Passiflora suberosa L. subsp. litoralis (Kunth) K.Port.-Utl. ex M.A.M.Azevedo, Baumbratz & Gonc .-Estev., Phytotaxa 53: 47. 2012.

Porter-Utley, Kristen, 2014, A revision of Passiflora L. subgenus Decaloba (DC.) Rchb. supersection Cieca (Medik.) J. M. MacDougal & Feuillet (Passifloraceae), PhytoKeys 43, pp. 1-224 : 69-76

2 b. Passiflora suberosa L. subsp. litoralis (Kunth) K.Port.-Utl. ex M.A.M.Azevedo, Baumbratz & Gonc.-Estev., Phytotaxa 53: 47. 2012. Figs 27 -28

Passiflora litoralis Kunth, Nov. Gen. Sp. 2: 138. 1817. Type: Peru. Lima: [Pativilca], “Patibilca”, A. Humboldt & A. Bonpland s.n. (holotype: P [P00307301, photograph seen]; isotype: B, destroyed).

Passiflora limbata Ten., Index Seminum Horto Bot. Neapol. 10: 12. 1839. Type: Cultivated in Naples Botanical Garden, Italy (holotype: NAP [photograph seen]).

Passiflora pseudosuberosa Fisch., Index Sem. (St. Petersburg) 9: 82. 1843. Type: Cultivated in St. Petersburg (Russia), originally from Brazil, Anon. s.n. (type material not seen, probably at LE).

Passiflora oliviformis Vell., Fl. Flumin. 9: pl. 83. 1831, as " olivaeformis ". nom. superfl., non Passiflora oliviformis Mill. 1768. Type: Brazil (no specimens extant; lectotype, designated here: Vellozo, Fl. Flumin. 9: pl. 83. 1831.)

Passiflora globosa Vell., Fl. Flumin. 9: pl. 85. 1831. Type: Brazil (no specimens extant; lectotype, designated here: Vellozo, Fl. Flumin. 9: pl. 85. 1831.)

Passiflora flexuosa Gardn. London J. Bot. 1: 174. 1842. Type: Based on Passiflora oliviformis Vell.

Cieca oliviformis (Vell.) M.Roem., Fam. Nat. Syn. Mon. 2: 144. 1846. Type: Based on Passiflora olivifornis Vell.

Cieca globosa (Vell.) M.Roem., Fam. Nat. Syn. Monogr. 2: 144. 1846. Type: Based on Passiflora globosa Vell.

Cieca litoralis (Kunth) M.Roem., Fam. Nat. Syn. Mon. 2: 145. 1846, as " littoralis ". Type: Based on Passiflora litoralis Kunth

Cieca pseudosuberosa (Fisch.) M.Roem., Fam. Nat. Syn. Monogr. 2: 146. 1846. Type: Based on Passiflora pseudosuberosa Fisch.

Cieca limbata (Ten.) M.Roem., Fam. Nat. Syn. Monogr. 2: 148. 1846. Type: Based on Passiflora limbata Ten.

Cieca flexuosa (Gardn.) M.Roem., Fam. Nat. Syn. Monogr. 2: 148. 1846. Type: Based on Passiflora flexuosa Gardn.

Passiflora suberosa var. divaricata Griseb., Bonplandia (Hanover) 6 (1): 7. 1858. Type: Panamá, E. Duchassaing s.n. (holotype: GOET [photocopy seen] [GOET009402]).

Passiflora suberosa subvar. argentea Mast., Trans. Linn. Soc. London 27: 630. 1871. Lectotype (designated by Hemsley 1888, pg. 480): Mexico. Puebla: Tehuacán, H. Galeotti 3663 (lectotype: K!; isolectotype: G!, [photocopies (2)], BR! [BR0000006943400]).

Passiflora suberosa var. longiloba Triana & Planch., Ann. Sci. Nat., Bot. 17: 157. 1873. Type: Colombia. “Tocayma”, J. Goudot s.n. (holotype: P [photograph seen]).

Passiflora suberosa var. longipes S. Watson. Proc. Amer. Acad. Arts 25: 149. 1890. Type: Mexico. Jalisco: in barranca near Guadalajara, 26 Sep 1889, C. G. Pringle 2966 (holotype: GH! [GH00065788] [photographs, AAU!, DUKE!, F!]).

Type.

