Miconia quinquenervia ( Miller 1768: 735 ) 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 : 110-114

25. Miconia quinquenervia ( Miller 1768: 735) Gamba & Almeda , comb. nov. Basionym: Melastoma quinquenervium Miller. Octopleura quinquenervia (Mill.) Triana (1871: 145) . Ossaea quinquenervia (Mill.) Cogniaux (1891a: 1064) . Clidemia quinquenervia (Mill.) Almeda (2004: 112–114) . Type: Exact locality not legible, Miller s.n. (holotype: BM-internet image!).

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Clidemia decurrens Beurling (1854: 127–128) . Type: PANAMA. Portobello, in silvis ad viam versus Panama, April 1826, Billberg 280 (holotype: S-internet image!; isotypes: F-internet image!, MO!).

Melastoma diversifolium Bonpland (1816: 138–140) . Clidemia diversifolia (Bonpl.) de Candolle (1828: 159) . Staphidium diversifolium (Bonpl.) Naudin (1851: 322) . Octopleura diversifolia (Bonpl.) Triana (1871: 145) . Ossaea diversifolia (Bonpl.) Cogniaux (1887a: 968) . Type: COLOMBIA. Habitat ad ripas fluvii Magdalenae, frequens juxta S. Bartholomé, Bonpland 1617 (holotype: P; isotypes: P-3 sheets, US).

Subshrub or shrub 0.5–4(–6) m tall with slender erect branching, bark green-brownish. Upper internodes [(2.6–)3.8–5.3(–11.1) cm] and cauline nodes terete, nodal line absent. Indumentum on branchlets, petioles, adaxial surface of young leaves, primary and secondary leaf veins abaxially, inflorescence axes, bracts, bracteoles, pedicels, hypanthia, calyx lobes, and calyx teeth densely to moderately composed of brownish sessile to thinstipitate-dendritic trichomes 0.1–0.2 mm long with short axes and few-moderate number of terete arms, sparsely intermixed with caducous elongate smooth trichomes 1–1.3 mm long. Leaves of each mature pair somewhat anisophyllous in size, the younger pairs being more or less isophyllous; the semiterete petiole 0.3–1 cm, deeply canaliculate adaxially, green-brownish; larger blades (10.2–)12–25 × (6.5–) 7.5–15 cm, smaller blades (4.5–)5–10.5 × (2.5–) 3–6 cm, broadly elliptic to ovate, the base rounded to abruptly attenuate and slightly oblique, narrowly long-decurrent on the petiole, the margin ciliolate-denticulate, the apex bluntly acute to short-acuminate, or obtuse, chartaceous; the adaxial surface flushed pink at the base of juvenile leaves, at maturity the adaxial surface rapidly becoming glabrate, or moderately to sparsely strigose with brownish elongate smooth trichomes 0.5–1.5 mm long, the primary and secondary veins on young leaves copiously covered with the general dendritic trichomes, becoming glabrate, the tertiary and higher order veins glabrous; abaxial surface occasionally flushed red or chocolate brown when dry, glabrate except for a few dendritic trichomes and resinous glands on the venules, the tertiary and higher order veins sparsely beset with caducous resinous slightly furrowed more or less stalked glands <0.05 mm long; 7- or 9-plinerved, including the tenuous marginals, innermost pair of secondary veins diverging asymmetrically from the primary vein 0.5–3.5 cm above the