Revision of Monstera (Araceae: Monsteroideae) of Central America Author Croat, Thomas B. 0000-0002-1804-0532 P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. & email: thomas. croat @ mobot. org; https: // orcid. org / 0000 - 0002 - 1804 - 0532 thomas.croat@mobot.org Author Cedeño-Fonseca, Marco 0000-0003-0119-0427 P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. & Botanischer Garten und Botanisches Museum Berlin, Freie Universität Berlin, Königin-Luise-Strasse 6 - 8, D- 14195 Berlin, Germany. & Herbario Luis Fournier Origgi (USJ), Centro de Investigación en Biodiversidad y Ecología Tropical, Universidad de Costa Rica, Apdo. 11501 - 2060, San José, Costa Rica. & Jardín Botánico Lankester, Universidad de Costa Rica, Apartado 302 - 7050 Cartago, Costa Rica. & email: marcovf. 09 @ gmail. com; https: // orcid. org / 0000 - 0003 - 0119 - 0427 marcovf.09@gmail.com Author Ortiz, Orlando O. 0000-0002-7805-0046 P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. & Botanischer Garten und Botanisches Museum Berlin, Freie Universität Berlin, Königin-Luise-Strasse 6 - 8, D- 14195 Berlin, Germany. & P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. & Herbario PMA, Universidad de Panamá, Estafeta Universitaria, Panama City, Republic of Panamá. & P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. & Coiba Scientific Station (COIBA AIP), Clayton, Panama City, Panama. & P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. & email: ortizopma @ gmail. com; https: // orcid. org / 0000 - 0002 - 7805 - 0046 * Corresponding author. e-mail: marcovf. 09 @ gmail. com & P. A. Schulze Curator, Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U. S. A. ortizopma@gmail.com text Phytotaxa 2024 2024-06-27 656 1 1 197 http://dx.doi.org/10.11646/phytotaxa.656.1.1 journal article 303607 10.11646/phytotaxa.656.1.1 5e8d8c9a-f245-4504-9c32-1d82dffcbb90 1179-3163 13213072 3. Monstera adansonii Schott, Wiener Z. Kunst 4: 1028. 1830 . — Dracontium pertusum Linnaues, Sp. Pl. 2: 968. 1753 . ( Figs. 5–7 ) Type: —Figs. LVI & LVII, ‘ Arum hederaceum , amplis foliis perforatis’, in R.P.C. Plumier, Description des Plantes de l’Amerique. 1693. ( lectotype designated by Madison (1977)) . Monstera friedrichsthalii Schott, Oest. Bot. Wochen. 4: 65 (1854) . TYPE:—[ Nicaragua ], [Niquinohomo], Friedrichsthal [ 1206 ] ( holotype W, destroyed). Neotype (designated here) : Nicaragua . Departamento de Masaya : Laguna de Apoyo , 11°56’N 086°02’W , 100–140 m , 20 sept. 1981 , Pedro P. Moreno 11142 (MO!, two sheets: 3189310 and 3614609). Monstera seemanii Schott, Oest. Bot. Zeit. 9:40 (1859) . TYPE:—[ Panama ], Insula Taboga , Seeman 1568 ( holotype K, isotype BM!). Terrestrial or nomadic vine, appressed-climbing or climbing habit. SEEDLINGS: bearing foliage leaves. JUVENILE PLANTS: root climbers; stems light-green or dark, smooth or slightly rough, cylindrical or flattened; internodes 3–5 cm long, 0.5–10.0 mm diam., usually 5 times longer than wide; petiole distinct, dark-green, smooth or slightly rough, 10–15 cm long, sheathed to base of the geniculum; petiole sheath deciduous or persistent; blades ovate-lanceolate, attenuate, cordate or truncate at base, acuminate at apex, subcoriaceous, 10–15 × 5–10 cm , not appressed to the phorophyte; fenestrations present or absent. ADULT PLANTS: root climbers; stem light-brown, light-green or dark-green, smooth or with small white pustules, sometimes with white dots, cylindrical or slightly flattened; cataphylls light green, whitish or yellowish-green, deciduous but leaving dry fragments on the peduncle; internodes 1–6 cm long, 1.5–3.5 cm diam.; anchor roots dark brown or blackish; feeder roots dark brown; petiole greenish, sometimes with white dots, smooth or with small white pustules, scarcely verrucose at basebase, 25–70 cm long, sheathed to basebase of the geniculum; petiole sheath deciduous, sometimes with fibrous residues; geniculum smooth, flattened or sunken adaxially, convex abaxially, 2–4 cm long; blades ovate, lanceolate-ovate or oblong, broadly cuneate to rounded, truncate or rarely cordate at base, acuminate at apex, coriaceous, drying blackish, reddish, yellowish or olive-green, 23–65 × 17–30 cm , 1.3–1.8 times longer than wide, undulate-decurrent on the geniculum (7–11 undulations of 0.5–1.0 mm wide); midrib flattened adaxially, convex abaxially, drying black or yellowish green on both surfaces; primary lateral veins 15–35 per side, emitting at a very low angle one or two robust secondary veins from near base or to ca. 1/3rd of the way from base of the primary veins, especially in the proximal 2/3rds of the blade, sunken to slightly sunken adaxially, raised abaxially, departing midrib at 45–80° (rarely at 90°), drying black or yellowish; secondary veins slightly prominent, reticulate towards the margin or parallel to the lateral nerves, undulate