-Sopkin 783 ( MO, TULV) ; Alto Mercedes - Carretera al March, 2200 m, 1–30 September 1939, Dryander 2433 ( US); ( Cali ) , La Elvira, Finca Zingara , km 18 de la carretera Cali-Buenaventura , km 4 vía a Dapa , cordillera Occidental , 3°30’N, 76°34’W, 1900 m, 18 April 1994, Giraldo-Gensisni 280 ( TULV, US); ( La Cumbre) , 1800–2200 m, 14–19 May 1922, Killip 5570 ( NY, US) ; Costa del Pacífico, Río Cajambre, Barco , 5–80 m, 21–30 April 1944, Cuatrecasas 17113 ( US) . COSTARICA. Monteverde Cloud Forest R., Border of Guanacaste, Puntarenas & Alajuela Provinces, Cordillera de Tilarán , along the Sendero El Valle , 1550–1640 m, 7 March 1986, Almeda & Anderson 5433 ( CAS) . Alajuela: (San Carlos), Villa Quesada, Caribbean Coast , 840 m, 28 February 1939, Smith 1732 ( NY) ; Approx, 17 km N, of Río Naranjo on the road to Upala , 300 m, 5 February 1976, Utley & Utley 4017 ( CAS, US); ( Alfaro Ruiz ) , La Peña, 1400 m , 2 January 1941, Smith 10019 ( NY); (Upala) , San Cristóbal, C.B. Guanacaste, Rincón de la Vieja, Cuenca del Pizote, Sendero El Pinal , 10°52.00’58.474"N, - 85°24’5.946"W, 500 m, 1 July 1998, Espinoza 1747 ( NY) . Bocas del Toro: On Chiriquí trail between Buena vista coffee finca and Cerro Pilón , 17 April 1968, Kirkbride & Duke 685 ( NY) . Cartago: Near and along crest of ridge Sof Alto Patillos (and a continuation of the latter), ca 6 km SE of Tapantí, 9°43’N, 83°46.5’W, 1700–1840 m, 9 October 1986, Grayum & Herrera 7727 ( CAS, MO) . Guanacaste: 10 km E of Quebrada Grande , 580 m, 18 August 1968, Davidse & Pohl 1220 ( ISC, US) ; La Tejona , Nof Tilarán, 600–700 m, 25 January 1926, Standley & Valerio 45843 ( US) . Heredia: Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , W Line, Quebrada el Sura, 5 m, Nof creek, 100 m, 30 September 1982, McDowell 276 ( NY, US); (Sarapiquí) , Z.P. La Selva, Cuenca del Sarapiquí , 10°26’0"N, 84°2’0.0001"W, 100 m, 8 February 2003, Boyle et al. 225 ( INB, MO, NY); (Sarapiquí) , La Selva de Sarapiquí , 200 m, 4 February 1966, Schnell 416 ( CR, US) ; La Selva, near Puerto Viejo , El Salto bridge, 90 m, 21 April 1972, Opler 749 ( US) ; Finca La Selva, The OTS Field Station on the Río Puerto Viejo just Eof its junction with the Río Sarapiquí , 100 m, 26 July 1980, Hammel 9330 ( CAS, NY) . Limón: ( Barra de Colorado ), 0 m, 3 March 1966, Schnell 530 ( US); (Talamanca) , Amubri, Camino entre Amubri y Soki, Siguiendo el Río Ñabri hacia Alto Soki , 9°29’50"N, 82°59’10"W, 150 m, 1 July 1989, Herrera 3096 ( CAS, CR) ; Drenaje de Río Valle Estrella, Montaña Andromeda, 26–28 October 1951, Shank & Molina 4540 ( US); ( Barra de Colorado ) , 0 m, 5 March 1966, Schnell 583 ( CR, US); (Baja Talamanca) , Crece en playón asoleado del Río Lari en su confluencia con el Río D'pari ,

