Sapindus saponaria subsp. saponaria

Franck, Alan R., 2024, Revision of Sapindus sect. Sapindus (Sapindeae, Sapindoideae, Sapindaceae), including the description of three new species, Phytotaxa 648 (1), pp. 1-71 : 36-39

publication ID

https://doi.org/ 10.11646/phytotaxa.648.1.1

persistent identifier

https://treatment.plazi.org/id/03C3F16E-0838-FFE5-22BE-97A4B6BFFB59

treatment provided by

Felipe

scientific name

Sapindus saponaria subsp. saponaria
status

 

8a. Sapindus saponaria subsp. saponaria View in CoL ( Figures 2 View FIGURE 2 and 13 View FIGURE 13 ).

Description:—Tree, to 13 m tall, bark surface texture finely roughened, sometimes becoming irregularly flaky, broken up into small scales, or fissured with the ridges broken up and blocky. Petiole 1–8(9) cm long, pale yellow to green, sometimes with brown speckling, glabrous to sparsely pubescent, unwinged or wing to 5 mm wide on one side; rachis unwinged or wing to 5 mm wide on one side, glabrous to sparsely pubescent; leaflets 4–12(14); petiolule 0–5 mm long, glabrous to moderately pubescent; leaflet blade lanceolate, ovate, elliptic, to obovate, strongly asymmetric to more or less symmetric, straight to falcate, the apex obtuse to acuminate, 3–12(32) cm long, 1.5–5(7.5) cm wide, 1.8–3.5(5) times longer than wide, adaxially drying pale green, green, pale brownish green, glabrous or sparsely pubescent along the midrib, the midrib pale green to pale yellow, glabrous to densely pubescent, the ridge 0.1–0.3 mm wide, 0.1–0.3 mm high, occasionally nearly flat or sunken, the blade not sunken or scarcely sunken along the midrib, secondary veins pale green to pale yellow, discolorous to nearly concolorous with the blade surface, tertiary venation prominulous, lightly discolorous to nearly concolorous with the blade surface, quaternary venation prominulous to obscure, nearly concolorous with the blade surface, abaxially drying pale green, glabrous to densely pubescent or short-pilose, midrib pale yellow, glabrous to densely pubescent, secondary veins pale yellow, secondary veins pale green to pale yellow, discolorous to nearly concolorous with the blade surface, tertiary venation prominulous, nearly concolorous with the blade surface, quaternary venation prominulous to obscure, nearly concolorous with the blade surface; exudates clear, inconspicuous to conspicuous abaxially on the blade surface. Petal without appendages (the base sometimes truncate to subsagittate with inrolled corners). Mature mericarp subglobose, 14–23 mm wide, seed 11–15 mm wide.

Distribution, habitat, and phenology:—Caribbean Islands, Mexico, Central America, South America, and USA (Florida) ( Fig. 3 View FIGURE 3 ); 0–2130 m, usually at lower elevations. Flowering (August) September–March (April, July) in the northern hemisphere, (January–) March–September (–December) in the southern hemisphere. Dry to humid, lowland to premontane, mostly open forests (Gentry 1942, MacBride 1956, Croat 1977, Wiggins 1980, Ferrucci 1991, Robbins 2001, Acevedo-Rodríguez 2005, Morales 2015), also locally cultivated and in some places probably naturalized due to local cultivation ( Standley & Steyermark 1949, Allen 1956, Parker 2008).

Notes:— Sapindus saponaria subsp. saponaria exhibits appreciable morphological variation, especially in the leaf rachis being winged to unwinged, the leaflets sessile to short-petiolulate, the blades straight and symmetric to falcate and asymmetric, and abaxially glabrate to densely soft-pubescent. Regionally, a particular morphology can be more common.

In Mexico, unwinged rachises with falcate, asymmetric leaflets tend to be more common than those with winged rachises and more symmetric leaflets. Native populations in Florida, USA nearly all have the leaf rachis conspicuously winged and the leaflets rather straight, symmetric, and abaxially uniformly sparse to densely pubescent. Much rarer in Florida are specimens with the rachis distinctly unwinged, the leaflets more falcate and asymmetric, and abaxially glabrate .

Plants of Central America generally all have unwinged rachises and falcate, asymmetric, abaxially glabrate to pubescent leaflets. In Brazil, plants typically have the rachis unwinged with falcate, asymmetric, abaxially pubescent leaflets although occasionally glabrate; less frequent are those with winged rachises with mostly straight and symmetric leaflets. In Africa , nearly all specimens have the rachis unwinged, leaflets falcate and asymmetric, and abaxially moderately pubescent. Only a few specimens from Cape Verde and Liberia have the rachis winged, the leaflets rather straight and symmetric, and abaxially pubescent. In Peru, one specimen (Cook & Gilbert 1446) has a surprisingly glabrous inflorescence rachis and glabrous pedicels with glabrate flowers, but otherwise the specimen appears consistent with S. saponaria subsp. saponaria .

Radlkofer (1932b) placed the specimens with mostly winged leaf rachises and mostly sessile, symmetric, ellipticlanceolate leaflets under forma genuinus , nom. inval. (= S. saponaria f. saponaria ). Those with primarily unwinged leaf rachises and more asymmetric, petiolulate narrowly ovate-lanceolate leaflets were placed in forma inaequalis by Radlkofer. Based on the examination here, there is considerable overlap between the two forms and it is not practical to recognize them. The lectotype illustration of S. saponaria cannot be clearly associated with either form, but it seems nearest to forma inaequalis , the lectotype having a narrowly winged rachis, asymmetric to nearly symmetric, narrowly lanceolate, and mostly petiolulate leaflets.

In the Caribbean Islands, Central America, Mexico, and Florida ( USA), S. saponaria subsp. saponaria nearly exclusively flowers September–March (Gentry 1942, Buswell 1945, Allen 1956, Croat 1977), very rarely in August or April. In South America, the flowering time is more inconsistent.

The petals lack appendages, as reflected in several illustrations ( Radlkofer 1932b: Fig. 14 View FIGURE 14 , Wiggins & Porter 1971, Fouilloy & Hallé 1973, Croat 1977, Reitz 1980, Wiggins 1980, Ferrucci 1991). A few detailed anatomical studies with illustrations of various tissues have been published ( Rock 1972, Albiero et al. 2001). Properties of the wood from Florida specimens were given by Sargent (1885).