when dry; collective veins slightly visible; fenestrations present or absent, rounded, ovate or ellipsoid when present, and distributed along the blade near the midrib, sometimes with filaments separating the fenestrations; margins entire or pinnatilobed (2–5 lobes per side), due to tearing of the fenestrations that extend to the margin. INFLORESCENCES on ascending and pendent stems, 4–8 simultaneously at the flowering season, arranged in the leaf axils or into cataphylls; peduncle smooth or slightly rough, 12–20 cm long; 1.4–1.6 cm diam.; spathe short or long acuminate, light green and yellowish green during development, white or yellowish green externally, white internally at anthesis, 13–20 × 5–14 cm , up to 5 cm longer than the spadix, slightly coriaceous or membranous, sometimes with revolute margins, completely open during anthesis or remaining convolute in the lower part, later deciduous or marcescent post-anthesis; spadix white or yellowish during development, cream-yellowish at anthesis, 10–15 cm long, 1.5–2.0 cm diam., (4.6)7.0–9.2 times longer than wide; basal sterile flowers with a yellow stigmatic secretion, 3–5 mm long; fertile flowers 4–6 mm long; stamens 1–6 mm long, with laminar filaments; anthers 1–2 mm long; ovary quadrangular or rectangular in longitudinal section, ribbed, 3–4 × 2–3 mm ; style compressed, pentagonal or hexagonal, 0.5–2.0 × 2–3 mm ; stigmatophore columnar, 0.5–1.0 mm long, absent (i.e. becoming unobservable) in dry specimens; stigma linear or linear-curved, with a transparent stigmatic secretion; berries creamy-white, yellowish green during development, pulp whitish; seeds dark green, 3–6 mm long. FIGURE 5. Monstera adansonii . Adult plant growing in Cerro Azul, Panama. Photo by M. Cedeño-Fonseca. (Not collected). FIGURE 6. Monstera adansonii . Adult plant growing in Buenos Aires, Costa Rica. M. Cedeño et al. 804 (USJ). Photo by M. CedeñoFonseca. Distribution and ecology: — Monstera adansonii subsp. laniata ranges from Nicaragua to Venezuela , Guyana , French Guiana, Suriname , Brazil , Trinidad & Tobago , the Lesser Antilles. In Central America it is very common in Tropical moist forest , whereas in Colombia it occurs most often in Tropical wet forest from sea level to 975 m . Phenology: —In Central America the species flowers and fruits throughout the rainy season (April to August), less frequently as early as February and as late as October. Fruiting occurs mostly September to December, less frequently as early as July. Discussion: —The species is a member of sect. Monstera and is characterized by its ovate-elliptic, usually perforate blades, usually less than twice as long as broad, unequal at base with one side truncate to subcordate and the other side cuneate to acute, by primary lateral veins at least on one side arising at an angle of more than 60° and with a peduncle that equals or exceeds the spadix. It is most easily confused with M. siltepecana but that species differs by typically drying darker brown, by having more perforations and by having the secondary lateral veins conspicuously reticulate. FIGURE 7. Monstera adansonii from San Jose, Costa Rica. (A) Developing inflorescence. (B) Open inflorescence, frontal and back views. (C) Mature infructescence, stylar plates detached. (D) Fertile flower; in lateral view (left), and longitudinal section (right). (E) Stylar plate with stigma (left) and one stamen (right). (F) Sterile flower; in lateral view (left), and longitudinal section (right). (G) Portion of juvenile plant. (H) Portion of adult plant. M. Cedeño et al. 1008 (USJ). Image from Cedeño-Fonseca et al. (2022) . Madison (1977) recognized M. adansonii as having three varieties with the typical variety being restricted to the West Indies, var. klotzschiana (Schott) Madison as being restricted to South America, throughout much of the Amazon basin from southern Venezuela and the Guianas south to Paraná State in Brazil to Amazonian Peru and Bolivia , mostly preferring secondary forest, and the var. laniata (Schott) distributed from Nicaragua to Peru , Venezuela , the Guianas, and Brazil . Subsequent to Madison’s, studies by Mayo and Andrade (2013) have recognized the three varieties recognized by Madison as distinct subspecies, M. adansoni i subsp. laniata , M. adansonii subsp. klotzschiana and M. adansonii subsp. blanchetii . For this review M. adansonii subsp. laniata is the only subspecies present in Central America. In addition, Monstera seemanii described from Panama is included under synonyms of this subspecies. Populations of M. adansonii considered as the subsp. laniata in central Panama and towards the Darien region , have a more closely related morphological similarity to M. adansonii subsp. blanchetii . However, this complex must be analyzed with more morphological details about the subspecies