Name
Vachellia aroma (Gillies ex Hooker & Arnott) Seigler & Ebinger, Phytologia 87:143. 2005. syn. Acacia aroma Gillies ex Hook. & Arn. Bot. Misc. 3:206. 1833.
Synonymy and types
Basionym: Acacia aroma Gillies ex Hook. & Arn. Bot. Misc. 3:206. 1833. - Acacia lutea (Mill.) Britton var. aroma (Gillies ex Hook. & Arn.) Kuntze, Revis. gen. pl. 3(2):47. 1898. - Vachellia lutea (Mill.) Speg. var. aroma (Gillies ex Hook. & Arn.) Speg., Bol. Acad. Nac. Ci. Córdoba 26:307. 1924. - Vachellia lutea (Mill.) Speg. f. aroma (Gillies ex Hook. & Arn.) Speg., Bol. Acad. Nac. Ci. Córdoba 26:313. 1924. - TYPE: ARGENTINA. a little way south of San Juan, Dec. 1823, J.Gillies s.n. (holotype: K).
Acacia moniliformis Griseb., Abh. Königl. Ges. Wiss. Göttingen 19:136. 1874. - Acacia lutea (Mill.) Britton var. moniliformis (Griseb.) Kuntze, Rev. gen. pl. 3(2):47. 1898 (as monoliformis). - Vachellia lutea (Mill.) Speg. var. moniliformis (Griseb.) Speg., Bol. Acad. Nac. Ci. Córdoba 26:303. 1924. - Vachellia lutea (Mill.) Speg. f. moniliformis (Griseb.) Speg., Bol. Acad. Nac. Ci. Córdoba 26:312. 1924. - TYPE: ARGENTINA. TUCUMAN: La Cruz, 20-24 Apr. 1872, T.G.Lorentz 190 (holotype: GOET).
Acacia michelii Rusby, Mem. Torrey Bot. Club 6:28. 1896. - TYPE: BOLIVIA. vic Cochabamba, 1891, M.Bang 1095 (holotype: K; isotype: NY).
Acacia aroma Gillies ex Hook. & Arn. var. cochlearis Griseb., Abh. Königl. Ges. Wiss. Göttingen 24:122. 1879. - TYPE: ARGENTINA. CÓRDOBA: north of San Javier, 18 Mar. 1876, T.G.Hieronymus 558 (holotype: GOET).
Vachellia lutea (Mill.) Speg. f. oocephala Speg., Bol. Acad. Nac. Ci. Córdoba 26:312. 1924. - TYPE: none cited.
Vachellia lutea (Mill.) Speg. f. thlipsacantha Speg., Bol. Acad. Nac. Ci. Córdoba 26:312. 1924. - TYPE: none cited.
Vachellia lutea (Mill.) Speg. var. aroma (Gillies ex Hook. & Arn.) Speg. f. leptocarpa Speg., Bol. Acad. Nac. Ci. Córdoba 26:305. 1924. - TYPE: none cited.
Vachellia lutea (Mill.) Speg. var. aroma (Gillies ex Hook. & Arn.) Speg. f. pachycarpa Speg., Bol. Acad. Nac. Ci. Córdoba 26:307. 1924. - TYPE: none cited.
Formal description
Large shrub to small tree 6 m tall. Bark dark gray to dark brown, smooth to shallowly furrowed. Twigs light to dark purplish brown, slightly flexuous, glabrous to densely puberulent. Short shoots sometimes present above the stipular spines particularly on slow growing branches, to 2 mm long, covered with acuminate stipules and old leaf bases, sometimes developing into long shoots. Leaves alternate, also occasionally clustered on the short shoots, 60-140 mm long. Stipular spines reddish brown, usually becoming light gray with age, symmetrical, terete to angular or rarely flattened, straight, stout, to 10(60)mm x 1.0-2.5 mm near the base, glabrous to puberulent at the base and sometimes throughout. Petiole adaxially grooved, 5-9 mm long, glabrous to lightly puberulent; petiolar gland solitary, located medially to just below the first pinna pair, sessile, circular to elongated, 0.7-1.8 mm long, apex usually depressed to rarely bulbous, sometimes puberulent. Rachis adaxially grooved, 30-135 mm long, usually lightly puberulent, a sessile, circular gland, 0.2-0.6 mm across, between the upper 1 to 3 pinna pairs. Pinnae (4)12 to 29 pairs per leaf, 18-31 mm long, 2.5-6.0 mm between pinna pairs. Petiolules 0.4-1.1 mm long. Leaflets 24 to 50(58) pairs per pinna, opposite, 0.4-0.9 mm between leaflets, linear, 1.5-4.1 x 0.5-0.8 mm, glabrous to lightly puberulent, lateral veins not obvious, only one vein from the base, base oblique, margins lightly ciliate, apex broadly acute to obtuse. Inflorescence a densely flowered globose head, 7-10 mm across, solitary (rarely 3) from the nodes. Peduncles 22-40 x 0.4-0.7 mm, usually lightly puberulent. Involucre 4- to 6-lobed, located at the base of the globose head, puberulent, persistent. Floral bracts spatulate, 1.3-1.9 mm long, apex puberulent, deciduous. Flowers sessile, pale yellow; calyx 5-lobed, 1.3-1.8 mm long, the lobes densely puberulent; corolla 5-lobed, 2.0-2.7 mm long, the lobes commonly puberulent; stamen filaments 2.5-4.0 mm long, distinct; ovary glabrous to lightly pubescent, on a stipe to 0.2 mm long. Legumes dark reddish brown, straight to slightly curved, flattened to elliptical in cross section, constricted between the seeds, oblong, 30-150 x 9-15 mm, coriaceous, reticulately striate, usually densely puberulent, eglandular, indehiscent; stipe to 6 mm long; apex acute, narrowing to a beak to 15 mm long. Seeds uniseriate, no pulp, light to dark brown, ovoid to nearly circular, strongly flattened, 5.3-7.3 x 4.0-6.2 mm, smooth; pleurogram nearly oval, 2.5-4.0 mm across. Flowers from September to February. Chromosome number: Not determined.