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Phytotaxa 179 (1) © 2014 Magnolia Press 99 700–800 m, 19 March 1964, Jiménez 1865 ( CR, F, NY); A 2 millas de Diamantes, Guápiles , 150 m, 9 August 1963, Jiménez 1048 (F, NY); (Siquirres) , R.B. Barbilla, Cuenca del Matina, Sendero Valle Escondido, 9°59.00’3"N, 83°28’W, 400–500 m, 21 July 1999, Mora-Castro 424 ( NY). Puntarenas: Above coffee fincas along Río Coto Brus , near Coton, 23 km Nof La Union (on Panama border), 9 August 1974, Croat 26606 ( US); Monteverde R., 1550 m, 13 January 1979, Lumer 1003 ( NY); Monteverde, Lower community, 1300 m, 29 November 1984, Haber 1082 ( NY); (Golfito), P.N. Corcovado, Península de Osa , Estación Agujas, alrededores de la estación, 8°31.00’52.786"N, - 83°26’0.991"W, 300 m, 6 November 1999, Mora et al. 653 ( NY); (Coto Brus) , R. de la Biósfera de la Amistad, cerca Estación Biológica Las Alturas de Coton , 8°57’0.3"N, 82°49’56.8"W, 1455–2100 m, 8 July 1994, Kress & Sawyer 94-4956 ( US) ; Lumber road along Fila Tigre, Sand Eof Las Alturas between Río Cotón and Río Quebrada Nochebuena , 1400 m, 14 February 1991, Almeda et al. 6587 ( CAS) . San José: Carara R., Hills at SW part of Montañas Jamaica ca 2.5 km NE of Bijagual de Turrubares, (lower slopes and small creek draining to Río Tarcolitos system) to 575 m, (top of ridge), 9°45’N, 84°33.5’W, 460–520 m, 26 June 1985, Grayum et al. 5497 ( CAS, MO); ( Alto La Palma ) , 1500 m, 17 October 1969, Schnell 1085 (F). ECUADOR . Azoques: On highway from Guayaquil to Machala , 1 km Sof Ponce Enriques & 2 km in from road on low hill, 70 m, 9 October 1979, Dodson et al. 9187 (F, MO) . Bucay: Chimbo River Valley, In western Cordillera , lower valley of Chimbo river , 1000 m, 1–30 June 1934, Rimbach 189 (F) . Carchi: Ascent up main river, Medium Cerro Golondrinas beginning at principal stream division, 0°52’N, 78°7’W, 2030 m, 2 December 1987, Hoover 2228 ( CAS, MO) ; Environs of Maldonado , 1450–1650 m, 1 June 1978, Madison et al. 4869 (F, QCA, US); (Mira) , Ndel Carmen, Camino a Chical , 0°17’N, 78°13’W, 2000–2200 m, 10 February 1992, Palacios et al. 9774 ( MO, QCNE); (Espejo) , Faldas del Cerro Golondrinas Hembra , 0°51’N, 78°7’W, 2300–2400 m, 20 August 1994, Palacios 12737 ( MO, QCNE); (Maldonado) , 1500 m, 1 March 1974, Harling & Andersson 12394 ( US) ; 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 3561 ( MO) . Cotopaxi: (Tenefuerste), Río Pilalo, km 52–53, Quevedo-Latacunga , 750–900 m, 2 October 1981, Dodson & Dodson 12002 ( MO, US); ( La Mana ) , Alongroadbetween Guayacán (13.1 km NE of La Mana ) and Montenuevo (N of Pucayacu ), 0.8 km Nof Pucayacu, 0°41’S, 79°6’W, 760 m, 20 March 1992, Croat 73259 ( MO) ; 3 km Eof El Palmar on road Quevedo- Latacunga , 800 m, 5 April 1980, Dodson & Gentry 10256 ( MO); (Latacunga) , Quevado, Tenefuerste, Río Pilalo , km 52–53, 750– 1300 m, 7 February 1982, Dodson & Gentry 12281 ( MO) . El Oro: 20 km W Piñas on road to Machala , 3°43’S, 79°49’W, 600 m, 3 February 1991, Kessler 2662 ( US) ; 15 km NW of Paccha on the road to Pasaje , 3°30’S, 79°40’W, 1800 m, 29 September 1983, Brandbyge 42364 ( MO, NY, QCA) ; Return trip from Hacienda Ingenio to Piedras over Cordillera Marcabeli, 666 ft, 21 June 1943, Little 6679 (F, US) ; Hacienda Daucay , 3°29’S, 79°45’W, 600 m, 12 October 1993, Cornejo 497 ( MO) . Esmeraldas: (San Lorenzo), R. Etnica Awá, Centro Guadalito , 1°15’N, 78°40’W, 80 m, 20–29 July 1992, Aulestia et al. 176 ( MO, QCNE) ; Lita-San Lorenzo road, 18 km Wof Río Lita Bridge (on old road below Lita), 6.6 km Wof bridge over Río Chuchubí , site near Alto Tambo , 0°52’11"N, 78°27’16"W, 425 m, 10 July 1998, Croat et al. 82628 ( MO) ; Cerro de Río Bravo de Cayapas , 0°41’N, 78°56’W, 250 m, 1 September 1980, Holm-Nielsen et al. 25559 ( MO, US); (San Lorenzo) , R. Etnica Awá, Centro Guadalito , 1°15’N, 78°40’W, 80 m, 20–29 July 1992, Aulestia et al. 47 ( MO, NY, QCNE) . Guayas: ( Teresita ), 3 km Wof Bucay, 270 m, 5–7 July 1923, Hitchcock 20414 ( NY, US) . Imbabura: ( Lita ), 501 m, 28 April 1949, Acosta Solís 12297 (F); (Cotacachi) , R. Privada Intag, Parroquia Intag, Comuna Santa Rosa de Pucurá , 0°22’N, 78°28’W, 1850 m, 6 March 2002, Caranqui et al. 520 ( MO, NY); (Cotacachi) , Carretera de Cotacachi a Nangulbí y Apuela , 0°20’N, 78°35’W, 3000 m, 11–14 August 1990, Rubio & Quelal 603 ( MO, QCNE) . Loja: (Loja), El Limo, Alamor-Cazaderos road, El Limo , 1000 m, 3 April 1980, Harling & Andersson 17842 (F, MO, NY, US) . Los Ríos: Bosque de Oro, Hacienda Las Balsas near Montalvo , 300–400 m, 27 July 1962, Javita & Epling 227 ( US) ; Río Palenque B.S., km 56 road Quevedo-Santo Domingo , 150–220 m, 4 November 1979, Schupp 82 (F) ; Río Palenque B.S., Halfway between Quevedo and Santo Domingo de los Colorados , 200 m, 6 February 1974, Gentry 9682 ( MO, US) ; Río Palenque B.S., km 47, Santo Domingo-Quevedo , 150–220 m, 2 August 1984, Dodson & Gentry 14826 ( MO) ; Río Palenque B.S., Halfway between Quevedo and Santo Domingo, 200 m, 4 October 1976, Gentry & Dodson 17986 ( MO) ; Trails of Río Palenque Biological Station , 0°35’37"S, 79°21’44"W, 215 m, 6 February 2009, Stern & Tepe 372 ( NY) ; Río Palenque B.S., Halfway between Quevedo and Santo Domingo, 200 m, 9 October 1976, Gentry & Dodson 18041 ( MO) ; Centinela ridge area, 12.5 km Eof Patricia Pilar, 1400 ft, 28 July 1980, Hansen et al. 7737 ( MO) . Manabi: Machalilla N.P., Zona de San Sebastián , ridgetop, 1°36’S, 80°42’W, 600–700 m, 21 January 1991, Gentry et al. 72518 (F, MO) ; Machalilla N.P., Zona de