Sapindus saponaria subsp. saponaria might be a relatively recent colonizer to Florida, since the last sea level highstand ca. 120,000 years ago left much of South Florida underwater where the taxon now occurs. Although documented by several specimens on Marco Island (Collier County, Florida, USA), S. saponaria is now thought to be extirpated from that island’s flora ( Wilder & Roche 2009) as well as from Key West (Monroe County, Florida) (Small & Mosier 5999). One cultivated collection from Manatee County, Florida ( White s.n.) has densely pubescent leaves with unwinged rachises, quite unlike any wild populations in Florida; this cultivated occurrence is more like some specimens from Mexico and Central America that are similarly densely pubescent with unwinged rachises.

Sapindus saponaria subsp. saponaria has been widely used for soap and ornament throughout its range. From one of the earliest written references on Sapindus from Hispaniola in the 1500s, Gonzalo Fernández de Oviedo y Valdés recorded its use as a soap (de los Rios 1851: 347, as “qüentas de xabon”, Radlkofer 1900). Humboldt ( MacGillivray 1833) reported the taxon being used to wash clothes outside of Cariaco, Venezuela during his visit in 1799. This particular subspecies may have been depicted by several early works that were probably using material from the Americas ( Clusius 1605: 42 [fructu 16], Bauhin 1650: 312, Piso 1648: 113, Tournefort 1700: 659 & tab. 440, Hill 1751: 408 & pl. 9, Lamarck 1819, 1823: pl. 307).

The ss variety of S. drummondii described by Hooker & Arnott (1838) is based on a Beechey specimen of S. saponaria subsp. saponaria that is in flower. Given its frequency along coastal southwestern Mexico and its flowering period, quite possibly the Beechey specimen was collected somewhere along his routes from San Blas (December 1827), Mazatlan (February 1828), to Acapulco (April 1828) ( Hooker & Arnott 1838).

Sapindus saponaria subsp. saponaria probably arrived in Africa during the colonial era ( Alpern 2008), and it has been cultivated there in many countries, e.g. Cape Verde, Congo, Liberia, Tanzania, Zimbabwe (specimens cited below; Davies & Verdcourt 1998). Material originating from the West Indies was said to be cultivated in Mauritius in the early 1800s ( Farquhar 1822). Specimens from the Pitcairn Islands were introduced from a single seedling imported from Valparaiso, Chile by George H. Nobbs in the 1853 ( Göthesson 1997, Kingston 2001, Erskine 2004). In 1881, it was introduced to Sri Lanka ( Trimen 1888). It was listed in cultivation in Bogor, Indonesia with an origin of “Amer. trop.” ( Teijsmann & Binnendijk 1866). Rock (1913) stated that American S. saponaria had been introduced to Hawaii, “growing about town [Honolulu]”. The subspecies was also cultivated in Tahiti ( Grant 4360), and possibly it is cultivated in the Cook Islands ( Sykes 2016). It was listed as a transplant in the Bahamas by Schoepf in 1784 ( Morrison 1911: 307), although otherwise Sapindus has never been documented in the Bahamas.

Sapindus saponaria subsp. saponaria has been targeted for eradication as a non-native on Santiago Island, Galápagos ( Gardener et al. 2010), though it is otherwise considered native to Isabela and said to be the largest forest tree of the Galápagos ( Stewart 1911). A forest fire impacted the stands of Sapindus on Isabela ( Gara et al. 1987, Nowak & Lawesson 1988).

Sapindus lippoldii from Cuba is treated as a synonym, following Acevedo-Rodríguez (2014), although it was an accepted species in POWO (2023). The original description was brief and did not explicitly compare it with S. saponaria , but the description of leaflets 10–35 cm long and 4–8 cm wide was likely thought to be a distinguishing feature ( Lippold 1974). Similarly large leaflets 20–32 cm long and up to 7.5 cm wide are found on several other specimens of S. saponaria subsp. saponaria from various other countries (e.g. Dugand & Jaramillo 2807, Felger 01-627, Hanson s.n., Spence s.n., Standley 66738, and Takeuchi s.n.). These larger leaflets appear to be confined to immature, sterile specimens of S. saponaria subsp. saponaria , as leaflets of mature stems are normally not larger than 15 cm long.

Leptostylus gibbulosus ( Coleoptera ) feeds on the seeds of S. saponaria subsp. saponaria in Colombia ( Hernandez-Jaramillo et al. 2012). In the USA, a tree in Miami-Dade County was nominated as a Florida state champion tree, measuring 12.8 m tall with a circumference of 1.5 m ( FDACS 2023). A chromosome count of 2 n =28 was reported for several collections of S. saponaria subsp. saponaria (Ferruci & Solis Neffa 1997, Rice et al. 2015).

Conservation Status:—This subspecies is very widespread with a large extent of occurrence. It should be categorized as Least Concern ( IUCN 2012).

