Name

Vachellia caven (Molina) Seigler & Ebinger, Phytologia 87:  148.  2005.
syn.  Acacia caven (Molina) Molina, Sag. Stor. Nat. Chili, 2nd ed. 163.  299.  1810.

Synonymy and types

Basionym:  Mimosa caven Molina, Sag. stor. nat. Chili, ed 1, 174. 1782.  Mimosa cavenia Molina, Eassai hist. nat. Chili, 338.  1789.  Acacia caven (Molina) Molina, Sag. stor. nat. Chili, 2nd. ed., 163, 299. 1810.  Acacia cavenia (Molina) Hook. & Arn., Misc. Bot. 3: 206.  1833.  Acacia farnesiana (L.) Willd.  var. cavenia (Molina) Kuntze, Revis. gen. pl.  3(2): 47.  1898.  Acacia farnesiana (L.) Willd. var. cavenia (Hook. & Arn.) Arechav., Anales Mus. Nac. Montevideo  1: 436.  1901.  Vachellia farnesiana (L.) Wight & Arn. f. cavenia (Molina) Speg., Bol. Acad. Nac. Ci. Córdoba 26: 297.  1923.  Acacia farnesiana (L.) Willd. f. cavenia (Hook. & Arn.) E. C. Clos., Bol. Minist. Agric. Nac. (Argentina) 28: 455.  1930. - TYPE:  CHILE.  Raneagna, Oct 1828, C. G. Bertero s.n. [lectotype, designated by Aronson (1992):  SG0].  NOTE:  We interpret the specific epithet cavenia to be a misspelling.

Acacia adenopa Hook. & Arn., Bot. Misc.  3: 206.  1833. - TYPE:  Islands of the Uruguay, J. Tweedie s.n. (holotype:  K).

Acacia aromatica Poepp., Mart., Fl. Bras.  15: 395.  1879. - TYPE:  CHILE.  Valparaiso, E. Poeppig 177 [lectotype, designated by Ebinger et al. (2000):  W].

Acacia farnesiana (L.) Willd. var. brachicarpa Kuntze, Revis. gen. pl.  1: 156.  1891. - TYPE:  none cited, from Argentina, Córdoba.

Acacia farnesiana (L.) Willd. var.  heterocarpa Kuntze, Revis. gen. pl.  3(2): 47.  1898. - TYPE:  ARGENTINA.  CÓRDOBA:  Dec 1891, O. Kuntze s.n. [lectotype, designated by Ebinger et al. (2000):  MO].

Acacia caven (Molina) Molina var. dehiscens Burkart ex Ciald., Darwiniana  25: 76.  1984. - TYPE:  ARGENTINA.  CÓRDOBA:  Ascochinga, 22 Sep 1936, E. G. Nicora 962 (holotype:  SI).

Acacia caven (Molina) Molina var. microcarpa (Speg.) Burkart ex Ciald., Darwiniana  25: 77.  1984. - Vachellia farnesiana (L.) Wight & Arn. f. microcarpa Speg., Bol. Acad. Nac. Ci. Córdoba  26: 301.  1924. - TYPE:  ARGENTINA.  FORMOSA:  Depto. Patino, Fortin Soledad, A. Krapovickas 1283 [lectotype, designated by Cialdella (1984):  SI; isotype:  LIL].

Acacia caven (Molina) Molina var. stenocarpa (Speg.) Burkart ex Ciald., Darwiniana  25: 78.  1984.  Vachellia farnesiana (L.) Wight & Arn. f. stenocarpa Speg., Bol. Acad. Nac. Ci. Córdoba 26:301. 1924. -  TYPE:  ARGENTINA.  MISIONES:  Depto. Candelaria, Santa Ana, A. Burkart 14734 [lectotype, designated by Cialdella (1984):  SI].

Acacia caven (Molina) Molina var. sphaerocarpa Burkart ex Aronson, Ann. Missouri Bot. Gard. 79: 964.  1992. - TYPE:  ARGENTINA.  CORRIENTES:  ca. 27° 27'S, 58° 46'W, 60 m. "alrededores de la ciudad de Corrientes, antiguo camino a Matadero, 50 m de la ruta, 17 Feb 1989, S. G.Tressens & A. Radovancich 3539  (holotype:  K; isotype:  CTES).

Acacia caven (Molina) Molina var. macrocarpa Aronson, Ann. Missouri Bot. Gard.  79: 965.  1992. - TYPE:  ARGENTINA.  SALTA:  Salta, Chicoana, El Carril, 19 Oct 1948, A. Burkart 17577 (holotype:  SI).

Vachellia farnesiana (L.) Wight & Arn. f. brachypoda Speg., Bol. Acad. Nac. Ci. Córdoba  26: 301.  1924. - TYPE:  not cited, from Argentina, Buenos Aires Prov., San Fernanda, Gualeguaychu, Ibicuy y chaco santefecino.

