Vachellia tortuosa (Linnaeus) Seigler & Ebinger, Phytologia 87: 168. 2005.syn. Acacia tortuosa (Linnaeus) Willdenow. Sp. pl. 4: 1083. 1806.
Synonymy and types
Basionym: Mimosa tortuosa L., Syst. Nat., 10th ed., 2: 1312. 1759. Acacia tortuosa (L.) Willd. Sp. Pl. 4: 1083. 1806. Poponax tortuosa (L.) Raf., Sylva tellur. 118. 1838. - TYPE: JAMAICA. without exact locality, Patrick Brown(e) s.n. (holotype: LINN, microfiche).
Acacia virescens DC., Cat. pl. horti monsp. 74. 1813. - TYPE: probably South American, the species was described from a plant grown in the Botanical Garden in Montpellier. The seeds for this plant were obtained from the Botanical Garden in Madrid. (holotype: G-DC, microfiche).
Acacia leucacantha Bertero, Spreng., Syst. veg. 3: 144. 1826. - TYPE: JAMAICA. C. G. Bertero s.n. [lectotype, designated by Ebinger et al. (2000): MO].
Acacia seifriziana León in León and Alain. Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 9: 8. 1950. - TYPE: CUBA. ORIENTE: low forest near Maisí, Jul 1938, León and Seifriz 18351 (holotype: HAC). NOTE: We have not been able to examine the type, however, based on the description, it appears referable to Acacia tortuosa, a conclusion also advanced by Bässler (1998).
Mimosa salinarum Rohr ex Benth., Trans. Linn. Soc. London 30: 501. 1875. - TYPE: none cited.
Shrub or small tree to 6 m tall. Bark dark gray to dark brown, shallowly furrowed. Twigs dark purplish brown to dark gray, strongly flexuous, usually pubescent. Short shoots commonly present above the stipular spines, to 10 mm long, covered with acuminate stipules and old leaf bases. Leaves alternate, also commonly clustered on the short shoots, 12-45 mm long. Stipular spines light to dark brown to purplish, sometimes becoming light gray with age, symmetrical, terete, straight, stout, to 40 (55) x 1.5 mm near the base, densely pubescent at least toward the base, glabrous. Petiole adaxially grooved, 3-7 mm long, usually densely pubescent with erect hairs; petiolar gland solitary, located medially to near the base of the lowermost pinna pair, sessile, elongated to circular, 0.3-2.0 mm long, apex depressed, glabrous. Rachis adaxially grooved, 5-40 mm long, usually densely pubescent with long, straight hairs, a sessile, circular gland, 0.3-0.4 mm across between the upper 1 to 2 pinna pairs. Pinnae 2 to 8 pairs per leaf, 12-23 mm long, 3.2-7.5 mm between pinna pairs. Petiolules 0.5-1.4 mm long. Leaflets 11 to 19 pairs per pinna, opposite, 0.8-1.7 between leaflets, oblong, (2.5) 3.0-5.5 x 0.9-1.5 mm, glabrous or nearly so, lateral veins more or less evident, only one vein from the base, base oblique, margins strongly ciliate, apex broadly acute to obtuse. Inflorescence a densely flowered globose head, 6-9 mm across, solitary to clusters of 2 to 5 on the short shoots. Peduncles 12-36 x 0.4-0.7 mm, lightly puberulent and usually with minute, red, deciduous glands. Involucre 4- to 6- lobed, located at the base of the globose head, glabrous to lightly puberulent, persistent. Floral bracts spatulate, 0.8-1.5 mm long, puberulent, deciduous. Flowers sessile, pale yellow; calyx 5-lobed, 1.1-1.8 mm long, the lobes pubescent; corolla 5-lobed, 1.5-2.4 mm long, the lobes pubescent; stamen filaments 3.5-4.5 mm long, distinct; ovary glabrous, on a stipe to 0.2 mm long. Legumes dark brown to black, straight to slightly curved, elliptical in cross section, constricted between the seeds, linear (65) 80-150 x 5-9 mm, coriaceous, strongly reticulately striate, pubescent, usually glandular with minute, reddish glands, indehiscent; stipe to 7 mm long; apex acuminate. Seeds uniseriate, no pulp, light brown, ellipsoid to ovoid, slightly flattened, 5.3-6.8 x 3.5-5.0 mm, smooth; pleurogram U-shaped to nearly oval, 2.5-4.0 mm across. Flowers intermittently throughout the year. Chromosome number: 2n = 26 (Atchison 1948).