Based on Passiflora litoralis Kunth

Description.

Sparsely to densely pubescent with unicellular or multicellular curved trichomes on leaf, petiole, stipule, stem and sepal 0.16-1.13 mm long, 0.02-0.03 mm wide, also minutely antrorsely appressed-puberulent on leaf, petiole, stem, stipule and sepal with unicellular, curved trichomes, 0.05-0.10 mm long, 0.02-0.03 mm wide. Laminas not peltate. Flowers borne in leaf axils or sometimes in indeterminate axillary or terminal inflorescences; inflorescences 2.0-4.0 cm long, associated reduced laminas 2.0-4.3 mm long, 1.5-3.1 mm wide. Pedicels 2.3-17.9 mm long, 0.2-0.7 mm wide, 2 per node; bract(s) absent or rarely with 1 or 2 narrowly ovate-triangular bracts present at (0.23-)0.42-0.88 of the distance from the base toward the apex of the pedicel, 0.4-1.1(-2.3) mm long, 0.1-0.2 mm wide, acute. Flowers 12.3-26.1 mm in diameter with stipe 0.2-7.5(10.1) mm long; hypanthium (3.0-)4.0-6.3(-7.1) mm in diameter; sepals 4.0-9.0(-10.8) mm long, 2.0-5.5 mm wide, abaxially and adaxially greenish yellow to very light greenish yellow (5GY 7/4, 8/4-8/2); coronal filaments in 2 series (very rarely 1 series), the outer 2.5-7.5 mm long, linear, sometimes capitellate, erect (ca. 70°) or slightly spreading (ca. 110°) or spreading (ca. 180°-220°), greenish yellow with yellow tips (5Y 8/10) or flushed with reddish purple (5RP 5/6-3/6) at base and greenish yellow at middle with yellow tips, ratio of outer coronal row to sepal length 0.39-0.95, the inner (1-)20-50(-53), erect to spreading slightly, greenish yellow or greenish yellow with yellow tips or greenish yellow flushed with reddish purple at base and yellow toward tips; operculum 0.7-2.3 mm long, greenish yellow or greenish yellow with a flush of reddish purple at base or reddish purple, margin white; nectary 0.1-0.9 mm high, 0.3-1.3 mm wide; limen recurved, erect or slightly inclined toward the operculum, greenish yellow or greenish yellow with reddish purple tip, limen floor greenish yellow or greenish yellow flushed with reddish purple; androgynophore 2.1-4.4(-6.1) mm long, 0.3-1.3 mm wide, greenish yellow or greenish yellow with a flush of reddish purple at base or greenish yellow with reddish purple spots and streaks; free portions of the staminal filaments 1.6-3.9 mm long, anthers with yellow pollen; styles 1.7-4.7(-5.8) mm long including stigmas. Berry 7.9-11.9 mm long, depressed globose to globose to ellipsoid. Seeds ca. 8-34.

Phenology.

Flowering and fruiting throughout the year.

Distribution.

In the New World tropics: Central America, Mexico, Argentina, Bolivia, Brazil, Colombia, Ecuador, Paragual, Peru, and Venezuela. Introduced in the Old World tropics: Africa, Asia, Australia, and the Hawaiian Islands. Growing in shrubs, trees or trailing on the ground in secondary successional areas, along the edges of semideciduous to deciduous, dry to moist tropical forests, both inland and near the seashore, 0-2800 m.

Discussion.

Passiflora suberosa subsp. litoralis has the widest geographic range of any species in supersection Cieca . In the New World, its range extends from northern Mexico, through Central America, to central Argentina. In these areas it may be confused with Passiflora pallida and Passiflora obtusifolia , which are sometimes similar vegetatively. The similarities and differences between these two species are discussed under their respective descriptions. The primary difference between Passiflora pallida and Passiflora suberosa subsp. litoralis is the hypanthium diameter, with that of Passiflora pallida rarely exceeding a width of 4.0 mm and that of Passiflora suberosa subsp. litoralis commonly 4.0 mm or wider. One of the more useful characters employed in separating Passiflora obtusifolia and Passiflora suberosa subsp. litoralis is the presence/absence of inflorescences. When mature, Passiflora obtusifolia bears flowers in long inflorescences (i.e., 5.3-18.3 cm) and Passiflora suberosa subsp. litoralis almost always lacks inflorescences; when Passiflora suberosa subsp. litoralis does possess inflorescences they are not as long (i.e., 2.0-5.0 cm).

