Miconia tetrandra (Sw.) D. Don ex G. Don

Miconia tetrandra (Sw.) D. Don ex G. Don View in CoL in J.C. Loudon, Hort. Brit. 174.1830. Melastoma tetrandrum Sw.,Prodr. 72. 1788. Cremanium tetrandrum (Sw.) Griseb.,Fl.Brit.W.I. 262.1860. Tetrazygia tetrandra (Sw.) DC.,Prodr. 3:172.1828. Acinodendron tetrandrum (Sw.) Kuntze, Revis.Gen.Pl. View in CoL 2:953. 1891. TYPE: JAMAICA:without definite locality (fl), O. Swartz s.n. (LECTOTYPE, initially designated by Howard & Kellogg, 1986, followed here by a second-step lectotypification): S, internet image! S-R-3485; ISOLECTOTYPES: BM, internet image! 001008090; C, internet image! 10014870; S, internet image! S-R-3484; SBT, internet image! 12607).

Melastoma farinulentum Sw. ex Wikstr.,Kongl.Vetensk.Akad.Handl.1827:65 View in CoL .1827. TYPE: GUADELOUPE:1827, ForsstrÖm s.n. (HOLOTYPE: S,internet image!16-2405;photo of holotype: A!) .

Miconia abeggii Urb.& Ekman, Ark. Bot. View in CoL 22A (17):35.1929. TYPE: HAITI: Massif du Nord, Port Margot , Morne Maleuvre, 800–1000 m 8 Dec 1924 (sterile), E. L . Ekman H2790 (LECTOTYPE, designated here: S!04-414) .

Shrub or tree to 20 m tall, with ± gray, vertically furrowed bark. Indumentum of multicellular, pale to dark ferrugineous, globular-stellate to stellate hairs. Young twigs 2–9 mm wide, rectangular, sometimes with a shallow longitudinal groove on the broader faces, becoming terete with age, with ± dense, globular-stellate to stellate hairs; internodes 0.6–7.5 cm long, nodal line present, clearly evident to ± obscure. Leaves opposite and decussate, isophyllous or nearly so, with petiole 7–73 mm long, with dense, globular-stellate to stellate hairs; blade 4.2–24 × 1.3–11 cm, 1.7–5 times longer than wide, ovate to elliptic (rarely oblong), coriaceous, apex acuminate to attenuate (or rarely acute), base acute to rounded or occasionally slightly cordate, margin plane to revolute, entire to occasionally obscurely toothed, the teeth (if present) to 0.2 mm, not associated with a multicellular, elongated, simple, eglandular hair; venation acrodromous, ± basal (but prominent pair of secondary veins diverging slightly above base), with prominent midvein and 2 pairs of secondary veins, 1 pair of secondary veins conspicuous and 1 pair inconspicuous, 2 conspicuous secondary veins positioned 1–17 mm from margin, and 2 inconspicuous secondary veins intramarginal to 3.5 mm from margin, tertiary veins percurrent, oriented subperpendicular to midvein, 1.5–11 mm apart, connected by quaternary veins, higher order veins reticulate, ± obscure; the midvein and major secondary veins moderately to slightly impressed, tertiary veins ± flat, remaining veins flat on adaxial surface; the midvein conspicuously raised, major secondary veins slightly to moderately raised, minor secondary veins ± flat, tertiary veins very slightly raised to flat, and higher-order veins flat on abaxial surface; adaxial surface dark green (brownish green to brown when dried), indumentum of dense to moderate, globular-stellate hairs when young, but quickly ± glabrous, but with a few such hairs persisting on major veins and sometimes also on lamina; abaxial surface with dense to moderate, pale to dark ferrugineous, globular-stellate to stellate hairs, and these 0.05–0.15 mm across, and not obscuring the