Miconia galeiformis and Miconia neei (Miconieae: Melastomataceae ...

Miconia galeiformis and Miconia neei (Miconieae: Melastomataceae), two new species from Bolivia JANELLE M. BURKE

AND

FABIÁN A. MICHELANGELI

Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458-5126, USA; e-mail: [email protected]; e-mail: [email protected]

Abstract. Two new species of Miconia from Bolivia are described: Miconia galeiformis and Miconia neei. Both species occur in Andean montane forests. Miconia galeiformis (sect. Chaenopleura) is distinctive within Miconia due to the presence of setose stems, petioles and abaxial leaf surface, and a large, globose stigma. Miconia neei bears unisexual flowers, and can be distinguished from similar species of Miconia sect. Cremanium in Bolivia based on the furfuraceous stems and young nodes, in addition to serrulate leaf margins. The documentation of dioecy in Miconia neei and the occasional occurrence of unisexual flowers in Miconia galeiformis adds to our burgeoning understanding of complex breeding systems within Miconia. Key Words: Bolivia, dioecy, Miconia section Chaenopleura, Miconia section Cremanium. Resumen. Se describen dos especies nuevas de Bolivia: Miconia galeiformis y Miconia neei. Ambas especies crecen en bosque montano húmedo de los Andes. Miconia galeiformis (secc. Chaenopleura) es una especie única en Miconia debido a sus tallos, pecíolos y el envés de las hojas setosos, y el estigma globoso. Miconia neei tiene flores unisexuales, y se distingue de otras especies de Miconia secc. Cremanium por sus tallos y nodos furfuráceos, y los márgenes de las hojas serrulados. El reporte de Miconia neei como especie dioica y la presencia ocasional de flores aparentemente unisexuales en Miconia galeiformis contribuyen a nuestro creciente conocimiento de los sistemas reproductivos en Miconia.

The flora of Bolivia remains one of the most poorly known in South America, with no modern study for all groups (Beck, 1998; Frodin, 2001). In recent years, different projects led by the Herbario Nacional de Bolivia in La Paz, the Museo de Historia Natural Noel Kempff Mercado in Santa Cruz, The New York Botanical Garden, and the Missouri Botanical Garden have attempted to either produce local floras or a comprehensive checklist for the entire country (see Nee, 2004). These efforts have increased the number of plant collections, but also highlighted the need for detailed monographic work. One group that is poorly known in Bolivia is the Melastomataceae Juss., and particularly the large and complex genus Miconia Ruiz & Pav. Most of the species of

Miconia from Bolivia were described by Cogniaux (1890a, b, 1891, 1893, 1896, 1907, 1913) based mostly on collections by M. Bang, T. C. J. Herzog, G. Mandon, and H. Rusby. Since then, only a handful of species of Miconia have been described from Bolivian material (Gleason, 1932a, b, 1941, 1950; Wurdack, 1972, 1988; Renner & Beck, 2003), mostly due to the fact that no researcher has actively worked on Bolivian Melastomataceae. Additionally, some species from the Peruvian Andes and western Brazil have been shown to occur in Bolivia (Foster, 1958; Renner, in prep.). Recent field work organized by Michael Nee (NY) has yielded a few new species of Miconia, two of which are described here.

Brittonia, 65(2), 2013 , pp. 171–180 © 2012, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

ISSUED: 1 June 2013

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Miconia, as currently defined, has over 1050 species, but clearly is not monophyletic, with several other genera embedded within (Goldenberg et al., 2008; Michelangeli et al., 2008). Preliminary phylogenetic data place these two new species in a broader Andean clade (Miconieae III sensu Goldenberg et al., 2008), an amalgamation of Miconia sect. Amblyarrhena, Chaenopleura and Cremanium, and several other genera. The most recent, comprehensive sectional classification system (Cogniaux, 1891) is not satisfactory, as demonstrated by Goldenberg et al. (2008); however we give the new species a sectional designation as a means to categorize the vast number of species in Miconia, pending a novel infrageneric classification system based on natural groups. Miconia galeiformis Jan. M. Burke & Michelangeli, sp. nov. Type: Bolivia. Cochabamba: Prov. Carrasco, gorge of Río Ivirizu, 6.6 km by road NE from Monte Punco, on road from Monte Punco to Sehuencas, 17˚32´ 59˝S, 65˚16´13˝W, 2662 m, humid forest in steep canyon, 4 May 2007 (fl), M. Nee, F. Michelangeli, O. Colque I., R. Flores S., D. McClelland & S. Stern 55340 (holotype: NY; isotypes: LPB, MO, USZ). (Figs. 1, 2) Diagnosis: Indument on petioles and abaxial leaf surface setose, hairs translucent. Stigma galeate.