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San Sebastián, ridgetop, Transect #6–8, 1°36’S, 80°42’W, 550 m, 21 January 1991, Gentry & Josse 72543 (F, MO); ( Jama ) , Cordillera de Jama (coastal range), Cerro Nueve , 15 km Eof Jama, Nof Río Jama, below microwave tower, 0°16’S, 80°12’W, 600 m, 16 December 1998, Neillet al. 11585 ( MO, QCNE) . Morona-Santiago: Loma San José Grande-Sendero San José-Río Bomboiza, Transecto 50 × 2 m, 2°38’S, 78°27’W, 1700–1750 m, 19 April 1991, Cerón 14330 ( MO); (Tumbes) , 17–18 km N, of Gualaquiza on road to Indanza , 1700–1800 m, 17 April 1985, Harling & Andersson 24248 ( CAS, NY) . Napo: (Archidona), Comunidad de Pacto Sumaco , sector SE, 0°38’56"S, 77°35’49"W, 1550 m, 25 April 1997, Alvarez et al. 1968 ( MO, QCNE) ; Carretera Hollín-Loreto, km 40–50, Alrededores de comunidad Huamaní y del Río Pucuno , 0°43’S, 77°36’W, 1200 m, 10–22 October 1988, Hurtado 606 ( MO); (El Chaco) , Margen derecha del Río Quijos, Finca "La Ave Brava" de Segundo Pacheco, 0°12’S, 77°39’W, 1800–1900 m, 7–10 September 1990, Palacios 5337 ( MO); (El Chaco) , Proyecto hidroeléctrico Coca Punto ST3, Margen derecha del Río Quijos , ca 10 km al Sde Reventador, 0°11’S, 77°39’W, 1500 m, 3–5 October 1990, Palacios 5823 ( MO, QCNE) . Orellana: (Orellana), Comunidad Indillama , a 30 min en deslizador del Puerto Pompeya aguas abajo por el Río Napo , 0°26’S, 76°31’W, 280 m, 6 May 2004, Reyes & Carrillo 538 ( MO) . Pichincha: Reserva ENDESA, 9 km Nof km 113 on Quito–Pto. Quito Highway , 0°5’N, 79°2’W, 750 m, 15 July 1986, Croat & Rodríguez 61491 ( US) ; Carretera Quito–Pto Quito , km 113, 10 km al Nde la carretera ppal, 0°5’N, 79°2’W, 800 m, 27 December 1983, Balslev & Balseca 4753 ( CAS, NY, QCA) ; Wet luxuriant forested slopes of barrancos between km 37 and 50 along Río Saloya (between Volcán Atacaso and Volcán Pichincha), 1830–2430 m, 22–23 April 1943, Steyermark 52584 (F, NY) ; Carretera Quito-Pto Quito, km 113, 10 km al Nde la carretera ppal, 0°5’N, 79°2’W, 800 m, 18 February 1984, Betancourt 93 (F, MO, NY); ( Quito ) , R. Ecológica Maquipucuna, Parroquia Nanegal, 0°10’N, 78°40’W, 1250 m, 9 January 1992, Quelal 90 ( MO, QCNE); ( Quito ) , El Recreo, Eggers 15635 (F); Via Calacali-Nanegalito , 0°3’N, 78°35’W, 2100 m, 24 January 1989, Palacios & van der Werff 3605 ( CAS, MO) ; R. Florística-Ecológica "Río Guajalito", Km 59 de la carretera antigua Quito-Sto Domingo de los Colorados , 3.5 km al NE de la carretera, estribaciones occidentales del volcán Pichincha , 0°13’53"S, 78°48’10"W, 1800–2200 m, 22 June 1985, Jaramillo & Zak 7779 ( MO, NY, US); (Santo Domingo de Colorado) , 0.5 km Nof town, 4 April 1943, Little 6166 (F, NY, US) ; km 40–51 on road Santo Domingo de los Colorados-Quito, Forested slopes along Río Pilatón , 0°22’S, 78°55’W, 1100–1400 m, 14 June 1973, Holm-Nielsen et al. 7165 ( NY, US) ; On ridge ca 10 km Eof Patricia Pilar, 0.3°S, 79.16°W, 300 m, 20 July 1978, Webster et al. 22772 ( MO) ; R. Florística-Ecológica Río Guajalito, km 59 de la carretera antigua Quito-Sto Domingo de los Colorados , 3.5 km al NE de la carretera, estribaciones occidentales del volcán Pichincha , 0°13’53"S, 78°48’10"W, 1800–2200 m, 29 June 1985, Jaramillo & Zak 7884 ( MO, NY) ; Las Palmeras, Quito-Sto Domingo old road, ca 59 km WSW of Quito, Pablo Feret trail above Río Guajalito , 0°18’S, 78°43’W, 1900–2000 m, 16 December 1990, Luteyn & Berg 14374 ( MO, NY); ( Maquipucuna ) , 5 km Eof Nanegal, Transect 3, 0°7’N, 78°37’W, 1550 m, 10 February 1991, Gentry & Valencia 73117 ( MO, US); ( Quito ) , Parroquia de San José de Minas, Sector Las Palmas, margen derecha del Río Cambugán , propiedad del investigador, Parcela permanente 10 × 1000 m, 0° 11’N, 78° 31’W, 1800–2200 m, 1 April 1999, Jiménez et al. 1125 ( MO, QCNE); ( Quito ) , R. Orquideológica El Pahuma, Carretera Calacalí-Los Bancos km 22, 0°1’42"N, 78°37’50"W, 2000 m, 19 October 1999, Rojas et al. 398 ( MO); ( Quito ) , San José de Minas, Cordillera paso Alto , cuenca del Río Cambugán , 0°11’N, 78°30’W, 1980 m, 27 December 2002, Fernández et al. 770 ( MO, QCNE) ; Chiriboga on road from Quito to Santo Domingo de los Colorados, 1800 m, 11 September 1939, Asplund 8703.bis ( MO, NY); ( Maquipucuna ) , 5 km Eof Nanegal, Transect 1, 0°7’N, 78°37’W, 1630 m, 9 May 1990, Gentry et al. 69926 ( MO) ; Road El Paraiso-Saguangal, 3 km from El Paraiso, 0°10’N, 78°46’W, 1500 m, 2 May 1982, Øllgaard et al. 37804 ( MO, NY); ( Nanegalito ) , 0°4’S, 78°41’W, 1600 m, 16 March 1967, Sparre 14886 ( MO) ; R. Florística-Ecológica "Río Guajalito", km 59 de la carretera antigua Quito-Sto. Dimingo de los Colorados , a 3.5 km al NE de la carretera, estribaciones occidentales del Volcán Pichincha , 0°31’S, 78°48’W, 1800–2200 m, 13 August 1986, Zak 1104 ( MO) ; R.N. Río Guajalito, Along main entrance from Quito-Chiriboga-Santo Domingo Road, 3.5 km from highway, departing highway at km 59, 0°13’53"S, 78°48’10”W, 1800 m, 9 February 1992, Croat 75025 ( MO); ( Quito ) , R. Ecológica Maquipucuna, Parroquia Nanegal , 0°10’N, 78°40’W, 1250 m, 9 January 1992, Quelal 8 ( MO, NY, QCNE) ; R. Maquipucuna, Vía Nanegalito-Marianitas , desde la casa por la trocha principal, 0°7’26"N, 78°37’44"W, 1300 m, 25 June 2009, Ulloa et al. 1755 ( MO) ; Along road between Nono and Nanegal NW of Quito , 17 km NW of Nono, (9 km SE of Tandayapa), 2000 m, 4 September 1976, Croat 38853 ( MO) ; Trail along Río Chictoa, tributary of Río Pilatón , 2–8 km NE of Quito-Sto, Domingo Road, 11 km Wof Tandapi, 1500–1600 m, 27 October 1974, Gentry et al. 12142 ( MO, US); R. Forestal ENDESA, Río Silanche : "Corporación Forestal Juan Manuel Durini", km 113 de la carretera

