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 40. Monstera pinnatipartita Schott, Oesterr. Bot. Wochenbl. 7: 197. 1857 . ( Figs. 67 , 68 ) Type: — VENEZUELA . Distrito Federal: Caracas, H.G . Reichenbach s.n. ( holotype W destroyed, see Riedl & Riedl-Dorn, 1988 ) . — VENEZUELA. Zulia : Distrito Perijá , ca. 13 airline km NE of intersection of the Maracaibo-La Fría Hwy. (Hwy. 6) and the Río Aricuaisá (near intersection of LAGOVEN picas 80-2 and 19), 40 m , 20 June 1980 , G. Davidse , A.C. González & R.A. León 18286 ( neotype MO !, isoneotype VEN, designated by Grayum (1997)) . [ Monstera dilacerata auctt. non ( Koch & Sello 1853 ) K. Koch (1855: 5) . (i.e. Epipremnum pinnatum (L.) Engl.): Standley, Fl. Panama 2(3): 31. 1944 (pro parte); Madison (1977: 57 , pro parte); Croat (1978: 207) .] Nomadic vine, appressed-climbing habit. SEEDLINGS: bearing foliage leaves. JUVENILE PLANTS: root climbers; stems dark green with white dots, smooth, cylindrical or flattened; internodes 3–6 cm long, 3–5 mm diam.; petiole distinct, dark green, smooth, 4–12 cm long, sheathed to base of the geniculum; petiole sheath persistent; blades lanceolate, subcordate to truncate at base, acuminate at apex, coriaceous, 10–15 × 3–8 cm , not appressed to the phorophyte; fenestrations absent or present. ADULT PLANTS: root climbers; stems smooth, cylindrical, or slightly dorsoventrally compressed, dark green with white dots; internodes 1–3 cm long, 1–3 cm diam., as long as wide; cataphylls light or dark, mottled or whitish, persistent or occasionally deciduous; anchor roots dark brown; feeder roots dark brown; petiole light or dark-green, mottled to whitish, smooth, 15–65 cm long, sheathed up to 3–6 cm before the geniculum or to base of the geniculum; petiole sheath persistent, involute; geniculum smooth, sunken adaxially, convex abaxially, 2–3 cm long; blades ovate to lanceolate-ovate, broadly cuneate to rounded or truncate at base, obtuse to short-acuminate at apex, subcoriaceous to coriaceous, drying yellowish, blackish or light brown, 20–70 × 20–35 cm , (1.2)1.7–2.1 times longer than wide, decurrent on the geniculum, decurrent portion 1–2 mm wide; midrib ribbed adaxially, convex abaxially, drying black or yellowish on both surfaces; primary lateral veins 6–20 per side, slightly sunken adaxially, prominent abaxially, departing midrib at 35°–75°, drying black or yellowish; collective vein not visible; fenestrations absent or present; margins deeply pinnatifid, 3–10 lobes per side, 1.5–5.0 cm wide, 1–3 veins per lobe. INFLORESCENCES on ascending stems, 1–3 simultaneously at flowering time, arranged in the axils of the leaves or into cataphylls; peduncle smooth, mottled, 10–25 cm long, 1.0– 1.5 cm diam.; spathe acuminate to long-acuminate, yellowish green externally during development, white yellowish externally and white internally at anthesis, overlapping basal margins, deciduous after anthesis; 12–20 × 6–9 cm , up to 7 cm longer than the spadix; spadix white during development, creamy-yellow at anthesis, 6–15 cm long, 1.3–2.5 cm diam., (4.4)5.5–9.1 times longer