Miconia aguilarii ( Kriebel & Almeda 2009: 207–211 ) 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 : 37-40

1. Miconia aguilarii ( Kriebel & Almeda 2009: 207–211) Gamba & Almeda , comb. nov. Basionym: Clidemia aguilarii Kriebel & Almeda. Type: COSTA RICA. Prov. Puntarenas: along gravel road to microwave tower above Golfito , 0.7 m above main road, 08°39’N, 83°11’W, 90 m, 15 September 1987, Croat 67598 (holotype: CAS!; isotype: MO).

Suffrutescent herb or shrub (0.3–)0.4–2(–3) m tall, densely or laxly branched, bark brownish. Upper internodes (0.6–)1.1–5.8(–8.5) cm long, terete like the cauline nodes, nodal line absent. Indumentum on branchlets, petioles, adaxial base of leaf surfaces, primary and secondary leaf veins, inflorescence axes, bracts, bracteoles, and pedicels densely to copiously composed of brownish subsessile or thin stipitate dendritic trichomes (0.1–)0.2–0.3(–0.42) mm long with short axes and few-moderate number of terete arms, sparsely intermixed with caducous elongate smooth trichomes up to 3 mm long. Leaves of each pair somewhat unequal in size; the semiterete petioles 0.4–2.5(–5.85) cm long, adaxially canaliculate; larger blades (6.3–)11–23 × (3.4–) 5–11.5 cm, smaller blades 2.5–10 × 1.7–7 cm, elliptic-ovate, the base attenuate to oblique and narrowly decurrent along the petiole, the margins ciliolate-denticulate, the apex acuminate, membranaceous to chartaceous; mature leaves (adaxial surface) glabrescent except for a few elongate smooth trichomes near the margins 0.5–1 mm long, the primary, secondary, tertiary and higher order veins glabrous; abaxial surface together with the tertiary and higher order veins with a sparse and caducous resinous indumentum of slightly furrowed more or less stalked glands 0.1–0.2 mm long; 5- or 7-(9-)plinerved, including the tenuous marginals, innermost pair of secondary veins diverging from the primary vein 0.5–2.8 cm slightly asymmetrically above the decurrent base, areolae 0.8–1.5 mm, adaxially the primary and secondary veins impressed, the tertiary and higher order veins slightly impressed, abaxially the primary and secondary veins elevated and terete, the tertiary and higher order veins slightly elevated. Inflorescences a pseudolateral dithyrsoid 2.25–6.95 cm long, including a peduncle (0.2–) 0.4–1.5 cm long, the central flower of some dichasia and of the main axis occasionally aborted, divaricately branched, borne on the upper leafless nodes, the rachis green; bracts 0.6–0.75(–1.2) × 0.15–0.25 mm, spatulate to subulate, spreading and somewhat concave, on both surfaces the indumentum sparse and caducous, densely intermixed with resinous slightly furrowed more or less stalked glands 0.1–0.2 mm long; bracteoles 0.2–0.6 × 0.1–0.3 mm, subulate, spreading, on both surfaces the indumentum dense, sparsely intermixed with resinous-glandular trichomes similar to those of the bracts; both persistent in flower and commonly deciduous in fruit. Flowers 4-merous, the central flower of each dichasium sessile to subsessile, the rest on pedicels 0.25–0.5 mm long. Hypanthia at anthesis 1.7–1.85(–2.1) × 0.8–1.15 mm, free portion of hypanthium 0.95–1.15(–1.4) mm long, urceolate, bluntly 8-ribbed, green-whitish, caducously resinous with slightly furrowed more or less stalked glands 0.03–0.05(–0.1) mm long, inner surface and torus ridged, sparsely and caducously resinous-glandular with glands like those of the hypanthium exterior. Calyx open in bud and persistent in fruit, light green; tube 0.25–0.35 mm long, adaxially with the same indumentum as the inner torus, abaxially as the hypanthium; lobes 0.2–0.4 × 0.5–0.8(–0.99) mm, broadly triangular, the margin vaguely undulate, the apex bluntly acute, caducously resinous-glandular on both surfaces; exterior teeth (0.1–) 0.2–0.25 mm long, bluntly triangular, inserted near the base of the calyx lobes and rarely equaling them, caducously resinous-glandular on both surfaces. Petals 0.65–0.85(–1.25) × 0.75–1 mm, obovate-oblong, with a minute projecting infra-apical tooth on the abaxial surface, the margin repand-undulate, the apex obtuse to slightly truncate, dull-greenish to white, densely and caducously papillose on both surfaces, reflexed at anthesis. Stamens 8; filaments 0.9–1.4 × 0.25 mm, light-green to white, glabrous; anther thecae 1.25–1.95 × 0.25–0.5 mm, linear-oblong and subulate, truncate-acuminate at the apex, opening by one dorsally inclined pore 0.11–0.15 mm in diameter, whitish to yellow at anthesis; connective yellow, its prolongation and appendage 0.25–0.35 mm long, the appendage deltoid-spatulate, bluntly acute to obtuse at the apex, densely and caducously gland-edged, also sparsely and minutely glandular toward the center and throughout the connective. Ovary 4-locular, completely inferior, 0.7–0.75 mm long at anthesis, the apical collar absent, the apex ca. 0.6 mm in diameter, somewhat depressed, deciduously glandular-puberulent; style 3.5–5.5 mm long, narrowed distally (i.e. tapering), white, glabrous; stigma capitellate at anthesis (truncate when dry). Berries mostly 3–5 × 2.5–3.5 mm when dry, globose and slightly oblate, initially white turning bright orange when ripe, hypanthium indumentum somewhat persistent at maturity. Seeds 0.28–0.33 × 0.16–0.19 mm, ovoid, angled, light-brown; lateral and antiraphal symmetrical planes ovate, the