Shrub or treelet, up to 8 m. Young stems terete; internodes solid, longitudinal ridges faint; nodal line absent. Indument of mixed hair types, setose, with translucent or yellow uncinate hairs up to 2.5 mm and furfuraceous with sessile, brown, dendritic hairs 0.1 mm wide, deciduous on stems. Leaves isophyllous; petiole 1.2–2.3 cm long, indument as stem but with denser setose hairs on abaxial surface; lamina 8–10.4×3.1–3.6 cm, lanceolate to elliptical, chartaceous, base round to attenuate, apex acuminate, margin finely serrulate, the teeth 0.4–0.7 mm long, linear, deciduous; veins slightly impressed on adaxial surface and raised on abaxial surface, brown to beige, secondary veins 2 pairs (including faint marginals), basally nerved, tertiary veins percurrent, arched at 45–50˚ angle from midvein, quaternary veins reticulate; adaxial surface dark green, indument on

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lamina strigose, hairs appressed, 1 mm long, indument on primary and secondary veins glandular, glands translucent, sessile; abaxial surface yellow-green, lamina glabrous, indument on veins of mixed hair types, as stem. Inflorescence a terminal panicle, 5–11.6 cm long, with or without auxiliary inflorescences; peduncles flattened, green or brown, indument sparsely furfuraceous of sessile, dendritic hairs; bracts 1×0.3 mm, linear, caducous; bracteoles 0.25–1×0.3–1 mm, linear, caducous before anthesis. Pedicels 0.5–0.75 mm long, furfuraceous, articulated at base. Flowers bisexual, 5merous, diplostemonous. Hypanthium 1.5– 2.8 mm long, campanulate to broadly campanulate, 1.5–3.8 mm wide at the torus, external surface puberulent of sessile, dendritic hairs, internal surface glabrous, androecial fringe absent. Calyx open in bud; tube 0.3–0.5 mm long at anthesis; lobes 0.2–0.6×0.3–0.6 mm, round, glabrous, pink; teeth vestigial to nearly equaling calyx lobes, terete in cross section, 0.2–0.3×0.4 mm, glabrous, green. Petals 1– 2.2×1.2–2 mm, ovate to orbicular, erect, white at anthesis (drying yellow), glabrous, base obtuse, apex entire or retuse, margin entire to erose (apical portion), densely granulous. Stamens isomorphic, radially spreading at anthesis; filament 2–2.3 mm long, puberulent of glandular hairs, white; anthers with 2 locules, thecae 1.5–1.9×1.1–1.3 mm, stout, opening by 2 truncate 0.3 mm pores, septum prolonged or not, pale yellow at anthesis, later turning brown; connective prolonged basally, appendages present or absent, when present, bilobed ventral appendage and 1 short dorsal appendage, yellow, puberulent with glandular hairs. Ovary 3-locular, 2/3 inferior, the free portion projecting 0.4×1.2 mm, conical, with sessile glands, apex present as a corona with 1 ring of glandular hairs; style 2.5–7.5 mm long, accrescent after anthesis, erect, white, puberulent with elongated, roughened hairs; stigma galeiform, 0.5–1.4×0.4–1.2 mm. Berries not seen. Chromosome number unknown. Distribution.—Humid montane forest, 2600–2700 m, in the department of Cochabamba, Bolivia. Phenology.—Flowering in May. Etymology.—The epithet is derived from the latin ‘galea’ for helmet, and refers to the helmetshaped form of the stigma.

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FIG. 1. Miconia galeiformis. A. Flowering branch (Nee 55321, NY). B. Detail of setose hairs on stem (Nee 55340, NY). Miconia neei. C. Flowering branch. D. Detail of pistillate inflorescence. Anthers are exerted, but thecae are collapsed and pore is not open (C, D from Nee 55339, NY).