Thickets, open pastures, successional fields, and on dry sites, from sea level to about 1100 m in extreme southern Florida and the Bahamas south through the West Indies to Colombia and western Venezuela (Clarke et al. 1989).
Vachellia tortuosa is a lowland species, occurring at or near sea level in southern Florida and the West Indies where it is the dominant and co-dominant of the woody vegetation in scrub thickets. Its presence in extreme southern Florida was probably the result of pre-Columbian introduction, as Ward (1968) found that this species was always associated with sites of human occupation. In South America it is also commonly a lowland species, but a few specimens were collected at elevations as high as 1100 m. Presently, this taxon is not known from Mexico or most of Central America.
Most of the variation in Vachellia tortuosa is related to the differences in leaves on short shoots, and the leaves originating from between the stipular spines on rapidly growing branches. The smaller leaves with fewer and shorter pinnae, and small, circular petiolar glands occur on the short shoots. More and longer pinnae, and a larger, elongated petiolar gland occur on the primary leaves. Also, young stems are commonly flexuous, but sometimes this is not very obvious on slow growing branches.
Vachellia tortuosa is closely related to V. farnesiana, and these taxa are sometimes considered conspecific. Both have short shoots just above the stipular spines, the bipinnately compound leaves generally have less than 7 pairs of pinnae, whereas the leaflets are relatively large (3-6 mm long) and closely spaced along the costa. Some traits, however, can be used for consistent separation, particularly mature fruits. In V. tortuosa the fruits are relatively long (80-150 mm) and narrow (5-9 mm), densely pubescent, and constricted between the seeds; whereas in V. farnesiana the fruits are shorter (30-90 mm), broader (10-18 mm), usually glabrous, inflated, and not constricted between the seeds. Without fruits consistent separation is still possible as in V. tortuosa the petioles, rachises, and spines are densely pubescent, the petioles, even on primary leaves, are less than 7 mm long, the leaflets are strongly ciliate, and the calyx and corolla lobes are densely puberulent. In V. farnesiana, in contrast, the plants are usually glabrous to only lightly pubescent, the petioles on the primary leaves are up to 12 mm long, the leaflets are not, or only slightly ciliate, and the calyx and corolla lobes are commonly glabrous. The petiolar glands of the smaller leaves of V. tortuosa, particularly those on the short shoots, are small and circular, similar to those in V. farnesiana. The leaves on rapidly growing branches, however, have petiolar glands that are much larger and usually elongated.
The only other species within its range with which Vachellia tortuosa could be confused is V. guanacastensis. However, the larger leaves with consistently 7-9 pinna pairs, and the pinnae with more than 20 leaflet pairs in V. guanacastensis, separates this species from V. tortuosa. In the past, V. tortuosa has also been confused with V. bravoensis (Turner 1959, Isely 1969). The circular petiolar gland between the lower pinna pair in V. schaffneri consistently separates this species from V. tortuosa.
Clarke et al. (1989) found that nearly all specimens (95%) of this taxon from Central and North America were weakly cyanogenic. Of the 75 specimens tested from South America almost all (96%) gave a positive reaction, but the reaction was weak and slow to develop. In some cases, living material of this species gives a rapid, positive reaction for cyanide.
Intermittently throughout the year.