decurrent base, areolae 1–1.5 mm, adaxially the primary, secondary, tertiary and higher order veins impressed, abaxially the primary, secondary, tertiary and higher order veins elevated and terete. Inflorescences a pseudolateral multiflorous dithyrsoid 2–4 cm long, including a peduncle 0.2–0.5 cm long or sessile, highly and divaricately branched from the peduncle apex or the base, borne on the upper foliar nodes, rachis bright pink; bracts and bracteoles 0.5–0.7 × 0.2 mm, subulate-apiculate, occasionally with two thick trichomes projecting from the base, pink, the bracts spreading, the bracteoles spreading to erect, persistent in fruit. Flowers 5-merous on pedicels 0.25–0.5 mm long. Hypanthia at anthesis 2.5–2.7 × 1 mm, free portion of hypanthium 1.3–1.5 mm long, suburceolate to campanulate, bluntly 10-ribbed, yellowish becoming bright pink, copiously to sparsely resinous-glandular with slightly furrowed more or less stalked glands <0.05 mm long, caducously intermixed with the dendritic and elongate trichomes, ridged on the inner surface and like the torus adaxially sparsely and caducously resinous-glandular. Calyx open in bud and persistent in fruit, pinkishgreen; tube 0.1–0.3 mm long, with the same vestiture of the torus adaxially and of the hypanthium abaxially; lobes 0.1–0.25 × 0.25 mm, broadly triangular, the margin vaguely undulate, the apex bluntly acute, glabrescent with few resinous glands; exterior calyx teeth 0.5–1 mm long including the apical trichome, bluntly conic and>3 aristate, the aristae brownish, inserted at the base of the calyx lobes and projecting beyond them, glabrescent. Petals 0.75–1.5(–1.7) × 0.75–1 mm, obovate-oblong, with a projecting infra-apical abaxial tooth, the margin entire, the apex rounded-obtuse, white, densely papillose on both surfaces, reflexed at anthesis. Stamens 10; filaments 1–1.5(–1.7) × 0.25 mm, white, glabrous; anther thecae 1.5–2.4 × 0.25–0.35 mm, linear-oblong and subulate, truncate to emarginate and short-acuminate at the apex, opening by one dorsally inclined pore 0.1–0.15 mm in diameter, cream to light yellow at anthesis; connective yellow, its prolongation and appendage 0.25–0.45 mm long, the appendage oblong-spatulate, obtuse at the apex, copiously gland-edged, the glands sessile to shortly stalked and rounded. Ovary 5-locular, completely inferior, ca. 1.2 mm long at anthesis, the apical collar absent, the apex 0.3 mm in diameter, somewhat depressed, caducously glandular-puberulent; style 5.5–6 mm long, narrowed distally (i.e. tapering), white, glabrous or rarely sparsely to copiously glandular-puberulent at the base; stigma expanded truncate to capitellate. Berries 4–5 × 4.5 mm when dry, globose-oblate, bright pink, then green, and ultimately ripening purple-black, the hypanthial indumentum somewhat persistent at maturity. Seeds 0.33–0.54 × 0.16–0.2 mm, ovoid, angled, light-brown; lateral and antiraphal symmetrical planes ovate, the highest point toward the chalazal side; raphal zone suboblong, ca. 10–20% larger than the corpus of the seed, extending along its entire