Additional specimens examined: BOLIVIA. Cochabamba: Prov. Chapare, along road to Tablas Monte, 1.9 km (by winding road) W from the junction with Villa Tunari-Cochabamba highway. 17˚09´56˝S, 65˚53´48˝W, 2695 m, 3 May 2007 (fl), Nee 55321 et al. (LPB, MO, NY, USZ).

Miconia galeiformis is a shrub or treelet growing in humid montane forest of Bolivia. The living plants have a distinctive green to red tinge on the hypanthium, and large, yellow stigmas. The flowers are

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FIG. 2. Miconia galeiformis. A. Habit, with detail of uncinate hairs. B. Node of stem. C. Inflorescence with pendant flowers and detail of sessile, dendritic hairs. D. Flower bud. E. Flower at anthesis. F. Longitudinal section of flower. G. Flower after anthesis with accrescent style. H. Petal. I. Two different anther morphs, from left to right: ventral, dorsal, lateral view. (All drawn from the holotype.)

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pendant at anthesis. The newly described species is most similar to M. mandonii Cogn. and M. flavescens Cogn., other Miconia species of Bolivia found in a similar habitat. The aforementioned species exhibit the same inflorescence structure and cernous flowers (Table I), though M. galeiformis can be distinguished based on the setose indument and stigma size. In Miconia galeiformis, the anthers are stout and open by two cleft-like pores at the apex, with a protruding septum. Triana (1871) placed species with similar anther morphology in Miconia sect. Chaenopleura. Cogniaux and later workers followed this precedent, and Miconia sect. Chaenopleura, as currently circumscribed, includes species with cuneiform or obovate anthers dehiscing by slits or slit-like apical pores (Judd, 2007); hence, we assign this species to section Chaenopleura. Phylogenetic data place M. galeiformis in a subclade consisting of M. flavescens, M. galeiformis, Cogn., and M. griffisii J. F. MacBr., which are part of a larger clade of high-Andean Miconia sect. Chaenopleura (unpublished data). Of the two specimens collected, we are fortunate to have pickled material available of both. One plant has perfect flowers (Nee 55340),

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with completely-formed ovules and pollen. The other collection (Nee 53321) has what appear to be fully-formed ovules, but the anthers are never extended. Upon dissection, no pollen was found within the thecae of the latter. Pollen was easily discovered in the former collection at the same stage of anthesis. As we only have two collections for investigation, we hesitate to make any statements regarding breeding system in this species. Dioecy is not common in Miconia, though, when present, it is usually expressed in the Andean species of Miconia sect. Cremanium (Almeda & Dorr, 2006; Wurdack, 1980). In addition, apomixis also occurs in Miconia (Goldenberg & Shepherd, 1998; Goldenberg & Varassin, 2001; Mendes-Rodrigues & Oliveira, 2012). The inconsistent occurrence of unisexual flowers suggests an intermediate evolutionary stage from synoecy to dioecy, or may be evidence of agamospermy, or both. Further investigation, along with sampling from numerous populations, will increase our understanding of the distribution of these phenomena in the genus. Miconia neei Jan. M. Burke & Michelangeli, sp. nov. Type: Bolivia. Cochabamba: Prov. Carrasco, gorge of Río Ivirizu, 6.6 km by road NE from Monte Punco, on road from

TABLE I COMPARISON OF MICONIA GALEIFORMIS TO SIMILAR SPECIES IN BOLIVIA

Miconia galeiformis

Miconia mandonii

Miconia flavescens

Leaf shape Petiole pubescence

Elliptical Densely setose, translucent

Branch morphology Branchlet indument

Straight

Lanceolate to elliptical Furfuraceous, hairs red or brown + line of setulose hairs on adaxial surface Flexuous

Setose, long, uncinate hairs + few sessile, dendritic hairs

Moderately furfuraceous, red, sessile-dendritic hairs

Absent

Present

Densely furfuraceous, red, sessile-dendritic hairs Absent

Chartaceous Symmetrical, attenuate to round Translucent, long setose hairs on veins 5 Pendant