There are three major morphological variants of Passiflora suberosa subsp. litoralis . In Mexico and Central America, Passiflora suberosa subsp. litoralis possesses shallowly trilobed leaves (commonly less than half the distance from the leaf outline to the leaf base) with the length of the central lobe often greatly exceeding that of the lateral lobes and an angle between the lateral veins that is frequently between 40° and 80°. The lateral lobes are also oblong to elliptic. The broadly capitate petiolar nectaries are commonly positioned on the distal half of the petiole, often over 0.60 the distance from the base to the apex of the petiole. The leaf bases are often cuneate to acute but rarely cordate. In Mexico and Central America, Passiflora suberosa subsp. litoralis is often found in high elevation (1000-3000 m) moist pine and oak forests along streams and rivers, but it may also occur in very dry forests with cacti (e.g., Cephalocereus forests of Tehuacán) and other species common in matorral vegetation (e.g., Tamaulipan matorral).

On the western side of South America (Colombia to Peru and Argentina), Passiflora suberosa subsp. litoralis possesses leaves very much like those of the Mexico/Central American variant, but the petiolar nectaries may be more discoid and are commonly positioned on the proximal half of the petiole. The lateral lobes are commonly distinctly ovate and diverge at an angle of 80-100°. The leaf bases are also distinctly cordate. In this region it is found in low (near sea level and on cliffs above the sea) to high (to 3000 m) elevation moist forests commonly along streams and rivers, but it also occurs in tropical dry forests.

On the eastern side of South America, in southeastern Brazil, the leaves are commonly trilobed but may also have unlobed, bilobed or trilobed leaves present on the same plant. The petiolar nectaries are commonly discoid and positioned on the proximal half of the petiole. The lateral lobes are ovate, but longer than those common in western South America, and commonly diverge at an angle of greater than 100°. The leaf bases are distinctly cordate. In Brazil, Passiflora suberosa subsp. litoralis is more common in coastal dunes and tropical dry forests, but it does occasionally occur in higher elevation moist forests as well. This variant is the only form of Passiflora suberosa subsp. litoralis found in the Old World. Laminar nectaries are commonly present in all three of the these variants.

In a recent manuscript, Milward-de-Azevedo et al. (2012) incorrectly designated Gardner 50 (BM) as a lectotype for Passiflora flexuosa . Gardner, in his manuscript (1842) was not publishing the new species, Passiflora flexuosa , but supplying a nomen novum for Passiflora oliviformis Vellozo. Therefore, the type of Gardner’s name is homotypic with Passiflora oliviformis Vellozo.

Selected specimens examined.