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 101 Quito-Pto. Quito, faldas occidentales, a 10 km al Nde la carretera principal, 0°5.00’N, 79°2’W, 650–700 m, 18 February 1984, Jaramillo 6351 (NY); R. Forestal ENDESA, ca 8 km Ede Pedro Vicente Maldonado, 0°5.00’N, 79°1’W, 780 m, 28 February 2004, Moran et al. 6944 (NY); Old road from Santo Domingo to Quito, 8 km Eof junction with new road (approximately 13 km E from La Union del Toachi), 0°15’55"S, 78°50’60"W, 1950 m, 10 February 2009, Stern & Tepe 386 (CAS, NY); Carretera Quito-Chiriboga-Empalme, km 92, desvio a Mulaulte, en borde de carretera y en quebrada de Mulaulte, 0°15’S, 78°50’W, 1200–1300 m, 13 December 1987, Zak & Jaramillo 3198 (MO, NY); (Quito), Chiriboga, R. Forestal "La Favorita", En la carretera vieja Quito-Sto. Domingo, Al borde del Río Saloya, Transecto de 50X 2 m, 0°12’S, 78°47’W, 1600–1800 m, 8 February 1990, Cerón & Iguago 8631 (MO, QCNE); (Quito), Chiriboga, R. Forestal "La Favorita", Minist. de Agriculatura, En la carretera vieja Quito-Sto. Domingo, Río Saloya, sendero hacia la cuchilla de los Yumbos, 0°12’S, 78°47’W, 1600– 1600 m, 7 December 1989, Cerón et al. 7977 (MO); (Quito), R. Orquideológica El Pahuma, Parroquia Nono, carretera Calacalí-Nanegalito, km 30, sendero hacia la cascada, pendiente 50%, Transecto 4, 0°1’N, 78°38’W, 1940 m, 13–14 January 1996, Freire et al. 1094 (MO); Road El Paraiso-Saguangal, 11 km from El Paraiso, 0°12’N, 78°46’W, 1200 m, 2 May 1982, Øllgaard et al. 37697 (MO, NY); R. Forestal ENDESA, Corporación Juan Manuel Durini, km 113 road Quito-Pto. Quito, about 4 km from take-off from main road, Protected area of the reserve, 0°7.785’N, 79°2.573’W, 530 m, 16 September 2001, Cotton et al. 1725 (MO, QCA); (San Miguel de los Bancos), Nanegalito-Mindo Road, 16.5 km SSW of Nanegalito, 0°1’14"S, 78°24’23"W, 1500 m, 21 July 1998, Croat 82782 (MO, NY); (Quito), R. Orquideológica El Pahuma, Parroquia Nono, carretera Calacalí-Nanegalito, km 30, Sendero hacia la cascada, pendiente 40%, Transecto 1, 0°1’N, 78°38’W, 1885 m, 23–25 December 1995, Freire et al. 1007 (MO). Tungurahua: (Banos), Los Llaganates, Colonia México, A 18 km del Topo, 1°21’S, 78°18’W, 1800–2000 m, 9 March 1995, Vargas & Sandoval 334 (MO, QCNE). Zamora-Chinchipe: ("Oriente") Eslopes of the cordillera, valley of the Río Negro, down to the Río Pailas (on the trail to Mendez), Near junction of rios Pailas and Negro, 6000–7500 ft, 20–24 August 1945, Prieto 4918 (MO, US). GUATEMALA. Alta Verapaz: Mountains Eof Tactic, on road to Tamahú, 1500–1650 m, 9 April 1939, Standley 71192 (NY); (Cubilquitz), 350 m, 1–31 August 1903, von Tuerckheim 8518 (NY, US). Izabal: (Cienaga), On Peten-Guatemala Road, 24 May 1971, Contreras 10832 (US). Peten: (Chinchila), In high forest 6 km from San Luis, W of Sebol Road, 11 October 1966, Contreras 6374 (NY, US); Cadenas Road, Wof km 142, 6 November 1966, Contreras 6529 (US); (La Cumbre), Eabout 4 km on Río Purula Road, in zapotal on top of hill, 19 September 1975, Lundell & Contreras 19879 (US). Quezaltenango: Volcán Zunil, 5600 ft, 7 August 1934, Skutch 964 (NY). San Marcos: (San Marcos), Finca Armenia, Rafael de Cuesta, 5000 ft, 6–7 July 1977, Dwyer 14407 (MO, NY). Suchitepequez: Finca Moca, 3300 ft, 20 October 1934, Skutch 1473 (NY). HONDURAS. Atlántida: Lancetilla Valley, near Tela, 20–600 m, 6–20 December-March 1927, Standley 54163 (US); Lancetilla Valley, near Tela, 20–600 m, 6–20 December-March 1927, Standley 55595 (US); Lancetilla Valley, near Tela, 20–600 m, 6–20 December-March 1927, Standley 52931 (US); Mancetilla Mountain, 100 m, 8 April 1970, Molina & Molina 25614 (F, US); Lancetilla near Tela, cerca del Río Lancetilla, arriba de la Estación Experimental, 15°43.00’0"N, 87°27’30"W, 40–180 m, 7 November 1988, MacDougal et al. 3321 (MO, NY); (Lancetilla), 50 m, 1–30 April 1947, Williams & Molina 13004 (NY). JAMAICA. Blue Mountains: (Portland), Stony River base camp, 1250 ft, 30 December 1973, Morley & Whitefoord 912 (MO). Ecclesdown: (Portland), 1400 ft, 25 August 1962, Adams 11573 (MO). St. Thomas: Along track between House Hill & Cuna Cuna Pass, 2000–2500 ft, 26 December 1969, Proctor 31156 (MO). Trelawny: 5 miles WNW of Troy on Crown Lands Road, 2100–2300 ft, 14 August 1965, Hespenheide et al. 1153 (MO). Westmoreland: Copse Mt, Woods, c 1 mile SW of RAT TRAP, 1200–1300 ft, 23 October 1960, Proctor 21475 (MO). MÉXICO. Chiapas: On the SE side of Volcán Tacaná above Talquian, 2200 m, 16 January 1973, Breedlove & Smith 31674 (DS, NY, US). Oaxaca: (Santa María), ca 6 km al Nde Sta. María por la vereda al Río Verde, al N del Río Corte, 16°56’30"N, 94°41’0"W, 220 m, 19 May 1987, Hernández 2467 (CAS, US). Veracruz: (Catemaco), Cumbres de bastonal, 500 m, 19 November 1974, Cedillo 427 (CAS, MEXU); (Minatitlán), Río Uxpanapa, cerca del límite con Oaxaca, 17°11’N, 94°8’45”W, 180 m, 27 September 1980, Wendt et al. 2774 (US); (Minatitlán), Lomas del Sdel Pob. 11, + 27 km al Ede La Laguna, cañón arriba de "La Cascada", 17°14’N, 94°30’W, 250 m, 26 April 1985, Wendt et al. 4839 (US); (San Andrés Tuxtla), Steep forested slopes on SE side of Volcán San Martín Tuxtla, 13 km N, of San Andrés Tuxtla, 18°33.00’30"N, 95°12’W, 1150–1350 m, 1 July 1982, Diggs et al. 2689 (NY). NICARAGUA. Rivas: Isla de Ometepe, Slopes of Volcán Maderas above coffee plantations above Balgue, 11°28.00’N, 85°31’W, 600–800 m, 14 September 1983, Nee & Téllez 28039 (MO, NY). PANAMA. Bocas del Toro: Changuinola Valley, 18 January 1924, Dunlap 363 (US); (Isla Colon), 1.5 km Nof La