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 37 highest point toward the chalazal side; raphal zone suboblong, nearly as large as the corpus of the seed, extending along its entire length, ventrally and longitudinally expanded, dark-brown; individual testa cells elongate, anticlinal boundaries channeled, undulate, with Ω- and U-type patterns; periclinal walls convex, low-domed, microrelief slightly punctate to smooth.

Additional specimens studied:— COSTA RICA. Guanacaste: (Nandayure), No protegida, Península de Nicoya , Bejuco , Pilas de Bejuco , Parche de bosque en cercanías de la carretera, 9°50'52"N, 85°21'27"W, 10 m, 10 March 1999, Rodríguez & Hurtado 4599 ( INB, MO); (Nandayure) , Área no protegida, Pacífico Norte, Bejuco , Pilas de Bejuco , Finca de Abel Rodríguez , 9°51’38"N, 85°21’33"W, 60 m, 13 July 1994, Estrada & Rodríguez 10 ( INB, MO) . Puntarenas: Steep forested and partially disturbed sites above Golfito along the road to the television tower, 50–500 m, 16 July 1977, Almeda et al. 3094 ( CAS) ; Nicoya Peninsula, Curú, Canyons and ridges towards Punta Georgia , Punta Blanca Trail , 9°46–48’N, 84°54–58’W, ca. 100 m, 23 August 1995, Sanders et al. 17706 ( CAS, CR) ; Steep forested and partially disturbed sites above Golfito along the road to the television tower, 50–500 m, 16 July 1977, Almeda et al. 3092 ( CAS) ; P.N. Corcovado, Ridges above Río Claro , 8°28’N, 83°35’W, 0–100 m, 24 November 1981, Knapp & Mallet 2209 ( CAS, MO); (Golfito) , P.N. Corcovado, Península de Osa, Estación Sirena, Sendero a Río Claro , arriba de la fila, 8°28’0"N, 83°35’0"W, 10 m, 14 October 1993, Aguilar 2529 ( CAS, CR); (Golfito) , P.N. Corcovado, Península de Osa, Estación Sirena, Sendero Pavo , 8°28’50"N, 83°35’30"W, 10 m, 23 July 1994, Aguilar 3507 ( CAS, CR, MO) ; Osa Peninsula, Road to Carate from Puerto Jimenez , ca. 8 km Sof Río Tamales, 185 m, 6 September 2005, Penneys & Blanco 1811 ( NY) ; R.B. Carara, Sector Quebrada Bonita , sitio área administrativa, 9°46’10"N, 84°36’30"W, 50 m, 13 September 1990, Zúñiga 256 ( CR, NY, MO); (Aguirre) , No protejida, Cuenca del Naranjo y Paquita , Manuel Antonio , Finca Anjalí , 9°24’16"N, 84°9’16"W, 150 m, 12 November 2006, Solano & Solano 3527 ( INB, NY) ; P.N. Manuel Antonio, Quebrada Camaronera , 9°23’N, 84°9’W, 1 m, 3 August 1990, Harmon 136 ( CAS, CR, MO) ; P.N. Corcovado, Ollas Trail , 8°27–30’N, 83°33–38’W, 0–150 m, 26 July 1988, Kernan & Phillips 717 ( CAS, CR, MO) ; P.N. Corcovado, Ollas trail, 8°27–30’N, 83°33–38’W, 0–100 m, 10 January 1989, Kernan & Phillips 883 ( CR, MO); (Golfito) , P.N. Corcovado, Península de Osa, Estación Sirena, alrededores de la estación y del Sendero La Olla , 8°28’0"N, 83°35’0"W, 10 m, 26 October 1990, Fonseca 9 ( CR, MO); (Golfito) , P.N. Corcovado, Península de Osa, Estación La Leona , junto a Río Madrigal , 8°27’0"N, 83°29’30"W, 100 m, 19 January 1993, Aguilar 1603 ( CR, MO) ; P.N. Corcovado, Osa Peninsula, 0 km to 2 km Wof the park headquarters at Sirena , 8°29’N, 83°36’W, 0–200 m, 5 July 1977, Liesner 2912 ( MO); (Golfito) , P.N. Corcovado, Península de Osa, Sendero Río Claro , 8°28’50"N, 83°35’30"W, 10 m, 12 September 1998, Aguilar 5542 ( INB, MO) ; Littoral woods Eof Punta Voladora, near entrance to Golfito Bay , 8.38°’N, 