Miconia radicans, Gamba, Diana & Almeda, Frank, 2014

Gamba, Diana & Almeda, Frank, 2014, Systematics of the Octopleura Clade of Miconia (Melastomataceae: Miconieae) in Tropical America, Phytotaxa 179 (1), pp. 1-174 : 114-117

26. Miconia radicans (Cogn. in Smith 1895: 288–289) Gamba & Almeda, comb. nov. Basionym: Clidemia radicans Cogn. in Smith. Type: COSTA RICA. Comarca de Limón, Llanos de Santa Clara, Río Jiménez , 650 ft, 11 April 1894, Donnell Smith 4789 (isotypes: BR-2 sheets-internet images!, US-internet image!) .

Suffrutescent herb or subshrub 0.2–1 m tall, occasionally rupicolous, suberect and sparingly branched. Upper internodes (1.6–3.8 cm long) and cauline nodes terete, nodal line present as a moderate ridge. Indumentum on branchlets, petioles, adaxial surface of young leaves, primary, secondary, tertiary and higher order veins abaxially, inflorescence axes, bracts, bracteoles, pedicels, hypanthia, calyx lobes abaxially, and exterior calyx teeth densely to moderately covered with brownish-translucent sessile-stellate trichomes 0.35–0.45 in diameter, superficially appearing simple. Leaves of each pair strongly dimorphic (1:6–8); the semiterete petioles 0.1–0.8 cm long, narrowly canaliculate adaxially, the channel obscured by the dense indumentum; larger blades 8.5–20 × 3–7.7(–9) cm, elliptic-oblong and somewhat falcate, the base obtuse to rounded and slightly oblique, the margin entire, the apex bluntly acute to rounded; smaller blades 1.5–2.7 × 0.9–4 cm, ovate to ovate-elliptic or subrotund, the base obtuse to cordate, the margin entire, the apex acute to short-acuminate; firm-membranaceous; mature leaves adaxially occasionally flushed red-purple on the surface and primary vein, glabrate, the primary, secondary, tertiary and higher order veins glabrous; abaxial surface glabrate; 3- or 5-nerved, including the tenuous marginals, with a moderate thick-callose vesicular structure at the base of the leaves abaxially where the innermost pair of secondary veins diverge from the primary vein (domatia?), areolae 1–2 mm, adaxially the primary, secondary, tertiary and higher order veins flat, abaxially the primary and secondary veins elevated and terete, the tertiary and higher order veins flat. Inflorescences an axillary few-flowered thyrsoid or dithyrsoid 3.5–8.3 cm long, typically including a peduncle 1.5–2.6 cm long, rarely sessile, somewhat divaricately branched from the peduncle apex, when sessile openly bifurcate from the base, solitary or paired in the axils of the smaller leaves at the upper nodes, the rachis pink-reddish to brown; bracts and bracteoles 0.3–0.5 × 0.5 mm, narrowly triangular, somewhat spreading, persistent to tardily deciduous in fruit. Flowers 4-merous on pedicels (0.3–) 0.5–1 mm long. Hypanthia at anthesis 2.6–2.75 × 1–1.5 mm, free portion of hypanthium 1 mm long, urceolate, constricted distally into a subcylindric