Distribution
Shrubby vegetation of pastures, successional field, edge of roads, sandy floodplains and other disturbed sites, thorn -scrub forests, and chaco scrub forest from near sea level to 3000 m in central and northern Argentina, Bolivia, western Paraguay and north to Peru and Ecuador.
Additional info
Vachellia aroma var. aroma is a wide-ranging species extending through most of the drier regions of southern South America. It is common in many areas where V. caven occurs, particularly in northern Argentina, Bolivia and Paraguay. Vachellia aroma is an invasive species, with many of the collections from pastures, successional fields, and roadsides. Like V. caven, this taxon seems to be characterized by a great deal of ecological variation. It occurs from near sea level to 3000 m, probably being limited by freezing temperatures. It also occurs from extremely dry Larrea associations to thorn scrub and dry forests to stream banks.
The only species in South America that Vachellia aroma var. aroma is usually confused with is V. macracantha. Both species have relatively large leaves with mostly more than 11 pinnae pairs, a characteristic limited to these two taxa and V. pennatula. Though superficially similar, numerous characteristics can be used for consistent separation. Vachellia aroma has flattened fruits that are constricted between the seeds; many of the peduncles are more than 25 mm long; the petiole glands are sessile; and the petiole, and rachis are puberulent with short, erect hairs that are less than 0.2 mm long. Vachellia macracantha, in contrast, has flattened fruits that are not constricted between the seeds, the peduncles are less than 25 mm long, the petiole gland is stalked, and the pubescence, if present, is of erect hairs that are more than 0.3 mm long.
Besides the characteristics listed above, short shoots are sometimes found in Vachellia aroma, while they are mostly absent in V. macracantha. These short shoots occur on slow growing branches were they develop just above the stipular spines in the axil of primary leaves. The primary leaves commonly have 12 to 29 pinnae pairs with relatively large leaflets. The leaves that develop from the short shoot are generally smaller, commonly with 4-15 pinnae pairs and leaflets than are less than 2.3 mm long. It is not uncommon for these short shoot leaves to lack a petiole gland, or the gland is smaller and more bulbous than on the larger leaves.
Throughout its extensive geographic range and habitats in which Vachellia aroma var. aroma occurs, this taxon is highly variable in some morphological characteristics. Extensive variation occurs in leaf size and pubescence, as well as in the overall size of the plants. Most of the specimens observed from east of the Andes Mountains, the vegetative parts of the plants were usually glabrous to lightly puberulent, particularly the petioles, rachises, petiolar glands and leaflets; while the primary leaves were longer, with more and longer pinna that had more leaflet pairs. Specimens from west of the Andes Mountains usually had densely puberulent petioles, rachises, leaflets and petiolar glands; while the primary leaves were smaller with fewer and shorter pinna that had fewer pairs of leaflets. Also, the information on the herbarium labels indicated that the specimens east of the mountains were on the average larger and more robust than those from west of the mountains.
It appears that Vachellia aroma var. aroma is only rarely cyanogenic. Aronson and Nash (1989) reported negative results for this species, while Aronson (1990) found some seedlings to be slightly to moderately cyanogenic. Of the 220 specimens test during the present study 215 gave a negative reaction even after the addition of emulsin. A weak positive cyanogenic reactions was observed in five specimens. Seigler and Conn (1982) found linamarin and lotaustralin to be the principal glycosides in this species.