- Chokoloskee, dry shell island, 5 Jun 1937, L.Arnold & E.West s.n. (FLAS);
- Chokoloskee Island, 11 Feb 1982, D.S.Correll & J.Popenoe 53232 (FTG);
- Fergusin River shell mounds, 28 Jun 1960, F.C.Craighead s.n. (FLAS);
- Chokoloskee, west of # 29, 31 Aug 1966, O.Lakela & F.Almeda 30151 (FLAS, FSU, GH, MINN);
- Chokoloskee Island, 4 mi. S of Everglades City, 25 Nov 1966, D.B.Ward 6021 (FLAS, FSU, GH, US);
- E of Homestead, pineland glade, 22 Mar 1965, O.Lakela 28372 (FSU);
- Everglades National Park, Bear Lake Road, 0.1 mi. W of entrance to old main Flamingo-Coot Bay Road, 25 Apr 1958, A.Traverse 642 (GH, TEX);
- Ramrod Key, 12 Jan 1952, E.P.Killip 41734 (US).
WEST INDIES AND BAHAMAS:
- Piccadilly, on flat lands near the coast, 2 Aug 1937, H.E.Box 945 (F);
- Langfords, 25 Apr 1937, H.E.Box 749 (F);
- Royal Bay, 16 Jul 1958, J.D.Sauer 2083 (WIS);
- English Harbour, 4 Apr 1956, A.C.Smith 10433 (US);
- Jun 1938, F.A.Squire 28 (A).
- beach at Bushiri Beach Resort, Oranjestad, 7 Aug 1993, J.E.Ebinger 25843 (EIU);
- beach at Bushiri Beach Resort, Oranjestad, 7 Aug 1993, J.E.Ebinger 25844 (EIU);
- beach at Bushiri Beach Resort, Oranjestad, 7 Aug 1993, J.E.Ebinger 25845 (EIU);
- beach at Bushiri Beach Resort, Oranjestad, 7 Aug 1993, J.E.Ebinger 25860 (EIU);
- at north shore near light house, 9 Aug 1993, J.E.Ebinger 25891 (EIU);
- north shore near Guadiri Kiri, 9 Aug 1993, J.E.Ebinger 25895 (EIU);
- NE of Saint Nicolaas, 21 Nov 1947, P.Jones 1423 (US).
- long grass savanna, 16 Oct 1944, J.S.Beard 375 (A).
- near Willemstad, 20-27 Mar 1913, N.L.Britton & J.A.Shafer 2956 (NY);
- "Waabi" Seroe pretoe, 16 Feb 1917, H.M.Curran & M.Haman 17 (A);
- "Waabi", 17 Feb 1917, H.M.Curran & M.Haman 32 (GH);
- sea level, 29 Mar 1927, E.P.Killip & A.C.Smith 21048 (A, GH);
- Jan Thiel, 22 Jan 1980, J.F.Morton s.n. (FTG, NY);
- near Weis Afo., 15 May 1969, W.P.Nagelkerken 161 (A, WIS);
- 5 miles S. of Otrabanda, 18 Aug 1935, D.Potter 5111 (F);
- Parea, suburb of Willemstad, 17 Aug 1958, J.D.Sauer 2283 (WIS);
- Groot Knip, West Punt, 3 Dec 1950, A.T.Semple 332 (US).