Formal description

Shrub or small tree to 7 (10) m tall.  Bark dark gray to brown, furrowed.  Twigs dark purplish brown to black, slightly flexuous, glabrous to densely puberulent.  Short shoots commonly present above the stipular spines, to 15 mm long, covered with acuminate stipules and old leaf bases.  Leaves alternate, also commonly clustered on spur branches, 25-55 mm long.  Stipular spines dark reddish brown, becoming light gray with age, symmetrical, terete, straight, stout, to 30(50) x 3.0 mm near the base, puberulent at the base and sometimes throughout.  Petiole adaxially grooved, 5-11 mm long, puberulent; petiolar gland solitary, located on the upper half of the petiole, sessile, elongated, 0.5-1.4 mm long, (sometimes circular on short shoot leaves), apex depressed, glabrous.  Rachis adaxially grooved, 5-45 mm long, usually densely puberulent, a sessile, circular gland, 0.3-0.4 mm across present between the upper 1 to 3 pinna pairs.  Pinnae 3 to 10 pairs per leaf, 12-22 mm long, 1.5-5.5 mm between pinna pairs.  Petiolules 0.4-1.4 mm long.  Leaflets 11 to 26 pairs per pinna, opposite, 0.5-1.4 mm between leaflets, linear, 1.0-4.0 x 0.4-0.9 mm, mostly glabrous, lateral veins not obvious, only one vein from the base, margins sometimes sparsely ciliate, apex acute.  Inflorescence a densely flowered globose head 7-10 mm across, solitary or in clusters of 2 to 10 on the short shoots.  Peduncles 6-11(16) x 0.4-0.6 mm, lightly to densely puberulent.  Involucre 4- to 5- lobed, located at the base of the globose head, puberulent, persistent.  Floral bracts spatulate, 0.7-1.1 mm long, puberulent, deciduous.  Flowers sessile, pale yellow; calyx 5-lobed, 1.0-1.5 mm long, the lobes usually puberulent; corolla 5-lobed, 1.8-2.5 mm long, the lobes usually puberulent; stamen filaments 3.0-4.5 mm long; distinct ovary glabrous, on a stipe to 0.1 mm long.  Legumes dark brown to black, straight, nearly circular in cross section, not  constricted between the seeds, oblong, 20-95 x 13-25 mm wide, coriaceous, lightly reticulately striate, glabrous, eglandular, indehiscent; stipe absent to 4 mm long; apex narrowing to a short beak to 10 mm long.  Seeds biseriate to irregularly arranged in a white to reddish pulpy material, light to dark reddish brown, ovoid to ellipsoid, slightly flattened, 4.3-7.2 x 3.3-5.4 mm, smooth; pleurogram U-shaped, 2.5-4.0 mm across.  Flowers in June to November. Chromosome number:  2n = 26 (Castronovo 1945).

Distribution

Shrubby vegetation of pastures, successional fields, stream banks and other disturbed sites, rocky slopes, thorn -scrub grasslands, open savannas, and inundated plains from near sea level to 3200 m in northern Argentina, southern Bolivia, Paraguay, Uruguay, southeastern Brazil, and the Mediterranean-type climate zone of Chile (Aronson and Ovalle 1989).

Additional info

Vachellia caven is an extremely wide-ranging species that probably originated in the warm temperate to subtropical biogeographical region known as the Gran Chaco of southern South America (Aronson and Ovalle 1989).  This region extends through much of north central Argentina, adjacent portions of Bolivia, a small part of the Matto Grosso of Brazil, and nearly half of Paraguay.  Vachellia caven also is common in the Central Valley of Chile in a number of natural ecosystems of both the semi-arid and subhumid portions of the Mediterranean-type climate of that country.  Ovalle et al. (1990) suggest that it is probably an invasive species that has become common by radical changes to the landscape brought about by European colonization.

This species is characterized by a great deal of ecological versatility and morphological variation. It occurs from near sea level to more than 3200 m in mountainous terrain, probably being limited by freezing temperatures. Vachellia caven is also known from near desert conditions in Chile to seasonally inundated savannas in Argentina and Paraguay.  Extensive variation in plant height is also reported, mostly due to land use, moisture, fire and past cutting.  Rarely, ants inhabit the thorns of this species.  One specimen (Mereles 1130 from Paraguay) had a small hole near the tip of the thorn.  The stipular thorns on this specimen were more than 70 mm long and 10 mm wide at the base.

This species is similar to Vachellia farnesiana and V. tortuosa in that it commonly produces short shoots from which clusters of relatively small leaves develop.  These leaves are usually much smaller than the leaves that develop on rapidly growing shoots, have smaller and fewer pinnae, smaller leaflets, smaller petiolar glands, and a rachis that commonly lacks a groove on the upper surface.  Although rarely are there more than 10 pinna pairs to a leaf, even on rapidly growing shoots, occasional individuals have a few more. Even on these specimens many of the leaves have fewer than 10 pinna pairs, and the leaves developing on the short shoot commonly have fewer than 6 pinna pairs.