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Gruta , 10 m, 29 January 1989, Peterson 6406 ( MO, US) ; Campamento Forestal , Boca Chica Changuinola ( IRHE) , 22 February 1980, Correa et al. 3547 ( MO, NY, PMA) ; Río Teribe between Q. Huron and Q. Schlunjik , 12 April 1968, Kirkbride & Duke 469 ( MO) ; Vicinity of Chiriquí lagoon, Fish Creek Mts. , 28 April 1941, von Wedel 2354 ( MO, NY) ; Río Terebe foorthills of Cerro Bonyic , 400–800 ft; 11 April 1968; Kirkbride & Duke 420 ( MO, NY) ; 29 July 1940, Wedel 192 ( NY) ; Al SE y NE del campamento Changuinola 1 del IRHE, 19 January 1980, Correa et al. 3386 ( MO, PMA, US) ; Palo Seco R., Carribbean slopes of Cerro Fábrega at foot of "Falso Fábrega" in Palo Seco Reserve, second Nmost tributary (on map) of Culubre River , Pavón Camp , 9°9’54"N, 82°40’45"W, 980 m, 19 March 2005, Monro & Cafferty 4723 ( INB, MEXU, MO, PMA) . Canal Zone: (Barro Colorado Island), Edge of wooded bank of stream near Fuertes House , Wpart of Island , 3 July 1940, Chrysler & Roever 4871 ( NY) . Chiriquí: (Jurutungo), De la Finca de Los Quetzales siguiendo el curso del Río Candela , luego hacia la derecha, para luego bajar por el río, 8°54’N, 82°43’W, 1900 m, 24 September 1996, Aranda et al. 3245 (F, MO) ; Forests and edges of cafetales along Río Chevo , 8°51’N, 82°47’W, 1150–1200 m, 13 October 1981, Knapp 1569 ( MO) ; ca 0.5 km Eof Cerro Pate Macho, headwaters of Río Palo Alto , 8° 47’N, 82° 21’W 1800–2100 m, 12 November 1981, Knapp et al. 2125 ( MO) ; Along road between Volcán and Serrano , 7 mi Nof Volcán, remnants of primary forest on steep slopes above Highway , 8°50’N, 82°38’W, 15 June 1987, Croat 66234 ( MO) ; Trail to Cerro Pate Macho, 8°50’N, 82°25’W, 1500–1900 m, 7 January 1983, Stein et al. 1247 ( MO); ( Boquete ) , Bajo Chorro , 6000 ft, 11 January 1938, Davidson 105 ( MO) ; Quebrada Velo , 1800 m, 8 July 1940, Woodson & Schery 267 ( MO) ; Vicinity of Boquete , Cerro Pate de Macho, SW slope, 8°46’N, 82°25’W, 1800–1950 m, 19 June 1987, Croat 66429 ( MO) ; Las Lagunas area , Wof El Hato del Volcán, 8°47’N, 82°40’W, 1400 m, 23 August 1982, Hamilton et al. 940 ( MO) ; On NW side of Cerro Pando cloud forest, 21 July 1971, Croat 15960 ( MO) ; Cerro Horqueta, 4500–5500 ft, 20 July 1968, Dwyer Lallathin 8773 ( MO) ; SE slopes and summit of Cerro Pate Macho, trail from Río Palo Alto , 4 km NE of Boquete, 1700–2100 m, 26 May 1981, Sytsma et al. 4861 ( MO) . Coclé: Area surrounding Rivera Sawmill , 7 km Nof El Cope, continental divide, 750–850 m, 24 November 1977, Folsom & Collins 6439 ( MO) . Colón: Forest and fields to 3 miles upstream from sea, Río Iguanita , 0–500 m, 4 April 1980, D'Arcy & Croat 13591 ( MO) ; Along the Río Escandalosa, 19 mi Eof the Transisthmian Highway on the road to Salamanca , 160–180 m, 28 March 1982, Huft & Knapp 1632 ( MO) ; Along the Río Escandalosa, 19 mi Eof the Transisthmian Highway on the road to Salamanca , 160–180 m, 28 March 1982, Huft & Knapp 1647 ( MO) . Comarca de San Blas: Headwaters of Río Nergala along continental divide, 350 m, 11 January 1985, de Nevers & Herrera 4520 ( MO) . Darién: Cerro Tacarcuna massif between summit camp on Wmost ridge and Pico Tacarcuna mostly elfin forest, ( Colombia on some maps), 1650–1800 m, 22 July 1976, Gentry et al. 16929 ( MO, US) ; 0.2 miles Eof Tres Bocas, along the shortest headwater of Río Cuasí , 28 April 1968, Kirkbride & Duke 1169 ( MO, NY) ; Cerro Campamento ( S of Cerro Pirre), 20–22 March 1968, Duke 15616 ( MO) ; Cerro Campamento ( S of Cerro Pirre), 20–22 March 1968, Duke 15582 ( MO, US) . Panamá: Gorgas Memorial Labs yellow fever research camp, ca 25 km NE of Cerro Azul on Río Piedras, 550 m, 20–22 November 1974, Mori & Kallunki 3310 ( MO) ; Serranía de Maje, In ravines and along seasonal streams among ridges Sof Ipetí 5 hours walk from Chocó village, 8°45’N, 77°30’W, 450–600 m, 1 April 1982, Knapp et al. 4546 ( CAS, MO) . Veraguas: Trail on ridge to summit to Cerro Tute, Cordillera de Tute , 1 km past Escuela Agrícola Altos de Piedras , Wof Santa Fe, 8°36’N, 81°6’W, 750–950 m, 15 December 1981, Knapp & Sytsma 2526 ( MO) ; Along base of Cerro Tute, just past Escuela Agrícola , up from Santa Fe , 8°32’N, 81°7’W, 1200 m, 10 September 1982, Hamilton et al. 1309 ( MO) ; Mts. 3-5 mi, Nof Santa Fe, 500–1000 m, 12 December 1971, Gentry 3022 ( MO) ; Along trail to summit of Cerro Tute, ca 3 km above Escuela Agrícola Alto Piedra near Santa Fe , 2600–2800 ft, 4 January 1981, Sytsma & Antonio 2985 ( MO) ; Vicinity of Escuela Agrícola Alto Piedra, near Santa Fe, Along trail to top of Cerro Tute, 3500 ft, 29 June 1980, Antonio 4937 ( MO, NY) ; Vicinity of Escuela Agrícola Alto Piedra, near Santa Fe, 3 mi beyond fork in road near the school toward Atlantic coast, Near trail to top of Cerro Tute, 2200–2400 ft, 26 January 1980, Antonio 3520 ( MO) ; 5 mi Wof Santa Fe on road past Escuela Agrícola Alto Piedra on Pacific side of divide, 800–1200 m, 18 March 1973, Croat 22988 ( NY); ( Santa Fe ) , Altos de Piedra, Trail leading circling around base of Cerro Mariposa (= Cerro Arizona ) about 1 km past the Escuela Agrícola , 3°30.70’N, 81°7.03’W, 800 m, 29 January 2005, Penneys & Blanco-Coto 1701 ( NY) ; Forested mountains Wof Alto de Piedras Wof Santa Fe , 3200–5600 ft, 8 September 1978, Hammel 4598 ( MO) ; Cerro Tute , ca 10 km NW of Santa Fe, <1000 m, 19 June 1975, Mori 6774 ( CAS, MO) ; Lower montane wet forest 6–7 km Wof Santa Fe on new road past agriculture school, 2900 ft, 16 February 1974, Nee 9775 ( MO) ; NW of Santa Fe, 2.6 km from Escuela Agrícola Alto de Piedra , 23 February 1975, Mori & Kallunki 4744 ( MO) ; Río Segundo Braso, 8 km beyond Escuela Agrícola Alto Piedra beyond Santa Fe , 750 m, 24 July 1974, Croat 25552