83.11°’W, 24 July 1977, Webster 22037 ( MO); (Golfito) , R.F. Golfo Dulce Los Mogos, 8°45’30"N, 83°22’30"W, 100–200 m, 5 September 1991, Aguilar 335 ( CR, MO) ; Along gravel road to microwave tower above Golfito , 0.7 km above main road., 8°39’N, 83°11’W, 90 m, 15 September 1987, Croat 67598 ( MO) ; R.N.A. Cabo Blanco, Península de Nicoya, Estación San Miguel , límite de la Reserva, 9°35’42"N, 85°7’13"W, 355 m, 10 February 1987, Alvarado 115 ( INB, MO) ; P.N. Corcovado, Sirena, Río Claro Trail , 8°27–30’N, 83°33–38’W, 0–150 m, 11 February 1988, Kernan 109 ( MO) ; Punta Catedral , ca. 7 km SE of Quepos, 9°22.5’N, 84°9’W, 20–70 m, 20 August 1985, Grayum et al. 5896 ( CAS, MO) ; R.B. Carara, Sendero Quebrada Bonita-Bijagual, Sitio Montaña Jamaica , 9°46’10"N, 84°33’50"W, 550 m, 8 December 1989, Zúñiga et al. 18 ( CAS, CR, MO) ; P.N. Corcovado, Golfito, Estación Sirena , al borde de bosque primario, 8°29’0"N, 83°36’0"W, 2 m, 1 October 1990, Saborio et al. 78 ( CAS, CR, MO); (Golfito) , P.N. Corcovado, Península de Osa, Sirena , 8°28’0"N, 83°35’0"W, 1–20 m, 15 June 1990, Maass 37 ( CAS, CR, MO) ; R.N.A. Cabo Blanco, Estación Cabo Blanco , 9°35’N, 85°6’W, 20–100 m, 4 November 1991, Chavarría 305 ( CAS, CR, MO) ; Punta Banco , 0–200 m, 22 August 1988, Chavarría-Díaz et al. 266 ( CR, MO) ; Corcovado N.P., Osa Peninsula, Primary forest on hills 0 km to 1 km Wof the park headquarters at Sirena , 8°29’"N, 83°36’"W, 0–200 m, 4 July 1977, Liesner 2834 ( MO) ; R.B. Carara, Estación Quebreda Bonita, 30 m, 29 September 1983, Chacón 1398 ( CR, MO); (Puntarenas) , R.N.A. Cabo Blanco, Península de Nicoya, Sendero Central , 9°35’0"N, 85°7’0"W, 0–200 m, 17 December 1993, Fernández et al. 1315 ( MO, INB); (Golfito) , P.N. Corcovado, Península de Osa, Estación Sirena , Alrededor de Sendero Ollas , 8°28’50"N, 83°35’30"W, 10 m, 6 February 1994, Aguilar 3076 ( CAS, INB); (Puntarenas) , Estación San Miguel , ca 2 km Sde Malpais, 9°35’40"N, 85°7’50"W, 0 m, 18 January 1996, Hammel 20091 ( CAS) . San José: (Turrubares), Valle del Tárcoles, SE de San Luis, entre Quebrada Zorrillal y Cerro San Luis , 9°50’20"N, 84°28’40"W, 600–950 m, 20 December 1995, Hammel & González 20033 ( CR, MO); (Ciudad Colón) , Zona Protectora El Rodeo, 9°54’N, 84°16’W, 600 m, 4 August 1992, Varela 241 ( CAS, CR, MO) . PANAMA. Chiriquí: (San Bartolo), Burica

38 Phytotaxa 179 (1) © 2014 Magnolia Press

GAMBA & ALMEDA

Peninsula , Near costa Rica border, 12 mi Wof puerto Armuelles, 400–500 m, 24 February 1973, Croat 22197 ( MO) ; Burica Peninsula , Quebrada Quanabanito, 1 km Wof the dam at la Represa, 20 February 1973, Busey 480 ( MO) ; Vicinity of Puerto Armuelles , 0–75 m, 28 July 1940, Woodson & Schery 855 ( MO) ; Burica Peninsula , Quebrada Tuco, 9 mi Sof Puerto Armuelles, 0–150 m, 21 February 1973, Liesner 171 ( MO); (San Bartolo) , Burica Peninsula , 20 km Wof Puerto Armuelles, 400–600 m, 22 February 1973, Busey 563 ( MO) ; Burica Peninsula , Quebrada Merida, 4mi S Puerto Armuelles, 0–100 m, 3 March 1973, Liesner 392 ( MO) . Los Santos-Veraguas : Mountains of southern Azuero Peninsula , near proposed route of road from El Cortezo ( Prov , Los Santos ), to Arenas (Prov, Veraguas), 0–5 km SW of El Cortezo, 305 m, 29 October 1978, Hammel 5461 ( CAS, MO) .