than wide; basal sterile flowers 3–5 mm long, with a rusty-red stigmatic secretion; fertile flowers 4–7 mm long; stamens 2–6 mm long, with laminar filaments; anthers 1.5–2.0 mm long; ovary rectangular in longitudinal section, ribbed, 4–6 × 3–4 mm ; style hexagonal, 2–3 × 3–5 mm ; stigmatophore columnar, 0.3–0.5 mm long; stigma linear, with an orange stigmatic secretion; berries with a yellowish-green stylar cap during development, mature stylar cap cream-white; pulp white; seeds oblong, black, 5–6 mm long. Distribution and ecology: — Monstera pinnatipartita ranges from El Salvador and Costa Rica to Panama and Colombia ( Magdalena , Bolívar, Huila , Valle, Risaralda , Nariño , Meta and Caquetá ), northern Venezuela , western Ecuador ( Esmeraldas , Pichincha , Los Rios , Guayas ) and Peru ( Loreto , Ucayali , San Martín , Huánuco ), at 0–600 m , rarely to 1000 m , in Tropical wet forest , Premontane wet and Lower montane rain forest life zones. Phenology: —Flowering has been recorded in January, September-November. Fruit in January, March-June and October. Discussion: —The species is a member of sect. Monstera . It differs from the other species of the genus by the light or dark green petiole, speckled to whitish, the persistent petiolar sheath with involute wings, the deeply pinnatifid leaf blade, and externally yellowish-white and internally white spathe. It can be confused with individuals of Monstera dissecta but that has pinnatilobed leaves with wider lobes ( 4‒10 cm vs. 1.5‒5.0 cm in M. pinnatipartita ). Monstera pinnatipartita is distributed throughout the Pacific slope, being the most frequent species, growing by estuaries, open areas and primary forest. This species develops hanging stems when the plant cannot continue its ascent, but these hanging stems are never reproductive. Moreover, pre-adult plants can develop inflorescences in ascending stems when they have fenestrate and not pinnatifid leaves ( Cedeño-Fonseca et al. 2022 ). For more details about the taxonomic comments and nomenclatural note see Cedeño-Fonseca et al. (2022) . Additional specimens examined: — EL SALVADOR . Jardín Botanico , Zona 2S , 13°40’N 089°15’W , 800 m , 12 Jul 1989 , R. Villacorta & S. Martínez 308 ( MO !) . COSTA RICA . Guanacaste : Hojancha, Puerto Carrillo , Finca Dyalá , Puerto Carrillo , Bosque situado en el nacimiento de la Quebrada Zapotal, 20 m , 26 December 1988 , (Fr.), L. Flores & C. Herrera 12 ( CR !, MO !) ; Hojancha , Puerto Carrillo , Along Quebrada Zapotal (not on current maps), from Finca Dyalá upstream to near summit of ridge (ca. 2 km ENE of Puerto Carrillo ), Península de Nicoya , 60 m , 30 April 1989 , (Fr.), M.H. Grayum et al. 9438 ( CR !, MO !) , Santa Cruz , Cuajiniquil , Cerro Vista al Mar , 750 m , 16 March 2008 , (Fr.), B. Hammel 24541 ( CR !) ; Puntarenas : Montes de Oro , San Isidro , Interamerican Highway km marker 122; patch of forest west of road, 100 m , 4 August 1985 , (Infer.), B.E. Hammel & J. Trainer 14369 ( CR !, MO !) ; Osa Sierpe , P.N. Corcovado, Llorona Forest , 75 m , 29 May 1988 , (Fr.), C. Kernan et al. 549 ( CR !) ; Secondary vegetation on former plantations and pasture and remnants of original tall evergreen forest on steep slopes and stream edges in the Cabo Blanco Nature Reserve , Southern tip of the Nicoya Península, 100 m , 1 December 1969 , (Fr.), W. Burger & R. Liesner 6607 ( CR !, MO !) ; Golfito , Puerto Jiménez, P.N. Corcovado, Near estación Corcovado, 10 m , 7 December 1989 , (Fr.), Merz 454 ( CR !) ; Golfito , Puerto Jiménez. P.N. Corcovado, Pavo forest, Poorly drained lowland forest, 70 m , 14 July 1988 , (Fr.), C. Kernan 693 ( CR !) ; Osa , Sierpe , Pasture and forest edge along Quebrada Banegas , ca. 4 km W of Rincón de Osa, 40 m , 8 October 1984 , (Fr.), M.H. Grayum 4118 ( CR !, MO !) ; Golfito , Puerto Jiménez , Along road between Rincón and Puerto Jiménez, 15 km S of Rincón, disturbed areas along road, 30 m , 4 March 1985 , (Fr.), T.B. Croat 59796 ( CR !, MO !) ; Golfito , Puerto Jiménez , P.N. Corcovado , Sirena Woods , 50 m , 30 April 1989 , (Fr.), C. Kernan 1062 ( CR !) ; Garabito, Tárcoles, P.N. Carara , Along Quebrada Bonita , Carara Reserve , 35 m , 25 July 1985 , (Fr.), M.H. Grayum et al. 5736 ( CR !, MO !) ; Isla del Caño , 25 July 1985 , (Infer.), R. Soto 2421 ( CR !) ; Osa , Sierpe , Ridge between Quebrada Banegas and Río Riyito , ca. 7 Km W of Rincón de Osa, 200 m , 8 October 1984 , (Fr.), M.H. Grayum 4085 ( CR !, MO !) ; Area del Faro en el lomo de la fila, Isla del Caño, 26 July 1985 , (Infer.), R. Soto 2457 ( CR !) ; Sendero al chorro, Isla del Caño , 13 March 1986 , (Fr.), R. Soto 2886 ( CR !) ; Golfito , Puerto Jiménez , Jiménez , cuenca inferior del Río Piro , 20 m , 16 September 1990 , (Fl.), G. Herrera 4300 ( CR !, MO !) ; Golfito , Puerto Jiménez , P.N. Corcovado, Sirena Woods , 50 m , 30 April 1989 , (Fr.), C. Kernan 1065 ( CR !, MO !) ; Osa , Bahía Ballena , Uvita , San Josecito , Faldas de la fila Alivio, Lado Pacífico de la fila Costeña, Finca Oro Verde , 300 m , 12 January 2000 , (Fr.), M. Blanco 1245 ( CR !) ; Golfito , Jiménez , Estacion Sirena , sendero Rio Claro , 10 m , 11 October 1993 , (Fr.), R. Aguilar 2456 ( CR !) ; Golfito , Jiménez , Alrededor de la Estacion Los Patos, 200 m , 7 June 1994 , (Fr.), R. Aguilar 3356 ( CR !) ; Golfito , Jiménez , Alrededor de la estación , 20 m , 11 September 1998 , (Fr.), R. Aguilar 5534 ( CR !, MO !) ; Osa , Sierpe , Isla del Caño , 0 m, 30 August 2000 , (Infer.), J. Azofeifa 26 ( CR !) ; Osa , Sierpe , Vicinity of Boscosa , at Quebrada Aguabuena , 100 m , 11 September 1996 , (Fl., Fr.), T.B. Croat 79247 ( CR !, MO !) ; Puntarenas , Cóbano , Estacion San Miguel , ca. 2 km S. de Malpais, 18 January 1996 , (Infer.), B.E. Hammel 20115 ( CR !) ; Golfito , Jiménez , Estacion Sirena , al borde del bosque primario, 2 m , 1 October 1990 , (Fl.), J. Saborio 100 ( CR !) ; Corredores , Ciudad Neily , Cerro Punta Gorda , 3 km, northwest of Punta de Burica, Mature rainforest, 200 m , 5 March 1973 , (Fr.), P. Busey 777 ( MO !) ; Osa , Bahía Ballena , Cerros de playa Dominical , 550 m , 30 Octubre 2016 , (Fl.), M. Cedeño et al. 946 ( USJ !) ; Golfito , Puerto