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neck, bluntly 8-ribbed, reddish, the indumentum sparse, ridged on the inner surface, glabrous, the torus adaxially sparsely glandular-puberulent becoming glabrous, the glands stalked or subsessile and rounded. Calyx open in bud and persistent in fruit, pink-red; tube 0.19–0.27 mm long, abaxially with the same vestiture as the hypanthium, adaxially minutely and sparsely glandular; lobes 0.3–0.5 × 1 mm, ovate, the margin entire, the apex rounded, adaxially glabrescent with a few papillae; exterior calyx teeth 0.29–0.34 mm long, bluntly triangular to subulate, callose, inserted at the base of the lobes and not projecting beyond them. Petals 1.5–2 × 1.4–1.5 mm, obovate, the margin entire, the apex rounded-obtuse to somewhat truncate, white, copiously and minutely papillose on both surfaces, slightly spreading to erect at anthesis. Stamens 8; filaments 1.5–2 × ca. 0.22 mm, white, glabrous; anther thecae 2 × 0.25–0.29 mm, linear-oblong, obtuse-emarginate at the apex, opening by one dorsally inclined pore 0.1 mm in diameter, cream to yellow; connective yellow, its prolongation and appendage 0.2–0.4(–0.5) mm long, the appendage oblong, obtuse at the apex, copiously gland-edged, the glands minute and rounded, mostly sessile, sparsely distributed throughout the connective. Ovary 4-locular, completely inferior, 1.6–1.75 mm long at anthesis, the apical collar absent, the apex 0.15 mm in diameter, somewhat depressed, glabrous to inconspicuously glandular-puberulent; style 4–4.5 mm long, tapering distally, glabrous; stigma truncate to expanded-truncate. Berries 3.8–4.2 × 4–5 mm when dry, globose-oblate, bright pink ripening bright blue, the hypanthial indumentum early caducous at maturity. Seeds 0.33–0.38 × 0.14–0.16 mm, ovoid, angled, light brown; lateral and antiraphal symmetrical planes ovate, the highest point toward the chalazal side or near the central part of the seed; raphal zone suboblong, nearly as long as the corpus of the seed, extending along its entire length, ventrally and longitudinally expanded, dark brown; individual cells elongate, anticlinal boundaries moderately channeled, undulate, with Ω- and U-type patterns; periclinal walls convex, low-domed to nearly flat, microrelief punctate.