considered by Mayo & Andrade (2013) . For more details about the taxonomic comments and nomenclatural notes see Cedeño-Fonseca et al. (2022) . Nomenclatural remarks: — Schott (1854) , in the original protologue of M. friedrichsthalii , only included the description and some taxonomic notes, however he did not include information on the locality or collector name. Subsequently, Schott (1860) in his Prodromus systematis Aroidearum , added that the plant comes from ” Guatemala ” and recognizes Friedrichsthal as the original collector. This information is again reiterated in the revision by Engler & Krause (1908) , where they cited the holotype specimen as follows: “ Guatemala , Niquinomo, Friedrichsthal n. 1206 - Kais. Herb. Wien.” Almost all the Central American herbarium material collected by Emanuel von Friedrichsthal contains labels designating that the specimens were collected in “ Guatemala ”; nevertheless, historical evidence ( Taracena-Arriola & Sellen 2006 ) and location notes included on the labels suggest that Friedrichsthal actually collected in several countries of Central America, not only in Guatemala . Furthermore, the locality cited by Engler & Krause (1908) indicates that the plant actually comes from Niquinohomo (as “Niquinomo”) in Nicaragua . It is well known that almost all of the original Araceae material housed in W (which was studied by Schott himself) was destroyed during World War II ( Riedl & Riedl-Dorn 1988 ), including the holotype of M. friedrichsthalii . Unfortunately, until now, it has been impossible to locate any original material of M. friedrichsthalii in the herbaria consulted. In the absence of original material, we propose the specimen Pedro P. Moreno 11142 (MO, two sheets: 3189310 and 3614609) as the neotype of M. friedrichsthalii . Additional specimens examined: — HONDURAS . Atlántida : Carlos Soto V. 749 ( TEFH !) ; Cirilo H. Nelson et al. 17472 ( TEFH !) ; Tela. m, C.H. Nelson & R. Andino 15438 ( TEFH !) ; Colón : 1.8 mi strip on the north bank of rio Guaimoreto between old bridge and opening of Laguna Guaimoreto 4.5 mi. NE of Trujillo on old road to Castilla, 15°57’30”N 085°54’30”W , 1 Aug 1980 , J.G. Saunders 519 ( MO !) ; Trujillo . Old Airport road to Castillo , 3 km east of Trujillo, 15°55’N 086°00’W , 50 m , 09 June 1980 , J.G. Saunders 320 ( MO !) ; Capuchin Site West , [ 1.8 mi. strip on the north bank of río Guaimoreto, between old bridge and opening of Laguna Guaimoreto, 4.5 mi. NE of Trujillo on old road to [Puerto] Castilla], 15°57’30”N 085°54’30”W , 0–20 m , 5 Aug 1980 , J.G. Saunders 524 ( MO !) ; La Paz : Guajiquiro . R.J. Evans 1705 Olancho : Along Río Olancho, W of main Tegucigalpa-Catacamas Highway, ca. 1 km up stream from and NW of Puente Boquerón, 8.6 miles SW of Catacamas, 6 miles SW of Santa María del Real, 14°45’N 086°00’W , 400 m , 04 February 1987 , T.B. Croat & D.P. Hannon 64107 ( AAU , B , MO , SEL , TEX ) . COSTA RICA . Alajuela : Atenas , Río Grande , Balsa de Atenas , ca 3 km southeast of Atenas on the future site of the Escuela Centroamericana de Ganadería, 400 m , 7 July 1975 , (Fr.), J. Utley & B. Utley 2618 ( CR !) ; San Carlos , La Tigra, 15 km WNW of Quebrada by air, 1 km W of Jabillos, Disturbed primary forest, 175 m , 29 April 1983 , (Fl.), R. Liesner & E. Judziewicz 15172 ( CR !, MO !); San Carlos , Cutris, 3 km south of Boca de Arenal in remnant forest patch along río San Carlos on Hacienda Boca Arenal, 100 m , 3 June 1986 , (Fr.), B.E. Hammel 15323 ( CR !, MO !); San Ramón , San Isidro, 1–3 km E of San Ramón coffee fincas, pasture, and stream, Near Waterfall, 1000 m , 13 April 1983 , (Fl.), R. Liesner 14251 ( CR !, MO !); San Ramón , Santiago, Finca Barranca, Bosque muy húmedo premontano, 900 m , 6 January 1984 , (Fl.), L. Gómez et al. 20762 ( CR !, MO !); San Carlos , La Tigra, 15 km WNW of Quebrada by air, 1 km W of Jabillos, Disturbed primary forest, 175 m , 29April 1983 , (Fl., Fr.), R. Liesner et al. 15139 ( CR !, MO !); Upala , Bijagua, Evergreen tropical wet forest formations and pastures on eastern slopes of Volcán Miravalles, West of Bijagua, near the río Zapote, 600 m , 11 February 1982 , (Fr.), W. Burger et al. 11663 ( CR !, MO !); San Carlos , Florencia, Directly across the river W of Muelle de San Carlos, 22 km NNE of Quebrada by air, 100 m , 8 April 1983 , (Fr.), R. Liesner 14085 ( CR !, MO !); Upala , Aguas Claras, P.N. Rincón de la Vieja, Sector de Ríos Aguas Verdes, 2 km S aguas arriba del puente, falda NE del Volcán Santa María, 600 m , 12 February 1991 , (Fl.), G. Rivera & C. Dennis 1062 ( CR !) ; San Carlos , Fortuna, San Carlos, Fortuna, R. B. Arenal Mundo Aventura, 255 m , 5 March 2004 , (Fr.), A. Rodríguez 8500 ( CR !) ; San Carlos , Fortuna, San Carlos, Fortuna, R.B. Arenal Mundo Aventura, 255 m , 18 March 2004 , (Fr.), A. Rodríguez 8557 ( CR !) ; San Carlos , Pital, San