Fruit size and shape is highly variable, with Spegazzini (1924), and more recently, Aronson (1992), describing six varieties of Vachellia caven based upon pod morphology (dehiscence, shape, size, mesocarp color and structure, and sutural ridges). These include var. caven with elongated, dark brown pod 5-8 cm long, with one sutural ridge; var. macrocarpa Aronson with elongated, reddish purple pods more than 8 cm long, with three distinct sutural ridges; var. stenocarpa (Speggazini) Burkart ex Cialdella with light brown, fusiform pods 3-4 cm long; var. microcarpa (Speggazini) Burkart ex Cialdella with subglobose pods 2-3 cm long by 1-1.5 cm wide, and a reddish mesocarp; var. sphaerocarpa Burkart ex Aronson with subglobose pods 2.l-6.5 cm long by 1.6-4.8 cm wide, and a white mesocarp; and var. dehiscens Burkart ex Cialdella with dehiscent pods.  No other morphological traits are consistently associated with the pod characteristics.
Baranelli et al. (1995) noted that the fruit are not always constant on a single tree.  Also, the geographic separation of these varieties, at the present, appears tenuous. It is the opinion of the present authors that these variations in pod morphology are too variable to serve as a basis for distinguishing varieties.

Molina (1810) originally described this taxon from specimens found in central Chile, and considered it to be closely related to V. farnesiana.  These two species are superficially similar, and, as a result, some subsequent authors have merged the two taxa (Hassler 1909, Spegazzini 1924).  The two taxa are distinct, and, except for possibly in southeastern Brazil, are not sympatric.  Both species have inflated pods, short shoots with numerous small leaves, and similar flowers and inflorescences. In V. farnesiana the petiole has a small, circular gland that is raised above the petiolar groove, leaves with 1-6 pinna pairs, and the leaflets with obvious lateral veins.  Vachellia caven, in contrast, has larger, elongated petiolar glands that are sessile in the petiolar groove, many of the leaves have more than 6 pinna pairs, and the leaflets lack obvious lateral veins.

Aronson and Nash (1989) found that a few herbarium specimens (5 of 40) released cyanide, while Aronson (1990) observed a slight cyanogenic reaction with some seedlings and mature trees of this taxon.  During the present study of the 158 herbarium specimens examined for hydrogen cyanide, nine were cyanogenic, five strongly so.

Flowering time

June-November.

Representative specimens

ARGENTINA:

Buenos Aires:

• island in Río Paraná, route 12, alt. 60 m, 22 Oct 1988, J.Aronson 7647 (MO);
• Los Talas, 1 Nov 1921, A.L.Cabrera 1826 (NY);
• Las Palmas, 13 Oct 1946, J.H.Hunziker 1709 (MO);
• Ciudad Universitaria, 24 Sep 1976, D.Seigler & J.Hunziker 10094 (ILL, MO);

Catamarca:

• Dpto. Belén, Londres, 18 Feb 1978, P.Arenas & D.Herrera 263 (MO);
• Dept. Andalgalá, along Río Andalgalá, 4 km N of Andalgalá, alt. 1200 m, 13 Sep 1972, P.Cantino 322 (GH);

Dep. Paclín, Cuesta de La Vina:

• Ruta 38, 20 Sep 1970, A.T.Hunziker 20697 (EIU);
• Andalgalá, 9 Sep 1915, P.Jorgensen 960 (UC);

Chaco:

• Margarita Belén, 2 Sep 1946, R.M.Aquilar 840 (NY);
• vicinity of Barranqueras, 12 Nov-15 Dec 1913, H.M.Curran 23 (US);
• Dep. San Fernando, Resistencia, 4 Jul 1944, A.Hunziker 5406 (EIU);
• Dep. San Fernando, entre Resistencia y Colonia Benítez, 11 Aug 1944, A.Hunziker 5512 (EIU);
• Fontana, 8 Aug 1938, T.Meyer 3198 (US);
• Presidencia de la Plaza, Jul 1937, A.G.Schulz 2003 (F);
• near Resistencia, 19 Nov 1976, D.Seigler 10164c (ILL);

Córdoba:

• 39 km W of Córdoba, above San Roque, alt. 790 m, 12 Sep 1989, J.Aronson 7884 (ILL, MO);
• Unquillo, 1926, C.Bruch s.n. (NY);
• Córdoba, 2 Jan 1880, C.Galander s.n. (G);
• Córdoba, 1874, G.Hieronymus s.n. (G, US);
• en la cercania de la ciudad, 10 Sep 1877, G.Hieronymus s.n. (NY);
• Dep. Colón, Sierra Chica, Mendiolaza, 5 Jun 1964, A.Hunziker 17466 (EIU);
• Dep. Calamuchita, Dique Los Molinas, 27 Jan 1969, A.Krapovickas & C.L.Cristóbal 14695 (WIS);
• Sierra de San Luis, Bajo de Velis, 25 Jan 1895, F.Kurtz 8492 (EIU);
• Bajo Grande, 9 Aug 1939, R.Maldonado B. 102 (NY);
• Dep. Colón, 10 Mar, 1944, C.A.O'Donell & J.M.Rodríguez 461 (F);
• vic. of Cosquín, 10 Sep 1915, J.N.Rose & P.G.Russell 21073 (NY, US);
• Río I, 29 Aug 1900, T.Stuckert 9511 (G);

Corrientes:

• Paso de la Patria, 6 Jan 1975, Cuezzo A., Ruiz T. & De Marco N. 11274 (CAS);
• vicinity of Goya, alt. 15-30 m, 2-10 Nov 1913, H.M.Curran 327 (US);
• Paso de la Patria, 12 Aug 1967, A.Krapovickas & C.L.Cristóbal 13019 (WIS);
• Dep. Mburucuyá, Santa Teresa, 18 Aug 1952, T.M.Pederson 1799 (GH, MO, NY, US);
• Dep. Empedrado, La Yela, 23 Dec 1954, T.M.Pederson 3072 (G, MO, US);
• Dep. Itati, Tuyuti, 19 Aug 1956, T.M.Pederson 3957 (MO, NY, UC, US);
• Dep. Empedrado, Ea. Saiach, 17 Aug 1989, A.Schinini 26594 (F, MO);
• Corrientes, 18 Oct 1976, D.Seigler & A.Krapovickas s.n. (ILL);
• Dep. Capital, alrededores de la ciudad de Corrientes, 17 Feb 1989, S.C.Tressens & A.Radovancich 3539 (MO);

Entre Ríos:

• Dept. Islas del Ibiquay, alt. 70 m, 22 Oct 1988, J.Aronson 7648 (MO);
• 2 km E of Gauleguaychu, alt. 45 m, 23 Oct 1988, J.Aronson 7652 (MO);
• 5 km S of Colón on ruta 14, alt. 20 m, 27 Mar 1989, J.Aronson 7882 (MO);
• San Izidro, Puerto Constanza al N, 18 Sep 1961, A.Burkart 22590 (US);
• Dep. La Paz, 20 km al N de La Paz, 16 Jan 1960, A.Burkart & N.M.Bacigalupo 21201 (SI);
• Isla del Puerto, 25 Nov 1946, T.Meyer 10466 (LIL);
• Dept. Uruguay, Laguna de la Negro, 19 Jul 1941, E.G.Nicora 3157 (SI);
• Dep. Federación, Barra del Mocoretá, 26 Mar 1967, T.M.Pedersen 8142 (A, UC, US);
• Colón, 2 Jan 1945, C.L.Schulz 365 (A);

Formosa:

• Parque Nacional Pilcomayo, alt. 110 m, 12 Mar 1989, J.Aronson 7841 (MO);
• Arroyo Quita, 60 km N of Resistencia, alt. 10 m, 1 Nov 1988, J.Aronson 7670 (MO);
• Dep. Formosa, 6 Jan 1945, A.Krapovickas 943 (LIL);
• Dep. Pilcomayo, Portenito, 12 Jun 1947, T.Morel 3057 (CAS);
• 2 km E de Portenito, 29 Jun 1947, T.Morel 3269 (MO);
• Dep. Pilcomayo, Puente Ceilo, 22 Sep 1947, T.Morel 3609 (CAS);
• Riacho Porteño, 7 Aug 1958, T.Morel 6084 (MO);
• Ruta 11, entre Dalmacio y Formosa, 24 Jan 1981, C.A.Petetin & A.M.Molina 1501 (G);
• Pilcomayo, 1906, J.Rojas 309 (A);

Jujuy:

• Villa El Perchal, alt. 2490 m, 7 Nov 1988, J.Aronson 7677 (MO);
• Valle de Humahuaca, access road to village of Jueya, alt. 2520 m, 7 Nov 1988, J.Aronson 7682 (MO);
• ca 5 mi. S of Jujuy, alt. 1400 m, 21 Oct 1971, T.Arnow s.n. (MO);
• Dep. Capital, Lozano, 23 Jul 1963, A.L.Cabrera 15837 (US);
• Dep. Santa Barbára, camino de El Fuerte a Palma Sola, 18 Sep 1976, A.L.Cabrera, N.M.Bacigalupo, R.Kiesling & M.E.Múlgura 27983 (F);
• Dep. Capital, outlet Laguna de Yala, 25 km NE of Jujuy, alt. 1300 m, 26 Sep 1938, W.J.Eyerdam & A.A.Beetle 22210 (G, UC);
• Dep. Capital, Río Chijra, 22 Oct 1963, H.A.Fabris 4636 (NY);
• Santa Bárbara, 7 Jul 1901, R.E.Fries 332 (G);
• H.Humbert 21138 (US);
• Dep. Capital, Río Salí, alt. 450 m, 12 Sep 1920, S.Venturi 908 (US);
• Dep. Ledesmia, Sierra de Calilegua, alt. 750 m, 3 Sep 1927, S.Venturi 5249 (CAS, F, GH, UC, US);
• near Jujuy, alt. 1200 m, 25 Feb 1936, J.West 6120 (UC);