Additional specimens studied:— COLOMBIA. Chocó: (San José del Palmar), San José del Palmar- Corondoto road, ca. 16–25 km Wof San José del Palmar , 4°42’N, 76°27’W, 300–366 m, 14 May 1984, Luteyn et al. 10482 ( MO, NY, US) ; Hwy Bolivar-Quibdó, Around km 155 by Río Atrato , 5°40’N, 76°22’W, 500 m, 11 March 1984, Juncosa 2427 ( MO) ; Along road between San José del Palmar and Novita , vicinity of Santa Rosa at end of existing road, 4°57’N, 76°30’W, 350 m, 11 May 1983, Croat 56612 ( MO) . Valle: (Buenaventura), Vereda Bellavista, P.N.N. Farallones, Sector Bajo Anchicayá, Zona en cercanías de la Draga , 3°36.906’N, 76°54.107’W, 225 m, 11 January 2013, Alvear et al. 1567 ( CAS, COL) . COSTA RICA. Bois de la vallée de Tuis, 600 m, 1 November 1893, Pittier 8123 ( US) . Alajuela: Cataratas de San Ramón , 20 February 1931, Brenes 13520 (F, NY) ; R.F. de San Ramón, Quebrada Cacical , 10°14’13"N, 84°36’22"W, 2 May 1987, Herrera et al. 588 ( CAS, MO) ; Alongroadto Colonia Virgendel Socorro, barrancaof Río Sarapiquí , 10°15’N, 84°10W, 700–800 m, 8 August 1979, Stevens 13536 ( CAS, F, MO) ; Cataratas de San Ramón , 1 May 1931, Brenes 13691 ( CR, F, NY); (San Ramón) , R.F.Camino, Entrada cerca a la Estación, 10°13’N, 84°37’W, 900 m, 4 May 1991, Gómez-Laurito & Mora 12056 ( CR, F) ; R.B. Monteverde, Río Peñas Blancas, Quebrada Azufre , 10°19’N, 84°42’W, 800 m, 22 December 1990, Bello 2598 ( INB, MO) ; Monteverde R., Peñas Blancas river valley, Atlantic slope rain forest, 10°20’N, 84°43’W, 800–900 m, 3 October 1986, Haber et al. 6077 ( MO) ; Monteverde Cloud Forest R., Peñas Blancas river valley, Atlantic slope rain forest, Farms of Jesus Rojas and Alejandro García , 10°20’N, 84°45’W, 900 m, 28 November 1986, Haber & Bello 6435 ( MO); ( Río Peñas Blancas ) , R.Monteverde, Vertiente Atlántica. Sendero a Finca de Jesús de Novo , 10°19’N, 84°44’W, 950 m, 24 July 1987, Haber & Bello 7364 ( CR, MO); (Grecia) , R.V.S. Bosque Alegre, Cuenca del Sarapiquí, Laguna Hule , 10°17’55.456"N, - 84°12’54.057"W, 700–800 m, 28 July 2002, Kriebel & Larraguivel 676 ( INB, MO); (San Ramón) , No protegida, Cuenca del San Carlos, Curso de plantas, 10°13’0"N, - 84°35’20.0001"W, 800–900 m, 11 August 2002, Kriebel 908 ( INB, MO) ; Peñas Blancas river valley NE of San Carlos, along road from Jabillos , 10°22’N, 84°38’W, 650 m, 29 June 1985, Hammel et al. 14076 ( MO); (Alajuela) , Virgen del Socorro, No protegida, Cuenca del Sarapiquí , 10°15’25"N, - 84°10’20.0001"W, 800 m, 21 July 2002, Kriebel 538 ( INB, MO) ; Cataratas de San Ramón (desecho babezas), 28 February 1931, Brenes 13553 ( CR, F) ; Cataratas de San Ramón , 19 February 1931, Brenes 13424 ( CR, F) . Cartago: Area near Pavones , about 15 km Eof Turrialba, 600 m, 7 May 1956, Williams 19715 (F, NY) ; 13 km Eof Turrialba on the hwy to Limón, Canyon of Río Chitaría , 9°55’N, 83°36’W, 750–800 m, 10 May 1983, Liesner et al. 15417 ( CAS) ; Vicinity of Pejivalle, 900 m, 7 February 1926, Standley & Valerio 47078 ( US); ( Turrialba ) , R.B. Barbilla, Cuenca del Matina, Sendero Las Quebradas , 9°57’48"N, 83°27’51"W, 500 m, 8 October 1999, Mora- Rojas & Rojas 511 ( INB, MO) . Guanacaste: (Río Chiquito de Tilarán), Río Negro valley, Atlantic slope, lower montane wet forest, 10°22’N, 84°52’W, 1400 m, 30 July 1986, Haber Bello 5837 ( MO) . Heredia: In forest along canyon of the Río Sarapiquí, about 10 km from Cariblanco, upstream from bridge on road to La Virgen del