epidermis. Hair tuft domatia absent. Inflorescences terminal, with vegetative shoots producing numerous nodes before becoming reproductive, paniculate cyme, 3–19 cm long, 3–12 cm across, with 4–9 major branch pairs, these square to rectangular in cross-section; proximal segment of lowermost inflorescence branches 5–60 mm long, with dense, globular-stellate to stellate hairs, the ultimate axes (pseudopedicels) very reduced to ± absent, and flowers congested (in 3-flowered glomerules) or ± slightly separated; peduncle 1–6.8 cm long, with similar indumentum; bracts 0.5–27 (–50) × 0.2–6 (–8) mm, not involucrate, sometimes intergrading with leaves, triangular or narrowly triangular to linear, with acute apex, usually deciduous along primary inflorescence axis, but otherwise ± persistent, with stellate to globular-stellate hairs; bracteoles 0.2–0.5 × 0.15–0.2 mm, triangular to linear, with acute apex, deciduous to persistent, with stellate to globular-stellate hairs. Flowers 4-merous, radial, with pedicel 0–1.2 mm long. Hypanthium 1.1–2 mm long, ± cylindrical, unlobed, often longitudinally ridged externally, not to only very slightly constricted above ovary; the free portion 0.55–0.85 mm long, 0.85–1.5 mm wide at the torus, the outer surface with dense stellate to globular-stellate hairs, the inner surface smooth, glabrous; androecial fringe absent. Calyx lobes 4, separate in bud (i.e., non-calyptrate), the tube 0.06–0.26 mm long, not tearing between lobes, the lobes 0.06–0.55 × 0.5–1.2 mm, shallowly triangular to nearly obsolete, the apex acute to obtuse, abaxially with hairs similar to those of the hypanthium, and adaxially with a few branched to globular hairs; calyx teeth present, 0.15–0.5 mm long, flattened-triangular, terminating in a short bump, with stellate to globular-stellate hairs. Petals 4, 1.2–2.2 × 0.8–1.5 mm, ovate to obovate or oblong, ± spreading, white, glabrous, the apex rounded, often with a shallow notch, the base narrowed, and ± clawed, the margin entire. Stamens 4, alternating with petals, geniculate near apex of filament; staminal filament 1.3–2.2 mm long, glabrous, white, anther 1.1–1.5 × 0.3–0.6 mm, ± obovate, straight, yellow, opening by 2 broad, gaping pores (together as wide as the anther or nearly so), with proximal portion of the anther sacs reduced, sterile, connective not prolonged below thecae, and not appendaged, glabrous. Ovary 4- locular, 0.8–1.4 mm long, 1.2–1.5 mm across, 2/3- to 4/5-inferior, apically ± conic, glabrous or with a few branched hairs, the apex with a cylindrical and apically lobed collar (encircling style), ovary apex without longitudinal ridges, and lacking crown, with axile placentation, the ovules numerous, borne on expanded placenta that extends into each locule; style 1.5–3.8 mm long, terete, not swollen, straight to curved, white, glabrous; stigma truncate to punctate, minutely papillose, ca. 0.1–0.15 mm wide. Berries 2.5–5.5 mm long (including inconspicuous calyx), 2.5–7 mm in diameter, subglobose or short-ovoid to ± globose, blue-black, with sparse to moderate globular-stellate hairs, the hypanthium not constricted above ovary. Seeds 0.9–1.7 mm long, angular obovoid, with flat, ± narrowly triangular to rectangular raphe, testa ± smooth; appendage absent. ( Fig. 1 View FIG .)