La Rioja:

• Chilecito, 12 Jan 1942, T.Meyer 4018 (F);

Misiones:

• 65 km N of Santo Tomé on route 14, Arroyo Espinillo, alt. 65 m, 27 Oct 1988, J.Aronson 7658 (MO);
• Dep. Posadas, costal del río, 28 Jul 1945, Bertoni 1775 (MO, UC);
• Dep. Candelaria, 17 Nov 1949, Bertoni 4684 (LIL);
• Dep. Bonpland, ruta 4, pasando 5 km Bonpland, 18 Jan 1975, Cuezzo A. 10909 (F, LIL);
• vicinity of Posadas, alt. 50-60 m, 5-7 Dec 1913, H.M.Curran 54 (US);
• San Javier, en cerrito de la costa, alt. 3-4 m, 24 Jan 1976, A.Krapovickas & C.L.Cristóbal 28886 (G);
• Santa Ana, 17 Aug 1912, Rodríguez 525 (F);
• Dep. Cainguas, Pto. Frances, 29 Jul 1949, C.J.Schwarz 7959 (GH);

Salta:

• 16 km S of Salta, alt. 1220 m, 5 Nov 1988, J.Aronson 7675 (MO);
• Cafayate, 20 Aug 1936, A.L.Cabrera 3789 (F, SI);
• Dpto. Rosario de Lerina, 5 Nov 1991, A.Charpin & L.Novara 23116 (G);
• Dpto. Santa Victoria, ca de S. Victoria, 10 Nov 1991, A.Charpin & L.Novara 23153 (G);
• Salta, 23 Sep 1901, R.E.Fries 550 (G);
• environs of Salta, 1967-68, J.Lavaque & A.Bolívar 463 (NY);
• Depto. Anta, camino de Las Lajitas a Rivadavia, 2 Oct 1974, P.R.Legname & A.R.Cuezzo 10426 (CAS, US, WIS);
• Manzano, alt. 2000 m, 2 Feb 1941, T.Meyer 3556 (NY);
• Santa Victoria, 2500-2800 m alt., 13/16 Dec 1988, L.J.Novara 8349 (G);
• Dpto. La Caldera, Cuesta del Gallinato, 10 Oct 1987, L.J.Novara 6977 (G);
• Dpto. Capital, Castellanos, alt. 1500-1800 m , 16 Sep 1988, L.J.Novara 8087 (G);
• Depto. Metán, Metán, 30 Mar 1945, C.A.O'Donell 2494 (A, NY);

San Luis:

• Dep. Capital, 2 km from Trapiche, alt. 930 m, 27 Mar 1990, J.Aronson 7907 (MO);

Santiago del Estero:

• Ojo de Agua. Sierra de Sumampa, 22 Oct 1979, B.G.Piccinini & J.Hilfer 4060 (G);

Tucumán:

• Tucumán, Nov 1892, O.Kuntze s.n. (MO, NY);
• La Cocha, 23 Feb 1941, T.Meyer 3897 (NY);
• Capital, alt. 470 m, Sep 1943, T.Meyer 5255 (US);
• 13.8 km SE of Amaicha del Valle on rd. to Tafi del Valle, alt. 2400 m, 4 May 1985, J.C.Solomon 13537 (ILL, MO);

Isla Martín García:

• Borques, Oct 1934, A.L.Cabrera 2913 (NY).

BOLIVIA:

Cochabamba:

• 14 km E of Cochahamba, 4 Dec 1975, C.Davidson 3720a (RSA);
• 13-15 km E of Cochabamba, L.J.Dorr & L.C.Barnett 6991 (ILL);
• public park area on Colina de San Sebastían in Cochabamba, alt. 2575 m, 14 Nov 1984, M.Nee 30353 (MO, NY);
• Cochahamba, alt. 2825 m, 19 Nov 1999, M.Nee 50517 (MO);
• Tarata, alt. 2760 m, 28 Apr 1995, N.Ritter 1950 (MO);

La Paz:

• valley separating Inquisivi and Sita, 22 Dec 1989, L.J.Dorr, L.C.Barnett, A.Canessa & M.Lewis 6797 (ILL, MO);
• 1 km N of Inquisivi, alt. 2150-2400 m, 16 Dec 1988, M.Lewis 882079 (ILL, MO);
• Prov. Murillo, Hacienda Huajchilla, 13 km SE of Calacoto, alt. 3000 m, 30 Jan 1983, J.C.Solomon 9449 (MO);
• Prov. Murillo, Mallasa (La Paz), alt. 3200 m, 7 Aug 1986, J.C.Solomon 15483 (MO, NY);