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 115 Socorro, 10 June 1982, Hammel et al. 12832 ( CAS); ( Colonia Virgen del Socorro ) , Ca, 5 km Eof Cariblanco, 850 m, 13 August 1974, Maas 1287 (F, US); ( Virgen del Socorro ) , Barranca del Río Sarapiquí, 700 m, 3 February 1984, Gómez-Laurito 9873 (F); ( Colonia Virgendel Socorro ) , Virgendel Socorro–Río Sarapiquí–Cariblanco , 600–800 m, 31 August 1983, Chacón & Herrera 1221 ( CAS, CR, F) . Limón: (Siquirres), Las Brisas de Pacuarito , 500 m, 17 April 1985, Gómez et al. 23375 ( CAS, CR) ; Rainforest slopes of Cerro Skopte just Wof Río Siori about 7 km beyond Coroma , 450–700 m, 18 February 1992, Almeda & Daniel 7031 ( CAS, CR) ; Flood plain of the Río Toro Amarillo ca. 6–7 km Sof Guápiles, Sof the suspension bridge, 420–440 m, 22 June 1975, Lellinger et al. 1840 (F) ; Cordillera de Talamanca, Ridge separating Quebrada Cañabral from Río Barbilla , and slope leading down to the latter, 10°2’N, 83°26’W, 200–400 m, 4 September 1988, Grayum et al. 8750 ( CAS, CR, MO) ; Ridge between Río Chirripó and Quebrada El Molinete, Wof new highway, 10°12’N, 83°54’W, ca. 400 m, 19 July 1984, Grayum et al. 3558 ( CAS, MO); (Talamanca) , Amubri, Camino entre Amubri y Cachabri, margen derecha del Río Labri , 9°29’40"N, 82°59’40"W, 100–200 m, 27 June 1989, Herrera 3033 ( CAS, CR) ; Along a stream by trail between, Sitio Rangalle and Cuen , Between Cerros Jacrón & Bitarkara , 700–900 m, 25 March 1983, Hazlett 5146 ( CAS, F, NY); ( Talamanca ) , P.N. La Amistad, Cuenca del Sixaola, Bratsl, Punto 10 NE de Laguna Sacabico, 9°37’22.15"N, - 83°16’43.53"W, 1000–1100 m, 27 July 2007, Solano et al. 4542 ( INB, NY, PMA); ( Matina ) , P.N. Barbilla, Cuenca del Matina, Colonia Puriscaleña , Sendero Cerro Azul , 9°59’44.9069"N, - 83°23’8.4987"W, 400 m, 7 March 2000, Mora 872 ( INB, MO) . Puntarenas: (Golfito), R.N.V.S. Golfito, Camino a las torres del ICE, cabecera del Río Sorpresa , 5 km al E, 8°39’25"N, 83°9’44"W, 300–400 m, 1 February 1992, Zamora et al. 1766 ( CAS, INB, MO) ; 4 km NE of Escuela Santa Constanza, Nof San Vito de Java, 760 m, 27 September 1968, Davidse & Pohl 1275 (F, ISC, US); ( Osa ) , R.F. Golfo Dulce, Península de Osa, Rancho Quemado , Tierra de Conservación , 8°43’0"N, 83°34’50"W, 200–350 m, 11 July 1993, Aguilar 1969 ( CAS, CR, MO); ( Osa ) , Rancho Quemado, Fila División, Entre Rancho Quemado y junta de quebrada, 8°41’40"N, 83°36’30"W, 350 m, 18 July 1991, Marín 25 ( CR, MO) ; Monteverde community, Windbreak next to road, 10°20’N, 84°50’W, 1450 m, 22 July 1986, Haber 5733 ( MO); (Osa Peninsula) , Corcovado N.P., 0 km to 1 km upstream from Los Chiles , 8°31’N, 83°31’W, 300–400 m, 10 July 1977, Liesner 3120 ( MO); (Osa) , R.F.Golfo Dulce, En el valle de Rancho Quemado, ca. 15 km al Wde Rincón, en el Sdel valle por la Quebrada Quebradona y Río Riyito , 8°40’0"N, 83°34’0"W, 200 m, 13 September 1990, Hammel et al. 17855 ( CR, MO) ; Eastern Osa Peninsula, more than 500 m from forest edge on the Ecoast of the Osa Penninsula , 8.70681°N, - 83.59142°W, 25 January 2003, Mayfield s.n. ( MO) . San José: (Pérez de Zeledón), Cuenca del Barú, Fila Tinamastes , por la carretera entre Dominical y San Isidro, 9°18’43"N, 83°46’19"W, 950 m, 9 September 1996, Hammel et al. 20440 ( INB, MO); ( Pérez de Zeledón ) , No protegida, Cuenca del Savegre, Savegre Abajo , 3 km Ny 500 m W, de Savegre Abajo, 9°26’59"N, - 83°50’55"W, 800 m, 7 February 2001, Rodríguez et al. 6998 ( INB, MO, NY); ( Tarrazú ) , Zona Protectora Cerro Nara, 9°29’40"N, 84°1’0"W, 840 m, 13 Mayo 1997, Estrada & Sánchez 784 ( CR, F, MO); (Tarrazú) , Los Llanos de Santa María, Filas Las Rejas , 9°32’30"N, 84°1’0"W, 700 m, 9 July 1997, Valverde et al. 84 ( CR, MO) . ECUADOR. Esmeraldas: Environs of Lita, on the Ibarra-San Lorenzo RR, 550–650 m, 9 June 1978, Madison et al. 5098 (F, QCA, US); ( Quinindé ) , Bilsa Biological Station, Mache mountains, 35 km Wof Quinindé, 5 km Wof Santa Isabel, Near the Station's shower, 0°21’N, 79°44’W, 400–600 m, 10 April 1995, Clark & Troya 677 ( MO, QCNE, US); ( Quinindé ) , Bilsa Biological Station, Montañas de Mache , 35 km Wof Quinindé, 5 km Wof Santa Isabel, Mature forest in property of Sr. Rios , Sof Station , 0°21’N, 79°44’W, 400–600 m, 9 May 1995, Clark & Watt 821 ( CAS, MO, NY, QCNE); ( Quinindé ) , Bilsa Biological Station, Montañas de Mache , 35 km Wof Quinindé, 5 km Wof Santa Isabel, Steep muddy slope near large waterfall SE of Station , upstream of Station shower, 0°21’N, 79°44’W, 400–600 m, 22 September 1994, Baas & Abbott 48 ( MO, NY) . Imbabura: ( Lita ), 501 m, 23 April 1949, Acosta-Solís 12195 (F) . Los Ríos: Río Palenque B.R., Km 56 Quevedo to Sto. Domingo , 220 m, 15 December 1971, McMahon 4280 ( US) ; Río Palenque B.R., km 56 Quevedo to Sto. Domingo , near creek SW of station, 220 m, 31 March 1972, Vrieze et al. 4312 ( NY, QCA) ; Río Palenque B.R., km 56 Quevedo to Sto, Domingo , 150–220 m, 5 September 1972, Dodson & McMahon 5099 ( MO, NY, QCA) . NICARAGUA. Zelaya: ca. 6.3 km Sof bridge at Colonia Yolaina and ca. 0.8 km Sof ridge of Serranias de Yolaina on road to Colonia Manantiales ( Colonia Somoza ), 11°36–37’N, 84°22’W, 200–300 m, 13 February 1978, Stevens 6418 ( CAS, MO) ; 6.3 km Sof bridge at Colonia Yolania on road to Colonia Manantiales of Nueva Guinea , 200–300 m, 13 February 1978, Vincelli 244 ( MO) . PANAMA. Bocas del Toro: Road from Fortuna Dam to Chiriquí Grande , 10 mi from Continental Divide , 8°51’N, 82°10’W, 400 m, 2 August 1984, Churchill 5931 ( CAS, MO) ; Along oleoducto road between contienetal divide and Chiriquí Grande , 8°50–55’N, 82°9–15'‘W, 350–400 m, 30 April 1985, Hammel 13692 ( MO) . Coclé: (El Copé), on slope and ridge Wof sawmill, 5 April