Carlos, Boca Tapada , Bosque Ancianos, 50 m , 23 May 2004 , (Fl.), D. Solano 1060 ( CR !) ; San Carlos, Pital, Cerros Chaparrón , 100 m , 8 July 2005 , (Fr.), D. Solano 2596 ( CR !) ; Along road between Cañas (Guanacaste) and Upala , 100 m , 25 June 1976 , (Infer.), T.B Croat 36382 ( MO !) ; Along road between Cañas (Guanacaste) and Upala , 100 m , 25 June 1976 , (Fl., Fr.), T.B. Croat 36382 ( MO !) ; Atenas , Banks of Río Cacao , 800 m , 2 January 1983 , (Fl., Fr.), L. Gómez 19565 ( MO !) ; San Carlos , La Fortuna, El Tanque, 9 July 1982 , (Fr.), E. Valerio 61 ( USJ !) ; San Carlos , La Fortuna, El Tanque, 10 July 1982 , ( Fr. ), E. Valerio 62 ( USJ !) , San Carlos , La Fortuna, El Tanque, 10 July 1982 , (Fr.), E. Valerio 63 ( USJ !) ; San Carlos , Florencia, San Luis, 280 m , 17 Marzo 2001 , (Fr.), C. Trejos 123 ( USJ !) ; Los Chiles ; Los Chiles ; Río Frio ; 40 m , 1 August 1949 , (Fr.), R. Holm & H. Hugh 780 ( USJ !) ; Lago Cote , Bosque Aledaño , 800 m , 23 June 1968 , (Fr.), L.A. Fournier et al. 1448 ( USJ !) ; San Carlos , La Palmera, Extremo este del Tajo de la comunidad, 200 m , 19 February 2000 , (Fr.), A. Pérez 33 ( USJ !) ; Atenas , Concepción , Río Grande , 480 m , 30 May 1981 , (Fr.), E. Valerio s.n ( USJ !) ; Alajuela , Orotina , Coyolar , 170 m , 1 agosto 1982 , (Infer.), R. Ocampo 3885 ( CR !) ; Cartago : Turrialba , Turrialba , CATIE, Confluencia del Rìo Tuis y el Rìo Reventazòn , 580 m , 15 June 1994 , (Fr.), G. Herrera 7180 ( CR !) ; La Unión , San Diego, Z.P. Cerros de La Carpintera , 1800 m , 28 September 2006 , (Fr.), A. Cascante 1586 ( CR !) ; Paraíso , Orori , Bosque primario en las faldas del Alto El Jaular, 1400 m , 20 July 1994 , (Fl., Fr.), K. Taylor 209 ( CR !) ; Turrialba , Ganadería , 10 July 1965 , (Infer.), T.B. Croat 257 ( MO !) ; Turrialba , Pavones , Cruce a La Suiza , En Plantación de Caña , 600 m , 12 December 1999 , (Fr.), M. Blanco 1009 ( USJ !) ; Turrialba , Turrialba , Jardín Botanico del CATIE , 616 m , 26 October 2015 , (Fr.), M. Cedeño 836 ( USJ !) ; Turrialba , Turrialba , Jardín Botanico del CATIE , 616 m , 8 December 1981 , (Fr.), E. Valerio 32 ( USJ !) ; Turrialba , Turrialba , CATIE , 675 m , 2 May 1951 , (Fr.), J. León 3388 ( USJ !) ; Turrialba , 20 July 1984 , (Fr.), C. Helfenberger s.n ( USJ !) ; Guanacaste : Liberia , Liberia , P.N. Rincón de la Vieja , Cordillera de Guanacaste , Sendero de la toma de agua, a 3 km de la estación, 1000 m , 17 September 1990 , (Fr.), G. Rivera 601 ( CR !) ; Nicoya , San Antonio , P.N. Barra Honda , Península de Nicoya , Los Mesones , cerca de los tanques de captación, 350 m , 21 August 1992 , (Fr.), M. Reyes et al. 8 ( CR !) ; Bagaces , Bagaces, 1 km W of Hacienda Palo Verde , 50 m , 10 July 1976 , (Infer.), J. Salomon 2437 ( CR !, MO !); Nicoya , San Antonio , R. B. Lomas de Barbudal , Valle del Tempisque , Estación Barra Honda , Sendero de la Flor , 300 m , 1 July 1994 , (Fl., Fr.), U. Chavarría 987 ( CR !) ; Abangares , Sierra, R.B. Monteverde, San Luis valley below comminity, premontane moist forest on Pacific slope, 1000 m , 9 May 1986 , (Fr.), W. Haber & E. Bello 5014 ( CR !, MO !); La Cruz, Santa Elena , Along rìo Cuajiniquil , ca. 1 km SE of Rabo de Mico , 30 m , 24 January 2003 , (Infer.), R. Espinoza et al. 11503 ( CR !, MO !); Santa Cruz , Santa Cruz, Cuenca del Tempisque, Bosque Nacional Diriá, margen del Río Enmedio de la casona a Río arriba, 300 m , 12 June 1998 , (Fr.), U. Chavarría & J. González 1868 ( CR !, MO !); Bagaces , Bagaces , Cuenca del Tempisque , Colecta Lomas Barbudal, Sendero colegio Bagaces, 0 m, 18 July 2000 , (Fr.), G. Vargas & E. Wehncke 2024 ( CR !, MO !); Along stream, ca. 11 km n of La Cruz , 0.5 km west of main road, 75 m , 2 February 1978 , (Fl.), R. Liesner 4840 ( CR !, MO !); Nandayure , Bejuco , Pacífico Norte , Bejuco , Pilas de Bejuco , Finca de Benigno Mayorga , 40 m , 26 July 1994 , (Fr.), A. Estrada & A. Rodríguez 91 ( CR !, MO !); Santa Cruz , Santa Cruz , Bosque Nacional Diría, 126 m , 25 January 2000 , (Fr.), L. Acosta 289 ( CR !, MO !); Bagaces , Bagaces , Ojo de Agua , Papayito , Balas de Cañón , 100 m , 22 August 2000 , (Infer.), L. Acosta 2569 ( CR !) ; La Cruz , La Cruz, Cerro El Hacha, Camino a Santa Cecilia, 300 m , 23 March 1992 , (Fl.), R. Espinoza 264 ( CR !) ; Santa Cruz , Santa Cruz , Fila Vista del mar, camino a las torres de comunicación, 300 m , 23 September 1996 , (Fr.), J. Gonzáles 1252 ( CR !, MO !); Tilarán , Tronadora , Rio Chiquito , Arenal , Zona Monteverde , 730 m , 3 March 1988 , (Fl.), W. Haber 8256 ( CR !) ; Nicoya , San Antonio , Los Mesones, cerca de los tanques de captación, 300 m , 21 August 1992 , (Fl.), M. Reyes 8 ( CR !) ; Tilarán , Tierras Morenas , Tierras Morenas , Rio Cabuyo , 685 m , 1 August 1994 , (Fl.), G. Rodríguez 298 ( CR !) ; Liberia , Curubandé, P .N. Rincón de la Vieja , sendero