Chartaceous to subcoriaceous Slightly asymmetrical, attenuate to round, basal nerved (rarely plinerved) Red or brown, sessile-dendritic hairs on midvein 4 (5) Erect to pendant

Subcoriaceous Slightly asymmetrical, round, short-plinerved Red sessile-dendritic hairs on veins 4 Pendant

Galeate

Galeate

Peltate

Petiole appendages Leaf texture Leaf base Abaxial leaf pubescence Merosity Flower posture Stigma shape

Elliptical to ovate Furfuraceous, hairs red Flexuous

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Monte Punco to Sehuencas, 17˚32´59˝S, 65˚16´13˝W, 2662 m, humid forest in steep canyon, 4 May 2007 (♀ fl), M. Nee, F. Michelangeli, O. Colque I., R. Flores S., D. McClelland & S. Stern 55339 (holotype: NY; isotypes: LPB, MO, USZ). (Figs. 1, 3) Diagnosis: Predominately glabrous species with ovate to elliptical leaves with serrulate margins, hollow stems and unisexual (imperfect) flowers.

Shrub or treelet, up to 4 m. Bark grey. Young stems quadrangular, then terete; internodes hollow (solid), longitudinal ridges blunt on young stems, occasionally 2 sulci present on mature stems; nodal line absent (present). Indument of young stems (especially nodes) furfuraceous with sessile, tan, dendritic hairs ca. 0.1 mm wide. Leaves isophyllous; petiole 0.6–3.5(–4.6) cm long, indument mostly glabrous to furfuraceous with dendritic hairs as stem; lamina 4.1– 19.5×1.8–6.3(–9.8) cm, ovate to elliptical (obovate), chartaceous, base round or slightly attenuate, the apex acuminate 4–9 mm, margin serrulate, the teeth 0.3–1.3 mm long, linear, deciduous; veins slightly impressed on adaxial surface and raised on abaxial surface, green to pink, secondary veins 2 pairs, including faint marginals, these more apparent on abaxial surface, basally nerved, tertiary veins percurrent (becoming reticulate distally), arched at 60–90˚ angle from midvein, quaternary veins obscure to articulate; adaxial surface green, indument on lamina absent to sparsely furfuraceous, with sessile, dendritic hairs (sessile translucent glands), indument on primary and secondary veins same as lamina; abaxial surface light green, lamina glabrous to sparsely glandular with minute, sessile, pigmented glands, indument on veins with sparse, sessile, dendritic hairs. Inflorescence a terminal panicle, 2.4–12.6 (–15.5) cm long, with auxiliary inflorescences; peduncles bluntly quadrangular, green to brown, indument furfuraceous or sessile, tan, dendritic hairs; bracts 0.5–1.2 ×0.2–0.3 mm, linear, caducous, beige; bracteoles ca. 0.5 × ca. 0.1 mm, linear, caducous, beige. Pedicels 0.2–0.4 mm long, densely furfuraceous. Flowers unisexual, 5-merous, diplostemonous. Hypanthium 1.3–2.4 mm long, campanulate, 2.0–3.2 mm wide at the torus,