MEXICO. Chiapas: Barranca La Venta at Cascada El Aguacero, 16 km W of Oxozocuautla on Hwy. 190 and at the end of the road to the river, 650 m, Mayfield, Hemple & Jack 977 (MEXU). Chihuahua: Guasaremos, Río Mayo, Gentry 2910 (F, GH, MO). Durango: Sierra Madre, Rose 3504 (US). Guanajuato: 5 km W de Iramuco, sobre el camino a Santa Ana Maya, 1950 m, Rzedowski 44847 (CHAPA, XAL). Guerrero: Taxco, Lyonnel 303 (US); close to mirador over Taxco on Mexico 95, 1790 m, Porter-Utley & Mondragón 343 (CICY). Hidalgo: Trancas, 13 km NE de Zimapán, Mpio. de Zimapán, 2000 m, Hernández 3695 (MEXU, MO). Jalisco: Guadalajara, Las Trancas, camino a Mascuala, Mpio. Ixtlahuacán del Río, 1600 m, Guerrero & Chazaro 283 (MO, TEX). México: Cerro de los Capulines, Palmar Chico, 1100 m, Matuda 31332 (US). Michoacán: Zitacuaro-Salto de Enandio, 1600 m, Hinton 13492 (GH, LL, NA, NY, PH, TEX, US). Morelos: Xochitepec, Lyonnel 1425 (US). Nayarit: W slope of Volcán San Juan at km 6 on road from Tepic to Jalcocotan, Mpio. Tepic, 900 m, Breedlove & Almeda 45168 (CAS). Oaxaca: Mesa del Calvario, Cerro de el Ramón, NE de el Rodeo, Mpio. Tepelmeme, Dist. de Coixtlahuaca, 2100 m, Tenorio et al. 9262 (US). Puebla: Jardín Botánico Helia Bravoh, road between Tehuacán and Zapotitlán Salinas., 1450 m, Porter-Utley & Mondragón 344 (CICY); Tehuacán, Purpus 1272 (G, MO, UC). Querétaro: Jalpan, 800 m, Arguelles 909 (CAS); 4-5 km SE de Ayutla, Mpio. Arroyo Seco, 720-760 m, Carranza 2819 (IEB). San Luis Potosí: 6.5 (rd) mi. S of Arista, in Chihuahuan Desert, on limestone hills, 5000 ft., Henrickson 6428 (TEX); ca. 8 km NW de Guadalcázar, 2000 m, Rzedowski 6673 (XAL). Sinaloa: 60 m W de la casa Ramón Cabrera, ejido Cuitavaca, a 35 km de Agua Caliente de Zevada, Mpio. Sinaloa de Leyva, Perez 71 (CAS, UC). Sonora: Santa Ana de Yecora, 850 m, Van Devender et al. 98-1420 (FLAS, MO). Tamaulipas: Las Yucas, along the road which bears W-SW from Village of Las Yucas for 2.2 miles, Mayfield et al. 871 (TEX). Veracruz: Mpio. Emiliano Zapata, entre El Palmar & El Roble, Castillo & Tapis 764 (Froad from Nautla to Tlapacoyan, Porter-Utley & Mondragón 332 (CICY, FLAS);); road from Nautla to Tlapacoyan, Porter-Utley & Mondragón 333 (CICY, FLAS); Colonia Revolucion, Mpio. Boca del Río, 10 m, Ventura 5394 (GH. MO, XAL). Yucatan: road between Chabihau and San Crisanto, Porter-Utley & Mondragón 413 (CICY, FLAS); road between Chabihau and San Crisanto, Porter-Utley & Mondragón 414 (CICY).

COSTA RICA. Cartago: Cartago, near the Mirador Ujarrás about 4.5 mi. SE of Paraíso, 1200 m, MacDougal 906 (DUKE); vicinity of Cartago, Standley 33363 (US). Puntarenas: Canton de Buenos Aires Rey Curre, camino a Sabana Mamey, Rojas & Rojas 69 (MO). San José: San Francisco de Guadalupe, 1500 m, Pittier 7151 (BR).

EL SALVADOR. Ahuachapan: San Benito, al N del mirador El Cerrito, Sandoval & Roman1361 (MO).

GUATEMALA. Baja Verapaz: Mpio. San Jerónimo. km 137 carretera La Cumbre-Salamá, 1030 m, Tenorio et al. 14839 (MEXU). Chimaltenango: between Chiquimula and La Laguna, 500-1000 m, Steyermark 30703 (F). Guatemala: 20 km NW of Guatemala City, 5000 ft., Andrews 541 (NY). Hueheutenango: vicinity of San Sebastián, 1600 m, Molina & Molina 26507 (F, U); along Río Selegua, opposite San Sebastián H., 2000-2100 m, Steyermark 50460 (F). Jalapa: Laguna de Ayarza, 8000 ft., Heyde & Lux 3777 (GH, M, MO, NY, US). Petén: Santa Elena, Walker 1344 (MO). Sacatepéquez: slopes of Volcán de Agua, N of Santa María de Jesús, 1800-2100 m, Standley 59341 (F). Sololá: mountain slopes above Lake Atitlán, about 3-5 km W of Panajachel, 2100 m, Williams et al. 25309 (F).

HONDURAS. Comayagua: Los Alpes on cordillera Montecillos, road to El Cedral, Molina 23340 (F).

NICARAGUA. Chontales: along road from Juigalpa NE toward La Libertad, ca. 17.4 km NE of Río Mayales, at ford of Río Bizcocho, 350-400 m, Stevens & Krudoff 4162-b (MO). Esteli: Loma Ocotecalzado (Mesas Moropotente), ca 11 km NE of Hwy. 1 at Estelí, 1260-1300 m, Stevens et al. 15609 (MO). Jinotega: along trail between Jinotega and Las Mesitas, W of Jinotega, 1100-1400 m, Standley 9717 (F). Madriz: lado E cerro Volcán Somoto ( Volcán Tepe Somoto), 1300 m, Moreno 2949 (MO). Matagalpa: along road between San Simón de Palcila and Mesa La Cruz, 1150-1220 m, Stevens et al. 18533 (G, HUA, MO). Nueva Segovia: La Tronquera, 660-700 m, Moreno 19449 (MO).