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Phytotaxa 179 (1) © 2014 Magnolia Press 111 length, ventrally and longitudinally expanded, dark-brown; individual cells elongate, anticlinal boundaries channeled, undulate, with Ω- and U-type patterns; periclinal walls convex, low-domed to nearly flat, microrelief inconspicuously punctate. Chromosome number: n=17.

Additional specimens studied:— COLOMBIA. Antioquia: (Nicocli), <50 m, 27 June 1946, Haught 4924 ( US); ( Turbo ) , Quebrada Isaías , Eof Turbo, 50 m, 8 July 1946, Haught 4946 ( US) ; Cauca Valley, vicinity of Puerto Valdivia, Roadside vegetation, 300 m, 19 August 1976, Gentry et al. 17927 ( COL, MO, NY) . Bolívar: Boca Verde, Río Sinú , 8.64°N, - 76.06°W, 13–14 February 1918, Pennel 4227 ( US) . Caldas: (Chinchiná), orilla Quebrada Campo Alegre , 1200 m, 17 February 1990, Aguirre et al. 12 ( COL) . Chocó: Río El Valle, Cerca a la desembocadura del Río Miniquía , entre éste y el Río Mutatá , 7 January 1973, Forero & Gentry 742 ( COL, F) ; Río Ciego , 17 June 1967, Duke 13290 ( NY) ; Trail from Unguía along Río Tigre toward base of Serranía del Darién , 200–300 m, 16 July 1975, Gentry & Aguirre 15208 ( MO, NY) . Cundinamarca: ( San Antonio ), 200 m, 14 June 1948, Haught 6241 ( CAS, F, NY, US) . Magdalena: P.N. Tayrona, Along mule trail from Pueblito to Calabazo , 11°19’N, 73°58’W, 250–450 m, 26 October 1972, Kirkbride 2565 (F, NY, US) ; Alto Río Frío, Cabeceras del Río Congo, Ciudad Antigua , por el camino al río, 10°59’N, 74°4’W, 1000–1100 m, 25 July 1989, Madriñán & Barbosa 497 ( NY) . Meta: P.N.N. Tinigua, Río Duda, Serranía Chamusa, Centro de Investigaciones Ecológicas La Macarena, Trocha a Puerto de Invierno , 350 m, Julio 1996, Stevenson 1818 ( COL) . Quindio: (Montenegro), Vereda La Montaña , finca El Reflejo, 1090 m, 30 October 1990, Vélez et al. 2611 ( COL) . Tolima: ( Mariquita ), 500 m, 10 November 1938, Haught 2429 ( CAS, US) . COSTARICA. Cartago: ( Atirro ), 600 m, April 1896, Smith 6547 ( US); (Jiménez) , No protegida, Cuenca del Reventazón, Pelibaye , Turrialba , camino a El Copal , del Humo al Copal , 9°47’0"N, 83°43’13"W, 760 m, 5 January 2010, Kriebel et al. 5451 ( INB, MO, NY) . Heredia: At confluence of Ríos Sarapiquí and Puerto Viejo, 100 m, 1–6 August 1968, Schnell 1015 (F, US) . Limón: Vicinity of Moin , 3 km Wof Limón, 19 July 1974, Utley & Utley 5548 ( CAS) ; Along Highway 32 from Turrialba to Limón, ca. 11 miles Sof Siquerres, 650 m, 13 August 1977, Croat 43352 ( CAS, MO) ; About 2 km Nof Bribri along banks of the Río Sand Box , 100 m, 17 July 1976, Utley & Utley 5495 ( CAS) ; ca. 7 km SSW of Home Creek on the road to Bribri , 200 m, 26 July 1977, Almeda et al. 3256 ( CAS, CR) ; Near Carmen Station on Indiana Branch , 30 m, 20 February 1926, Standley & Valerio 48361 ( US) ; Goldengrove , drenaje de Río