The two varieties of Vachellia aroma (aroma and huarango) are easily separated since var. aroma is a large shrub or small trees with an wide distribution throughout much of southern South America. Variety huarango, in contrast, is a low dense shrub usually less than 1.5 m tall, and is commonly broader than tall with a restricted distribution, occurring in Ecuador and Peru.
Flowering time
September-February.
Representative specimens
ARGENTINA:
Catamarca:
- Miraflores, 9 Dec 1942, H.H.Bartlett 19207 (MICH);
- Miraflores, 9 Dec 1942, H.H.Bartlett 19219 (MICH, US);
- 6 km SW of Andalagalá on rd. to Belén, 23 May 1943, H.H.Bartlett 20213 (F, G, MICH, NY, UC, US);
- Miraflores, 14 Jan 1947, J.Brizuela 119 (CAS);
- 18 km SSW of Tinogasta, alt. 1300 m, 13 Jan 1956, J.P.Hjerting, E.Petersen & K.Rahn 2292 (MO);
- Andaejula, alt. 1070 m, 10 Dec 1915, P.Jörgensen 1427 (G, MO, UC);
- Dep. Santa Rosa, San Pedro, 15 Nov 1946, M.R.Malvárez 62 (F);
- Dept. Pomán, national highroad 62, about half-way between Nat. road 60 and Pomán, T.M.Pedersen 11820 (A);
- 6 km W of Andalgalá on road to Belén, 13 Nov 1974, B.Vuilleumier 1052 (US);
Chaco:
- Laguna Blanco, 22 Dec 1944, R.Aguilar 49 (NY);
- Dep. Resistencia, Margarita Belén, 7 Jan 1948, R.Aguilar 1209 (NY);
- Dep. Napalai, Las Cuchillas, 15 Jan 1947, P.Buratovich 724 (CAS);
- vicinity of Barranqueras, alt. 35-40 m, 12 Nov-15 Dec 1913, H.M.Curran 25 (US);
- entre Castelli y Pto. Lavalle, 13 Oct 1973, H.E.Erb 301 (CAS);
- entre Tirol y Río Araza, 17 May 1947, T.Meyer 8611 (A, NY);
- Laenz Peña, 18 Jun 1944, T.Rojas 11215 (A, US);
- Puerto El Colorado, Aug 1944, T.Rojas 12094 (A);
- Enrique Urien, 15 Oct 1945, C.L.Schulz 760 (WIS);
- Col. Benítez, May 1947, A.G.Schulz 2011 (MO);
- Feb 1917, P.Jörgensen 2116 (G, US);
Córdoba:
- Sierra de Guasapampa, Serrezuela, H.H.Bartlett 19576 (G, MICH, UC, US);
- Sebastián Elcano, 31 Mar 1943, H.H.Bartlett 19814 (MICH, US);
- Monte, at San José, 13 km W of Villa Dolores, 8 Jul 1943, H.H.Bartlett 20603 (G, MICH, UC, US);
- Unquillo, 1926, C.Bruch s.n. (NY);
- Dep. Cruz del Eje, 7 Nov 1945, A.R.Cuezzo 899 (F);
- N voir San Javier, 18 Mar 1876, G.Hieronymus 558 (F, US);
- Dep. Capital, Observatorio, 9 Mar 1944, C.A.O'Donnell & J.M.Rodríguez 256 (F);
- Dep. Cruz del Eje Soto, 1 Apr 1944, C.A.O'Donnell & J.M.Rodríguez 623 (A, F);
Corrientes:
- Dep. Capital, Corrientes, 25 Mar 1947, M.R.Malvarez 1440 (CAS);
- Dep. Capital, Corrientes, 26 Jan 1944, T.Meyer 6063 (A);
- at Río Empedrado on the main rd. from Corrientes S, 23 Oct 1951, T.M.Pedersen 1302 (US);
- Dep. Empedrado, Estancia "Las Tres Marias", 10 Jan 1952, T.M.Pedersen 1428 (MO, NY, US);
- Barrancas del Río Paraná, 6 Jul 1944, T.Rojas 11529 (A);
- Arryo Riachuelo, 27 Jun 1974, A.Schinini, C.Quarin, M.M.Arbo & S.G.Tressens 9428 (MO);
- Puerto Empedrado, 28 Feb 1950, G.J.Schwarz 9876 (US);
- El Sombrero, Dep. Empedrado, 9 Mar 1950, G.J.Schwarz 10083 (CAS);
Formosa:
- Riacho San Hilario, 15 Nov 1986, C.L.Cristóbal, A.Krapovickas & A.Schinini 2149 (A, F, UC);
- Dep. Pilcomayo, 3 km de Riacho Negro, 4 Aug 1947, T.Morel 3821 (MO);
- Ruta 86 at km 55, Dep. Pilcomayo, 9 Feb 1948, T.Morel 4617 (CAS);
- Dep. Pilcomayo, Ruta 11 al km 101, I.Morel 4286 (MO);
- Dep. Pilcomayo, La Frontera, 8 Nov 1949, I.Morel 8840 (WIS);
- Pirane, Chacros, 30 Nov 1945, S.Pierotti 4154 (NY);
Jujuy:
- 2 km S of Santa Rita along banks of Río Lavayén, alt. 520 m, 3 Oct 1971, L.Arnow 3512 (MO);
- La Esperanza Estate, San Pedro de Jujuy, 31 May 1943, H.H.Bartlett 20311a (G, NY, MICH, UC, US);
- Calilegua, 1 Jun 1943, H.H.Bartlett 20394 (G, MICH, US);
- Estes Hacienda, 40 km S of Jujuy, alt. 1250 m, 5 Oct 1938, W.J.Eyerdam & A.A.Beetle 22434 (G, UC);
- Pampa Blanca, 12 Nov 1980, A.L.Cabrera, S.Botta, A.M.Ciadella & A.Rotman 31922 (MO);
- Cuesta de las Lajitas, 13 May 1981, A.L.Cabrera, S.Botta, N.Deginani & F.O.Zuloaga 32743 (MO);
- R. de Sora irrigation channel, 14 Nov 1978, S.A.Renvoize 3481 (MO, NY, US);
- Sierra de Calilegua, alt. 750 m, 14 Oct 1927, S.Venturi 5376 (F, GH, MO, RSA, US);
- Dep. San Pedro, Santa Claria, alt. 650 m, 18 Oct 1929, S.Venturi 9741 (A, G, MO, NY);
La Rioja:
- Ilta, alt. 700 m, 8 Oct 1944, M.P.Gómez 88 (A, F);
- Sierra de Los Llanos, 6 km al SE de Chamical, alt. 700/800 m, 20 Feb 1964, A.T.Hunziker & T.E.Di Fulvio 16977 (MO);
- Chilecito, alt. 1100 m, 12 Jan 1942, T.Meyer 4023 (GH);
- Vinchina, alt. 1500 m, 17 Jan 1942, T.Meyer 4035 (GH, US);
- Guandacol, alt. 950 m, 3 Feb 1942, T.Meyer 4214 (A, F);
- La Puntilla, Chilecito, 31 Jul 1927, L.R.Parodi 7852 (G);
- R. Laenz Peña, 18 Jun 1944, T.Rojas 11215a (A);
Salta:
- Fundo Campos del Norte, 20 km S of Salta, alt. 1120 m, 5 Nov 1988, J.Aronson 7672 (MO);
- Dep. Orán, Río Pescado, 13 Oct 1961, Ahumand, Vaca & Legname 2136 (G);
- Coronel Moldes, vicinity of the dam (El Dique), 15 Feb 1943, H.H.Bartlett 19707 (G, MICH, US);
- Coronel Moldes, 28 May 1943, H.H.Bartlett 20266 (MICH, US);
- Coronel Moldes, 28 May 1943, H.H.Bartlett 20267 (MICH, US);
- Dep. Capital, 30 Jan 1941, J.B.Borrea 2 (GH, NY, UC);
- Dep. Orán, Río Pescado, 11 Jul 1946, O.Borsini 621 (CAR);
- Mun. Cafayate, on rd. to La Punilla, alt. 5400 ft., 27 Dec 1987, J.S.Bricker, L.Landrum & F.Grifo 164 (MO);
- Dep. San Carlos, Ruta 40, Río Las Vinas, 8 Nov 1978, A.L.Cabrera, S.Botta, C.Ezcurra, A.M.Ragonese & M.V.Avila 29710 (MO);
- at junction of Río Tarija and Río Bermejo, alt. 800 m, 4 Nov 1938, W.J.Eyerdam & A.A.Beetle 22850 (GH, MO, UC);
- Dep. Chicoana, Los Los, 7 May 1975, A.Krapovickas, A.Schinini & J.M.González 304 (MO, WIS);
- 14 km WNW de El Tunal, alt. 600 m, 4 Apr 1980, A.Krapovickas & A.Schinini 35721 (NY);
- Dep. Oran, camino de Aguaray a Pocitos, a 4 km de Aquaray, 18 Mar 1972, R.Legname & A.R.Cuezzo 9105 (G);
- Metan, Feb 1873, P.G.Lorenty & G.Hieronymus 1219 (F);
- Quebrachal, 23 May 1947, F.E.Luna 135 (NY);
- Anita, 23 May 1947, F.E.Luna 155 (G);