PANAMÁ. Unknown Province: Grisebach s.n. (GOET). See specimen listed under cultivated material..

ARGENTINA. Catamarca: Sierra de Ancasti (Falda E), entre El Alto & el dique de Coyogasta, 950 m, Hunziker & Cocucci 17284 (F, NY). Chaco: Fontana, Meyer 2634 (F). Corrientes: Dep. Mburucuyá, 15 km NO de Mburucuyá, camino a Descabezado, Krapovickas & Mroginski 22226 (G, MO). Entre Ríos: Dep. Diamante, Punta Gorda, Bacigalupo & Deginani 16 (HUA). Jujuy: Dep. Santa Bárbara, NE of Libertador, ca. 20 km S of Palma Sola on the road to El Sauzal, 850 m, Taylor et al. 11489 (MO). Misiones: Dep. Cainguás, Ruta Prov. 7, camino de Aristóbulo del Valle a Jardín América, 4 km de del Valle, 270 m, Morrone et al. 629 (MO). Salta: Dep. Capital, 20 Km E de Salta, Ruta Salta a Gral. Güemes, Krapovickas & Schinini 30408 (C, F, G, US). Tucumán: Dep. Alberdi/Cocha, along Ruta 9 from Juan B. Alberdi to Balcosna, 7 km below of dique Escaba, 670 m, Till 10248 (MO).

BOLIVIA. Chuquisaca: Prov. Tomina, Monteagudo 64 km hacia Sucre, 1400 m, Beck 6350 (MO). Cochabamba: Prov. Campero, a 26 km de Aiquile rumbo a Peña Colorada, 2240 m, Saravia 522 (MO). La Paz: Viciniis Lorata, San Pedro, Larecaja, 2550 m, Mandon 612 (BM, G, GH, NY, S). Santa Cruz: Prov. Cordillera, Camiri, 900 m, Cardenas 4707 (US); Prov. Andrés Ibáñez, Jardín Botánico de Santa Cruz, 12 km E of center of Santa Cruz on road to Cotoca, 375 m, Nee 40425 (NY). Tarija: Prov. Cercado, Bañado del Paray, 450 m, Steinbach 13066 (F).

BRAZIL. Bahia: Mun. de Victoria da Conquista, 4.7 km south of center of city of Victoria da Conquista, along highway, Eiten & Eiten 10892 (US). Ceará: Maranguape, Serra de Maranguape, Trinta et al. 1280 (R). Distrito Federal: Santa Teresa, unknown collector s.n., 1888 (R). Goiás: Goyaz, S le figau central de la province, Glaziou 21461 (G). Mato Grosso Do Sul: Assentamento Tamarineiro, Mpio. Corumbá, Pott 1812 (NY). Minas Gerais: Lema de Caldas, Henschen & Regnelli III640 (MO, S). Paraná: Agua Branch (Mpio. Adrianópolis), 250 m, Hatschbach & Silva 51299 (C, MO, US). Pernambuco: Tapera, Pickel 465 (R). Rio De Janeiro: Climita Boa Vista, Rio de Janeiro, Brade & Freire 24384 (R). Rio Grande Do Sul: Estrada Oscar Marcelino Cardoso próximo a Faz. Renato Johan, Banhado Grande, Viamão, Abruzzi 814 (F). Santa Catarina: Mpio. Florianópolis, Canavieras, Ilha de Santa Catarina, 1-5 m, Smith & Reitz 12264 (NY, R, US). São Paulo: Botucatu, Rubião Júnior, Branzer 703501 (U).