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external surface with sessile, pigmented glands and sessile, dendritic hairs, yellow proximally, grading to red distally; internal surface ridged, androecial fringe absent. Calyx open in bud; tube 0.3–0.6 mm long at anthesis; lobes ca. 0.25×ca. 0.25 mm, round, glabrous, white, slightly longer than calyx teeth; teeth vestigial to deltoid, 0.2–0.5×ca. 0.1 mm, indument as hypanthium, green. Petals 0.8–1.6 × 0.8–1.2 mm, obovate to broadly obovate, erect, white at anthesis (drying white to yellow), base attenuate (cuneate), apex retuse, margin sinuous, both surfaces granulose. Functionally staminate flowers with stamens isomorphic, spreading radially; filament 1.8–2.3 long, glabrous, white; anthers with 2 locules, thecae 1–1.5× 0.6–0.7 mm, obovoid, opening by 1 ventrally-orientated to truncate 0.5–0.7 mm pore, septum not prolonged, white at anthesis, drying white to yellow; connective prolonged 0.3–0.5 mm into a dorso-basal spur, white to drying yellow, glabrous. Ovary 3-locular, free portion projecting 0.5×1 mm, conical, glabrous, strongly ridged, corona absent, apex with roughened hairs; style vestigial or up to 1 mm long, recurved or erect, white, glabrous; stigma capitate, 0.3–0.5 mm. Ovules present, not fully-formed. Functionally pistillate flowers with stamens isomorphic, spreading radially to incurved at anthesis; filament 1–1.6 mm long, glabrous, white; anther locules collapsed, thecae 0.8–1.2 × 0.4– 0.5 mm, subcuneiform, pore 0.2 mm, ventrally-orientated, not opening, septum not prolonged, white at anthesis, drying white to yellow; connective prolonged 0.3 mm into a dorso-basal spur, white to drying yellow, glabrous. Ovary as in staminate flowers; style 2.5–4.8 mm long, erect, white, puberulent; stigma broadly capitate, 0.8–1.1 mm wide. Ovules present. Berries 2.1–3.9×2.9–4.1 mm, globose, blue, indument partly persistent when mature, often subtended by 1 bracteole. Distribution.—Humid montane forest, 2300–3200 m, in the departments of Cochabamba, La Paz and Santa Cruz , Bolivia. Phenology.—Flowering May to August; fruiting July to February. Etymology. —This species name is dedicated to Dr. Michael Nee, who collected the majority of the specimens from which this species was described. We must also credit him with grouping these specimens at NY, and recognizing them as a cohesive, yet

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FIG. 3. Miconia neei. A. Flowering branch with detail of serrulate leaf margin. B. Node of stem. C. Dichasium with detail of dendritic hairs. D. Pistillate flower at anthesis with detail of calyx. E. Longitudinal section of pistillate flower. F. Petal. G. Stamens of pistillate flower in (from left to right) lateral, ventral and dorsal view). H. Staminate flower at anthesis and longitudinal section of staminate flower. I. Stamens of staminate flower in (from left to right) lateral, ventral and dorsal view). (A–G drawn from the holotype; H–I from Nee 52448, NY; J–L from Vargas 4150, NY.)

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undescribed, species. Michael Nee has significantly contributed to our knowledge of the Bolivian flora through several years of collecting effort and the ongoing publication of the Flora of the Amboró National Park. Additional specimens examined: BOLIVIA: Cochabamba: Prov. Carrasco, Old hwy (via Antigua) between Santa Cruz and Cochabamba, near border of Santa Cruz, 17˚50´3.6˝S 64˚41´36.2˝W, 2500 m, 1 Aug 2002 (♂ fl), Clark 6751 & Rodríguez (LPB, US); Serranía Siberia, 20–35 km W of Comarapa (Prov. Santa Cruz), on the old Cochabamba-Santa Cruz road (Hwy 4), 17˚54´S 64˚29´W, 2000 m, 14 Jan 1990 (fr), Dorr 7012 & Barnett (LPB, NY, US), Dorr 7028 & Barnett (LPB, MO, NY, US); sector Siberia, a 200 km de Cochabamba en la carretera a Santa Cruz, 2550 m, 27 Dec 1982 (fr), Fernández-Casas 7831 (NY); 57 km from Comarapa, on road to Cochabamba, 2560 m, 6 Feb 1978 (fr), King & Bishop 7665 (NY); along hwy from Cochabamba to Comarapa, ca. 8 km by road SE of turnoff to Pojo, 17˚45´30˝S 64˚49´W, 2560 m, 10 Jul 1998 (♂ fl ), Nee 50010 & Atha (LPB, MO, NY); 28 km al NE de Comarapa por el camino entre Santa Cruz y Cochabamba (20 km en línea recta al NE de Comarapa), 17˚49´S 64˚41´W, 2450 m, 10 Feb 1987 (fr), Solomon 15969 & Nee (LPB, M, MO, US). Prov. Chapare, Incachaca, 2600 m, 9 May 1971 (♂ fl), Cardenas s.n. (US); along road to Tablas Monte, 0.4 km (by winding road) W of turnoff from Cochabamba-Villa Tunari hwy, 17˚10´ 17˝S 65˚53´33˝W, 2845 m, 3 May 2007 (♀ fl), Nee 55306 et al. (LPB, MO, NY, USZ). La Paz: Prov. Murillo, 32 km