- Coronel Moldes, alt. 1100 m, 31 Jan 1941, T.Meyer 3551 (G, NY);
- Dep. Capital, alt. 1100 m, 8 Jan 1941, T.Meyer 3572 (GH, NY);
- Dep. Orán, El Tabacal, alt. 310 m, 18 Feb 1943, T.Meyer 4526 (F);
- Dep. Orán, Campamento Y. P. F. Río Pescado, 28 Feb 1943, T.Meyer 4768 (F);
- Chicoana, 3 May 1947, T.Meyer 12130 (MO);
- Dpto. Vaqueros, Arroyo Chaile, 21 Oct 1986, V.Nuñez 34 (NY);
- Metán, 30 Mar 1945, C.A.O'Donnell 2502 (NY);
- Orán, 14 Apr 1945, C.A.O'Donnell 3162 (A, NY);
- Orán, La Maroma, 23 Jan 1945, S.A.Pierotti 67 (A, F, NY);
- Orán, San Ignacio, 11 Apr 1945, S.A.Pierotti 1201 (NY, UC);
- Orán, Río de las Piedras, 3 Nov 1911, Rodríguez 24 (A);
- Dep. Campo Santo, Yaquiasmé, alt. 700 m, 4 Nov 1975, M.M.Schiavone, A.R.Cuezzo, M.R.Figueroa & P.R.Legname 11763 (CAS);
- between Cafayate and Tolombón, 19 Dec 1970, B.Timmermann 70127 (TEX);
San Juan:
- Ullun, 20 Jun 1943, H.H.Bartlett 20497 (G, MICH, NY, UC, US);
- ravines at Dique Nivelador, Marquesado, 22 Jun 1943, H.H.Bartlet 20524a (G, MICH, US);
- Marayes, 13 Nov 1945, A.R.Cuezzo 1024 (F);
- Pachaco, 89 km W of San Juan, alt. 1200 m, 9 Jan 1956, J.P.Hjerting, E.Petersen & K.Rahn 2226 (MO);
- Dep. Valle Fértil, Ruta Nac. 510, 15 km NE de Marayes, 27 Nov 1984, R.Kiesling, M.Múlgura & M.Ponce 4898 (NY);
- Ullun, alt. 820 m, Nov 1941, A.P.Rodrigo 2828 (F);
San Luis:
- Alto Pencoso, Feb 1914, Bruch & Carette (NY);
Santa Fe:
- Vera, 24 Nov 1939, M.Birabén 277 (NY);
- Guadalupe, 27 Nov 1939, M.Birabén 259 (NY);
- Dep. 9 de Julio, 29 Dec 1944, alt. 250 m, A.Krapovickas 783 (A);
- entre San Cristóbal y Crespo, 27 Dec 1983, J.Pensiero 1350 (MO);
Santiago del Estero:
- Dep. Lumampa, 25 Nov 1944, B.Balegno 108 (A);
- between Brea Pozo and Río Huayco Hondo, 23 Mar 1943, H.H.Bartlett 19755 (G, MICH, UC, US);
- Vilmes, Dep. de Robles, 12 Jun 1943, H.H.Bartlett 20472 (G, MICH, UC, US);
- Dep. Robles, alt. 170 m, 18 Jan 1945, P.García 732 (A, NY);
- La Banda, 24 Jan 1945, P.García 793 (A, NY, UC);
- Quebracho Coto, 15 Mar 1947, F.E.Luna 40 (NY);
- Beltran, 28 Nov 1940, R.Maldonado 523 (F, GH, NY);
Tucumán:
- camino a San Andrés, 4 Feb 1945, A.R.Cuezzo 27 (A);
- Tucumán, Nov 1892, O.Kuntze s.n. (F, MO, NY);
- Sala Ancajulio, 14 Dec 1944, D.Olea 95 (A, NY);
- Sala Ancajulio, 15 Dec 1944, D.Olea 112 (A, NY, UC);
- Tafi Viejo, San Javier, Ruta Prov. 340, alt. 1100 m, 4 Nov 1978, S.A.Renvoize 3362 (MO, NY, US);
- Francas, alt. 780 m, 12 Oct 1925, Schreiter 68587 (A, F);
- Famailla, S Rafael, Río Colorado, alt. 400 m, 5 Nov 1946. B.Sparre 654 (S);
- Trancas, Río Vipos, alt. 600 m, 19 Nov 1946, B.Sparre 779 (S);
- Tapia, 27 Nov 1976, D.Seigler & F.Vervoorst 10103 (ILL);
- Marino, 9 Mar 1944, L.A.Varela 21 (A);
- Tapia, alt. 700 m, 18 Sep 1921, S.Venturi 1345 (MO);
- Tapia, alt. 750 m, 26 Sep 1929, S.Venturi 9510 (MO);
BOLIVIA:
Cochabamba:
- Capinota, alt. 2400 m, 10 Jun 1921, E.Asplund 4143 (S);
- Cochabamba, 1891, M.Bang 1095 (MO, NY, PH);
- recolectado en el km 309 de Santa Cruz, 10 Oct 1977, G.Hartshorn & E.Meneces 2029 (NY);