Veraguas: ( Isla de Coiba), Campamento de La Salina , subida hacia Río Negro, 5 m, 21 October 1995, Aranda et al. MB1345 ( CAS, MO) .

Illustration:— Kriebel & Almeda (2009): 208, fig. 1.

Common names and documented uses:— None recorded.

Habitat, distribution and ecology:— Miconia aguilarii is known from the Pacific slope of Costa Rica and Panama ( Fig. 11 ), at 0–600(–950) m, where it grows in the understory of primary or secondary rain forest, and along banks of rocky streams. Among the collections studied, it has been considered locally common to occasional. In Costa Rica, it is widely distributed, ranging from the Nicoya Peninsula on the northern Pacific coast, through the central Pacific lowlands in El Rodeo Protected Ciudad Colón Zone, south to the Osa Peninsula. In Panama it is known from few collections in a region ranging from the Burica Peninsula south to Coiba Island, and at the border between Los Santos and Veraguas in the mountains of southern Azuero Peninsula.

Phenology:— Collected in flower from January through March and from June through October; in fruit from July through February.

Etymology:— The specific epithet honors Costa Rican botanist Reinaldo Aguilar, whose explorations of the Osa Peninsula have contributed much to our knowledge of that region's rich rain forest flora.

SYSTEMATICS OF THE OCTOPLEURA CLADE OF MICONIA

Phytotaxa 179 (1) © 2014 Magnolia Press 39

Discussion:— Miconia aguilarii was long overlooked and commonly determined as Ossaea quinquenervia or Clidemia quinquenervia , synonyms of what is here treated as Miconia quinquenervia . Within the Quinquenervia subclade both are part of a closely related group of species that includes M. reitziana ( Fig. 1 ). Within this subclade, M. aguilarii and M. quinquenervia are vegetatively very similar, especially in characters of pubescence (rustyasperous indumentum mixed with simple trichomes) and leaves (margins ciliolate-denticulate, base decurrent on the petiole). Their inflorescence architecture is also identical. The consistent differences between these two species include flower merosity (4-merous in M. aguilarii vs. 5-merous in M. quinquenervia ), exterior calyx teeth (not setose vs. setose), and the color of immature hypanthia (green-whitish vs. yellowish becoming bright pink). The most outstanding difference is the color of the mature berries (bright orange vs. purple-black) ( Kriebel & Almeda 2009).

In this study, additional specimens belonging to M. aguilarii were found among collections of M. quinquenervia received on loan, particularly from the Burica Peninsula in Chiriquí Province ( Panama, Pacific side), a locality from which M. aguilarii had not been reported previously. Although M. quinquenervia is more widespread than M. aguilarii (ranging from Honduras south through Central America, to Colombia, Venezuela and Ecuador), its restriction to the Caribbean slopes of Central America is consistent with its reported distribution in Costa Rica and Panama, where it is also only known from the Caribbean slope ( Kriebel & Almeda 2009).

This species, which is common in the understory of tropical forests, appears to be tolerant of deep forest shade. Accordingly, it may flower when less than 1 m tall and consequently the labels on at least seven collections examined describe the habit as herbaceous. This is common among other species in the Octopleura clade, including M. reitziana , which is also apparently tolerant of deep shade and has been described as a suffrutescent herb.

When rehydrated buds of immature flowers where examined, it was found that in at least four (A.C. Sanders et al. 17706, CAS!), the undeveloped stamens ranged from nine to 11. Further studies of flower development in M. aguilarii would clarify if these stamens are aborted before anthesis, or if the number of stamens is variable within the species. On mature flowers, the number of stamens was consistently eight.

Conservation status:— Based on IUCN criteria (AOO), this species would be considered Endangered EN B2ab(iii). However, it occurs in many protected areas, warranting a status of Least Concern LC. In Costa Rica it is protected in the Cabo Blanco Absolute Reserve, Curú Wildlife Refuge, El Rodeo Protected Zone, and Carara, Manuel Antonio and Corcovado National Parks ( Kriebel & Almeda 2009). In Panama it is only protected in the Coiba National Park.

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 11. Distributions of Miconia aguilarii, M. alatissima, M. albertobrenesii, and M. alboglandulosa.