- Las Matas de Farfán, 8 Aug 1978, A.H.Liogier 9029-1 (NY);
- En manigua, Hatillo Palma Montecristi, alt. 50-100 m, 8 Jun 1976, A.Liogier & P.Liogier 24613 (NY);
- 5-8 km E de Boca de Cachón, alt. 0-10 m, 29 May 1987, T.Zanoni, R.García & J.Salazar 39565 (FLAS);
- 1 km N of Jimani on road to Boca de Cachón, 16 Jul 1988, H.D.Clarke & R. García 35 (EIU, ILL);
- 1 km N of Jimani on road to Boca de Cachón, 16 Jul 1988, H.D.Clarke & R. García 36 (EIU, ILL);
- 1 km W of Boca de Cachón on road to Tierra Nueva, 16 Jul 1988, H.D.Clarke R. García 38 (EIU, ILL);
- 1 km W of Boca de Cachón on road to Tierra Nueva, 16 Jul 1988, H.D.Clarke & R. García 39 (EIU, ILL);
Prov. Monte Cristi:
- 3 km N of Villa Elisa on road to Agua de Palma, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 49 (EIU, ILL);
- 3 km N of Villa Elisa on road to Agua de Palma, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 50 (EIU, ILL);
- 4 km N of Villa Elisa on road to Agua de Palma, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 51 (EIU, ILL);
- 4 km N of Villa Elisa on road to Agua de Palma, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 52 (EIU, ILL);
- 4 km N of Villa Elisa on road to Agua de Palma, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 53 (EIU, ILL);
- 4 km N of Villa Elisa on road to Agua de Palma, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 54 (EIU, ILL);
- 3 km N of Monte Cristi on road to Isabel de Torres, alt. 100 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 55 (EIU, ILL);
- 4 km N of Monte Cristi on road to Isabel de Torres, alt. 70 m, 17 Jul 1988, H.D.Clarke & F.Jiménez 56 (EIU, ILL);
- S of Monte Cristi, 23-24 Oct 1946, R.A.Howard & E.S.Howard 9603 (US);
- en la Reserva Cientifica "Dr. Orlando Cruz Franco" 8 km N de Villa Elisa, alt. 120 m, 21 Aug 1985, J.Pimentel & R.García 511 (MO);
- 10 km NW of Santiago on Carretera Duarte, alt. 150 m, 17 Jul 1988, H.D.Clarke and F.Jiménez 45 (EIU, ILL);
- 10 km NW of Santiago on Carretera Duarte, alt. 150 m, 17 Jul 1988, H.D.Clarke and F.Jiménez 46 (EIU, ILL).
- Morne-a-Cabrits, 18 Jul 1924, E.L.Ekman 967 (A, F, LL).
- St. Catherine Parish, opposite Horse Cave, 23 Jun 1968, F.A.Barkley & R.Proctor 38474 (FTG);
- St. Andrew Parish, Port Royal, sea level, 19 Jul 1963, M.R.Crosby, H.A.Hespenheide & W.R.Anderson 794 (F);
- Clarendon Parish, near Springfield, 19 Mar 1984, F.W.Davis s.n. (UCSB);
- St. Catherine Parish, 12 miles from Kingston, 20 Feb 1970, W.T.Gillis 8917 (FTG);
- Delta of Hope River, 24 Sep 1906, W.Harris 9386 (F);
- Port Henderson, 15 Dec 1966, R.D.Henry J582 (WIS);
- St. Catherine Parish, road from Port Henderson to Fort Clarence, alt. 1-200 ft., 26 Aug 1965, H.A.Hespenheide, E.B.Hespenheide, J.S.Calver & R.E.Ricklefs 1605 (FLAS, MO, TEX);
- Port Royal, 18 Dec 1890, A.S.Hitchcock s.n. (MO);
- St. Catherine, between Green Bay and Fort Clarence, 11 Aug 1963, D.M.Porter 1030 (GH);
- 0.8 mi due northeast of Grants Pen., alt. 300-500 ft., 22 Jan 1979, G.R.Proctor 37973 (FTG);
- Lime Cay, 18 Oct 1985, D.R.Stoddart & S.M.Head 9040 (US);
- W of airport and lighthouse, E.West & L.E.Arnold 461 (GH);
- east end of Palisadoes, 4 Jul 1952, E.West, L.E.Arnold & C.James 408 (FLAS);
- west of airport and lighthouse, 5 Jul 1952, E.West, L.E.Arnold & C.James 461 (FLAS).
- low hills at Carr's Bay, 19 Mar-16 Apr 1979, R.A.Howard, E.S.Howard & P.G.Howard 18930 (NY, TEX, US);
- Brausby Point, 20 Jan 1907, J.A.Shafer 109 (CM, F, US).