- Cochabamba, at base of Cerro de San Pedro, alt. 2625 m, 31 Mar 1963, D.Ugent, V.Ugent & M.Cárdenas 4570 (NY, WIS);
- Prov. Campero, Palca, 13 km S of Aiquile, alt. 2000 m, 11 Apr 1963, D.Ugent & M.Cárdenas 4879 (G, WIS);
Chuquisaca:
- 2.5 km S of Camargo, alt. 2370 m, J.Aronson 7796 (EIU);
- Prov. Nor-Cinti, Camargo, 26 km hacia Villa Abecia, alt. 2410 m, 24 Mar 1979, St.G.Beck 692 (NY, US);
- Prov. Oropeza, de Sucre pasando Yotala, alt. 2650 m, 15 Oct 1984, St.G.Beck 8884 (NY);
- Prov. H. Siles, por el camino de Padilla a Monteagudo, alt. 1680 m, 20 Sep 1980, G.Mühlbauer A (NY);
La Paz:
- Prov. Murillo, Abajes, 26 Jun 1921, E.Asplund 4229 (S);
- Prov. Murillo, La Paz-Calacete 13 km hacia Río Abajo, alt. 3100 m, 16 Nov 1980, St.G.Beck 4033 (MO, NY);
- Prov. Larecaja, Consata 26 km hacia Sorata, alt. 1850 m, 16 Dec 1981, St.G.Beck 4956 (NY);
- Stadtteil La Florida, alt. ca. 3200 m, 4 Mar 1982, T.Feurerer & X.Menhofer 10141 (NY);
- 1-3 km N of Inquisivi, alt. 2400-2550 m, 29 Mar 1989, M.Lewis 35383 (MO);
- 3 km N of Inquisivi, alt. 2200-2350, 30 Mar 1989, M.Lewis 35415 (EIU, MO);
- Inquisivi, hike from the Río Khatu Bridge, 3 km N of Inquisivi up the old mule trail to the upper edge of semi-arid forest, alt. 2400 m, 16 Dec 1988, M.Lewis 882080 (EIU, MO);
- Prov. Inquisivi, Río Khatu Bridge, 2 km N of Inquisivi, alt. 2050-2200, 12 Mar 1988, M.Lewis, M.Nee & J.Solomon 8889 (MO);
- Prov. Larecaja, vic. Sorata, Oct 1859, G.Mandon 735 (NY);
- Prov. Inquisivi, Sita road N Inquisivi, alt. 2400m, 29 Sep 1997. J.Müller, A.Berg & J.Heinricks 6377 (MO);
- Prov. Inquisivi, between Yamora & Micayani, 4 km SE of Inquisivi, alt 2550-2650 m, 14 Jan 1989, M.Nee 37582 (EIU, NY). Prov. Murillo, 2 km SE of Mecapaca, alt. 3000 m, 17 Jul 1981, J.C.Solomon 5850 (MO);
- Prov. Murillo, 2 km SE of Mecapaca, alt. 3000 m, 17 Jul 1981, J.C.Solomon 5851 (MO);
- Prov. Inquisivi, 10 km al N de Inquisivi por el camino a Suri, alt. 2100-2200 m, 12 Mar 1988, J.C.Solomon 18124 (MO);
- roadside, Monteagudo, alt 1250m, 10 Apr 2000, K.Wendelberger 159 (MO);
Potosí:
- cerca de Millarea, 2350 m alt., 18 Feb 1979, A.Ceballos, A.Charpin, J.Fernández Casas & E.Valdes-Bermejo 319 (G);
- Prov. Charcas, Est. Sucusuma, sobre la Qda. Sanja Mayu. alt. 2000 m, 29 Feb 1992, B.Mostacedo 310 (MO); (gland is wrong)
Santa Cruz:
- Prov. Cordillera, Santa Cruz ca. 200 km hacia el Sud, Proyecto Abapó Izozóg, 12 Mar 1981, St.G.Beck 6429 (NY);
- Prov. Cordillera, Santa Cruz ca. 200 km hacia el Sud, Proyecto Abapó Izozóg, 12 Mar 1981, St.G.Beck 6460 (NY);
- Prov. Cordillera, Charagua, Finca Itaguazurenda, 16 Sep 1982, A.L.Cabrera & M.M.Gutíerrez 33628 (MO);
- Río Piray, near Santa Cruz city, 15 Nov 1989, D.C.Daly & S.Strickland 6227 (MO, NY, US);
- Curuyugui, 50 km SE of Santa Cruz on Río Parapetí, alt. 350 m, 24 Oct 1991, A.Gentry, R.Foster & M.Peña 75167 (MO);
- Perforación, alt. 500 m, 28 Oct 1991, A.Gentry & R.Forest 75362 (MO);