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1978, Hammel 2434 ( CAS, MO) . Colón: Trail from Alto Pacora to Cerro Brewster, 9°18’N, 79°16’W, 700 m, 18 November 1985, de Nevers et al. 6229 ( CAS, MO) . Darién: Cerro Pirre, 762–1372 m, 9 August 1967, Duke & Elias 13790 ( MO) ; Trocha desde "Rancho Frío 2" hasta un poco más abajo de "Rancho Frío 1", Cerro Pirre, 13 October 1989, Palacios et al. 503 ( CAS, PMA) ; P.N. del Darién, Slopes of Cerro Mali, head waters of Sbranch of Río Pucuru , ca. 22 km Eof Pucuru, 8°4.5’N, 77°14’W, 1300–1400 m, 21 October 1987, Cuadros et al. 3935 ( CAS, MO) ; P.N. del Darién, Along Sbranch of Río Pucuro , forest and ridge Sof river & up river from old village of Tacarcuna , ca. 18 km Eof Pucuro, 8°4’N, 77°16’W, 600–800 m, 21 October 1987, Hammel et al. 16389 ( CAS, MO) ; Cercanías de "Rancho Frío 2", trocha hacia la cima de Cerro Pirre, 12 October 1989, Palacios & Polanco 468 ( CAS, PMA) ; Bank above the Río Paca , 26 June 1959, Stern et al. 594 ( MO, NY, US) ; Middle slopes on W side of Cerro Pirre, 7°57’N, 77°46’W, 550–760 m, 28 June 1988, Croat 68878 ( MO) . Panamá: (Distrito de Capira), Cerro Trinidad, Límite Edel P.N, Campana , 8°44’30"N, 79°57’45"W, 800–900 m, 24 June 1994, Galdames et al. 1169 ( CAS, US) ; Gorgas Memorial Labs yellow fever research camp, ca 25 km NE of Cerro Azul on Río Piedras, 550 m, 20 November 1974, Mori & Kallunki 3291 ( MO) ; Trail to top of Cerro Pelado, 1000 m, 16 June 1979, Antonio 1108 ( MO) . Veraguas: (Guabal), Río Dos Bocas , about 16 km NW of Santa Fe, ca. 500 m, 30 March 1975, Dressler 5013 ( CAS, US) .