hacia el cráter, 1004 m , 1 June 2011 , (Fl., Fr.), L. Vargas 4573 ( CR !, MO !); Bagases , Mogote, P.N. Rincón de la Vieja , colecta camino a San Jorge, 3 km , de la casona Sata María, 400 m , 17 February 1991 , (Fr.), G. Rivera 1149 ( CR !) ; Disturbed primary forest and open area along Río Higuerón near agricultural experimentation area near Taboga, 0–100 m , 29 June 1977 , (Fl., Fr.), R. Liesner 2709 ( MO !) ; Tilarán, Tronadora , Monteverde , 3.5 km N Santa Elena on road to San Gerardo, 0.5 km N of junction road and Rio Negro, Lower montane wet forest, 1540 m , 20 August 1988 , (Fl.), W. Haber 8619 ( CR !) ; Along Río Las Flores (identified on Tierras Morenas quadrangle as ”Río Flores”) 450 m , 25 January 1985 , (Fr.), M.H. Grayum 4913 ( MO !) ; Abangares , Cuenca del Abangares , Zona Protectora Abangares , Sector Ecomuseo hasta Río AguasClaras , 300–500 m , 20 June 1997 , (Fr.), U. Chavarría 1737 ( MO !) ; Finca La Pacifica along Rio Corobici 13 December 1977 , (Infer.), D. Janzen 10890 ( MO !) ; P.N. Rincón de la Vieja , sendero de la toma de agua, a 3 km de la estación 17 September 1990 , (Fr.), G. Rivera 601 ( USJ !) ; Guanacaste , Cañas, Porozal, Disturbed primary forest and open area along Río Higuerón near agricultural experimentation area near Taboga , 50 m , 29 June 1977 , (Fr.), R. Liesner et al. 2717 ( CR !, MO !) ; Heredia : Sarapiquí , Puerto Viejo , E.B. La Selva, Finca La Selva , En las parcelas de sucesión, 9 September 1983 , (Fr.), I. Chacón 1339 ( CR !) ; Sarapiquí , La Virgen , E. B. La Selva, E.B. La Selva, Along road leading to the reserve, 100 m , 16 August 1987 , (Fr.), J.F Smith & E. Frost 478 ( CR !) ; Sarapiquí , Las Horquetas, Along entry road to Finca La Selva , Puerto Viejo de Sarapiquí (from Puerto Viejo-Las Horquetas road), just before La Selva boundary, 5 m , 2 June 1985 , (Fr.), M.H. Grayum 5322 ( CR !, MO !); Sarapiquí , Puerto Viejo , Near the junction of the Río Puerto Viejo and the Río Sarapiquí , 100 m , 15 July 1980 , (Fr.), B.E. Hammel 9215 ( CR !) ; Sarapiquí , Las Horquetas , Just n of Las Horquetas along road to Puerto Viejo, 40 m , 19 July 1984 , (Fr.), M.H. Grayum 3563 ( CR !) ; Santo Domingo , Tures , Lote y charral por el Río Tures , ca. 1 km al sur de San Francisco de San Isidro de Heredia, 1250 m , 29 March 2004 , (Fr.), B.E. Hammel 22936 ( CR !) ; Sarapiqui , 2 km south of La Virgen on Puerto Viejo-San José road, 14 June 1981 , (Fl., Fr.), B.E. Hammel 10886 ( MO !) ; La Selva Biological Station . 100 m , 20 June 1984 , (Fr.), B. Jacobs 2441 ( MO !) ; Sarapiqui , La Selva Biological Station 100 m , 8 June 1983 , (Fr.), I. Chacón 898 ( MO !) ; Finca La Selva , 100 m , 4 August 1980 , (Fl.), B.E. Hammel 9436 ( MO !) ; Finca La Selva , 100 m , 23 May 1980 , (Fr.), M.H. Grayum 2848 ( MO !) ; Finca La Selva , 100 m , 19 June 1981 , (Fr.), B.E. Hammel 10890 ( MO !) ; La Selva Biological Station , 100 m , 25 June 1984 , (Fl., Fr.), B. Jacobs 2569 ( MO !) ; Near La Selva Biological Station , 100 m , 4 July 1984 , (Fr.), B. Jacobs 2655 ( MO !) ; Sarapiquí , La Virgen, Reserva Biológica La Tirimbina , En el parqueo del Lodge, 280 m , 1 July 2013 , (Fr.), J. López 327 ( USJ !) ; Heredia , Sarapiquí , Horquetas , Estación Biológica La Selva , 50 m , 3 November 2018 , (Fr.), M. Cedeño & M. Chaves 1495 ( USJ !) ; Limón : Pococí , Guacimo , Forested area near Guasimo along river, 22 April 1970 , (Fr.), E. Valerio s.n ( CR !) ; Puntarenas : Corredores, Corredor, Forests on main ridge and NE slopes of Fila de Cal , between San Vito and Ciudad Neilly , 550 m , 13 September 1985 , (Fr.), M. Grayum et al. 6049 ( CR !, MO !); Montes de Oro , San Isidro, Interamerican Highway km marker 122; patch of forest west of road, 100 m , 4 August 1985 , ( Fr. ), B.E. Hammel & J. Trainer 14367 ( CR !, MO !); Golfito , Puerto Jiménez , P. N. Corcovado , Estación Sirena , 5 m , 15 June 1990 , (Fl.), N. Obando 61 ( CR !) ; Osa , Palmar , Sitio Arqueológico , Finca 6 , Unos 5,5 Km SO en linea recta de Palmar Sur, Camino del portón a los montículos principales, 100 m , 11 October 2011 , (Fr.), A. Quesada et al. 3369 ( CR !) ; Garabito, Tárcoles, P.N. Carara. Along Quebrada Bonita, Carara Reserve , 35 m , 25 July 1985 , (Fl., Fr.), M.H. Grayum et al. 5715 ( CR !, MO !); Golfito , Golfito, Along short-cut road to Golfito from Villa Briceño on Interamerican Hwy, W side of Fila Gamba, ca. 6 km from Golfito airport, 80 m , 6 March 1985 , (Fr.), T.B. Croat & M.H. Grayum 59904 ( CR !, MO !), Puntarenas , Paquera , Reserva Absoluta Cabo Blanco , Estación Cabo Blanco , Límite de la Reserva , Borde de bosques y potreros, 80 m , 7 November 1991 , (Fl., Fr.), U. Chavarría 320 ( CR !) ; Puntarenas , Guacimal, Monte Verde area, valley of Río San Luis just south of Monte Verde; from 1/ 2 km below waterfall to base of waterfall, 1050 m , 23 June 1985 , (Fr.), B.E. Hammel & J. Trainer 14016 ( CR !, MO !); Puntarenas , Paquera , P.N. Cabo Blanco , Secondary forest , 30 to 50 years old, 10 m , 30 April 1994 , (Fr.), M. Blanco 2257 ( CR !) ; Osa , Sierpe, Valley of Laguna Chocuaco , ca. 9 km W of Rincón de Osa, 200 m , 8 October 1984 , (Infer.), M.H. Grayum et al. 4074 ( CR !, MO !); Garabito , Tárcoles, P. N. Carara, Cuenca del Tárcoles, Quebrada Bonita sector, 25 m , 11 May 2009 , (Fr.), L. Vargas et al. 3779 ( CR !, MO !); Golfito , Golfito , P.N. Piedras Blancas , 100 m , 6 June 2000 , (Fl.), L. Acosta et al. 1514 ( CR !) ; Garabito , Jacó , 7 m , 5 September 2001 , (Fr.), A. Ruíz et al. 545 ( CR !) ; Puntarenas , Monteverde, Monteverde, 6 km south of Santa Elena on road to highway, Los Cerros, ridge between Río Guacimal and Río Lagarto, 950 m , 19 July 1991 , (Fl., Fr.), W. Haber & W. Zuchowski 10769 ( CR !, MO !), 8 km N of Barranca, 1 km N of Miramar turn off. W side of km 123 on the Interamerican Hwy. Quebrada Negros, 25 m , 30 April 1983 , (Fl.), R. Liesner et al. 15133 ( CR !, MO !); Puntarenas , Guacimal, R.B. Monteverde, San Luis, Monteverde, Río Guacimal, Bosque húmedo premontano, 700 m , 24 June 1988 , (Fl.), E. Bello et al. 21 ( CR !) ; Puntarenas , Paquera, Península de Nicoya , Curú , Río Curú mouth, Pasture on flats W of ranch headquarters, 0 m, 14 August 1995 , (Fr.), A. Sanders et al. 1755 6 ( CR , USJ , MO ) ; Parrita , Parrita, Forest along Río Paquita , 2 m , 13 August 1936 , (Infer.), C. Dodge & V. Goerger 9763 ( CR !, MO !); Curu , 5 November 1986 , (Fr.), R. Soto 3149 ( CR !) ; Golfito , Puerto Jiménez. P.N. Corcovado, Sirena, Ollas Trail , 0 m, 28 May 1988 , (Fl.), C. Ketnan et al. 538 ( CR !) ; Golfito , Golfito , Refugio Nacional de Fauna Silvestre Golfito , Cerro Nicuesa , 345 m , 7 April 1994 , (Fl.), G. Rivera & G. Herrera 2268 ( CR !) ; Golfito , Jiménez, Alrededor de la estación, 20 m , 11 September 1998 , (Fr.), R. Aguilar 5530 ( CR !, MO !); Puntarenas , Lepanto , Reserva Karen Mongense , Sendero al Mirador viejo ; colectado a orillas del sendero; 400 m , 9 May 2003 , (Fl., Fr.), E. Alfaro 4342 ( CR !) ; Puntarenas , Cóbano , Sendero Central , 17 December 1993 , (Fl.), A. Fernández 1320 ( CR !) ; Puntarenas , Lepanto , Jicaral , San Ramón de río Blanco , Cerro Escondido , Bosque alterados remanentes y áreas abiertas, en la cuenca de la Quebrada Pérez, 317 m , 8 May 2003 , (Fl., Fr.), J. Gonzales 3679 ( CR !) , Puntarenas , Cóbano , Estacion San Miguel , ca. 2 km S. de Malpais, 0 m, 18 January 1996 , (Fl.), B.E. Hammel 20090 ( CR !) ; Buenos Aires , Potrero Grande , Bomba de Cedro , por el Río Cabagra , 200 m , 3 March 2000 , (Infer.), B.E. Hammel 22101 ( CR !) ; Golfito , Jiménez, Rio Piro , Playa Carate , 10 m , 15 September 1990 , (Fl., Fr.), G. Herrera 4298 ( CR !, MO !); Golfito , Jiménez , Estacion Sirena , 1 m , 1 September 1990 , (Fl.), J. Quesada 52 ( CR !, MO !); Between Palmar Sur and Piedras Blancas , 20 m , 28 February 1976 , (Fr.), T. Croat 32915 ( MO !) ; La Llorona , 0– 200 m , 8 August 1978 , (Fr.), D. Janzen 11043 ( MO !) ; Golfito , Jiménez , Estacion Sirena , Sendero Naranjos , 10 m , 27 July 1994 , (Fr.), R. Aguilar 3522 ( CR !) ; Aguirre , Quepos, Escuela Finca Palma al lado de Rio Naranjo , 10 m , 26 July 1995 , (Fl.), M. Chavarría 929 ( CR !, MO !); Golfito , Jiménez , Albergue Cerro de Oro , Rio El Nino , 100 m , 18 March 1995 , (Fl., Fr.), E. Fletes 104 ( CR !) ; Golfito , Jiménez, Alrededores de la Estacion Sirena, Bosque primario, 10 m , 14 April 1995 , (Fr.), B. Gamboa 152 ( CR !, MO !); Golfito , Jiménez , Rio Claro , 5 m , 6 August 1988 , (Fr.), C. Kernan 784 ( CR !) ; Golfito , Jiménez , Quebrada La Ignasia , del puente para el manglar, Finca de Pedro Vaca , 3 m , 20 June 1998 , (Fl.), M. Lobo 224 ( CR !) ; Golfito , Jiménez, Estacion Sirena, Recolectado a orilla del Rio Pavo , 10 m , 16 April 1995 , (Fl.), A. Picado 176 ( CR !, MO !); Monte Verde área , valley of Río San Luis just south of Monte Verde , in woods along the river, 1000–1200 m , 18 June 1985 , (Fl.), B.E. Hammel 13935 ( MO !) ; Northeastern slopes of Fila de Cal , 500–620 m , 12 July 1985 , (Fl.), M.H. Grayum 5649 ( MO !) ; Swampy area near Interamerican Highway in vicinity of Piedras Blancas , 28 February 1978 , (Infer.), T.B. Croat 32954 ( MO !) ; Puntarenas , Secondary vegetation on former plantations and pasture, 0–200 m , 1 December 1969 , (Fr.), W. Burger 6657 ( MO !) ; Osa Peninsula , 19 August 1984 , (Fr.), A. Gentry 48693 ( MO !) ; Monteverde , San Luis river valley below community on Pacific slope, 1000 m , 9 May 1986 , (Fl.), W. Haber 5010 ( MO !) ; Canton OSA , along road between Rincón and Boscosa, 50 m , 11 September 1996 , (Fr.), T.B. Croat 79251 ( MO !) ; Buenos Aires , Potrero Grande , Las Vueltas , Centro Turístico Los Chucuacos , 295 m , 25 July 2015 , (Fr.), M. Cedeño 804 ( USJ !) ; Puntarenas , Monteverde, Reserva Biológica San Luis, Bosque secundario cercano a la rivera del Río Guacimal, 673 m , 6 December 2013 , (Fr.), P. Juárez et al. 497 ( USJ !) ; Reserva Biológica Cabo Blanco , 6 August 1993 , (Fr.), R. Soto s.n ( USJ !) ; Parrita , Plantación vieja de palma africana, 10 m , 20 Enero 2001 , (Fr.), C. Morales 1496 ( USJ !) ; Camino del ICE de Santa Cruz a Vista de Mar , 22 July 1985 , (Fr.), H. Pittier 23671 ( MO !) ; Puntarenas , At Lindora near Monteverde , 29 September 1970 , (Fl.), H. Kennedy 571 ( CR !) ; San José : Puriscal , Chires, P.N. La Cangreja, Along Río Negro , E of Santa Rosa, 320 m , 21 July 1988 , (Fl.), M.H. Grayum et al. 8608 ( CR !, MO !); Alajuelita , San Antonio , Z.P. Cerros de Escazú , Alrededores de Cerro Rabo de Mico , 2200 m , 22 September 1989 , (Infer.), G. Vargas et al. 765 ( CR !) ; Playa Dominical , Baru, and Tinamastes (along road to San Isidro del General), 250 m , 20 November 1975 , (Fr.), M. Burger 10165 ( CR !, MO !); Mora , Colón , Z.P. El Rodeo , Reserva de la Universidad para la Paz , 500 m , 28 January 1994 , (Fr.), A. Cascante et al. 198 ( CR !) ; Región de El General , 850 m , 1 March 1940 , (Fr.), A. Skutch 4824 ( CR !) ; Turrubares , San Juan de Mata , No protegida, Cuenca del Tárcoles , San Luis de Turrubares , finca de Melvin Chavarría, 1200 m , 5 October 2004 , (Infer.), A. Soto 173 ( CR !) ; Pérez Zeledón Río Nuevo, R.F. Los Santos, Savegre Abajo , Márgenes del río División , 550 m , 11 November 1998 , (Fr.), A. Estrada et al. 1899 ( CR !) ; Mora , Colón , Z.P. El Rodeo, Bosque de la Universidad para la Paz , 1020 m , 13 December 2001 , (Fr.), A. Quesada et al. 925 ( CR !) ; Mora , Colón , Z.P. El Rodeo, Finca el Rodeo , camino a la Universidad para la Paz , 500 m , 27 April 1994 , (Fl.), A. Ruíz 440 ( CR !) ; Mora , Tabarcia , Zona Protectora Cerros de Escazú , cuenca del río Negro , 2 km en línea recta al noreste de la plaza de Palmichal en remanentes de bosque a la orilla del río, 1283 m , 19 August 2010 , (Fr.), J. Sánchez & R. Chacón 2166 ( CR !) ; Pérez Zeledón , Daniel Flores , San Isidro de El General , Repunta , Finca de Miguel Quesada , 700 m , 1 August 1993 , (Fl.), R. Aguilar 2077 ( CR !) ; Mora , Colón , Camino a Piedras Negras, sabanas poco antes de llegar al Río Jaris , 500 m , 1 November 2003 , (Fr.), R. Kriebel 4020 ( CR !) ; Turrubares , San Juan de Mata , San Pablo de Turrubares , 100 m , 8 December 2004 , (Fr.), D. Santamaría 331 ( CR !) ; Along Interamerican Highway , 28 February 1976 , (Infer.), T.B. Croat 32942 ( MO !) ; Puriscal , Along main road between Mercedes Sur and Zapotal , 900 m , 1 January 1985 , (Fr.), M.H. Grayum 4688 ( MO !) ; San José , Carmen , Barrio Escalante , 2 November 1984 , (Fr.), E. Valerio 121 ( USJ !) ; Curridabat , Curridabat , Granadilla , 30 August 1980 , (Fr.), E. Valerio s.n ( USJ !) ; Curridabat , Curridabat , Granadilla , 18 September 1984 , (Fr.), E. Valerio 116 ( USJ !) ; Curridabat , Curridabat . Granadilla. 28 July 1980 , (Fr.), E. Valerio s.n ( USJ !) ; Mora , Colón , Zona Protectora El Rodeo , Ladera Sur-Oeste de la Fila Diamante , 880 m , 25 October 2011 , (Fr.), L. Ríos 2 ( USJ !) ; Montes de Oca , San Pedro, San Pedro, 17 July 1981 , (Fr.), E. Valerio s.n ( USJ !) . PANAMA . Bocas del Toro : Rambala and vicinity , 08°55’06”N 082°09’18”W , 50 m , 24 April 1988 , S.A. Thompson 4921 ( CM , MO ) ; Chiriquicito to 5 miles S along Río Guarumo , 08°55’N 082°10’W – 09°00’N 082°11’W, 05 June 1967 – 07 June 1967 , W.H. Lewis et al. 2012 ( MO !) ; Canal Area : Old Fort San Lorenzo , 09°18’06”N 080°00’05”W , 0-15 m , 06 November 1965 , E.L. Tyson 2217 ( IBE , SCZ ) ; Along road to radar station on Semaphor Hill. 1km NW of Summit Garden. Secondary tropical moist forest, 09°04’31”N 079°38’52”W , 100 m , 07 Oct 1973 , M.H. Nee 7291 ( MO !) ; Barro Colorado Island . Fairchild Point [Salud Point], 09°10’20”N 079°50’04”W , 0–5 m , 20 Sep 1968 , T.B. Croat 6225 ( MO !) ; Summit Gardens , 09°03’52”N 079°38’58”W , 75 m , 25 March 1970 , T.B. Croat 9061 ( MO !) ; Chiriquí : Jardin Botánico de la UNACHI , 08°23’03”N 082°27’10”W , 24 m , 28 August 2018 , O. Ortiz et al. 3388 ( MO !) ; Vicinity of Puerto Armuelles , 08°17’00”N 082°52’00”W , 0–75 m , 28 July 1940 – 31 July 1940 , R.E. Woodson & R.W. Schery 903 ( MO !) ; Burica Peninsula.Along stream; Quebrada Mellize, 6 miles ( 10 km ) south of Puerto Armuelles , 08°11’N 082°53’W , 0–150 m , 05 March 1973 , R.L. Liesner 503 ( MO !) ; 1 mi W of airport at Puerto Armuelles , 08°15’00”N 082°52’30”W , 0–5 m , 17 Feb 1973 , T.B. Croat 21914 ( MO !) ; Coclé : Along El Valle de Antón , roadside near town, 08°37’N 080°07’W , 23 Apr 1977 , J.P. Folsom & J. Kauke 2777 ( MO !) ; El Jordanal , Rio Indio Arriba , 08°40’10”N 080°07’01”W , 570–840 m , 23 Jul 2001 – 25 Jul 2001 , M.B. Jorge 5–130 ( PMA !) ; Colón : Along tributary of Quebrada Anchca . 