- 1906, I.Boldingh 1908 (NY).
- Isla Palominos, 2 Jul 1966, E.L.Little,Jr. 21563 (NY, US);
- 1 mile N of Fajardo on hwy. 987, 25 Jul 1988, H.D.Clarke 78 (EIU, ILL);
- 1 mile N of Fajardo on hwy. 987, 25 Jul 1988, H.D.Clarke 79 (EIU, ILL);
- l.5 miles N of Fajardo on hwy. 987, 25 Jul 1988, H.D.Clarke 81 (EIU, ILL);
- 17 May 1978, J.D.Griffis 30-13 (NY);
- Salt Pond, Jun 1967, R.K.Wadsworth 69 (A).
- Pasia Hall Estate, 6 Oct 1918, W.C.Fishlock 200 (NY).
TURKS AND CAICOS ISLANDS:
- Salt Caly, 17 Jun 1979, D.W.Buden 39 (FTG);
- just SE of Black Salina, 22 Dec 1975, D.S.Correll 46642 (F, FTG, MO, NY);
- coppiced area on north edge of town, 9 Sep 1978, D.S.Correll 50127 (FTG);
- Salt Cay, in drainage area from ridge running NW of village, 11 Sep 1978, D.S.Correll 50168 (FTG, NY);
- between Commissioner's house and airfield, 22-27 Apr 1954, C.B.Lewis 22 (GH).
VIRGIN ISLANDS (U.S.):
- Cottan Grove, 17-25 Mar 1923, N.L.Britton, E.G.Britton & J.F.Kemp 53 (NY);
- Robin Bay, 6 Jan 1980, F.R.Fosberg 59137 (FTG, US);
- Estate South Gate, Christiansted, 25 Apr 1983, A.H.Liogier 34180 (NY);
- Buck Island Reef National Monument, 22 Jun 1966, E.L.Little,Jr. 21530 (NY, US);
- Buck Island Reef National Monument, 22 Jun 1966, E.L.Little,Jr. 21553 (US);
- Bassin, 18 Jan 1897, J.J.Ricksecker 34 (NY);
- roadside, 23 Jan 1896, A.E.Ricksecker 240 (F, GH, NY);
- East End Road after Cramer Park, 21 Aug 1963, D.Seligson 24 (A). St. John:
- East End Quarter, Center Line Road (Road 10), km 10.8, alt. 80 m, 28 Jan 1991, P.Acevedo Rdgz. & A.Siaca 4074 (F, US);
- Coral Bay Quarter, Road 107, Km 1.5, near John's Folly, 12 Feb 1993, P.Acevedo Rdgz. & A.Siaca 5241 (MO, US);
- Charlotte Amalie, 17-18 Jan 1899, C.F.Millspaugh 444 (F);
- Long Bay, Jan 1877, B.von Eggers s.n. (WIS).
- swamp between El Jagua Hunting Club on Río Jagua and El Congor Hill, alt. 2 m, 10 Feb 1935, A.A.Hunter & P.H.Allen 479 (MO).