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 103 ( MO) ; Forest along Río Gatu above Chitra Bajo , NW of Laguna La Yeguada, 8°42’N, 80°28’W, 650 m, 14 January 1986, McPherson 8014 ( MO, US) ; Valley of Río Dos Bocas, 11 km from Escuela Agrícola Alto Piedra (above Santa Fe ) on road to Calovebora , 450 m, 30 August 1974, Croat 27518 ( MO) ; Vicinity of Escuela Agrícola Alto Piedra near Santa Fe, 0.3 mi beyond fork in road near school toward Atlantic coast, along trail to top of Cerro Tute , 1050–1150 m, 29 November 1979, Antonio 2919 ( MO) ; Vicinity of Escuela Agrícola Alto Piedra, near Santa Fe along trail to top of Cerro Tute , 2200 ft, 4 April 1980, Antonio 4034 ( MO) . PERÚ. Cajamarca: ( La Palma), 10 km NW de Chirinos, 5°25’S, 78°53’W, 1780 m, 5 February 1988, Gentry et al. 61219 (F, MO, NY, US); (La Palma) , 10 km NW de Chirinos, Transect 1, 5°25’S, 78°53’W, 1780 m, 4 February 1988, Gentry & Diaz 61119 ( MO) .

Huánuco: (Leoncio Prado), Hermilio Valdizán, La Divisoria, 1600 m, 18 April 1980, Schunke-Vigo 11287 ( MO, US) . Junín: (Puerto Bermúdez), 375 m, 14–17 July 1929, Killip & Smith 26503 ( NY); ( Puerto Bermúdez ) , 375 m, 14–17 July 1929, Killip & Smith 26425 (F, NY); ( San Nicolás ) , Pichis Trail , 1100 m, 4–5 July 1929, Killip & Smith 25974 ( NY) . Loreto: (Alto Río Itayá), Forest of San Antonio , 145 m, 7 October 1929, Williams 3465 (F) . Pasco: (Oxapampa), Huancabamba, Cordillera Yanachaga, W slope at base of highest peaks, Side road to NE, halfway between Oxapampa and Huancabamba, Sector Santadela, ridge SE of Cooperativa 3 de Mayo, 10°27’S, 75°26’W, 2000–2500 m, 18–21 September 1984, Foster et al. 11200 (F); ( Oxapampa ) , Headwaters of Río Tunqui, Trail to Chuchurras-Palcazu , 10°14’S, 75°28’W, 1850 m, 2 January 1984, Foster et al. 7762 (F, MO, NY); ( Oxapampa ) , Palmazú , 10°32’S, 75°23’W, 2100 m, 19 September 1984, Smith et al. 8507 ( MO) . VENEZUELA. Aragua: (Chuao), El Medio, Forests of Valle en El Medio , 600–800 m, 15 March 1926, Pittier 12123 ( US) . Barinas: (Bolivar), Altamira, La Gallineta , El Celoso , Near feldspar mine, Forest and road near waterfall and mouth of tunnel leading to dam, 8°50’N, 70°35’W, 1500–1700 m, 6 June 1988, Dorr et al. 5447 ( NY, US) . Carabobo: En las laderas arriba de las cabeceras del Río San Gián, arriba de Los Tanques y La Toma , entre Quebrada No. 2 y Quebrada de los Verros , al Sde Borburata, 750–900 m, 29 March 1966, Steyermark & Steyermark 95245 ( NY, US, VEN) . Mérida: 22.7 km Eof Santo Domingo on the road to Barinas, 1585 m, 27 January 1993, King et al. 10559 ( CAS, F, MO, US) ; Rich forest above Hacienda Agua Blanca above La Azulita , 1375–1920 m, 25 April 1944, Steyermark 56085 (F, NY) ; Along highway between Merida and La Azulita, 9 km above Plaza Bolivar in La Azulita, SE of La Azulita, 8°44’N, 71°26’W, 1590 m, 8 August 1982, Croat 54840 ( MO) ; Vertientes del Río Capaz, arriba de La Azulita, 2100–2400 m, 1 September 1966, Steyermark & Rabe 97117 ( NY, US, VEN); ( Rangel ) , Mucuchíes, Quebrada de Los Chochos , unos 2 km antes de la aldea El Carrizal, 1520 m, 27 June 1966, Ruiz-Terán 3348 ( US) . Portuguesa: (Ospino), P.N. Guache, Sector Santa Rosa de Guache, Orilla de Quebrada intermitente, 9°30’42.03"N, 69°30’37.93"W, 14 April 1996, Licata et al. 704 ( MO) ; 15 km al Ede Chabasquén, 67 km al NNW de Guanare , 9°26–27’N, 69°54–55’W, 1450–1520 m, 29–31 October 1982, Steyermark et al. 126873 ( MO, US, VEN) . Trujillo: 13 km ESE of Bocono, 1 km Wof Guaramecal, 9°11’N, 70°9’W, 1600 m, 16 March 1982, Liesner et al. 12864 ( MO); ( Boconó ) , P.N. Guaramacal, Sector Occidental Qda. Honda, Parcela de estudio fitosociológico No. 21, UTM 359185E, 1012298N, 1880 m, 14–15 November 2001, Cuello et al. 2168 ( MO) ; 13 km ESE of Bocono, 1 km Wof Guaramecal, 9°11’N, 70°9’W, 1600 m, 16 March 1982, Liesner et al. 12883 ( MO); ( Boconó ) , P.N. Guaramacal, Montane forest near Quebrada Honda, UTM: 19–359029E, 1011815N, 1900–2000 m, 28–29 December 