- Puerto Suárez, 14 Oct 1972, G.Hatschback & P.Scherer 30466 (CM, MO, MU, NY);
- Prov. Cordillera, Bañados del Izozóg, Cachari, 5 km al sur de Cachari, alt. 350 m, 10 Mar 1991, G.Navarro & I.Vargas 270 (MO);
- Prov. Andrés Ibañez, along Río Pirai, 1 km NW of El Torno;
- alt. 495 m, 17 Jan 1987, M.Nee 33598 (MO, NY);
- Prov. Andrés Ibañez, along Río Pirai, 1 km N of La Guardia, alt. 460 m, 20 Jan 1987, M.Nee 33676 (MO, NY);
- Prov. Ichilo, W side of Río Surutu across from Huaytu, alt. 350 m, 2 Aug 1987, M.Nee 35498 (MO, NY);
- Prov. Florida, 5 km SE of Mairana on road to Samaipata, alt. 1550 m, 4 Feb 1988, M.Nee 36169 (MO, NY);
- Prov. Andrés Ibañez, along Río Chore-Chore, 7 km SE of Palmer del Oratorio, alt. 380 m, 9 Dec 1988, M.Nee 37127 (MO, NY);
- Prov. Chiquitos, SE side of Robore, alt. 250 m, 23 Nov 1989, M.Nee 37874 (MO, NY);
- Proc. Cordillera, 5 km N of YPFB gas plant on the Río Grande, alt. 350 m, 2 Dec 1989, M.Nee 37929 (MO, NY);
- Prov. Vallegrande, SW side of Los Sitanos, alt. 1350 m, 27 Dec 1989, M.Nee 38487 (MO, NY);
- Prov. Sara, W side of Río Pirai, alt. 275 m, 7 Nov 1990, M.Nee 39717 (NY);
- Prov. Ichilo, Parque Nacional Amboro, alt. 285 m, 10 Nov 1990, M.Nee 39857 (NY);
- Prov. Andrés Ibañez, 12 km SE of Comunidad Don Lorenzo;
- alt. 300 m, 17 Nov 1990, M.Nee 39978 (MO, NY);
- Prov. Warnes, 13 km ENE of Colonia Okinawa 1, alt. 250 m, 2 Dec 1990, M.Nee 40170 (MO, NY);
- Prov. Andrés Ibañez, along Río Pirai, 1.5 km NW of El Torno;
- alt. 490 m, 24 May 1991, M.Nee 40553 (CM, MO, NY);
- Prov. Florida, Quebrada Yungas, alt. 1825 m, 2 Jun 1991, M.Nee 40713 (MO, NY);
- Prov. Vallegrande, 4 km SW of El Trigal, alt. 1600 m, 8 Mar 1988, M.Nee & J.Solomon 36522 (MO, NY);
- Prov. Vallegrande, 5.5 S of Vallegrande, vic. Santa Rosita, alt. 2050 m, 31 Dec 1988, M.Nee & I.Vargas C. 37467 (MO);
- Prov. Vallegrande, 1 km S of highest point on Mataral to Trigal rd., alt. 1700 m, 24 Dec 1989, M.Nee & I.Vargas C. 38306 (MO, NY);
- Prov. Cordillera, a 6 km en linea recta, al NO de Charagua, alt. 800 m, 12 Apr 1990, R.C.Quevedo S. 75 (MO);
- Prov. Caballero, 2.5 km N of Tambo, alt. 1525 m, 4 Feb 1984, G.Schmitt & D.Schmitt 11 (MO, NY);
- Prov. Ichilo, Parque Nacional Amboro, alt. 400 m. 24 Aug 1985, J.C.Solomon 14016 (MO);
- Bueno Vista, alt. 500 m, 17 Apr 1921, J.Steinbach 5611 (A, F, NY);
- Prov. Florida, Cerro La Negrao, alt. 1400 m, J.Steinbach 8173 (A);
- Prov. Vallegrande, Huasacañada, 5 km S of Vallegrande, alt. 2050 m, 10 Oct 1988, I.G.Vargas 10 (MO, NY);
- Prov. Cordillera, La Brecha, alt. 400 m, 3-5 Jul 1991, I.G.Vargas & E.Tapia 1047 (MO, NY);
- Cordillera Prov., Ibasiri, orilla del Río Parapeti, alt. 390 m, 4 Apr 1992, I.G.Vargas C. & V.Gallo 964 (MO);
- Missiones Guarayos, alt. 250-300 m, Oct 1926, E.Werdermann 2649 (MO);
Tarija:
- Prov. Cercado, cerca San Luis, 15 May 1986, E.Bastíqn 1305 (NY);
- Prov. Gran Chaco, 15 km hacia el N de Yacuiba, alt. 630 m, 23 Sep 1985, Bech, Michel & García 11501 (NY);