4 km E of Buena Vista, 3 km N of cement plant, 09°18’N 079°39’W , 80 m , 04 November 1973 , M.H. Nee 7787 ( MO !) ; Mosquera , forest, 09°13’30”N 080°05’30”W , 15–50 m , 17 September 1974 , P.J. Maas & S.A. Mori 1749 ( MO !) ; Forest and forest edge from Portobelo Highway to 40 km up Río Guanche. [Coordinates on original label: 09°30’ N 079°40’ W], 09°30’N 079°38’W - 09°31’N 079°41’W, 0–50 m , 30 August 1981 , S. Knapp 1023 ( MO !) ; Portobelo. 2–4 km up the Río Guanche from the Portobelo Highway, 09°30’N 079°39’W – 09°30’N 079°40’W, 0–50 m , 03 October 1981 , S. Knapp & R. Chazdon 1407 ( MO !) ; Darién : 1–4 miles N of Pucro , 08°00’07”N 077°33’00”W – 08°03’00”N 077°35’00”W, 0–100 m , 22 June 1967 , J.A. Duke 13051 ( MO !) ; Puerto Santa Dorothea (=Puerto Piñas), near Piñas Bay, 07°35’N 078°11’W , 0 m, 21 July 1962 , J.D. Dwyer 2299 ( MO !) ; Area de Manejo Especial de Bahia Piñas , bosque cercano a Aceite , 07°37’41”N 078°11’10”W , 87 m , 30 June 2018 , O. Ortiz et al. 3011 ( MO !, PMA !); Area de Manejo Especial de Bahia Piña , b osque cercano a Aceite, 07°37’41”N 078°11’10”W , 87 m , 30 June 2018 , O. Ortiz et al. 3012 ( MO !, PMA !); Area de Manejo Especial de Bahia Piñasm bosque seundario circundanate al hotel Tropic Star Lodge , 07°35’11”N 078°12’38”W , 0 m, 30 June 2018 , O. Ortiz et al. 3028 ( MO !, PMA !); Vicinity of gold mine at Canal , 07°45’N 077°41’W , 500–600 m , 26 Jul 1976 , T.B. Croat 37604 ( MO !) ; Río Pirre , 07°55’N 077°44’W , 14 Jul 1971 , T.B. Croat & D.M. Porter 15487 ( MO !) ; Los Santos : Tonosi . Azuero Peninsula, along trail between Jobero and headwaters of Río Pedregal, 07°20’N 080°35’W , 300–700 m , 28 April 1976 , T.B. Croat 34464 ( MO !) ; Along road between Tonosí and Jobero, 07°20’30”N 080°30’00”W , 50–80 m , 27 April 1976 , T.B. Croat 34455 ( MO !) ; Panamá : Sendero ecológico del Centro de Ciencias v Arte. EXPLORA, J. Vergara 4 ( MO !, PMA !); Panamá . Panama City , at Lefevre Park, 08°58’00”N 079°32’00”W , 28 Jun 1940 , H.H. Bartlett & T. Lasser 16337 ( MO !) ; Cerro Jefé , 09°14’02”N 079°22’30”W , 700–1000 m , 12 Apr 1972 , T.C. Plowman & A. Weil 3159 Cerro Jefé , 09°14’02”N 079°22’30”W , 700–1000 m , 12 Apr 1972 , T.C. Plowman & A. Weil 3156 San Blas : Moli-ya , tierra firme frente a Isla Playón Chico, camino principal al Sur. [Coordinates on original label: 09º20’N, 078º13’W], 09°19’20”N 078°13’58”W, 0–50 m , 17 septiembre 1987 , H. Herrera 371 ( MO !) ; Coastal rocks between Puerto Obaldía and Puerto Armila , 08°40’N 077°24’W – 08°40’N 077°28’W, 0–300 f, 29 Apr 1980 , W.G. D’Arcy 13683 ( MO !) ; Veraguas : Along road from Guabala on Panamerican Hwy. and Soná, between Quebrada de Piedra and Puerto Vidal , ca. 1 km W of Puerto Vidal, 08°03’10”N 081°37’54”W , 37 m , 07 March 2015 , T.B. Croat 106239 ( HUA , MO , PMA ) ; Calobre . Colectado hacia Chitra, encalle que va hacia los pozos termales, 08°30’50”N 080°54’17”W , 564 m , 18 May 2017 , E. Campos 940 ( MO !, PMA !); Canazas . Ca. 1 km above Cañasas [Cañazas] on road to Los Valles, 08°19’59”N 081°12’43”W , 230 m , 14 July 1976 , T.B. Croat 37048 ( MO !) ; Montijo . Isla de Coiba, catival entre campamento de Playa Blanca y el Barco Quebrado. 17NMU2513, 07°23’01”N 081°40’07”W – 07°20’16”N 081°41’32”W, 0–50 m , 26 Nov 1994 , A. Espinosa et al. 727MB ( MO !, PMA !); Isla de Coiba , catival entre campamento de Playa Blanca y el Barco Quebrado. 17NMU2513, 07°23’01”N 081°40’07”W - 07°20’16”N 081°41’32”W, 0–50 m , 26 Nov 1994 , A Espinosa et al. 716MB ( MO !) ; Isla de Coiba , en cativales de Barco Quebrado, 07°20’16”N 081°41’32”W , 10 m , 26 May 1998 , A. Ibáñez et al. 653AI ( MO !, PMA !); P.N. Coiba . Jicarón. Río más largo de la isla. Orilla del río. 17NMU 1202, 07°15’23”N 081°47’50”W , 5 m , 09 Jul 2004 , A. Ibáñez et al. 3719AI ( MO !) ; Isla Coiba . Ensanada de Santa Cruz , 07°37’N 081°45’W , 2 May 1995 , C Galdames et al. 2367 ( MEXU !, MO !); Isla Coiba . Camino desde Barco Quebrada hasta el Campamento Playa Blanco , 07°20’16”N 081°41’32”W – 07°23’01”N 081°40’07”W, 0–50 m , 03 Sep 1995 , J.E. Aranda et al. 2339 ( MO !, PMA !); Cerro Hoya , subiendo por Cobachón, 07°16’20”N 080°39’15”W , 350 m , 21 abril 1997 , J. Deago et al. 475 ( MO !, PMA !); Isla de Coiba , entre el campamento de playa Blanca y el de Barco Quebrado, 17NMU2614, 07°22’54”N 081°39’18”W , 28 January 1994 , S. Castroviejo et al. 7253 MV ( MO !) ; Santa Fe . Vicinity Santa Fé, along road between Santa Fé and Calovebora, 1.7 miles past Alto Piedra School, 1.5 miles beyond Quebrada Cosilla (previously referred to as Río Primero Braso), 08°31’28”N 081°07’50”W , 570 m , 13 July 1994 , T.B. Croat & G.H Zhu 76811A ( MO !) ; Sona . Isla Canales de Tierra. Playa del Laboratorio. 17NMU 3656, 07°44’53”N 081°34’26”W , 2 m , 29 Jan 2002 , A. Ibáñez & N. López 1493 ( MO !, PMA !) .