Reventazón, 15 m, 23 October 1951, Shank & Molina 4375 ( EAP, US); (Piuta) , Adjacent to Limón, 5 m, 18 September 1968, Davidse & Pohl 1249 ( ISC, US) ; Hamburg Finca, On the Río Reventazón below Cairo , 55 m, 19 February 1926, Standley & Valerio 48677 ( US) ; Approximately 4.5 km NE of Bribri and 15 km Sof Cahuita along road to Bribri , 100 m, 12 March 1978, Utley 6048 ( CAS) ; ca 5.5 km Wof Limón on the road to Siquirres, Along the road, 27 July 1977, Almeda et al. 3261 ( CAS, CR) ; Río Catarata, 1 to 3 km Nof Bribri, Río Sixaola drainage, 9°38’N, 82°50’W, 20–200 m, 9–10 September 1978, Burger & Antonio 10977 ( CAS, F) . ECUADOR. Napo: Río Zumino (Sumino), tributary of the Río Napo, ca 5 km NE of Santa Rosa, 8 August 1968, Lugo 202 (F, NY, US) ; Río Napo between Coca ( Puerto Francisco de Orellana) and Armenia Vieja , - 0.5°S, - 77.03°W, 250–350 m, 15 February 1974, Harling & Andersson 11986 ( MO, US) ; Sacha Lodge, 3 km NW of the village Anangu , near the Napo river., - 0.5°S, - 76.43°W, 200 m, 5 June 1995, Clark et al. 969 ( MO) . Pastaza: Río Pastaza, between Destacamento Chiriboga and Apachi Entza, 2°20–35’S, 76–77°55–8’W, 285 m, 24 July 1980, Øllgaard et al. 35173 ( CAS, US) . HONDURAS. Gracias a Dios (La Mosquitia): (Ahuas Bila), 200 km SW de Puerto Lempira, orilla del Río Wankí , Coco o Segovia , 100 m, 5–13 May 1985, Nelson & Cruz 9345 ( CAS, UNAH) . NICARAGUA. Jinotega: Macizos de Peñas Blancas , 1400 m, 5 May 1976, Neill 7127 ( CAS) . Zelaya: Sector de Negro Wás, entre El Empalme y Rosita, 13°45’N, 84°25’W, 200 m, 5 September 1984, Ortiz 2123 ( CAS, HULE); ( Siuna ) , Wany , 27 August 1982, Ortiz 47 ( CAS, HULE) ; Río Kusulí , 13°24’N, 85°19’W, 200 m, 4 March 1981, Stevens & Moreno 19258 ( CAS, MO) ; Caño Matis (El Cianuro) cruce la carretera Wany-Siuna , ca 2 km Wde Siuna, 13°40’30"N, 85°45’40"W, 26 October 1982, Grijalva & Burgos 1578 ( CAS, HULE) . PANAMA. Bocas del Toro: Lincoln Creek , 28 June 1921, Carleton 87 (F, US) . Canal Zone: Pipeline road near Río Agua Salud , 21 October 1972, Kennedy & Redemsky-Young 1833 ( MO, NY); ( Barro Colorado Island ) , 18 September 1982, Schmalzel & Schupp 955 ( CAS, MO) ; Las Cascadas Plantation, near Summit , 2 December 1923, Standley 25688 ( US) ; Vicinity of Madden Dam, 90 m, 8 October 1939, Allen 2009 ( MO, US); ( Barro Colorado Island ) , 25 August 1929, Bangham 374 ( US) ; Along Changuinola River, 8 January 1924, Dunlap 296 ( US); ( Barro Colorado Island ) , Shoreline between Dock and Dump Cove, 13 September 1968, Croat 6050 (F, MO) ; George W. Green Park, Eof Summit Garden, in Madden Forest Preserve, Pacific Water Shed, near Continental Divide , 91 m, 10 August 1961, Welch 19632 ( NY) ; Pipeline Road , 6 km along road, 9°13’N, 79°43’W, 100–200 m, 27 October 1980, Systma 1940 ( CAS, MO) ; P.N. Soberanía, Madden Forest , las Cruces