2000, Dorr & Stergios 8750 (F, MO, NY, US); ( Boconó ) , P.N. Guaramacal, Sector Quebrada Honda-Pele Ojo, El Santuario , 1850–1950 m, 1–31 January 2001, Stergios & Caracas 18892 ( NY) . Yaracuy: (Bruzual), Montaña de María Lionza, Quebrada Quibayo , desde abajo hasta casi la cumbre, 10°6–7'‘N, 69°55’W, 250–1000 m, 14 March 1981, Steyermark et al. 125045 ( US, VEN) . Zulia: (Mara), Cuenca del Río Guasare, alrededores del Destacamento Guasare #1 ( La Yolanda ), en las laderas del cerro ca 5 km al SSE del Destacamento , entre el caño Indio y la fila arriba de su orilla izquierda, 10° 52.00’10"N, 72°29’30"W, 700–800 m, 7 December 1982, Bunting et al. 12711 ( NY); (Mara) , Vicinity of Río Guasare, between Rancho 505 and Cerro Yolanda , 10°53–56’N, 72°26–28’W, 200–270 m, 29 May 1980, Steyermark et al. 122879 ( MO) .

Illustration:— Berry et al. 2001: 485, fig. 404; Gleason 1958: 263, fig. 87; Wurdack 1973b: 719, fig. 64.

Common names and documented uses:— Belize: “mountain sirin” (Gentle 7174, CAS!) ; “sirin” ( Gentle 4281, CAS!) . Ecuador: “palo hueso” (Pichincha, Jiménez 1125, MO!) ; “quiliyuyo” (Morona-Santiago, Cerón 14330, MO!) “taja sangre” (Manabí, Gentry 72543, F!, MO-2 sheets!) ; “yuturi caspi” (in Quechua , Orellana, Reyes 538, MO!) . México: “jeepe” (Oaxaca, Hernández 2467, US!) . Perú: “tiri” (Pasco, Smith 8507, MO!) .

Habitat, distribution and ecology:— This is a widespread locally common species of primary and secondary rain forest, cloud forest and swamp forest, or disturbed sites, on steep slopes or near streams, in the understory or forest margins, throughout southern Mexico and Mesoamerica and in two Caribbean islands ( Jamaica and

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Hispaniola), south to Colombia, Venezuela (including the Venezuelan Guayana), Ecuador and Perú ( Fig. 15 ), at 0–2500(–3000) m. In México it has been collected in the southern states of Chiapas, Oaxaca and Veracruz, south to Belize and throughout the lowlands (except for El Salvador) and the Pacific and Atlantic slopes of Central America. In the Caribbean islands of Jamaica and Hispaniola ( Liogier 2000) it is less frequent at low to median elevations. In Colombia it has been collected in the Sierra Nevada de Santa Marta, the three Andean Cordilleras, including the Biogeographic Chocó premontane areas (Pacific and Andean regions), and south to Putumayo. Its range extends east to Venezuela in the valleys of the Aragua state to the Venezuelan Guayana (known from Capibara, Tamatama, Río Atabapo, San Carlos de Río Negro) ( Berry et al. 2001). Further south M. neomicrantha extends throughout Ecuador and western Perú to Oxapampa. It is not known from any island in the Pacific or from Panama’s Barro Colorado Island ( Croat 1978).

In Costa Rica Corapipo altera , Manacus candei , and Tachyphonus delatrii have been reported to feed on M. neomicrantha fruits ( Boyle 2006), as well as Semnornis frantzii ( Wheelwright et al. 1984) . In a montane forest in Colombia (Santuario de Fauna y Flora Otún-Quimbaya), Myadestes ralloides has also been reported to consume berries of M. neomicrantha ( Kessler-Rios & Kattan 2012) .

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

Etymology:— The specific epithet refers to the small flowers of this species.

Discussion:— This species is distinguished by its superficially glabrous appearance, commonly caudate foliar apex, and prominently 8-ribbed berries that are white at maturity. It is closely related to M. albertobrenesii and M. boekei , both of which have a dense conspicuous indumentum and larger flowers. Miconia alboglandulosa has occasionally been confused with M. neomicrantha , perhaps because of the foliar shape and overall appearance. However, the inflorescence architecture of the former consists of a cluster of cymes (vs. dithyrsoids) and M. neomicrantha lacks the white-translucent furrowed glands that are so typical of the leaves and floral organs of M. alboglandulosa .