COLOMBIA. Antioquia: Mpio. de Liborina, km 4 of road Liborina-Sabanalarga (32 km before Sabanalarga), 920 m, Zarucchi et al. 7248 (HUA, MO). Caldas: al lado de la carretera entre La Felisa & Manizales, 1400 m, Escobar & Brand 2059 (MA). Cauca: Cordillera Central, Vertiente Oriental, Mpio. de Inzá, Parque Arqueológico de San Andrés, 1700-2000 m, Idrobo & Weber 1368 (US). Cundinamarca: Mpio. Apulo (Rafael Reyes), Vereda El Portillo, 9 km de Viotá, 455 m, Escobar et al. 3002 (HUA). Magdalena: Sierra Nevada de Santa Marta, SE slopes, hoya del Río Donachui, below the village Donachui near the river, 1350-1230 m, Cuatrecasas & Romero 24402 (US). Norte De Santander: between Chinácota and La Esmeralda, 1000-1300 m, Killip & Smith 20887 (GH, US). Quindío: Mpio. Pijao, carretera a Caicedonia, 2 km antes de Barragán, Fca. Las Acacías, 1140 m, Arbelaez et al. 970 (HUA). Santander: N slope of Mesa de los Santos, 1000-1500 m, Killip & Smith 15029 (GH, NY, US); between Surata and California, Killip & Smith 16834 (GH, NY, US). Tolima: Doima, 700 m, Haught 2436 (US). Valle De Cauca: Cordillera Occidental, vertiente occidental, Hoya del Río San Quinini, 1200 m, Cuatrecasas 15371 (F).

ECUADOR. Chimborazo: cañon of the Río Chanchán near Huigra, 4000-4500ft., Camp 2945 (F, NY, US). Esmeraldas: Atacames, near Esmeraldas, Barclay 764 (BM). Galápagos: Santa Cruz, near the Caseta, 800ft., van der Werff 1420 (CAS, NY, U). Guayas: on the property of Richard Zeller near the village of Loma Alta, located about 10 km NE of the coastal village of Valdivia, N of Santa Elena peninsula, Río Valdivia drainage, 100 m, Anderson 2480 (MO); Guayaquil, Cerro Santa Ana, Asplund 15217 (AAU, B, NY, R, S). LA Chorrera: on Cerro Tiandeagote, 800 m, Jativa & Epling 13 (UC). LOJA: Bosque Petrificado Puyango, quebrada Cochurco, 350 m, Cornejo, Cornejo & Bonitaz 4034 (MO). Los Ríos: Hacienda Clementina between Babahoyo and Montalvo, 20 m, Sparre 17922 (S). Manabi: P.N. Machalilla, Agua Blanca, hasta cerro Las Goteras, 380 m, Josse 688 (AAU).

PARAGUAY. Caaguazú: Tavaí, 1 km S of Hospital, Zardini 7744 (MO, TEX); Ruta 2, km 98, Zardini & Aguayo 10551 (MO). Concepción: Arroyo Tagatiya-Misión, Zardini & Tilleria 38858 (MO). Guaira: Mbocayaty-Melgarejo, 2 km E of Mbocayaty on Arroyo Gerbasia Gallery Forest, Zardini & Tilleria 32349 (MO). Neembucu: Humaitá, Schulz 7770 (F). Paraguarí: Parque Nacional Ybycuí, Hahn 1951 (HUA, MO).

PERÚ. Amazonas: Prov. Chachapoyas. 9 km below and W of Chachapoyas on the road to Caclic, km 501, 2000 m, Hutchison & Bennett 4514 (F, NY, UC, US). Ancash: Prov. Huacho, Sayán, road to Acobamba, 700-900 m, Weigend & Dostert 97/116 (F, MSB). Arequipa: Prov. Caravelí, Lomas de Atiquipa (= km 591 Pananerica Sur), 150-750 m, Weigend & Forther 97/922 (MSB). Cajamarca: Prov. Contumazá, road Contumazá to Chilete, 2500 m, Weigend et al. 97/456 (MSB). Callao: Callao & Loma, Didrichsen 4398 (C). Cusco: Prov. La Convención, 139 km de Cusco en Quellomayo, subiendo hacia la “ceja”, entre Santa Teresa & Chaullay, 1200-2600 m, Nunez & Motocanchi 8782 (MO). La Libertad: Prov. Cajabamba, road from Cajabamba to Cajamarca, 8 km from Cajabamba, 2300 m, Weigand et al. 97/318 (MSB). Lima: Prov. Lima, Loma de Amancae, S of Lima near Pachacamac, 120-410 m, Gentry 16475 (AAU, MO, NY). Piura: Prov. Huancabamba, Procedencia, Porculla, km 38, 1800 m, Quiroz 2349 (TEX). Tumbes: Prov. Zarumilla, Matapalo, El Cauco-Campo Verde parcela de evaluación permanente “E”, 700 m, Díaz et al. 7426 (MO).