Santa Cruz:

• Curuyuqui, along Río Parapeti, alt. 350 m, 26 Oct 1991, A.Gentry & M.Peña 75269 (MO);
• Prov. Florida, 5 km S of Mataral on rd. to Vallegrande, alt. 1400 m, 15 Dec 1990, M.Nee ?? (ILL, NY);
• Tara, alt. 450 m, 8 Oct 1924, J.Steinbach 6582 (F);

Tarija:

• Prov. Avilés, Chocloca, 6 km SW al camino, alt. 2000 m, 25 Mar 1979, G.Beck 716
  (US);
• Prov. Cercado, alt. 2230 m, 26 Mar 1979, G.Beck 798 (US);
• Prov. Cercado, 8 km NW an der Hauptstrasse von Tarija, alt. 1700 m, 1 Jan 1980, T.Feuerer 7583a (NY);
• Cercado, Rancho Santa Ana, 17 km E of Tarija, 3 Sep 1987, T.Killeen 2691 (F, NY);
• O'Connor, Rancho Huayco, 8 km N of road to Villamontes W of Entre Ríos, 4 Sep 1987, T.Killeen 2693 (F, NY);
• La Capilla, alt. 1270 m, 6 Sep 1971, C.A.Nuesser 29 (GH);
• Prov. Cercado, flood plain of Río Guadalquivir opposite Tarija, alt. 1800 m, 30 Sep 1983, J.C.Solomon 10913 (MO);
• Prov. Arce, 5.5 km N of Mamora, alt. 1500 m, 27 Apr 1983, J.C.Solomon 10181 (MO, UC);
• Prov. Cercado, 40 km E of Tarija-Padcaya road, on road to Entre Ríos (near Abra Condor), alt. 2500 m, 1 May 1983, J.C.Solomon 10281 (MO, UC).

BRAZIL:

Mato Grosso:

• Mun. Porto Murtinho, Estrada Fazenda Jereré, alt. 90 m, 22 Oct 1985, R.L.Loureiro 161 (NY);
• 20 km E de Porto Murtinho, 23 Oct 1980, J.M.Pires & P.P.Furtado 17287 (NY, US);
• Pantano Grande, Rio Pardo, RS, 2 Oct 1984, M.Sobral & Y.Folz 3061 (F, MO);
• Pantano Grande, 1 Oct 1988, M.Rossato 4624 (G, NY);

Río Grande do Sul:

• 98 km W of Cacapava do Sul, alt. 120 m, 25 Mar 1989, J.Aronson 7877 (MO);
• Jaguarão, 7 Feb 1950, I.Gilberto M. 61 (F, GH, MO);
• 1835, M.Isabelle s.n. (G);
• L. Luiz das Minos, 4 Feb 1949, A.Lehnem 3603 (SI);
• Barra do Quaraí, 14 Oct 1971, J.C.Lindeman, B.Irgang & J.F.M.Valls 8442 (F);
• Rincão do Meio, Pelotas, 19 May 1959, J.C.Sacco 2696 (F).

CHILE:

Atacama:

• Copiapo Prov., Tierra Amarilla, 13 kms SSE of Copiapo, alt. 500 m, 7 Feb 1989, J.Aronson 7728 (MO);
• desert of Atacama, Sept-Oct 1890, T.Morong 1115 (NY, US);
• Dep. Copiapo, Tierra Amarilla, alt. 500 m, Sep 1924, E.Werdermann 410 (F, CAS, G, MO, NY, UC);

Colchagua:

• Estación Quinta, 9 Mar 1939, M.R.Espinosa 27 (CAS);
• Fundo Lolol, Lolol, 26 Nov 1970, J.P.Simon 377 (GH, MO, NY, UC);

Coquimbo:

• Elqui Prov. ca. 30 km S of La Serena, alt. 280 m, 5 Feb 1989, J.Aronson 7717 (MO);
• Elqui Prov. along access rd. to Tololo Observatory, alt. 700 m, 6 Feb 1989, J.Aronson 7720 (MO);
• 6 km S of Corral Quemado on road to Ovalle, alt. 600 m, 29 Oct 1990, T.G.Lammers, C.M.Baeza P. & P.Peñailillo B. 7581 (F, MO);
• 20 Aug 1917, C.I.Skottsberg 904 (F);
• Depto. Elqui, Los Chiches, 3 km W of Vicuna, road to La Serena, alt. 600 m, 14 Sep 1939, R.Wagenknecht 18430 (F, G, MO, UC);

Maule:

• Linares Prov., 7 km N of Linares, alt. 200 m, 20 Sep 1989, J.Aronson 7896 (MO);

Santiago:

• 80 Km W of Santiago, in cow pasture near Le Ovalle, 18 Nov 1988, J.Aronson 7692 (ILL);
• cow pastures of Fundo Sauce, road to Cuesta de Chacabuco, alt. 640 m, 18 Nov 1988, J.Aronson 7696 (ILL);
• Santiago, Nov 1921, Bro. Claude-Joseph 1490 (US);
• 1856-1857, Santiago, P.Germain s.n. (G);
• C. San Cristóbal, 14 Sep 1959, E.S.Gersh 51 (WIS);
• San Carlos de Apoquindo, Sep 1983, L.González 4970 (MO);
• waste places near Santiago, 22 Sep 1900, G.T.Hastings 49 (UC, US);
• base of mts. E of Santiago, alt. 2000 ft., 23 Nov 1900, G.T.Hastings 195 (NY, UC, US);
• Cuesta de Barriga, between Marruecos and Los Cerrillos, alt. 700-1000 m, 3 Nov 1948, E.P.Killip & E.Pisano 39648 (US);
• El Principal, Río Clarillo, near Puente Alto, 17 Oct 1971, L.R.Landrum 855 (MICH);
• southern edge of Santiago, 27 Jan 1966, G.F.Ledingham 4450 (NY);
• Mahu, cerros de Puente Verde, camino a Colina, 20 Sep 1964, M.Mahu 703 (CAS);
• Santiago, 22 Oct 1927, G.Montero 131 (F, MO);
• Capilla de Caleu, Comuna de Tiltil, alt. 900 m, 13 Apr 1969, T.Plowman 2678 (GH, US);
• Cerro San Cristóbal, 16 Sep 1917, C.I.Skottsberg 967 (NY);

Valparaíso:

• Quillota, 1879, D.Bertero 1830 (G);
• Quillota Prov., Cuesta de Chacabuco, alt. 740 m, 3 Feb 1989, J.Aronson 7705 (MO);
• Quillota Prov., Ochoa, alt. 490 m, 3 Feb 1989, J.Aronson 7706 (MO);
• about 4 km E of Lliu in Cajón de la Pataguilla, alt. 400 m, 23 Dec 1951, P.C.Hutchinson 191 (UC, US);
• near Valparaiso, Jun 1885, H.H.Rusby 953 (NY, US);
• Cajón de San Pedro, W side of Campana de Quillota, alt. 500 m, 19 Jan 1936, J.West 5193 (UC);

PARAGUAY:

Boquerón:

• cerca y lqa Sur de Mariscal Estigarribia, 23 Oct 1980, J.F.Casas & J.Molero 4431 (G, MO, NY);
• Mcal Estigarribia, Mar 1984, F.Mereles 1130 (G):
• Tte. Ochoa, 18 Feb 1987, F.Mereles 2272 (G);
• Espartillar, linea 118 hasta se cruce con la linea 10. a 60 km de la 7a. Division de Infantería, 25 Oct 1986, L.Molas 1424 (ILL, MO);
• roadside, km 120 along Ruta Trans-Chaco, 26 Aug 1986, N.Soria & G.Schmeda 1150 (US);
• Tte. 1o Alfredo Stroessner, 2 km al sur del cuartel, alt. 200-210 m, 9 Oct 1987, R.Spichiger, L.Ramella, F.Mereles, N.Soria & L.Spinzi 2088 (G);

Caaguazú:

• Trans les campos, Sep 1876, B.Balansa 1449 (G);
• cerca y al Sur de Yhú, no lejos de la desviación a San Joaquín, 13 Dec 1982, J.F.Casas, W.Hahn & A.Schinini 7506 (G, NY);
• viciniis Caaguázu, Feb 1905, E.Hassler 9085 (G, NY, UC);
• alredores, 26 Aug 1985, G.Hatschbach & A.C.Ceryi 49810 (G, MO);
• Arroyo Cambey, 10 Nov 1990, E.Zardini & U.Velázquez 23881 (MO);

Central:

• 4.4 km W of Asunción on Ruta Trans-Chaco, alt. 20 m, 30 Oct 1988, J.Aronson 7667 (MO);
• Lambare, 22 Jan 1984, W.Hahn 1925 (ILL, MO);
• prope Sapucay, 1913, E.Hassler 11843 (G, MO, NY, US);
• Luque, Compania Tarumandy, propiedad Nueva Asunción, 30 Aug 1991, F.Mereles 4040 (MO);
• 1888-90, T.Morong 751 (F, MO, US);
• Mariano R. Alonso, 28 Jul 1984, M.Ortíz 345 (G);
• Ruta Concepción-Pozo Colorado, 18 Feb 1990, R.Palacios 1850 (MO);Jardín Botánico y Zoológico, Trinidad, Reserva Natural, Sep 1991, B.Pérez 1111 (ILL, MO);
• Tavarory, Arroyo Avai, 9 Nov 1990, E.Zardini & C.Velázquez 23542 (ILL, MO);
• Mariano Roqie Alonso-Limpio, 4 km NW from entrance to Tavavory, 8 Jun 1992, E.Zardini & L.Guerrero 31953 (ILL, MO);
• Lake Yabeby, near Cerro Pe, 3 Aug 1991, E.Zardini & H.Velázquez 28164 (ILL, MO);