después de la cumbre, bajando por el Valle de Zongo, 2000 m, 20 Aug 1979 (fr), Beck 2209 (LPB, M, MO, US); Valle de Zongo, arriba de Jarca, 2100 m, 31 May 1980 (♀ fl), Beck 3606 (LPB, US); Valle de Zongo, vic. of Escuela Cambaya, 28.3 km N of La Cumbre, 16˚07´S 68˚05´W, 2560 m, 2560 m, 10 May 1990 (♀ fl), Luteyn 13609 & Dorr (AAU, CAS, LPB, M, NY, US); Río Zongo valley, 27 km below dam at Lago Zongo, 16˚08´S 68˚06´W, 2600 m, 10 Oct 1982 (fr), Solomon 8477 (LPB, MO, US); Valle del Río Zongo, a lo largo del Río Jachcha Cruz, 32.2 km al N de la cumbre, 16˚07´S 68 04´W, 2200 m, 22 Oct 1987 (fr), Solomon 17224 et al. (LPB, MO, US); Valle del Río Zongo, 28.5 km al N de la cumbre (cerca de la Escuela Cambaya), 16˚08´S 68˚06´W, 2400 m, 29 Jul 1988 (fr), Solomon 18695 (LPB, M, MO, US); Valle de Zongo, vic. of Escuela Cambaya, 28.3 km N of La Cumbre, 16˚07´S 68˚05´W, 2700 m, 10 May 1990 (♀ fl), Solomon 19039 et al. (LPB, MO, US). Santa Cruz: Prov. Caballero, P.N. Amboró, Cerro Bravo, cerca Comarapa, 2600 m, 20 Jun 1995 (♂ fl), Jardim 2036 & Abbott (MO, US, USZ), 22 Jun 1995 (♂ fl), Jardim 2078 (MO, US, USZ); 6 km (by air) N of Comarapa, along road to Cerro Bravo and Tinqui Laguna, 17˚51´42˝S 64˚31´54˝W, 2325 m, 3 Aug 2003 (♂ fl), Nee 52448 et al. (LPB, NY); along Comarapa-Cochabamba hwy, 2.5 km (by road) NE of El Empalme, 12.5 km (by road) NW of Torrecillas, 17˚49´30˝S 64˚41´18˝W, 2520 m, 7 Aug 2003 (♂ fl), Nee 52558 & Mendoza (LPB, NY); P.N. Amboró, proximidades del Cerro Bravo a 10 km al N de Comarapa, alrededores de la Parcela permanente, 17˚49´41˝S 64˚33´ 01˝W, 2400 m, 12 Nov 1995 (fr), Vargas 4150 et al. (LPB, NY, US, USZ). Prov. Florida, Comarapa Rd, 28 km from

TABLE II COMPARISON OF MICONIA

Miconia neei Anther pore number Flower sex Fruit color Leaf shape Leaf abaxial pubescence Nodal line Furfuraceous indument on young stems Leaf margin

NEEI TO SIMILAR GLABRATE SPECIES IN

Miconia cremophylla

BOLIVIA

Miconia amabilis

Miconia theizans

1

2

1

4

Imperfect Blue Elliptical to ovate (obovate) Sessile pigmented glands and few sessile-dendritic hairs Usually absent, faint when present Present

Perfect Green-blue Elliptical to obovate Densely branched, white, stellate hairs

Perfect White Large and ovate Densely furfuraceous with sessile-dendritic hairs along all veins

Perfect Blue, purple Variable, elliptical to ovate Glabrous except for pigmented-sessile glands

Very pronounced

Absent

Absent

Absent

Present

Absent

Teeth on serrulate margins are short and few

Entire

Few dentations to entire

Finely serrulate

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Comarapa, 2650 m, 15 Aug 1991 (♀ fl), AcevedoRodríguez. 4619 et al. (LPB, US); 7 km (by air) NE of Mairana, along S limit of P.N. Amboró, entering from Mairana, 18˚04´S 63˚55´W, 2200 m, 2 Jun 1991 (♀ fl), Nee 40634 (LPB, NY, US); 5.5 km by road N of Campamento la Yunga of P.N. Amboró, 8.5 km (by air) NE of Mairana, 18˚04´S 63˚43´W, 2270 m, 6 Jun 1998 (♂ fl), Nee 49641 (MO, NY).

based on pore number. With sterile material, leaf margin type and presence or a nodal line are useful diagnostic characters. Miconia galactantha Naudin is also dioecious, but has subcoriaceous leaves and almost entire leaf margin.