VENEZUELA. Aragua: Edos. Aragua & Miranda, bosque nublado de Loma de Hierro, 1350 m, Colella & Morales 731 (VEN). Bolívar: in valley on road from El Valle to La Miranda, Pittier 11970 (G, NY, US). Distrito Federal: on the old road from Caracas to La Guayra, between Bell Vista & Sanchorquiz, 1300-1450 m, Pittier 9567 (GH, NY, US, VEN). Falcón: Cerro Santa Ana, ascensión del lado S desde el pueblo de Santa Ana, 600-700 m, Steyermark & Braun 94621 (F, NY, US). Lara: al S de Quibor hacia Cubiro, Dist. Menez, 800 m, Steyermark et al. 110061 (NY, VEN). Mérida: between Sabana Grande and Baruta, 1000 m, Alston 5457 (BM); Colinas de Carrizal, 20 km de Caracas, Morillo 2820 (NY). Maracay: SW del Valle de Caracas, Colinas de Bello Monto, Edo. Miranda, 1100 m, Ramírez & Lopez 3273 (VEN). Monagas: Ladera S de Cerro San Bonifacio, en Bella Vista, 4 kms arriba del empalme con carretera Caripe-Teresén, 750-900 m, Bunting 2645 (GH). Sucre: Los Cocoteros (Via Casanay), Dist. Ribero, Cumana & Ceequea 4229 (WIS). Tachira: Sierra El Casadero, along hwy. between Las Dantas and Las Adjuntas, 850 m, Steyermark et al. 120174 (MO, NY).

SOUTH AFRICA. Kwazulu-Natal: Colony, District Alesandra, Station Dumisa, 400 m, Rudatis 1225 (BM); Denison Residence, Rutemaritburg, Weigend 2190 (M).

UGANDA. Kampala: Kyadondo, Mengo, Kyambogo, 1200 m, Rwaburindore 1735 (MO, US).

INDIA. Uttar Pradesh: Mothranwala, Dehra Døn, Parker s.n., 27 November 1927 (UC).

SRI LANKA. Uva: road between Bandarawela and Haputale, just below Kahagalla tea factory, Koyama et al. 16035 (AAU).

INDONESIA. Java: Cibodas, Nitta 15054 (MO).

SINGAPORE. Nassim Road, Togashi 6211611 (AAU).

TAIWAN. Chia-I: vicinity of Lianyun waterfall along the Tsengwen Hsi river, 300 m, Bartholomew & Boufford 6176 (US). P’ing-Tung: Kenting National Park, Oluanpi park, 5 m, Lammers 8488 (MO, US).

AUSTRALIA. Northern Territory: Nightcliff, Darwin, Arnhem Land Aboriginal Reserve, Specht 160 (US).

FIJI. Makondronga: Makondronga Island, 60 m, Degener & Orndonez 13801 (NY). NGAU: Shore of Herald Bay, in vicinity of Sawaieke, 0-30 m, Smith 7923 (NY, UC, US). OVALAU: E of Lovoni Valley, 100-300 m, Smith 7288 (NY, UC, US).

FRENCH OVERSEAS TERRITORY. New Caledonia: Au pied de l’Ouen Toro pres Nouméa, Baumann 6059 (UC).

FRENCH POLYNESIA. Society Islands: Tahiti, Papeete, Crete est de la Tipaerui, sentier du Mt. Marau, 1200 m, Florence 9735 (US).

SAMOA. Motootua, Upolu, Whistler W5368 (BM).

UNITED STATES. Hawaii: Honolulu Co.: O’ahu, on steep ridges above 'Aiea, Iltis H-610 (US).

SPAIN. Canarias: Isle of La Palma. Santa Cruz, Hausen 66 (C).

CULTIVATED MATERIAL. United States: Florida, grown in University of Florida Greenhouse (Gainesville, Alachua Co., Florida) from material collected in 1998 on an island off the coast of Panama, Porter-Utley P-64 (FLAS); Florida, grown in University of Florida Greenhouse (Gainesville, Alachua Co., Florida) from material collected in Sonora Mexico by T.R. Van Devender et al. (Van Devender et al. 98-1420), Porter-Utley P-58 (FLAS).