Specimens examined:—Sloane herb., H.S. 97: 127 ex Plukenet (BM); Sloane herb., H.S. 101: 122 ex Plukenet (BM); Hort.Cliffortianus(BM);LINN 514.2 (LINN); “Accepi Londini”, 1738, Ens s.n. (LE). ANTIGUA &BARBUDA. Antigua: road to St. Philips, 13 February 1913, Rose et al. 3444 (NY). ARGENTINA. Formosa: E de Puesto Porteño, 3 March 1947, Morel 2436 (NCSC). BELIZE. Orange Walk District: Orange Walk, [ca. 1926], Winzerling s.n. ( US). Cayo: Branch Mouth, below El Cayo, 19 February 1931, Bartlett 11959 (MICH); Little Cocquericot, 25 April 1933, Lundell 4140 (MICH); Rio Grande river, 24 February 1933, Schipp 1122 (MICH); El Cayo, Mar-Jun, Chanek 162 (MICH); Cayo Distr., Chaa Creek, 8 September 1993, Arvigo et al. 788 (MO). BOLIVIA. Franz Tamayo: Parque Nacional Madidi, rio Tuichi, 8 December 2005, Araujo-M. et al. 2777 (MO). La Paz: S. Yungas, San Bartolome, 1-22 July 1939, Krukoff 10265 (MICH). Santa Cruz: Nuflo de Chavez, vicinity of Perserverancia, banks of Rio Negro, 14° 45’ S, 62° 45’ W, 250 m, 15 September 1990, Lewis 37668 (EAP); El Cerrito, 61 km S of Concepción, 21 February 1995, Abbott 16329 (FLAS); San Ramón, 23 April 1995, Abbott 16674 (FLAS); Santa Cruz de la Sierra, 27 July 2002, Justiniano & Putz 144 (FLAS). Sud Yungas: km 21.5 from Chulumani toward La Asunta, 4 August 1991, Acevedo Rdgz. et al. 4444 ( US); Chicaloma, 6 February 1996, Beck 22564 ( US). BRAZIL. Villa, 8 February 1879, Lorentz s.n. (VT). Acre: upper Rio Jurupary, 15 July 1933, Krukoff 5215 (MICH); basin of Rio Purus, territory of Acre, near mouth of Rio Macauhan, 21 August 1933, Krukoff 5578 (MICH); Do Acre, Rio Branco Municip., estrada para Sena Madureira km 40, 23 October 1980, Cid & Souza 3014 (MICH). Ceará: Jati, VPR Ipê, 7° 41’ 57.98” S, 38° 59’ 51.86” W, 433 m, 30 October 2014, Fontana & Silva 8745 (RB). Espirito Santo: Cachoeiro de Itapemirim, 27 April 1972, Sucre & Soderstron 9002 (RB). Maranhão: 35 km from Bacabal, 1 October 1980, Daly et al. D397 (MICH). Mato Grosso: Serra das Araras, Fazenda Currupira , 30 March 1996, Dubs 2153 (Z). Mato Grosso do Sul: Faz Paraná e Faz Bacuri, 19 June 1998, do L.G. Salvador et al. 60 (NY). Minas Gerais: Carangola, Chácara dos Fragas, perímetro urbano, Bairro Santa Emília, May 1989, Leoni 782 (RB). Paraíba: Sousa, Margem do Rio do Peixe, 6° 45’ 5.09” S, 38° 20’ 58.26” W, 245 m, 17 July 2014, Pimentel & Fontana 132 (RB). CHILE. Tarapacá: Azapa, east of Arica, 15 March 1952, Hutchison 476 ( US). BURUNDI. Bujumbura Mairie: Route Busumbura-Bugarama, March 1950, Germain 6659 (PNH); Bujumbrua, 10 February 1967, Lewalle 1585 [cultivated] (MO). CAPE VERDE. Ilha de Santiago, “ 10/12/1955 ”, Grandvaux Barbosa 5917 [cultivated] (MO). COLOMBIA. Antioquia: Mpio. Frontino, corregimiento Nutibara, cuenca alta del Río Cuevas, 19 July 1987, Sánchez et al. 1508 (COL); Frontino Municip., 14.5 km O de Nutibara, 12 July 1988, Callejas et al. 6636 (MICH); Municip. Medellín - Antioquia, ciudad Universitaria, 18 November 1994, Fonnegra & Roldán P. 5259 ( US); ibid., 11 January 1995, Roldán & Fonnegra 2263 ( US). Atlántico: entre Baranoa 7 Usiacurí, 50-100 m, 13 January 1941, Dugand & Jaramillo 2807 (COL). Bolívar: Torrecilla, Killip & Smith 14663 (F); Cartagena, Isla Tierrabomba, 4 September 1989, Cuadros V. 4374 ( US). Boyacá: Soatá, 6 September 1938, Cuatrecasas 1071 ( US). Huila: Algeciras (San Juanito), 38 km S of Neiva, 3 December 1942, Fosberg 19237 (MICH, US). Norte de Santander: región de Sarare, La Cubuya, 24 October 1941, Cuatrecasas et al. 12591 (COL). Valle: Plana del Valle del Cauca, entre Cabuyal y La Solorza, 27 May 1943, Cuatrecasas 14463 ( US); Las Juntas del Dagua, and Guaca Valley, [s.d.], Lehmann 7733 ( US); carreterra from Buga to Darien, past Mediacanoa, 17 May 1982, Murphy 399 ( US); Río Cauca Valley , 17 December 1987, Gentry et al. 59542 (MO); Sevilla, Paila Arriba, 17 May 1979, Cuadros V. 728 ( US). COSTA RICA. Guanacaste: Quebrada Azul, 29 January 1926, Standley 46448 ( US); Comelco Ranch, Bagaces, 12 December 1970, Hartshorn 957 (F). Puntarenas: Palmar Norte de Osa, 100 ft., 27 March 1950, Allen 5515 (EAP); ibid., 0 m, 23 December 1950, Allen 5722 (EAP); near Palmar Norte, 30 m, 10 December 1951, Allen 6319 (EAP); Puntarenas Osa, Palmar Norte, 27 October 2001, Aguilar 6637 ( US). CUBA. Guantánamo: Bayate, Cuato bank, 28 June 1915, Ekman 6167 (MICH). La Habana: Near Havana, 4 April 1905, Curtiss 707 (LY, MPU); near Havana, banks of Almendares River, November 1915, León 5751 (NY). Pinar del Rio: base of Sierra Guane, 26 November 1911, Shafer 10563 (MO). Santiago de Cuba: 29 November 1981, Elenevskaya s.n. (MW). DEMOCRATIC REPUBLIC OF CONGO. Équateur: Eala, 1930, Corbisier-Baland 1305 [cultivated] (MO); ibid., 1930, Corbisier-Baland 1416 [cultivated] (MO). Kongo Central: Buami-M’Vuazi (Terr. Thysville), 25 November 1957, Dubois 202 (MO); Forêt (vieux nzumbu) de Kolo (Vuazi-Territoire de Thysville), 26 November 1957, Dubois 203 (MA, MO); Nkokodia (Kikongo), Nzumbu de Kolo (M’Vuazi), “ 3/6/1968 ”, Dubois 309 [cultivated] (MO).Mongala: Sumba, Peco, Quimanete, July 1926, Gossweiler 9192A ( US). DOMINICA. [1780s], de Ponthieu (BM). St. Joseph: Macoucheri Estate, 7 January 1966, Chambers 2512 (CLEMS, P). DOMINICAN REPUBLIC. Barahona: 7 km SE de la Ciudad de Barahona, 3 December 1981, Mejía & Pimentel 18385 (FTG). La Altagracia: Bayahibe, 14 May 1984, Zanoni 30057 (USF). La Romana: Isla Catalina, 16 December 1986, Zanoni et al. 37181 (FLAS). Perdenales: E de Oviedo, 19 February 1990, Zanoni 44097 (FLAS). San Pedro de Macorís: Boca de Soco, 9 September 1983, McDaniel 27172 (FLAS). ECUADOR. Galápagos:Albemarle Island [Isabela], Villamil, 23 August 1905 –1906, Stewart 1947 (MO, US). Guayas: Balao, April 1892, Eggers 14621 (L); Naranjal, Reserva Ecológica Manglares - Churute, 15 August 1992, Cerón 20222 ( US); Guayaquil, Bosque Protector Cerro Blanco, 19 December 1995, Núñez & Yagual 221 ( US). Imbabura: between Carchi, Guajara and Guayupe, 5 December 1980, Holm-Nielsen & Jaramillo 28892 ( US). Loja: 4.7 km N of Catamayo on old road to Chuqiribamba, 10 July 1989, Dorr & Valdespino 6587 ( US); Tunnel N of Catamayo on old road to Chuqiribamba, 10 July 1989, Dorr & Valdespino 6589 ( US); S of Solanda, 14 February 1993, Harling & Ståhl 26379 (MO); cerca a Catamayo, 22 March 1994, Cornejo & Bonifaz 2170 ( US); Bosque Petrificado Puyango, Sur de Puyango, 30 May 1995, Cornejo & Cornejo 3920 ( US). Manabí: Comuna Agua Blanca, 1 May 1985, Bonifaz 1070 ( US). EL SALVADOR. Ahuachapán: San Francisco Menéndez, Vivienda de Enrique Castaneda, 750 m, 13° 49’ N, 89° 56’ W, 23 May 1997, Sandoval ES01597 (EAP). Chalatenango: 3 km al w de Citalá, 4 April 1997, Villacorta 2537 (F). La Libertad: Centro Nacional de Agronomía, Santa Tecla, 3000 ft., 28 September 1958, Allen 6962 (EAP); Mun.Antiguo Cuscatlan, Laeras de la Laguna, 24 November 1988, Curz s.n. (F). San Miguel: Laguna de Olomega, alt. 75 m, 20 February 1922, Standley 20995 ( US). San Salvador: San Salvador, November 1921, Calderón 251 ( US). Santa Ana: vicinity of Santa Ana, 30 January 1922, Standley 20445 ( US). Sonsonate: vicinity of Sonsonate, 220-300 m, 18-27 March 1922, Standley 22284 ( US); Finca San Ignacio, 16 km N of Chalchupa, 1 April 1958, Allen & Armour 6822 (EAP); carretera a Metapán, entre la Hda. El Milagro y la Hda. San Cayetano, 600 m, 25 October 1993, Linares & Martínez 851 (EAP). FRENCH GUIANA. Mana: Mana, January 1856, Sagot 953 (P). FRENCH POLYNESIA. Windward Islands: Tahiti, Mamao garden, 29 October 1930, Grant 4360 [cultivated] (A). GRENADA. St. George: St. Georges, December 1904, Broadway s.n. ( US). GUADELOUPE. Pointe Noire, 1895, Duss 3590 (NY). GUATEMALA. Below Chili, 9 November 1916, Popenoe 713 ( US). Alta Verapaz: along Río Sebol, between Sebol and Carrizal, 18 April 1942, Steyermark 45753 (F). Guatemala: Sanarate, 30 December 1906, Kellerman 6015 (F). Jutiapa: camino a Finca El Platanar,Asunción Mita, 14° 12’ 53” N, 89° 36’ 24” W, 538 m, December 2005, Morales 3378 (EAP). Peten: Uaxactun, 19 March 1931, Bartlett 12195 (F, MICH); Tikal National Park, Arroyo Corriental below Aguada Naranjal, 19 January 1961, Contreras 1837 (EAP); Tikal National Park, Tikal, 12 January 1962, Lundell 17051 (EAP, F); Tikal National Park, Aguada Corriental, 15 January 1964, Contreras 3843 (F); Laguna Petexbatun, 2 April 1964, Lundell 181971 (F); San Benito, 110 m, 14 October 1964, Meneres 56 (EAP); Paso Caballos, 50 m, 1965, Ortíz 104 (EAP); Tikal National Park, Tikal, 17 March 1966, Contreras 5637 (EAP); south of Paso Caballos, 6 April 1966, Contreras 5660 (EAP, F); Tikal, Parque Nacional, 8 January 1969, Tún Ortíz 26 (EAP, F); Santa Elena, 7 January 1970, Tún Ortíz 521 (F, MICH); El Rosario, 28 January 1970, Ramirez S. 21 (EAP); Santa Elena, en orillando el camino para San Francisco, 2 September 1970, Tún Ortíz 1340 (EAP, F, MICH); Santa Elena, en orillando el camino para La Libertad , 9 October 1971, Tún Ortíz 2000 (EAP, F); El Bucute, ca. 4 km de la aldea de Tikal, Parque Nacional de Tikal, 13 February 1972, Tún Ortíz 2280 (F, MICH); San Luis, 29 April 1976, Ventur 231 (F); San José Petén, Reserva campesina de Bioitza’ de San José, 31 May 1996, Ucan et al. 9292 (MICH). Quiché: 1942, Ignacio Aguilar 1423 (F). Retalhuleu: Retalhuleu, 27 February 1939, Standley 66738 (F); vicinity of Las Delicias, south Retalhuleu, 22 February 1941, Standley 88073 (F). San Marcos: Finca El Porvenir, on Potrero Matasán, Volcán Tajumulco, 12 March 1940, Steyermark 37595 (F). Suchitepéquez: south of Alotenango Farm, 7 mi. south of Tiquisate, 19 June 