Distribution and ecology.— Miconia tetrandra is widespread in the Caribbean region, occurring on eastern Cuba, Hispaniola ( Haiti and the Dominican Republic), Jamaica, Puerto Rico, Guadeloupe, Dominica, and Grenada. On Cuba M. tetrandra is restricted to the mountains of the northern Oriente region, while on Hispaniola the species grows in the Cordillera Septentrional, Massif du Nord/Cordillera Central, and the Massif de la Hotte. In Jamaica it grows in the Blue Mountains and John Crow Mountains, while in Puerto Rico it is widespread, occurring in the Sierra de Luquillo, Sierra de Cayey, and the Cordillera Central. The species grows in cloud forests, moist, broadleaved, montane forests, and moist forests of Pinus occidentalis Sw. , from 100–1380 m.

Phenology.— Flowering specimens have been collected from January through June,and also in December.

Iconography.—An illustration of Miconia tetrandra was published in Little et al. (1974). An unpublished watercolor illustration of the species, under the name Acinodendron tetrandrum , is available online (Britton & Home, Popular Flora of Puerto Rico, Flora Borinqueña, t. 426; accessed online: www.plantillustrations.org).

Additional specimens examined. CUBA. Prov. Guantánamo: in Cuba Orientali, precise locality unknown,1856–1857, Wright 186 (MO); Monte Verde, Jan–Jul 1959, Wright 1227 (MO, NY). Prov. Holguín: Sierra de Nipe, Mpio. MayarÍ, Alto de la Torre, 732 m, 19 May 2004, Bécquer et al.82488 (FLAS); MayarÍ, S of Sierra de Cristal, 28 Dec 1955, Alain & LÓpez Figueiras 4719 (GH); Sierra de Nipe, at the foot of Loma Mensura, 450 m, 12 Jul 1919, Ekman9743 (NY,US); Sierra de Nipe, Loma Mensura, 725 m, 15 Oct 1919, Ekman 9910 (NY); MayarÍ,S slopes of El Cristal, 2–7 Apr 1956, Alain et al.5516 (GH). DOMINICA. in sylvis ad La Sois, Apr 1882, Eggers 749 (MO, mounted and accessioned photocopy); coastal rd between Calibishie and Woodford Hill Estate, 5 Jun 1964, Ernst 1558 (GH, US); midway along rd between Pointe Ronde and Milton Estate, 350 m, 10 Apr 1940, Hodge & Hodge 2671 (GH); La Chaudiere, valley of Hampstead River, ca. 2 mi from mouth, 100 m, 10–14 May 1940, Hodge & Hodge 3522 (GH); along rd from Pont Casse to Castie Bruce, 15°22.91 ʹ N, 61°20.40 ʹ W, 590 m, 29 May 2000, Penneys 1300 (FLAS, NY,US). DOMINICAN REPUBLIC. Prov. Azua: Las Lagunas, 800 m, 17 Jun 1926, Ekman H6407 (GH,NY, US). Prov. Duarte: Cordillera Septentrional, Loma Quita Espuela,subiendo por Los Bracitos hasta el firme, 19°21 ʹ N, 70°9 ʹ W, 300–800 m, 6 May 1993, Bastardo et al.38 (MO);Cordillera Septentrional, Reserva CientÍfica Loma Quita Espuela,El Valle, en las Márgenes del RÍo El Valle, 19°22 ʹ N, 70°09 ʹ N, 550 m, 13 Aug 1992, García et al. 5144 (FLAS, JBSD);Cordillera Septentrional, Mpio. San Francisco de MacorÍs, Loma Quita Espuela, 19.35029°N, 70.14893°W, 750 m, 12 Feb 2016, Majure & Clase 6055 (FLAS, JBSD,NY); Cordillera Septentrional, San Francisco de MacorÍs, Reserva CientÍfica Loma Quita Espuela, en la entrada a la Loma La Canela, 19°24 ʹ 44 ʺ N, 70°09 ʹ 57 ʺ W, 600 m, 21 Nov 2002, Veloz & Monegro 2914 (FLAS, JBSD). Prov. Peravia-Prov. San Cristobal border: Mahoma, 12 km NE de Rancho Arriba (en la carretera a Piedra Blanca) y 4km S hasta Mahoma y RÍo Mahoma, 18°43.5 ʹ N, 70°22 ʹ W, 800 m, 12 Aug 1982, Zanoni et al.22506 (JBSD,MO,NY, US). Prov. de San Juan: Cordillera Central, Parque Nacional José del Carmen RamÍrez, 14 km al N del poblado Arroyo Cano en el lugar llamado “Los FrÍos”, 400 m al E de la caseta de la Dirección Nacional de Parques, 18°53 ʹ N, 71°03 ʹ W, 1380 m, 1 May 1986, García & Pimentel 1295 (FLAS, JBSD); Cordillera Central, near Los FrÍos along jeep rd from Arroyo Cano, just below “Forestry House” in Parque Nacional José del Carmen RamÍrez, 1250 m, 2 Jun 1992, Judd & Skean 6701 (FLAS, JBSD); Cordillera