In Miconia neei, the anthers open by one broad pore, which is ventrally inclined. The filaments are flattened with an inflection at the filament midpoint. Based on theses staminal features, we assign this species to Cogniaux (1891) sect. Cremanium, the second largest section in Miconia. The imperfect flowers of Miconia neei retain vestigial organs, similar to other dioecious Miconia (Almeda & Dorr, 2006). To date, only 37 (out of ca. 1050) Miconia species have been categorized as dioecious, including six from Bolivia. Preliminary phylogenetic data place M. neei in a clade with other dioecious species from Bolivia: M. boliviensis Cogn., M. cyanocarpa Naudin and M. polygama Cogn. (unpubl. data). With increased sampling and resolution, we hope to interpret the evolution of dioecy in Andean Miconia in a phylogenetic context. There are ample collections of Miconia neei from across Bolivia to be able to assess its morphological variability. The species is polymorphic, with a notable suite of characters divided between two geographic areas: the specimens from La Paz and Cochabamba (Prov. Chapare) have larger leaves (8–19.5 cm), can be more elliptical to obovate and lack the red tinge in the hypanthium at anthesis. The specimens from Cochabamba (Prov. Carrasco) and Santa Cruz have small to medium leaves (4.1–10.3 cm), usually ovate, with a yellow and red hypanthium. In addition, the La Paz plants are taller, up to 8 m, while the specimens from Cochabamba and Santa Cruz are up to 4 m. Based on vegetative features, this species is most similar to M. amabilis Cogn., M. cremophylla Naudin or M. theizans (Bonpl.) Cogn. (Table II). With flowering material, the species can be distinguished

Acknowledgments We thank Michael Nee for his support during field work in Bolivia and for bringing to our attention the second species described here. We are grateful to Carmen Ulloa and Stephan Beck for reviewing the manuscript. We also wish to thank Bobbi Angell for her illustrations. This research was supported by NSF through the PBI Miconieae project (DEB-0818399).

Literature Cited Almeda, F. & L. J. Dorr. 2006. Miconia amilcariana (Melastomataceae: Miconieae), a new species from the Venezuelan Andes with notes on the distribution and origin of dioecy in the Melastomataceae. Proceedings of the California Academy of Sciences 55: 89–124. Beck, S. G. 1998. Floristic inventory of Bolivia–An indispensable contribution to sustainable development. In W. Barthlott & M. Winiger [eds.], Biodiversity, 243–267. Springer-Verlag, Berlin. Cogniaux, C. A. 1890a. An enumeration of the plants collected by Dr. H. H. Rusby in South America, 1885–1886.-XII. Bulletin of the Torrey Botanical Club 17: 90–94. ———. 1890b. An enumeration of the plants collected by Dr. H. H. Rusby in South America, 1885–1886.XI. Bulletin of the Torrey Botanical Club 17: 53–60. ———. 1891. Melastomaceae. In A. de Candolle & C. de Candolle [eds.], Monographie Phanerogamarum, 1–1256. G. Masson, Paris. ———. 1893. On the collections of Mr. Miguel Bang in Bolivia. Memoirs of the Torrey Botanical Club 3: 28–32. ———. 1896. New Melastomaceae collected by Miguel Bang in Bolivia. Bulletin of the Torrey Botanical Club 23: 15–17. ———. 1907. An enumeration of the plants collected by Dr. H. H. Rusby in South America, 1885–1886.-IV. Bulletin of the New York Botanical Garden 4: 354–361. ———. 1913. Die von Dr. Th. Herzog auf seiner zweiten Reise durch Bolivien in den Jahren 1910 und 1911 gesammelten Pflanzen. Mededeelingen van's Rijks Herbarium 19: 76–81. Foster, R. C. 1958. A catalogue of the ferns and flowering plants of Bolivia. Contributions of the Gray Herbarium 184: 1–223. Frodin, D. G. 2001. Guide to the standard floras of the world. Cambridge University Press, Cambridge.

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