Illustration:— Fig. 22 .

Common names and documented uses:— Ecuador: “blueberry” (Vrieze et al. 4312, NY!).

Habitat, distribution and ecology:— Rare to common, usually found on steep riparian rocky banks in rain forests and cloud forests, from Nicaragua south through southern Central America to Colombia and Ecuador ( Fig. 16 ), at (12–) 100–1450 m. In Nicaragua it is only known from the former department of Zelaya. It is more common throughout Costa Rica and Panama. In Colombia it has only been collected in Chocó; in Ecuador it is known from the Pacific Andean slope.

Phenology:— Collected in flower from December through October; in fruit throughout the year.

Etymology:— The specific epithet comes from the Latin radic (= a root), and might refer to the numerous aerial roots on the type specimens.

Discussion:— This species is distinguished by its prevailingly rupicolous habit, pronounced foliar dimorphism, and dense sessile-stellate indumentum. These features are also present in its closest relative M. biolleyana , but in M. radicans the foliar dimorphism is more striking (1:6–8 vs. 1:2), and the blade shape is different (elliptic-oblong and falcate vs. broadly ovate to subrotund). The inflorescences, although architecturally similar are fewer flowered in M. radicans than they are in M. biolleyana .

The thickened callose structure produced at the abaxial foliar base might be an acarodomatium, although it is less prominent than in M. biolleyana and no insects or eggs of any kind where found in the specimens studied. This domatium appears to be both taxonomically and phylogenetically significant since it is present in the trio of closely related species including M. biolleyana , M. erikasplundii , and M. radicans .

Conservation status:— This species would be considered Endangered EN B2ab(iii) based on IUCN criteria ( AOO). However, because it occurs in many protected zones, a status of Least Concern LC is justified. In Colombia it is protected in Farallones National Park ( Valle ). In Costa Rica it is protected in the San Ramón Forest Reserve and in the Monteverde Biological Reserve (Alajuela) ; in the Barbilla and La Amistad National Parks (Limón) ; in the Golfo Dulce Forest Reserve and the Golfito Wild Life Reserve (Puntarenas) ; in the Cerro Nara Portected Zone. In Ecuador it is protected in the Mache-Chindul Ecological Reserve (Esmeraldas) ; and in the Río Palenque Biological Station (Los Ríos) . Protected in Panama in Darién National Park (Darién) .

Figures

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

FIGURE 22. Miconia radicans. A. Habit, with detail of abaxial foliar base. B. Flower. C. Stamen, abaxial view (left), lateral view (right). D. Berry. E. Seed. Based on Zamora 1766, CAS. Drawn by Sean V. Edgerton.