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Trail, 3.6 mi Nof Gamboa Road turn-off, 9°26’20"N, 79°37’20"W, 140 m, 23 July 1994, Croat & Zhu 77060 ( CAS, MO); ( Barro Colorado Island ) , Lutz creek near Donate Trail , 100–300 m, 8 October 1968, Croat 6792 ( MO) . Colón: Nof Diamante , ridge NW of abandoned mine on Quebrada de la Mina, 9°24’N, 79°35’W, 600–700 m, 11 January 1984, Churchill & de Nevers 4216 ( CAS, MO) ; 1–2 km from the Portobelo Highway up the Río Guanche , 9°30’N, 79°40’W, 0–50 m, 17 February 1982, Knapp 3641 ( CAS, MO); (Peluca) , ca 27 km from Transistmian Highway on road to Nombre de Dios, Trail to Río Boquerón and along bank, 25 February 1973, Kennedy 2635 ( CAS, MO) ; Along Río Boquerón near N.1 (manganese mine), Eof Salamanca, 9°35’N, 79°32’W, 50 m, 3 July 1982, Knapp et al. 5804 ( CAS, MO) ; Forest and forest edge from Portobelo Highway to 4 km up Río Guanche , 9°30’N, 79°40’W, 0–50 m, 30 August 1981, Knapp 990 ( CAS, MO) ; Along Río Boquerón above Mina Boquerón #1 (manganese mine), main valley of Río Boquerón , 9°20’N, 79°35’W, 100–200 m, 13 December 1981, Knapp & Sytsma 2448 ( CAS, MO) ; Río Guanche , 9°30’N, 79°39’W, 0–75 m, 16 October 1980, Systma 1712 ( CAS, MO) . Darién: Mannene to the mouth of the Río Cuasí , 28 April 1968, Kirkbride & Bristan 1496 ( NY) ; Subiendo por el Río Peresénico desde la Estación de Pirre , 28 September 1989, Palacios et al. 402 ( CAS, PMA) ; De la estación de Cruce Mono hasta la quebrada Ipelisa (límite del parque), 28 August 1990, Aranda et al. 1517 ( CAS, PMA) . Panamá: Forests between Juan Mina, Canal Zone, and El Virgía , 20–80 m, 11–12 January 1911, Pittier 2401 ( US) ; Isla Majé, Bayano , Trocha C, campamento del G.M. L, 20 August 1976, Garibaldi 208 ( NY, PMA, US) ; Area around Torti Arriba , 31 August 1977, Folsom 5153 ( CAS, MO) ; Patch of forest along Panamerican Highway, 5 mi Wof bridge over Bayano lake , 200 m, 14 October 1978, Hammel & D'Arcy 5109 ( CAS, MO) ; 3.4 mi Eof Cañasas, checkpoint on PanAm. Hwy. in selectively cut forest ca. 4.5 km from road, Panamá-Darién border, foothills of Serranía de Cañas , 8°52’N, 78°15’W, 100 m, 12 September 1981, Knapp 1160 ( CAS, MO) ; Along stream 1 mi Nof Summit Gardens towards Gamboa , 11 February 1981, Systma 3497 ( CAS, MO) ; Canal Area, Boy Scout camp near Lake Madden , 100 m, 29 August 1982, Hamilton & Stockwell 1149 ( CAS, MO) ; Serranía de Majé, ridges Sof Choco village of Ipetí, Río Ipetí drainage system, 8°47’N, 78°27’W, 500–600 m, 11 December 1981, Knapp & Sytsma 2365 ( CAS, MO) ; Cañitas near dam site, just Sof Panamerican Highway, 9°10’N, 78°52’W, 150 m, 19 September 1982, Hamilton & D'Arcy 1415 ( CAS, MO); (Piriatí) , Sof Pan-Am Highway , 9°0’N, 78°30’W, 200–400 m, 8 August 1982, Hamilton 538 ( CAS, MO) ; Serranía de Majé, Trail along Río Ipetí , near confluence with Río Ambroino , 8°57’N, 79°32’W, 100 m, 29 January 1984, Churchill & de Nevers 4475 ( CAS, MO) ; Río Ipetí , Sof Panamerican Highway, 9°3’N, 78°25’W, 100 m, 17 September 1982, Hamilton & D'Arcy 1329 ( CAS, MO) ; Hills Sof Guacuco, 8 km Eof Ipetí, 8°55’N, 78°20’W, 300 m, 18 September 1982, Hamilton & D'Arcy 1353 ( CAS, MO) ; Pipeline Road , 10 to 15 km Nof Gamboa, 9°10’N, 73°45’W, 50 m, 16 September 1984, de Nevers 3892 ( CAS, MO) . San Blas: Ailigandi area, Along trail from ocean to waterfall on river, 0–61 m, 7 October 1978, Hammel & D'Arcy 4989 ( CAS, MO) ; Tierra firme frente a Isla Miria Ubigandup, camino Digole , 9°26’N, 78°54’W, 0–20 m, 19 July 1987, Herrera 265 ( CAS, PMA) ; Ailigandi cutover area along river feeding town water supply, At base of and on large (ca 50 m tall) limestone outcrop approximately 3 km inland, 8 October 1978, Hammel & D'Arcy 5014 ( CAS, MO) . VENEZUELA. Portuguesa: (Guanare), 17 km NW of Tucupido by road, future basin of Represa Tucupido and lower forested, Nslope of Fila Las Palmas , 9°1’N, 69°57’W, 200–400 m, 30 October 1982, Davidse et al. 21455 ( CAS, MO, NY) . Zulia: (Bolívar), Cuenca del Embalse Burro Negro (Pueblo Viejo): sector entre Quirós-El Pensado y el pie de Cerro Socopo, en el área aprox, 10 km en línea recta al Ede Churuguarita, 250–300 m, 5–8 August 1980, Bunting 9508 ( NY); (Mara) , La Yolanda, Cuenca del Río Guasare , alrededores del Destacamento Guasare N. 1, 10°52’10"N, 72°29’30"W, 200–250 m, 9 November 1982, Bunting et al. 11977 ( CAS, US) .