- Barranguilla, Las Flores-La Playa, alt. 0-2 m, 13 Sep 1959, J.Custrecasa 24290 (F, US);
- fruiticetum arido litoral y playas maritimas arenosas entre Puerto Colombia y Salgar, alt. 1-3 m, 23 Jul 1943, A.Dugand & R.Jaramillo 3201 (US);
- alrededores de Galapa y Baranoa, 27 Jul 1943, A.Dugand & R.Jaramillo 3267 US);
- Puerto Colombia, Barranquilla and vicinity, Jan 1928, Bro. Elias 458 (NY, US);
- Barranquilla, Bro. Elias 1509 (F);
- Puerto Colombia, Jan 1932, Bro. Paul 911 (F, US);
- Barranquilla District, 18 Apr 1974, T.Plowman 3527 (US);
- Valle de Soatá, alt. 1800 m, 17 Sep 1938, J.Cuatrecasas 1960 (F, US);
- Municipio Barrancas, near campamento Tabaco, 24 Nov 1980, O.Arboleda, P.Bunch & J.V.Loh 319 (MO);
- near Maicao, savanna fields along the road to Carraipia, alt. 50 m, 29 Nov 1959, J.Cuatrecasas & R.Romero Castañeda 25489 (F, US);
- entre Manaure y Riohacha, 20 Oct 1963, C.Saravia T. 2918 (NY, US);
- 8.7 km de Uribia, rumbo a Maicao, 30 Mar 1962, C.Saravia & D.Johnson 345 (US);
- 10 km de Maicao, rumbo a Uribia, 31 Mar 1962, C.Saravia & D.Johnson 361 (US);
- Hacienda Miraflores, 15-20 km S of Santa Ana, alt. 4500 feet, 21 Feb 1944, E.L.Little, Jr. 7312 (US);
- Municipio Santa Marta, Parque Nacional Tayrona, Bahía Concha, Feb 1989, L.Atehortua 8 (EIU);
- Parque Nacional, Isla de Salamanca, Los Cocos, 29 Sep 1971, J.Daniel 132 (WIS);
- Santa Marta, thorn forest back of beach, 18 Aug 1944, F.R.Fosberg 22110 (NY, US);
- 49 km W of Puebloviejo, 30 Oct 1983, C.D.Johnson 3169 (MO);
- Codazzi, 7 Sep 1938, O.Haught 2322 (US);
- 0.5 km E of Aeropuerto Simon Bolívar and 14 km S of Santa Marta, alt. 20 m, 1 Nov 1972, J.H.Kirkbride, Jr. 2611 (US);
- Santa Marta region, 17 Jan 1930, S.J.Record 49 (F, GH, NY);
- 21 Mar 1948, R.Romero 963 (US);
- Isla de Salamanca, en Mahoma, entre la carretera y el mar, 10 Dec 1966, R.Romero-Castañeda 10496 (MO);
- sea level, Santa Marta, Sep 1898-1901, H.H.Smith 297 (CM, F, MO, NY, UC, WIS);
- Northern slope of Sierra Nevada de Santa Marta, Jul 1932, W.Seifriz 613 (US);
Norte de Santander:
- vicinity of Cucutá, alt. 215 m, 22-23 Mar 1927, E.P.Killip & A.C.Smith 21010 (NY);
State not known:
- road from Malambo to Sabanagrande, A.Dugand G. 32 (F).
- Tacagua, May 1953, L.Aristeguieta 1719 (US);
- Dept. Vargas, Parroquia Catia La Mar, 3 Apr 1986, A.Castillo 2015 (MO, NY);
- Municipio Vargas, Catia La Mar, 30 May 1987, A.Castillo 2460 (MO);
- Municipio Vargas, Catia La Mar, 28 Nov 1987, A.Castillo 2655 (MO);
- Municipio Vargas, Catia La Mar, 28 Jan 1988, A.Castillo 2791 (MO);
- Municipio Vargas, Catia La Mar, 16 Mar 1988, A.Castillo 2804 (MO);
- Cabo Blanco, 11 Jun 1917, H.M.Curran & M.Haman 951 (GH, NY, US);
- 10 km NE of Carayaca, alt. 420 m, 17 Dec 1973, G.Davidse 5100 (MO, NY), near Maiquetía, along road, 21 Apr 1922, H.Pittier 10294 (GH, US);