Throughout its extensive range, this species is uniform in floral and fruit characters; all the flowers examined were 4-merous (except for two 5-merous populations discussed below). Anther shape, the glandular anther appendages and the white color of mature berries were consistent for all collections studied throughout its range. Although Wurdack (1973a) synonymized O. caudata and O. tetragona under M. neomicrantha he noted that two micromorphological variants occur in Central America. One has somewhat tetragonal cauline internodes, leaf apices generally only shortly and gradually acuminate, and petals usually 1 mm or less wide and externally moderately squamate. This variant mostly occurs at 0–700 m and matches the Caribbean and lowland continental Atlantic slope material (including the Sierra Nevada de Santa Marta populations in Colombia). The other variant usually has terete branchlets, leaf apices more or less long-acuminate to caudate, petal width 1.3–1.7 mm and rather glabrescent abaxially. These collections are from more montane areas in Central America, at (1100–) 1500–2400 m; but the modal differences that appear to have some elevational correlation in Central America breaks down in South America. Furthermore, the squamate-stellate vegetative and floral indumentum of this species varies greatly in quantity throughout the range as does foliage shape. Some collectors have also reported the petals to have red apices ( Ecuador, Rimbach 66, 189, 226, F!, US!, NY!) or red bases and spots ( Jamaica, Morley 912, MO!). This color anomaly appears to be rare.

Indumentum characters can be quite variable in this species. At least three micromorphological variants are evident. The typical indumentum in this species is squamate-stellate composed by hyaline lepidote trichomes with only partially fused radii (see Fig. 213 in Wurdack 1986). This indumentum which is present throughout a plant is occasionally very sparse. These typical individuals are present throughout the geographic and elevational range of the species. However, in South America some populations have a puberulent indumentum similar to that in M. variabilis that is composed of minute brownish dendritic trichomes with short axes and few-moderate number of terete arms instead of the squamate-stellate trichomes on vegetative parts, hypanthia, and petals. These plants are known from southern Colombia (Nariño and Putumayo) through the lowlands and Andean slopes of Ecuador (Carchi, Esmeraldas, Imbabura, Napo, Pastaza, Pichincha, and Los Ríos), at (150–)200–2000(–3000) m. Superficially they look like typical M. neomicrantha because the indumentum details are only visible with the stereomicroscope or with a powerful hand lens in the field.

In addition, a rare variant known from five specimens has similar stellate-squamate general indumentum like that of typical M. neomicrantha . However the trichomes are stellate and brownish-translucent in color. Due to their coloration they are visible to the naked eye, especially on the leaves abaxially. This variant occurs from the

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 105 southern provinces of Ecuador (Morona-Santiago and Zamora-Chinchipe) to the department of Cajamarca in Perú, at 1700–2300 m. These populations are rather isolated both elevationally and geographically. More typical M. neomicrantha occurs at lower elevations .

In the past, some inividuals represented by a few collections that are genetically distinct (based on DNA sequence data) from M. neomicrantha , have been determined as this species. These are gabresecent or with the typical but sparser indumentum of M. neomicrantha and with a well-developed membrane at the point where secondary veins diverge from the midvein at the abaxial base of the leaf blade (evident mite-domatia). These individuals have been collected in the low elevation Andean forests of Ecuador (Pichincha) disjunctly to northern Perú and then south to Oxapampa in Pasco, Perú at 530–2500 m. The highest elevational records for these specimens are in Perú (2000–2500 m) and the lowest in Ecuador (530–800 m). In the phylogenetic analysis performed here one of these individuals was included. In the resulting cladogram this taxon branches apart from M. neomicratha , in a basal position of the clade containing it. This variant may represent a new species ( Fig. 1 ). The only character that would differentiate it from typical M. neomicrantha is the presence of an evident mitedomatium. This distinction is not taxonomically practical due to an occasional presence in typical M. neomicrantha of a fusion of trichomes to form a conical chamber in the same area where the named mite-domatia occurs. For this reason, a taxonomic evaluation of this species must be postponed until more collections come to light. The available material for study showed that individuals with mite-domatia occur sympatrically with the typical forms of M. neomicrantha .

The membrane-like structure and associated cavity formed at the point where the secondary and primary veins diverge from one another is surely some kind of mite domatium. In several specimens with these structures, mite eggs were found in small numbers but the mites and eggs rarely survive the drying process. Eggs were also found on specimens with a fusion of trichomes to form a conical chamber in the area where the secondary veins diverge from the primary vein.

The floral merosity of M. neomicrantha is also variable in this species but it is only evident in a few specimens from disjunct areas. First, in the departments of Junín (Killip & Smith 26503, 26425, 25974, NY!) and Loreto (Williams 3465, F!) in Perú, the cited specimens of M. neomicrantha have 5-merous flowers and are rather glabrous on vegetative and floral parts and without the evident mite-domatia described above. Nevertheless, their leaf morphology (with a broadly obtuse base) and anther morphology is consistent with typical M. neomicrantha . Other specimens that probably have 5-merous flowers were examined from Veraguas province in Panama (Hammel 4598, CAS!, and Mori 6774, CAS!, MO!). These plants which are only in fruit have 10-ribbed hypanthia and likely 5- merous flowers. Their morphology is also consistent with all the other collections of M. neomicrantha examined for this study, although these plants are squamate-glabrescent, without mite-domatia. They also have a visible nodal line, the foliar bases are acute to attenuate, and the inflorescences are multiflorous. The area of Panama where they were collected is well known, and the fact that there are only two collections showing this variation indicates that it is probably rare.

No specimens of this species were examined from the Caribbean Island of Hispaniola or from the Venezuelan Guayana (state of Amazonas in southern Venezuela) for this study. However this species appears in the flora treatments for these regions ( Berry et al. 2001, Liogier 2000), and in other treatments it has been reported from Hispaniola ( Almeda 2009, Wurdack 1980).

Conservation status:— This species would be considered Vulnerable VU B2ab(iii) based on IUCN criteria ( AOO). Astatus of Least Concern LC is warranted because it occurs in many protected sites throughout its range. Protected in Belize in the Bladen Nature Reserve (Toledo) . In Colombia it is protected in the Sierra Nevada de Santa Marta National Park (Magdalena) ; in La Planada Nature Reserve (Nariño) ; in the Otún-Quimbaya Sanctuary of Fauna and Flora (Risaralda) ; in the Farallones National Park and in the Finca Los Sauces private protected area (Valle). In Costa Rica it is protected in the major Natural Parks and Reserves. In Ecuador it is protectd in the Awá Indigenous Reserve (Esmeraldas) ; in the Intag Private Reserve (Imbabura) ; in the Río Palenque Biological Station (Los Ríos) ; in Machalilla National park (Manabí) ; protected in the major Parks and Reserves in Pichincha Province. In Venezuela it is protected in the Guache and the Guaramacal National Parks (Portuguesa and Trujillo respectively) .

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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.