Illustration:— Bonpland (1816): Pl. 59.

Common names and documented uses:— Ecuador: “paichi panga”, for curing fevers (Clark et al. 969, MO!). Panama (Barro Colorado Island): “fruta de pava” ( Croat 1978).

Habitat, distribution, and ecology:— Uncommon to locally common in gallery forests, semideciduous forests, rain forests margins, river or stream banks, pasture margins and disturbed sites, often in deep or partial shade from Honduras south through southern Central America to Colombia, Venezuela (Portuguesa, Zulia), and Ecuador (Napo, Pastaza) ( Fig. 16 ), at 0–700(–1400) m. In Nicaragua it occurs in the “zona Atlántica” ( Almeda 2001). In Costa Rica it is mostly restricted to the Caribbean slope. In Panama, it is known principally from tropical wet forests in the Canal Zone, all along the Atlantic slope, and in Panama and Darién, on the Pacific slope; also known from humid forests in Panama (Panama City), Colón and Darien ( Croat 1978). In Colombia it is found throughout the Andes (Western, Central and Eastern Cordilleras), from low to premontane elevations, and in the Sierra Nevada de Santa Marta.

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Phytotaxa 179 (1) © 2014 Magnolia Press 113

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

Etymology:— The specific epithet refers to the five noticeable longitudinal foliar nerves.

Discussion:— This species is characterized by the conspicuous decurrent leaf base, multiflorous dichasial thyrsoid inflorescence, and setose calyx teeth. Its sister species is M. aguilarii , which is strikingly similar vegetatively (including the decurrent leaf base), but differs in floral characters (merosity and hypanthium color at anthesis and maturity). Both species are allopatric geographically (see M. aguilarii discussion).

As mentioned under the discussion of M. neocoronata , M. quinquenervia is very similar to this species as well. The distinctions between M. neocoronata and M. quinquenervia are discussed in detail in Almeda (2004). Another closely related species is M. reitziana , from which it is easily distinguished vegetatively by its lack of decurrent leaf bases. The inflorescence architecture is also different (not multiflorous groups of modified dichasia), and the elongate smooth trichomes are more abundant in M. reitziana , and completely lacking on the abaxial foliar surface of M. quinquenervia . Both share similar setose exterior calyx teeth, and the glands at the edges of the dorso-basal connective appendage.

Along with M. aguilarii , M. neocoronata and M. reitziana , this species is conspicuously anisophyllous. However, the anisophylly does not seem to be consistent in every mature leaf pair and the younger leaves are isophyllous.

The rusty-asperous general indumentum of dendritic trichomes in M. quinquenervia resembles that of M. bensparrei , which was considered to be its closest relative by Wurdack (1980). This type of indumentum which is also present in M. palenquensis and M. incerta , and in smaller versions in M. sessilis , M. variabilis , and some populations of M. neomicrantha , is common among closely related species in Octopleura , but not exclusive to these taxa. Therefore, the taxonomic importance of this character is best appreciated when used along with other diagnostic features like seed characters, when assessing relationships among species groups within the Octopleura clade.

Conservation status:— Vulnerable VU B2ab(iii). Althought this species is widely distributed geographically, it is not known from many protected areas. In Colombia it is known from Tayrona National Park (Magdalena) and Tinigua National Park (Meta). In Panama it is common in the Canal Zone and protected in the Chagres and Soberanía National Parks.

Figures

FIGURE 16. Distributions of Miconia quadridomius and M. veraguensis, M. quinquenervia, M. radicans, and M. reitziana.