1942, Steyermark 47780 (F). Zacapa: vicinity of Zacapa, 7-16 October 1940, Standley 74151 (F). GUYANA. East Berbice-Corentyne: 20 mi. SE of New Amsterdam, 18 August 1955, Little, Jr. 16704 ( US). HAITI. Grand’Anse: 8 km este de Anse-a-Veau en la carretera costear a Miragoane, 26 January 1984, Zanoni et al. 28747 (NY). HAWAII. Kauai: Koloa District, Lawai Valley, 27 January 1994, Flynn 5506 [cultivated] (BISH). Oahu: Honolulu, Waikiki, 246 Kapili Street, 28 Oct 1939, Neal s.n. [cultivated] (BISH); Honolulu, 597 King Street, 3 December 1942, Price s.n. [cultivated] (BISH); Kamahameha School Native Garden, 29 December 1951, Bowers s.n. [cultivated] (BISH); Honolulu, Manoa Vallely, 23 August 1963, Hanson s.n. [cultivated] (BISH); Honolulu, Makiki District Park, 9 December 1985, Lau 1720 [cultivated] (BISH); Bishop Museum, July 1987, Takeuchi s.n. [cultivated] (BISH); Honolulu, Bishop Museum, 24 March 1995, Staples 1001 [cultivated] (BISH). HONDURAS. Choluteca: vicinity of Choluteca, 31 Oct-9 November 1949, Standley 24583 (F). Colón: Rio Selen, 7 mi. E of Trujillo, 3 February 1981, Saunders 1011 (F). Comayagua: El Banco, 13 March 1945, Valerio Rodriguez 2338 (F); Ojos de Agua, 8 January 1981, Nelson et al. 6978 (MO). Cortés: vicinity of La Lima, 11-20 April 1947, Standley & Chacón P. 7167 (F). El Paraiso: Lizapa, 19 December 1948, Williams & Molina R. 14906 (F). Francisco Morazán: Zamorano, 22 November 1946, Williams & Molina R. 10810 (F, MICH); vicinity of Zamorano, 26 November 1946 - 9 January 1947, Standley 1716 (F); El Zamorano, 6 December 1948, Standley 15417 (F); Tegucigalpa, 28 September 1950, Molina R. 3252 (F); drainage of the Rio Yeguare, 3 January 1955, Molina R. 5382 (F). Yoro: Maloa District, near Aguan River, 13 February 1927, Record & Kuylen H.51 ( US). JAMAICA. St. Catherine: Salt Island, 30 November 1962, Adams 11902 (MO); Salt Island Lagoon, 23 April 1964, Powell & Lewis 1607 (MICH). LIBERIA. Western: Kolahun Distr., Karmadhun, 6 November 1947, Baldwin, Jr. 10158 (MO). MARTINIQUE. Fort de France, 1883, Duss 614 (NY). MEXICO. Herb. Beechey (E, K); Plantae mexicanae Liebmann, 1841-1843, Liebmann 2998 (F). Baja California Sur: between Agua Caliente & Miraflores, 6 May 1931, Wiggins 5667 (MICH); 1.5 km al NNE del poblado Boca de la Sierra, 23.402 N, 109.818 W, 250 m, 7 March 2014, Domínguez Cadena 4166 (SD). Campeche: Tuxpeña, 12 December 1931, Lundell 1049 (F, MICH); ibid., 23 March 1932, Lundell 1406 (F, MICH); Campo Experimental Forestal Tropical El Tormento Km. 5 arr. Escárcega a Candelaria, 13 January 1966, Hernández X. et al. ES-755 (F); Punta Zacatal a Puerto Rico, Carmen, 23 March 1985, Chan 4967 (F); Mun. Tenabo, Km 42 carretera Campeche-Mérida, 21 April 1992, Simá & Campos-Rios 1351 (F). Chiapas: Escuintla, 26 November 1936, Matuda 602 (MICH); 1942-1943, Wonderly 61 (MICH); Mojarra, Tonala, 27 November 1947, Matuda 17194 (F); Santa Rosa, Tonala, 17–18 December 1947, Matuda 17338 (EAP, F); along the Tanate River at Habenal, paraje of Mahbenchauk, Mun. Tenejapa, 4 October 1965, Breedlove 12885 (F); 32 km N of Ocozocoautla along road to Mal Paso, 19 October 1965, Raven & Breedlove 13609 (F, MICH); paraje of Kulak’tik Mun. Tenejapa, 25 November 1965, Breedlove 14200 (F); Mpio. Yajalón, Cerro Azufre, 25 October 1982, Méndez Ton & Shilom Ton 4902 (F); Campamento Bajada del Macho, Mun. Ocozocoautla, 5 May 1983, Calzada 9999 (F); arriba de la E.T.A., mpio. Yajalón, 5 September 1983, Méndez Ton & Shilom Ton 6598 (F); 2 km al S de Bonampak, 27 September 1984, Martínez S. 7903 (MICH); 2 km al N de Tanhi Perla camino a Monte Líbano-Ocosingo, 11 April 1986, Martínez S. 17639 (F); 13 km NW of Tumbo, 17° 8’ N, 91° 43’ W, 610 m, 26 September 1988, Stevens & Martínez S. 25888 (BRIT). Colima: Los Reyes, Cuyutlán, 5 August 1972, Díaz Luna 3347 (MICH); Rancho El Jabali, 22 km NNW of Colima, 5 January 1991, Sanders et al. 10247 (MICH). Guerrero: 2 mi. S of Acahuizatla, 16 June 1954, Floyd & Ryan 137 (MICH); Lomas del Alambrado, camino de herradura Lidice-Las Moneras, Mpio. Teloloapan, 18 May 1982, Martínez S. et al. 638 (F); Mpio. Ocosingo, Ribera del Rio Lacantún, frente al Rancho Puerto Rico, 170 m, 22 May 1999, Sinaca C. 2596 (EAP). Jalisco: east of Barra de Navidad, 7 April 1951, McVaugh 11838 (MICH); near highway to Autlán, 4 mi. N of Bahía Navidad, 8 November 1960, McVaugh 20848 (MICH); Etzatlán, 15 January 1968, Moran 14707 (MICH); “Centro de Investigación y Experimentación de la UNAM” 8 km E of Chamela, 8-10 December 1970, McVaugh 25108 (MICH); Loc. Est. de Biol. “Chamela”, 12 January 1978, Magallanes 926 (MICH); Arroyo Chamela, E of Hwy. 200 km 63.5, 11 January 1983, Bullock 1292 (F); 6 km al O de Río Samaria, Mpio. Cárdenas, 6 March 1983, Ramamoorthy et al. 1749 [cultivated] (EAP). Mexico: Cañitas, Distr. Temascaltepec, Hinton 5023 (F). Michoacan: 40 km S of Arteaga and ca. 20 km N of Playa Azul, 25 February 1965, McVaugh 22542 (MICH). Morelos: Mpio. Puente de Ixtla, 4.4 km al oeste de Los Tanques, 18° 29’ 4.5” N, 99° 20’ 6.2” W, 1252 m, 25 March 2013, Ramírez et al. 2970 (MEXU). Nayarit: Mpio. Nayar, Arroyo de Los Negros, 4 March 1992, Arcadia-Alvarez 186 (MICH); Mpio. San Blas, 11 km al NE del Izote camino a Palapita, 23 January 1994, Flores-Franco et al. 3345 (MICH). Nuevo Leon: Hacienda Vista Hermosa, 27 June 1939, White 1605 (MICH); between Las Ajuntas & Vista Hermosa, 25 August 1939, Muller 2999 (MICH); El Cercado, 25 mi. S of Monterrey, 9 July 1953, Manning & Manning 53175 (FLAS). Oaxaca: near Ixtepec, 11 January 1945, Alexander 323 (MICH); vicinity of Mitla, 7 July 1971, Messer 104 (MICH); Las Animas, Ejido B. Juárez, 12 km carret. Tuxtepec, Valle Nal, 21 Oct 1959, Comisión de Dioscóreas 617 (MEXU); Mun. San Juan Huichico, Cuautemoc, 18 October 1975, Marquez & Yong 431 (F); Dominguillo, 11 April 1978, Smith F-63 (MICH); Distrito Juchitan, Mpio Chichihua, 16.5 km NE de Santiago Ixtaltepec, 17 January 1984, Tenorio L. 5216 (F); Mun. Del Barrio Dto. Juchitán, 3 km al O de Lagunas, 16° 49’ N, 95° 4’ W, 26 February 1988, Torres C. & Cortes A. 11820 (BRIT [ex NLU]); Mun. San Pedro Juchatengo, Dto. Sola de VEga, 4 km al N de San Pedro Juchatengo, 16° 22’ N, 97° 05’ W, 780 m, 26 February 1988, Campos V. 1528 (EAP); Dto. Juquila, mpio. San Pedro Juchatengo, 11 km al S de Ojo de Agua, 1000 m, 20 January 1999, Reyes-García & Gordon B019/05 (EAP); Distr. Tehuantepec, Mpio. San Pedro Huamelula, Rio Zimatán, Rancho la Hamaca de Petatengo, 15° 56’ 41” N, 96° 1’ 32” W, 150 m, 5 Mar 2000, Elorsa C. 3011 (MEXU). Puebla: near Coxcatlan on Cerro Ajuereado, Ju l1961, Smith, Jr. et al. 4102 (F). Queretaro: 1 km al suroeste de Tangojo, mpio. de Landa, 750 m, 11 October 1989, Rubio 1212 (MEXU). Quintana Roo: Cozumel, 18-19 February 1899, Millspaugh 1471 (F); sur del entronque a Mérida, entre Carrillo Puerto 7 Chetumal, 10 January 1980, Jéllez y Cabrera 1220 (MO). Revillagigedo Islands: Clarion Island, March 1889, Townsend s.n. ( US); Clarion Island, near the beach, 26 April 1925, Mason 1561 (F, MO, NY); Clarion Island, Sulphur Bay, 23 March 1932, Howell 8360 (F); Clarion Island, 19 February 2017, Rebman et al. 32138 (BRIT, SD). San Luis Potosi: 8 December 1891, Pringle 5093 (MICH); near los Canos, 15-21 October 1902, Palmer 225 (F); Rascon, 19–22 June 1905, Palmer 674 (F, MO); El Salto Falls, 5 mi. w El Mecca, 16 June 1951, Burdine & Blume 516 (BRIT); 3 km W de El Naranjo, municip. Cuidad de Maíz, 22 March 1956, Rzedowski 7362 (MICH); Mpio. Cd. Valles, Rancho Tinaja, 5.5 mi. S of Cd. Valles, 22 May 1981, Fryxell & Anderson 3552 (MICH). Sinaloa: Mazatlan, April 1910, Rose et al. 13844 (F); ibid., 5 April 1910, Rose et al. 14062 (F); ibid., March 1931, Gonzalez Ortega 6789 (F); Rio Fuerte, 21 November 1933, Gentry 914 (BRIT, MICH); near Alcoqonque, 11 December 1936, Shreve 7796 (F); along road to Eldorado, 34 mi. SW of Culiacán, 12 December 1957, Alava & Cook 1469 (MICH); Rio Humaya, Culiacan, 28 January 1962, Breedlove 1529 (MICH); 24.6 mi. N of Mazatlan on Mexico 15, 29 January 1962, Breedlove 1570 (MICH). Sonora: Alamos, 27 January 1899, Goldman 293 (F); Alamos, March 1910, Rose et al. 12990 (F); San Carlos Bay, 8 July 1921, Johnston 4346 (F); Quiricoba, Distr. Alamos, 12 November 1933, Gentry 728M (MICH); 15 mi. NW of Guayamas, 20 March 1934, Shreve 6555 (MICH); Caramechi, Rio Mayo, 3-10 December 1934, Gentry 1170 (F); Alamos, Rio Fuerte, 6 July 1936, Gentry 2273 (F); Nacapuli Canyon, 12 January 1982, Daniel 1967 (ASU, MICH); Alamos Mpio., upper crossing of Rio Cuchujaqui, 14 December 1988, Charlton et al. 2722a (MICH); Municipio de Guaymas, Cañon Las Barajitas, 28° 3’ 3.6” N, 111° 11’ 1.7” W, 90 m, 6 January 1996, Felger et al. 96-22 (BRIT); Cañon La Balandrona, N side of Sierra El Aguaje, 28° 4.8’ N, 111° 4.29’ W, 19 December 2001, Felger et al. 01-627 (MEXU). Tamaulipas: 19 mi. N of Aldama, 5 mi. S of San Rafael, 11 December 1959, Johnston 4952 (MICH); road to Tula, 5-6 mi. NW of Ocampo, 25 February 1961, King 3925 (F, MICH). Veracruz:Atoyac, December 1882, Kerber 197 (MICH); Island of Juan Ramirez, 8-9 March 1910, Palmer 452 (F); Laguna Tamiahua, 40 mi. S of Tampico, “3/5/39”, LeSueuer 271 (F); Ejido de Manzanares, 1-4 km NW of Campo Experimental de Hule, 1 February 1944, Vera Santos 2764 (MICH); near Rancho ojo de agua, 31 October 1948, Langman 3573 (EAP); Mpio. Tierra Blanca, Acahual, “ 10/12/1967 ”, Martinez-Calderon 1589 (F, MICH); Mun. Emiliano Zapata, Plan del Rio, 28 January 1972, Dorantes 486 (F); San Andrés Tuxtla, Laguna Encantada, 31 January 1972, Beaman 5575 (BRIT, F, MICH); Conejos, municipio Puente Nacional, 50 m, 23 November 1972, Ventura A. 7446 (BRIT, MICH, SD); La Mancha, municipio de Actopan, 10 m, 15 January 1973, Ventura A. 7725 (BRIT); Pachuquilla, municipio Puente Nacional, 150 m, 26 January 1973, Ventura A. 7747 (BRIT); Mata de Caña, municipio de Puente Nacional, 13 March 1973, 100 m, Ventura A. 7988 (EAP); 15 km al NE de Coatepec, caminos a Jaloomulco, “ 06-11-1973 ”, Cedillo T. 21682 (F); Isla de Agaltepec Lado SW, Catemaco, 25 April 1974, Ponce C. & Alvarez del C. 306 (F); Mun. Coatepec, 3 km E de Tuzamapa, 17 November 1974, Vazquez 2234 (F); La Ceiba, municipio de Puente Nacional, 100 m, 28 January 1976, Ventura A. 12359 (EAP); Colonia “Coronel A. Tejeda”, Mpio. de Catemaco, 28 Mar 1980, Barajas & Angeles 64 (MEXU); Mun. Actopan, Tramo Almolonga a Trapiche del Rosario, 28 April 1980, Calzada 5938 (F); Mun. Las Choapas, Rancho Gavilan, 17 May 1980, Calzada 6074 (F); N of Juan Rodríguez Clara, along Sayula-Tierra Blanca highway, Mun. Juan Rodríguez Clara, 6 December 1981, Nee 23793 (F); junction of road to Tenenexpan and old free hwy. Mex 140, 3 km SW of Santa Fe, 23 February 1984, Nee & Taylor 29671 (F); Mpio. San Andres Tuxtla, Estación de Biología Tropical Los Tuxtlas, 18° 34’ N, 95° 04’ W, 200 m, 9 November 1984, Ibarra Manriquez et al. 2106 (BRIT [ex NLU]); LOTE 67, Estación de Biología Tropical Los Tuxtlas, 18° 34’ 36” N, 95° 04’ 09” W, 200 m, 6 Nov 1985, Ibarra Manríquez 2713 (MEXU); ibid., 7 Nov 1985, Ibarra Manríquez 2744 (MEXU); Sierra de Tantima, ladera N. Mpio. Tantima, 21° 18’ 25” N, 97° 49’ 34” W, 500 m, 17 Mar 2001, Durán E. & Rincón G. 1272 (MEXU). Yucatán: [s.d.], Gaumer 396 (F); Calotmul, [s.d.], Gaumer 2339 (EAP, F, MICH); ibid., Gaumer 1737 (EAP, F, MICH); 1895, Millspaugh 44 (F); Izamal, 23 January 1895, Millspaugh 177 (F, MICH); Pocoboch, 1917-1921, Gaumer 24198 (F); Thien-Welden Dzibilchatun Survey, cenote Xlacah, 9 January 1979, Bradburn & Darwin 1183 (F); Tixcacaltuyub, Municipio Yaxcaba, 29 November 1980, Vargas R. 257 (F); Xul, junto a ruinas KOM, Oxkutzcab, “ 05/06/1984 ”, Sanabria & Simá 394 (F). NICARAGUA. Chontales: Patacón, vicinity of Juigalpa, 160 m, June 1947, Standley 9197 (EAP); Valley of Río Cuapa, 31 January 2008, Stevens et al. 26865 (MO). Granada: Granada, December 1869, Levy 287 (P). Managua: near Managua, [s.d.], Garnier 3001 (F, MICH); Managua, [s.d.], Garnier 3023 (F). PANAMA. Colón: near Madden Dam, along Azote Caballo Road near Alahuela, 27 November 1934, Dodge 16559 (MICH). Herrera: vicinity of Chitre, 26 November 1938, Allen 1098 (EAP). Panama: between Matías Hernández and Juan Díaz, 21 January 1924, Standley 31928 ( US); vicinity of Pacora, 5 November 1939, Allen 2030 (EAP, US); camino al Leprosorio de Palo Seco, 12 December 1975, Correa & Mendieta 2017 (EAP). PARAGUAY. Central: Aregua, 7 April 1981, Vavrek & de Molas 232 ( US). PERU. 1778-1788, Ruiz & Pavon 12/22 (MA). Cajamarca: cerca a Tamborillo, 13 July 1982, Ferreyra & Sánchez 19596 ( US); edges of Colasay, 11 July 1993, Shonle 137 (FLAS). Cusco: La Convención, Santa Ana, 23 June 1915, Cook & Gilbert 1446 ( US); Iza’ Cunyaco, March 1967, Vargas C. 17133 ( US); Distr. Santa Ana, Esmeralda, 21 June 2003, Calatayud et al. 1607 ( US); La Convención, Distr. Echarate, 20 August 2003, Suclli & Farfán 1199 ( US); La Convencion, Papelata, 11 May 2005, Calatayud et al. 2949 (FLAS); La Convención, Distr. Maranura, 29 November 2006, Valenzuela et al. 8260 ( US); La Convención, Distr. Echarate, Papelpata, 25 May 2007, Calatayud et al. 4145 ( US). Huánuco: Huanuco, 28 April 1923, Macbride 3536 ( US). Junín: San Tarma, La Merced, 27 June 1948, Ferreyra 3711 ( US). Lima: San Miguel, Urubamba Valley, 31 May 1915, Cook & Gilbert 1014 ( US); Yangas, 12 October 1951, Soukup 4232 ( US). SanMartin: San Martin Distr., Valley of San Martin, 15 August 1937, Belshaw 3243 (MICH). PITCAIRN ISLANDS. Pitcairn Island, 1931, Ross s.n. [cultivated] (AK); Pitcairn Island, in village, 14 June 1934, St. John 15021 [cultivated] (BISH); ibid., 15 June 1934, Fosberg 11331 [cultivated] ( US); ibid., 30 December 1934, Chapin 967 [cultivated] (A, BISH); Pitcairn Island, 1934, Ross s.n. [cultivated] (AK). PUERTO RICO. Coamo: vicinity of Coamo Springs, 12 February 1922, Britton et al. 6010 (NY). SAINT LUCIA. [1785–1811], Anderson s.n. (BM). SENEGAL. Dakar: Dakar, Hann, 22 April 1948, Adam 1118 (MO). Fatick: Fatick, Ndangane, 2 May 1948, Adam 1232 (MO). SEYCHELLES. Jardin Botanique de Mahé, January 1982, Friedmann 4043 (SEY). SRI LANKA. Central: Bot. Garden Peradeniya, 450 m, 18 October 1979, Kostermans 27882 (BR, L). SURINAME. Commewijne: Spieringshoek, 30 April 1916, Samuels 69 (P). TANZANIA. Kagera: Bukoba, Minziro Forest Reserve, Itara area, 28 April 2001, Festo et al. 1385 (BRIT, MO). Kigoma: Gombe National Park, south of Kasakela Valley, 19 March 1996, Mbago et al. 1524 (MO). Morogoro: Ilonga T.T.C. Kilosa, 26 August 1977, Mhoro 2553 [possibly cultivated] (MO). Mwanza.: Bukumbi Chiefdom, Kashishi Parish, 12 February 1953, Tanner 1214 (MICH, MO). Tanga: Muheza Distr., 27 May 2000, Mwangoka & Maingo 1322 (BRIT, MO). TRINIDAD & TOBAGO. Tobago: December 1909, Broadway 3389 (L). Trinidad: 18 April 1920, Britton 1982 ( US). U.S. VIRGIN ISLANDS. St. Croix: Bethlehem, 17–25 March 1923, Britton et al. 127 (NY). USA. Florida: South Florida, [1800s, presumed Oct-January as specimen is in flower], Chapman s.n. (A, USCH). Broward County: woods, Davie, 30 November 1931, Hammerstein s.n. (FLAS); Hollywood, 16 March 1933, Hammerstein s.n. [cultivated] (FLAS); nursery at Davie, 6 September 1962, Donn s.n. [cultivated] (FLAS). Collier County: Marco, S. Florida, [1800s], Chapman 41 ( US); Turner River, 2 mi. NE of Chokoloskee Island, 14 March 1933, O’Neill 7621 (A, NY); Whitney River, 20 February 1940, Scull s.n. (FLAS); Marco Island, Goodland Point, 16 April 1947, Brass 18082 (ARCH); ibid., 14 October 1957, Brass 29037 (ARCH); Marco Island, route 92 to shores of Caxambas Bay, 1 May 1965, Lakela 28514 (FSU, USF); Chokoloskee, 20August 1965, Lakela & Laker 29112 (USF); Marco Island, 2 May 1975, Killmar 6 (FTG); Choskolukee Island, 14 April 1977, Harriman 13540 (ASU); Daniel’s Point, Fakahatchee Strand State Park, 25 October 1997, Hattaway s.n. (SEL); ibid., 26 March 2003, Bishop & Alvarez FS0333 (USF); Fakahatchee Island, 25 September 2010, Wilder et al. 31656 (SWF); ibid., Wilder et al. 31737 (SWF); ibid., Wilder et al. 31739 (SWF); ibid., Wilder et al. 31743 (SWF); ibid., Wilder et al. 31744 (SWF). DeSoto County: Arcadia, 16 December 1932, Mowry & West s.n. [cultivated] (FLAS). Lee County: vicinity of Marco, 25 February 1916, Standley 12780 ( US); 15 April 1933, Baker s.n. (FLAS); Mound Key, 21 June 1929, Buswell s.n. (FTG); 13 January 1933, Buswell s.n. (FTG); Mound Key, 20 October 1969, Simmons s.n. (FTG); ibid., 11 August 1973, Todd 7 (FLAS); Pine Island, Randell Research Center, 17 September 2022, Majure et al. 9557 (FLAS). Miami-Dade County: Elliott Key, 16 April 1886, Sargent s.n. (A); between Miami and Coconut Grove, 26 Nov-20 December 1913, Small & Small 4798 (DUKE, FLAS, GH, TENN); Brickell Hammock, 1 May 1920, Rehder 836 (A); Miami, 25 April 1943, Buswell s.n. (FTG); Gumbo Limbo Trail, Royal Palm Hammock, 9 July 1962, Taylor 18 (NCU); Totten Key, 15 October 1962, Cooley et al. 9339 (FSU, GH, USF); Elliott Key, 16 May 1969, Andorfer 33 (USF). Chicken Key, 28 July 1987, Guala 734 (FTG); Simpson Park, 2 December 2008, Woodmansee s.n. (ASU). Monroe County: Key West, [February-March 1846], Rugel s.n. (GH); Cape Sable, June [1880s-1910s], Curtiss s.n. (A); Chatham Bend, 9 March 1905, Eaton 1317 (ECON); Pumpkin Key, April 1911, Simpson s.n. (A); Coot Bay, E of Cape Sable, November 1912, Simpson s.n. (A); Key West, 20 March 1915, Small & Mosier 5999 (MICH, NY); Key Largo, 1 April 1931, McFarlin 4674 (MICH); Key Largo, 10 February 1939, Scull s.n. (FLAS); ibid., 24 August 1946, Buswell s.n. (FTG); ibid., 18 March 1947, M uenscher & Thorne 18371 (GEO); ibid., 6 March 1952, Caldwell 8764 (DUKE, FLAS); North Key Largo, 21 July 1956, Brizicky & Stern 552 (FLAS); Flamingo, 19 October 1961, Craighead s.n. (FLAS); Bear Lake Road, 8 May 1965, Lakela et al. 28612 (USF); Flamingo, Everglades National Park, 29 December 1965, Wooten & Sullivan 429 (FSU) [flowering]; Pavilion Key, 20 August 1969, McJunkin 407 (FTG); Everglades National Park, along Bear Lake Trail, 30 May 1970, Gillis 9305 (FTG, GH, USF); Everglades National Park, Snake Bight Trail, 10 May 1975, Hill 2949 (FTG, NCU, VT); Key Largo, Dynamite Docks Road, 24 September 1996, Coile et al. 8625 (PIHG); Dagny Johnson Key Largo Hammock Botanical State Park, 12 November 2023, Franck & Ross 6196 (FLAS). Orange County: Orlando, 23 August 1929, Williams s.n. [cultivated] (FLAS). Pinellas County: St. Petersburg, 20 November 1940, Spence s.n. [cultivated] (FLAS). VENEZUELA. Falcon: Colina District, rio Macoruca, 12 February 1977, Steyermark & Gonzales 113767 (U); Municip. Unión, carretera Churuguara - Santa Cruz de Bucaral, 15 January 2006, Meier & Forbes 12748 ( US). Guarico: between El Sombrero & La Democracia, 4 February 1935, Archer 3031 ( US). Monagas: Guarapiche Forest Reserve, 0–100 m, 7 May 1966, Breteler 5146 (EAP). Trujillo: Parque Nacional Guaramacal, Fila de Agua Fria, Jan-February 1996, Stergios & Zambrano 16682 ( US). ZIMBABWE. Harare: Salisbury, 23 December 1969, Biegel 3232 [cultivated] (MO). Manicaland: Gordon Avenue, Umtali, 15 November 1967, Chase s.n. [cultivated] (MO).

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