- vic. of Cabo Blanco, 23 Mar 1922, H.Pittier 10235 (US);
- Municipio Vargas, Parroquia Catia la Mar, 9 Jun 1987, N.Ramírez 2194 (MO, NY);
- Municipio Vargas, Parroquia Catia la Mar, 30 Jun 1987, N.Ramírez 2249 (MO, NY);
- Municipio Vargas, Parroquia Catia la Mar, 30 Jun 1987, N.Ramírez 2250 (MO, NY);
- Municipio Vargas, Parroquia Catia la Mar, 5 Apr 1988, N.Ramírez 2496 (MO);
- Municipio Vargas, Parroquia Catia la Mar, 23 Mar 1990, N.Ramírez 2704 (MO);
- Catia La Mar, alt. 20 m, 3 Jan 1940, L.Williams 12260 (F, US);
- Puerto Cabello, 10 Jul 1982, C.D.Johnson 2399-82 (MO);
- Puerto Cabello, 12 Jul 1984, C.D.Johnson 3344-84 (MO);
- Bahía Boca de las Piedras, Peninsula de Paraquana, 17 Oct 1945, D.S.Correll 12318 (GH);
- 10 km S of Guaibacoa, 18 Sep 1983, C.D.Johnson 2823 (MO);
- Distrito Silva, 4.7 kms N de Boca de Aroa, alt. 2-3 m, 1 Sep 1974, J.A.Steyermark & B.J.Manara 110532 (MO, NY);
- Dtto. Iribarren, donde se cruza con el camino de Agua Viva - Carretera Río Claro, 3 Oct 1981, C.Burandt, Jr. & D.Decker V0979 (F);
- 18 km E of Puente Torres, alt. 500 m, A.Gentry, G.Morillo & B. de Morillo 11007 (EIU, MO);
- 15 km W of Barquisimeto, alt. 530 m, 13 May 1982, C.E.Hughes 153 (MO);
- Barquisimeto, alt. 565 m., 16 Mar 1943, E.P.Killip 37073 (US);
- along Carretera 7, 16 km NE of Quibor, 1 May 1971, M.Nee & S.Mori 4097 (MO);
- road El Tocuyo-Humocaro Bajo, 6 Jan 1929, H.Pittier 13097 (A, MO, F, NY, US);
- near Humocaro Bajo, 6 Jan 1929, H.Pittier 13113 (A, MO, NY, US);
- 30 Km SW of Mérida along road to San Cristóbal, alt. 1100 m, 6 Nov 1963, F.J.Breteler 3261 (US);
- Brecenio Hacienda, 23 miles SE of Mérida, alt. 3740 ft., 6 Feb 1931, E.Reed 549 (US);
- Peracal, alt. 438 m, 23 Mar 1967, L.Cárdinas 240 (MO);
- Carretera frente a Lobatera, 10 Dec 1970, L.C. de Guevara 1251 (F);
- just S of La Mulata, alt. 300 m, 13 Nov 1979, J.A.Steyermark, R.Liesner & A.González 120233 (MO);
- Dtto. Mara, vía entre Carrasquero y Playa Bonita, en el trayecto entre la carretera Carrasquero-Mara-caibo y km 12 hacia Playa Bonita, alt. 0-50 m, 11 Jul 1981, G.S.Bunting 10038 (NY);
- Dtto. Mara, vía entre Carrasquero y Playa Bonita, alt. 0-50 m, 11 Jul 1981, G.S.Bunting 10043 (NY);
- a lo largo de una vía con rumbo al norte, entre km 13 de la carretera Maracaibo-Los Puertos de Altagracia, alt. 50-100 m, 19 Oct 1982, G.S.Bunting, N.Galué, E.Nucette & O.Huber 11846 (NY);
- Region Goajira, Carretera San Rafael del Moján, 1 km de Puerto Guerrero, Mpio. San Rafael, Distrito Mara, 3 Jun 1989, O.Zambrano, D.Pacheco & G.Sthormes 1920 (MO);
State not known:
- Tovar, 1854-5, A.Fendler 348 (GH, MO);
- La Vela de Coro, Hubada, 4 Apr 1917, H.M.Curran & M.Haman 501 (GH, NY, US);
- (on one label: between La Villa and El Puente (US);
- La Vela de Coro, 1 Apr 1917, H.M.Curran & M.Haman 432 (GH, US).