Concepción:

• Sep 1892, O.Kuntze s.n. (NY);
• 1901-2, E.Hassler 7227 (G);
• Sep 1877, P.Lorentz 1194 (G);
• Río Napeque, Aug 1988, F.Mereles 1374 (G);
• Concepción, Aug 1985, F.Mereles 2066 (G);

Cordillera:

• Río Salado, camino de Limpio a Emboscada, 13 Jun 1984, M.M.Arbo, M.S.Ferrucci, A.Schinini & R.O.Vanni 2623 (F, G, MICH);
• San Bernardino Campana Cierro Cua, 26 Jul 1988, I.Basualdo 1590 (CM, G, MO);
• near Luque, 15 Jan 1974, J.Conrad & W.Dietrich 2203 (MO);
• San Bernardino, Jul 1988, F.Mereles 1250 (G);
• Confluence of Río Paraguay and Río Salado, 23 Dec 1989, E.Zardini & U.Velázquez 17414 (ILL, MO);
• Río Salado basin, 7 km NE of Emboscada, 23 Dec 1989, E.Zardini & M.Velázquez 17462 (ILL, MO);
• Confluence of Río Paraguay and Río Salado, 18 Nov 1989, E.Zardini & U.Velázquez 16627 (ILL, MO);
• Confluence of Río Paraquay and Río Salado, 21 Jul 1990, E.Zardini & M.Velázquez 22285 (ILL, MO);

Guairá:

• Mbocyaity-Melgarejo, Cerro Naville, 6 Jul 1992, E.Zardini & L.Guerrero 32499 (ILL, MO);

Paraguarí:

• Estero del Ypoa, 10 km E of Nueva Italia on Arroyo Canabe, 23 Jun 1990, E.Zardini & C.Velázquez 21681 (ILL, MO);
• Estero del Ypoa, 10 km E of Nueva Italia on Arroyo Canabe, 8 Dec 1990, E.Zardini & R.Velázquez 25239 (ILL, MO);

Presidente Hayes:

• crece en monte cercano al Río Pilcomayo, 4 Jan 1980, P.Arenas 1571 (MO);
• 22 km W of Pozo Colorado, alt. 190 m, 15 Mar 1989, J.Aronson 7842 (MO);
• 22 km W of Pozo Colorado, alt. 190 m, 15 Mar 1989, J.Aronson 7843 (MO);
• Río Verde, cerca de Pozo Colorado, 24 Oct 1980, J.F.Casas & J.Molero 4481 (G, MO, NY);
• Chaco-i, 19 Jul 1987, A.Krapovickas 41865 (A, F, G, MO, UC, WIS);
• zona inundable del Río Paraguay, Oct 1988, F.Mereles 1625 (G);
• Río Confuso, Aug 1990, F.Mereles 3958 (G);
• Río Confuso, Oct 1988, F.Mereles 1575 (G);
• Tinfunké, Estancia Brusquetti, 20 Dec 1989, F.Mereles & P.Geissler 3330 (G);
• Estigarribia, 11 Dec 1987, A.Schinini & R.Palacios 25934 (G, MO);
• 6 km de Concepción, 14 Aug 1987, N.Soria 1817 (G, MO);
• Estancia Tinfunqué, Puesto Clorinda-i, alt. 100-150 m, 20 Sep 1987, R.Spichiger, L.Ramella, F.Mereles, N.Soria & L.Spinzi 2016 (G);
• Puerto Militar, Río Paraguay, 9 Dec 1989, R.Vanni, L.Ferraro & S.Ferrucci 1220 (G, MO);

State not known:

• In regione fluminis Capibary, 1898-99, E.Hassler 4411 (G);
• vicine San Estanislao, E.Hassler 4284 (F, G);
• Dep. Pilar, Humaita, 12 Dec 1950, T.Meyer 16171 (LIL);
• E.Hassler 1677 (G);
• E.Hassler 123 (G);
• Exploration of the River Plata, Capt. T.J. Page, Paraguay Expedition 1856, E.Palmer s.n. (US).

URUGUAY:

San José:

• Barra de Sta. Lucía, Apr 1927, G.Herter 669 (F, G, HBG, MO, NY, WIS);
• Santa Lucía, 1 Nov 1886, W.E.Safford s.n. (US).

UNITED STATES:

California:

• Santa Clara Co.  persisted for a number of years, roadside, Holsclaw and Gilman Roads, Gilroy, 10 Jul 1967, M.H.Murphy s.n. (CDA).

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