Central, Jose del Carmen Ramirez National Park,above “Los Frios,” on jeep rd from Arroyo Cano (N of Guanito),trail from forest house to Arroyo Palo de Quello, 1250 m, 2 Jun 1992, Skean & Judd 3275 (JBSD, MSC). Prov. San José de Ocoa: Cordillera Central, Municipio Rancho Arriba, Seccion Mahoma, Comúnidad Florencio, orillas del RÍo Florencio; 18.70046°N, 70.35976°W, 916 m, 17 May 2019, Majure 7888 (FLAS, JBSD). Prov. Santiago: Cordillera Central,Mpio.San José de las Matas, sección, Mata Grande, Parque Nacional Armando Bermúdez, en la Loma de la Barranca, 1370 m, 20 Mar 1999, Clase & Peguero 871 (FLAS,JBSD);Cordillera Central,Mpio.San José de Las Matas, en la carretera que va desde Diferencia a Los Ramones, 5 km antes de llegar a Los Ramones, Parque A. Bermúdes, 475 m, 18 Jun 1999, Clase & Peguero 1337 (FLAS, JBSD);Cordillera Central, Mpio. San José de Las Matas, Los Ramones, trillo La Hispaniola, 19°34 ʹ N, 70°05 ʹ W, 200–280 m, 10 Oct 2002, Clase et al. 3438 (FLAS,JBSD);Cordillera Central, 16–17 km al SO de Pedregal de San José de las Matas, en Mata Grande en las márgenes del RÍo Sape Malo al NO de la caseta de la Dirección Nacional de Parques, 19°14 ʹ N, 70°58 ʹ W, 28 Jun 1988, Pimentel & Cabral 1086 (FLAS, JBSD);Cordillera Central, en el poblado rural “Palacio”, aprox. 2–3 km de La Manacla en el camino a Los Ramones, 19°15 ʹ N, 71°09 ʹ W, 740–760 m, 24 May 1988, Zanoni et al.41155 (JBSD,MO, NY, US). Prov. Santiago Rodriguez: Cordillera Central,Monción, Lagunas de Cenobi, ca. 1100 m, 15 Jun 1929, Ekman H12868 (US);Cordillera Central,along rd between La Leonor and El Aguacate, 3 km W of La Leonor, 600 m, 30 May 1992, Judd et al.6671 (FLAS, JBSD). GRENADA: Grand Etang, 31 Mar 1897, Broadway s.n. (MO, specimen at F without indication of collector); Grand Etang, Mar 1937, Broadway 7897 (NY); ad Grand Etang, Dec 1889, Eggers 6403 (US); near Grand Etang, 4–10 Mar 1979, Howard & Howard 18822 (A, NY);vicinity of Grand Etang, 1600–1800 ft, 30 Oct–11 Dec 1957, Proctor 16991 (A). GUADELOUPE: Gourbeyre, Morne Gabelin, 300–600 m, 1894, Duss 3469 (F, MO, MO [as “3649” perhaps transposed numerals], NY, US);St. Rose, 28 Apr 1938, Questel 848 (US);Goyave, 20 m, 4 May 1938, Questel 2353 (US). HAITI. Dept. du Nord-Est: Massif du Nord,Mont-Organisé, 650 m, 1 Jun 1926, Ekman H6243 (US); Vallière, top of Morne Salnave, 1125 m, 1 May 1928, Ekman H9934 (US). Dept. du Sud: Massif de la Hotte, western group, Tiburon, Morne Sentier, 750 m, 26 Aug 1928, Ekman H10585 (NY). JAMAICA. Portland Parish: near Portland-St.Andrew border, W slope of Silver Hill, 4300–4500 ft, 25 Jul 1966, Anderson & Sternberg3258 (DUKE,GH,MICH,US);Vinegar Hill, 4300 ft, 25 Jun 1896, Harris 6408 (F,NY);Blue Mountains,Hardwar Gap,along “Shelter trail,” ca. 1200 m, 21 May 1987, Judd 5418 (DUKE, F, FLAS,GH, IJ, MO,NY); near St. Andrew Parish border,Blue Mountains,Hardwar Gap, along “Shelter Trail,” ca. 1200 m, 24 May 1987, Judd5470 (FLAS, IJ, MO);Blue Mts., Trafalgar, 9 May 1916, Perkins 1186 (A, GH); E slope of the John Crow Mts., 1.5–2 mi SW of Ecclesdown, 1500–2000 ft, 14 Jan 1955, Proctor 9811 (A, NY, US); NW slope of Joe Hill, 1000–2250 ft, 20 Apr 1955, Proctor 10092 (A, NY, US); along trail to waterfall N of Hardwar Gap, 3800–4000 ft, 8 May 1961, Proctor 22270 (GH, MICH); Vinegar Hill, 1300 m, 9 Mar 1906, Shreve s.n. (NY). St. Andrew Parish: Blue Mountains, 1.3 to 2 mi N (above) Newcastle on rd to Hollywell and Hardwar Gap,from 18°4.688 ʹ N, 76°43.158 ʹ W (at 1.3 mi) to 18° 4.955 ʹ N, 76°43.531 ʹ W (at 2 mi),and also at Hardwar Gap,along Oatley Trail, at 18°5.268 ʹ N, 76°43.651 ʹ W, 1200–1300 m, 11 Jan 2011, Judd et al.8301 (FLAS, IJ, NY). PUERTO RICO: Naguabo, Bo. RÍo Blanco, Caribbean National Forest, along 1.5 km stretch up RÍo Sabana S of closed portion of Rt 191,480– 600 m, 29Feb 1991, Axelrod & Chavez4119 (NY);Caribbean National Forest,along Rt 191,just S of jct with rd to summit of El Yunque, 18°17 ʹ N, 65°47 ʹ W, 600 m, 4 Jan 1987, Boom 6880 (MO); Caribbean National Forest, along Hwy 988 between Hwy 191 and Hwy 983, 100–200 m, 12 Jan 1987, Boom & Marshall 7084 (MO, NY); Caribbean National Forest, along hwy 191 just N of landslide, 18°17 ʹ N, 65°47 ʹ W, 600 m, 16 Mar 1990, Boom 9793 (NY); Caribbean National Forest, Hwy 191 ca. 1 mi N of landslide, 18°17 ʹ N, 65°47 ʹ W, 600 m, 4 Jan 1992, Boom 10270 (NY); Mt. Alegrillo, 900 m, 3 Apr 1913, Britton et al.2594 (F, NY, US); Catalina-Yunque trail, 500 m, 23–26 Feb 1923, Britton & Bruner 7659 (NY); without definite locality, 26 Nov–4 Dec 1963, Duke 7062 (MO);along RÍo Cuby, off Rt 191 through Luquillo Mts., 21 Mar 1966, Evans 70 (A); El Yunque, 600 m, 1926, Gleason & Cook M-118 (NY); N side, Luquillo Mts., 13 Apr 1899, Heller & Heller 1068 (F,NY); El Yunque, Jul 1933, Horn 9 (NY); Luquillo National Forest, along Rt 186 where it crosses RÍo Espirita Santo,S of El Verde, 410 m, 14 Jun 1991, Judd & Judd 6028 (FLAS); Maricao State Forest, 800 m, 30 Mar 1964, Liogier 10754 (NY); Maricao State Forest, 800 m, 31 Mar 1964, Liogier 10781 (NY,US); Luquillo Mts., near Pico del Este, 1050 m, 3 Aug 1982, Liogier et al.33323 (MO, NY); Luquillo Mts., El Yunque, 900 m, 16 Mar 1983, Liogier et al.34045 (MO,NY); Luquillo Mts.,Hwy 191,km 21.4, 2000–2500 ft, 10 Jul 1966, Little21616 (NY); Luquillo Mts., Hwy 191, km 17.3, 2000–2500 ft, 10 Jul 1966, Little21619 (GH, NY, US);Caribbean National Forest, El Verde Research Station, LTER Plot, 18°20 ʹ N, 65°45 ʹ W, 350 m, 26 Jan 1990, Moestl & Atwell 52 (NY); Sierra de Luquillo, Caribbean National Forest, Rd 191, km 19, 650 m, 20 Mar 1987, Proctor 43197 (US); Sierra de Naguabo, Loma Icaco, 210–675 m, 24 Jul 1914, Shafer 3445 (NY); Sierra de Luquillo, in monte Jimenes, Jun 1885, Sintenis 1564 (US); Sierra de Naguabo, ad los Ranchones, 30 Oct 1986, Sintenis 5439 (NY); Lares, in sylva ad Buenos Aires, 29 Jan 1887, Sintenis 6104 (F, MO, MSC, NY, US); Sierra de Cayey, Carite Forest Reserve, Mpio.de Patillas, Barrio Muñoz Rizera,Rd 184 at km 18.6,18°05 ʹ 68 ʺ N, 66°02 ʹ 49 ʺ W, 596 m, 15 May 1996, Skean & Proctor 3743 (MICH);Mpio.de Patillas, property of Tropic Ventures, at km 16.1 of Rt 184, 500 m, 8 Mar 1987, Taylor & Scott 7418 (DUKE); Cayey, Bo. Farallón, Carite State Forest, Rd 179, ca. 2 km S of intersection with Rd 184, 18°06.824 ʹ N, 66°04.725 ʹ W, 750 m, 6 Jul 1904, Trejo et al.2702 (F, FLAS); at km 15.7 on Palmer to Florida rd through Luquillo Mts., 7 Apr 1964, Wagner 521 (A);along Rt 191 at km 16,7 on S side of Luquillo Mts., 21 Apr 1968, Wagner 1473 (A,DUKE);Guavate Forest, 2200 ft, 15 Feb 1962, Woodbury s.n. (NY);El Yunque, 21 Jan 1967, Woodbury s.n. (NY).

Although widespread in the Caribbean region, no geographically correlated patterns of morphological variation were discerned. The name Miconia abeggii , based on a single specimen (the type) from the Massif du Nord, Haiti, with leaves larger than is typical for the species, represents a rapidly growing vegetative shoot. This specimen falls within the pattern of variation of the species, when numerous specimens (including rapidly growing, vegetative shoots) are considered.The specimen is here designated as the lectotype of M.abeggii because it is very likely that a specimen once existed at B (now destroyed); additionally other specimens may exist.

Howard and Kellogg (1986), when designating the lectotype of Melastoma tetrandrum , did not specify (in their publication) which of the two duplicates at Stockholm (S) was chosen. Thus, a second-step lectotypification (see Turland et al. 2018: Art. 9.17) is required, and we designate S-R-3485 as the lectotype, in agreement with the annotation of E. A. Kellogg.