Jiménez , Camino a la Estación Biológica Piro , 23 m , 31 Mayo 2016 , (Fr.), M. Cedeño et al. 891 ( USJ !) ; Parrita , En una plantació vieja de palma africana, ca. 4 km después de parrita, 10 m , 20 Enero 2001 , (Fr.), C. Morales 1498 ( USJ !) ; San José : Mora , Colón, Z.P. El Rodeo, Bosque de la Universidad para la Paz, Bosque Premontano Húmedo , Fila Diamante , 850 m , 13 December 1993 , (Fl.), A. Cascante et al. 90 ( CR !) ; Mora , Colón, Ciudada Colón, Finca El Rodeo, Camino Universidad para la Paz, 650 m , 27 April 1994 , (Infer.), V. Nilsson & A. Ruiz 441 ( CR !) ; Mora , Colón, Z.P. El Rodeo, Reserva de la Universidad para la Paz, Bosque húmedo premontano, 500 m , 2 May 1994 , (Fr.), V. Nilsson 462 ( CR !) ; Puriscal , Chires , Santa Rosa de Puriscal , Bosque primario y secundario en las faldas de Fila La Cangreja , 500 m , 6 January 1993 , (Fr.), J. Morales 1019 ( CR !) ; Puriscal , Chires , Mastatal de Puriscal ; bosque primario en la Quebrada Grande, por la fila y el rio, 200 m , 28 May 1994 , (Fr.), J. Morales 2817 ( CR !, MO !) ; Puriscal , Chires , Cerro Pelon , charrales y bosque primario remanente en la cima del cerro y cerca del antiguo camino a Quepos, 920 m , 20 April 1995 , (Fr.), J. Morales 3890 ( CR !) . PANAMA . Canal Area : Drowned forest of Quebrada Bonita , 09°20’04”N 079°35’51”W , 70–80 m , 20 Dec 1934 , C.W. Dodge & P.H. Allen 17126 ( MO !) ; Barro Colorado Island , edge of lake, 09°09’N 079°51’W , 0–5 m , 4 Jul 1931 , D.E. Starry 17 ( MO !) ; Barro Colorado Island , 09°09’N 079°51’W , 10–100 m , 20 Aug 1927 , L.A. Kenoyer 179 ( US !) ; Barro Colorado Island . Snyder-Molino Trail, 09°09’30”N 079°50’20”W , 10– 150 m , 30 Sep 1931 , O.E. Shattuck 27 ( MO !) ; Barro Colorado Island . Frank Lutz Trail, 09°09’49”N 079°50’17”W , 0–50 m , 29 Sep 1931 , O.E. Shattuck 6 ( MO !) ; Barro Colorado Island , Gatun Lake , 09°09’N 079°51’W , 0–120 m , 18 Nov 1925 – 24 Nov 1925 , P.C. Standley 41037 ( US !) ; Barro Colorado Island . Shoreline of Lighthouse Cove, north of #8 Front Light, 09°10’18”N 079°51’22”W , 0–5 m , 12 Feb 1969 , T.B. Croat 7906 ( MO , RSA , US ); Barro Colorado Island , Barbour Trail, 09°09’30”N 079°49’20”W , 10–100 m , 26 Sep 1968 , T.B. Croat 6485 ( MO !) ; Barro Colorado Island , William Morton Wheeler Trail , 09°09’20”N 079°51’10”W , 10–170 m , 30 Mar 1970 , T.B. Croat 9224 ( MO !) ; Barro Colorado Island . Burrunga Point, 09°08’18”N 079°50’29”W , 0–5 m , 17 May 1968 , T.B. Croat 5605 ( MO !) ; Barro Colorado Island . William Morton Wheeler Trail, 09°09’20”N 079°51’10”W , 10–170 m , 17 Jun 1970 , T.B. Croat 10909 ( MO !) ; Barro Colorado Island , clearing at laboratory, 09°09’45”N 079°50’30”W , 50 m , 15 Jan 1969 , T.B. Croat 7251 ( MO !) ; Barro Colorado Island . Snyder-Molino Trail, 09°09’30”N 079°50’20”W , 50 m , 12 Mar 1969 , T.B. Croat 8570 ( MO !) ; Barro Colorado Island . Lutz Trail, 09°09’49”N 079°50’17”W , 0–50 m , 11 Jan 1969 , T.B. Croat 7138 ( MO !) ; Madden Forest (Parque Nacional Soberanía) Las Cruces Trail , 3.6 mi N of Gamboa Road turn-off, 09°06’20”N 079°37’20”W , 140 m , 23 July 1994 , T.B. Croat & G. Zhu 77066 ( MO !) ; Chiriquí: Just W of the Fortuna Camp , 08°44’N 082°15’W , 1400–1600 m , 12 Sep 1977 , J.P. Folsom et al. 5358 ( MO !) ; NE del campamento Fortuna (Hornito), sitio de presa, después de excavaciones geológicas hasta la finca Santamaría , 08°45’N 082°14’W , 1000–1200 m , 15 agosto 1976 , M.D. Correa et al. 2465 ( MO !) ; NE del campamento de Fortuna ( Hornito sitio de presa). Camino hacia la finca Landau , 08°45’N 082°15’W , 1000–1200 m , 24 Sep 1976 , M.D. Correa et al. 2688 ( MO !) ; Burica Peninsula . Primary forest; San Bartolo Limite, 12 mi. (20 km) west of Puerto Armuelles , 08°18’N 082°58’W , 400–500 m , 24 Feb 1973 , R.L. Liesner 204a ( MO !) ; Cerro Colorado , along road above San Félix, 29 km above bridge over Río San Félix (7.9 km above turnoff to Escopeta), 08°32’07”N 081°49’11”W , 1500 m , 14 July 1976 , T.B. Croat 37080 ( MO !) ; Coclé : Near Sawmill above El Copé , Atlantic drainage east of sawmill , 08°40’00”N 080°35’30”W , 20 Jun 1978 , B.E. Hammel 3518A ( MO !) ; Colón : La Macha , 08°59’04”N 080°32’35”W , 20 m , 18 Aug 2001 , J.A. Mendieta M. 10–65 ( MO !, PMA !) ; La Macha , 08°59’04”N 080°32’35”W , 20 m , 18 Aug 2001 , J.A. Mendieta 10–208 ( MO !, PMA !) ; Along road between Portobelo and Nombre de Dios, 1.2 mi. beyond the junction of the road to Isla Grande. [Coordinates on original label: 09º36’N, 079º35’W], 09°34’N 079°34’W , 05 April 1980 , T.B. Croat 49800 ( MO !) ; Darién : Parque Nacional del Darién , along S branch of Río Pucuro; on ridge in forest E of old village of Tacarcuna; ca. 18 km E of Pucuro, 08°04’N 077°16’W , 600–900 m , 23 Oct 1987 , B.E. Hammel et al. 16449 ( MO !) ; About 10 miles S of El Real on Río Pirre (House no. 22), 08°01’N 077°44’W , 10 August 1962 – 11 August 1962 , J.A. Duke 5461 ( MO !) ; Upper Río Tuquesa , 08°26’N 077°28’W - 08°34’N 077°42’W , January 1973 - May 1973 , J.M. Gustave Le Clézio 14 ( MO !) ; Ensenada del Guayabo , 16–19 km SE of Jaqué , 07°25’N 078°03’W , 0–100 m , 17 Jan 1981 , N.C. Garwood 1025 ( MO !) ; Serrania de Majé . Reserva Privada Chucanti . Cerro Chucanti . Camnino a la cascada, 08°47’31”N 078°26’51”W , 699 m , 06 April 2018 , O. Ortiz 3171 ( MO !, PMA !) ; Moist tropical forest on the upper reaches of Río Tupisa (in and around village of Barranquillita ). [Coordinates on original label: 09º00’N, 78º00’W], 08°23’N 077°23’W , 09 February 1985 , S. Kane 33 ( MO !) ; Trail from Canglón-Yaviza road to Río Chucaraque , 7.7 miles E of Canglón , 08°20’08”N 077°48’10”W , 50 m , 06 March 1982 , S. Knapp & J. Mallet 3945 ( MO !) ; 112 miles from Bayano Dam Bridge . vicinity of Canglon , approximately one mile beyond concrete bridge in Canglón toward Yauisa, 08°19’30”N 077°50’00”W , 50 ft , 14 May 1980 , T. Antonio 4551 ( MO !) ; Pinogana. Along headwaters of Río Tuquesa, ca. 2 km air distance from the Continental Divide , in vicinity of upper gold mining camp of Tyler Kittredge , 08°33’30”N 077°29’00”W , 600 m , 25 August 1974 , T.B. Croat 27119 ( MO !) ; Los Santos : Above Guanico River ; forest on hills west of river, 07°20’N 080°30’W , 550–650 m , 4 January 1989 , G. McPherson 13500 ( MO !) ; Panamá : Panamá & Comarca de San Blas , Valle de Madroño ; ca. 10 road miles north of La Margarita (by Chepo); in forest just South of and on continental divide along main trail to Cangandi, 09°19’N 079°08’W , 350–450 m , 21 February 1986 , B.E. Hammel & G. McPherson 14525 ( MO !) ; Serranía de Majé . At top of ridge at headwaters of the Río Ipetí Grande , 08°51’N 078°34’W , 100 m , 25 January 1984 – 26 January 1984 , H.W. Churchill & G. de Nevers 4365 ( MO !) ; Serranía de Majé. Trail along Río Ipetí Grande, below confluence with Río Aqua Fria and Charco Rico. Alt. 300 m. 8º55’N, 78º31’W, 08°55’N 078°31’W , 300 m , 23 Jan. 1984 , H.W. Churchill & G. de Nevers 4314 ( MO !) ; Altos de Cerro Azul . Bosque cercano a Comasa , 16 May 2013 , Orlando Ortiz & et al. 1320 ( MO !, PMA !) ; m, T.B. Croat 68681 ( MO !) ; 6.5 kms from Pan American Highway along road to Cerro Campana Elevation 700 m ; 8º41’N; 79º58’W, 08°41’N 079°58’W , 700 m , 13 February 1986 , W.S. Hoover 1317 ( CM , MO ) ; Capira . Cerro Campana, sendero desde el mirador a la cima, 08°41’N 079°55’W , 700–1000 m , 12 Jan 1995 , C. Galdames & C.E. Guerra 1889 ( MO !) ; Chepo . Area around Torti Arriba , 08°56’01”N 078°25’25”W , 50–200 m , 31 August 1977 , J.P. Folsom 5160 ( MO !) ; Along trail between Río Majé and Quebrada Brava , 09°06’21”N 078°45’36”W , 60 m , 04 May 1976 , T.B. Croat 34653 ( MO !) ; Panamá . Parque Nacional Soberanía , Camino de Cruces , 09°07’16”N 079°38’37”W , 150 m , 27 Apr 1999 , A. Florpan et al. 4247 ( MO !) ; 5 miles above Interamerican Highway on road to Cerro Azul , 09°07’37”N 079°23’00”W , 240 m , 26 July 1970 , T.B. Croat 11516 ( MO !) ; San Blas : Trail from dock to Mandinga Airport to village of Cangandí.[Coordinates on original label: 9º24’N, 79º24’W], 09°27’00”N 079°05’00”W - 09°27’30”N 079°06’00”W , 0–30 m , 10 December 1985 , G. de Nevers et al. 6445 ( MO !) ; Trail east of Cangandi-Mandinga airport road , 2–5 miles south of Mandinga airport, 09°25’N 079°05’W – 09°27’N 079°05’W , 27 October 1967 , J. Duke 14820 ( MO !) ; Puerto Obaldia . Along the coast NE of Puerto Obaldía towards Colombian border. Tropical dry forest, strong sea winds, 08°40’N 077°24’W , 0 m, 18 Apr 1982 , S. Knapp & J. Mallet 4689 ( MO !) ; Along the coast NE of Puerto Obaldía towards Colombian border . Tropical dry forest, strong sea winds, 08°40’N 077°24’W , 0 m, 18 Apr 1982 , S. Knapp & J. Mallet 4682 ( MO !) ; Veraguas : Montijo . Parque Nacional Coiba , Jicarón , cima de la isla, bosque maduro, 07°16’04”N 081°37’57”W , 390 m , 27 Jan 2004 , A. Ibáñez et al.2443AI ( MO !) ; Parque Nacional Coiba . SE de Isla de Jicarón, río Arriba, 07°15’23”N 081°47’50”W , 0–320 m , 26 Apr 1997 , C. Galdames et al. 3862 ( PMA !) ; Sona. Isla Canales de Tierra . En la parcela 1. Bosque maduro. 17NMU 3657, 07°44’53”N 081°34’26”W , 30 m , 26 Feb 2002 , A. Ibáñez & A. Camarena 1700 ( MO !) ; Bahia Honda . Isla Canales de Tierra , entre playa Naranjo y punta, 07°44’N 081°35’W , 02 July 2001 , S. Castroviejo et al. 16120 ( MO !) .