- Prov. Arce, 85 km hacia Bermejo, antes de La Mamora, alt. 1530 m, 22 Nov 1986, St.G.Beck 14099 (NY);
- Prov. Arce, 5.5 km N of Mamora, alt. 1500 m, 27 Apr 1983, J.C.Solomon 10177 (MO, UC);
- Prov. Arce, 5 km S of comunidad Guayavillas, alt. 1800-1900 m, 5 May 1983, J.C.Solomon 10507 (MO);
- Prov. Cercado, floodplain of Río Guadalquivir, opposite Tarija, alt. 1800 m, 30 Sep 1983, J.C.Solomon 10913 (CTES);
State not known:
- Songo, Nov 1890, M.Bang 864 (F, MO, NY, US);
- Isapuri, 1 Oct 1901, alt 1550 ft., R.S.Williams 748 (NY);
- Nov 1911, alt. 2450 m, O.Buchtien 3190 (G, HBG, NY);
- Cotana, Nov 1911, O.Buchtien 3189 (US);
ECUADOR:
Guayas:
- Island of Puna, 24 Aug-8 Sep 1936, G.W.Barclay 448 (F);
- Engunga, 1 May 1980, M.Morgan & J.Garcia 11815 (EIU);
PARAGUAY:
Boquerón:
- Misión Santa Rosa, Nov 1987, P.Arenas 1756 (NY);
- 20 km de Mariscal Estigarribia, 20 Feb 1986, I.Basualdo 836 (MO);
- Piste Transchaco, pK 391, 16 May 1984, F.Billiet & B.Jadin 3090 (MO);
- Piste Transchaco, pK 467, 18 May 1984, F.Billiet & B.Jadin 3102 (MO);
- cerca y la Sur de Mariscal Estigarribia, 9 Oct 1980, J.F.Casas & J.Molero 4433 (MO, NY);
- Neuland, Parque Natural, en el borde del paleocauce, 29 Sep 1992, R.Degen & R.Mereles 2791 (MO);
- Filadelfia, 26 Nov 1982, W.Hahn 814 (MO);
- Filadelfia, 6 Dec 1983, W.Hahn 1844 (EIU, ILL, MO);
- Filadelfia, 7 Oct 1979, A.Schinini & E.Bordas 18191 (MICH);
- Campo Loro, 60 km NE of Filadelfia, Chaco Paraquayo, 8 Nov 1988, G.Schmeda 1171 (NY, US);
Central:
- Estero del Ypoâ, 10.5 km S of Villeta on Arroyo Pikysyry, 22 Jun 1993, E.M.Zardini & V.Jara 36420 (MO);
Chaco:
- Chaco Boreal, Mision Santa Rosa, 18 Aug 1993, T.L.Gragson 199 (MO);
- Parque Nacional Defensores del Chaco Madrejón, 15 Aug 1983, W.Hahn 1616 (MO);
- Cerro León, 2 Oct 1979, A.Schinini & E.Bordas 18024 (MO);
- Campo Loro, 60 km NE Filadelfia, 8 Nov 1988, G.Schmeda 1171 (F);
Nueva Asunción:
- Fortín Nueva Asunción, 24 Mar 1986, D.R.Brunner 1645 (MO);
- Parque Nacional Tte. Agripino Enciso, km 656, Ruta Trans-Chaco, 26 Jun 1984, E.L.Little, Jr. 40065 (MO, NY);
- 50 km, N de Mariscal Estigarribia, 26 Jun 1984, E.L.Little, Jr. 40075 (MO);
- Tte Enciso Nat. Park, 22 Sep 1981, M.Vavrek 363 (MO, NY, US);
Presidente Hayes:
- 10 km W of Ftn. Avalos Sanchez, alt 220 m, 15 Mar 1989, J.Aronson 7845 (MO);
- Km 256, 10 Oct 1987, I.Basualdo 1311 (CM, F, MO, NY, US);
- Villa Hayes, 5 Mar 1984, W.Hahn 2185 (EIU, MO);
- Villa Hayes, 17 Aug 1984, L.Peréz 278 (MO);
- Transchaco, camino Asunción-Filadelfia, a 10 km de Filadelfia, 12 Oct 1987, E.Zardini & I.Basualdo 3557 (MO);
- Pilcomayo River, 1888-1890, T.Morong 931 (F, MO, NY, US);
PERU:
Amazonas:
- Prov. Chachapoyas, rd. from Balsas to Huanabamba, shores of Río Maranón, alt. 880 m, 28 Feb. 1976, T.Plowman 5585 (G);
Ancash:
- dry slopes across Rio Santos from Mancos, Callejón de Huaylas, 8 Apr 1970, C.E.Smith, Jr. & J.Bas 4872 (US);
Apurímac:
- Prov. Abancay, Cunyac, alt. 1800 m, 22 Dec 1975, R.Chávez 3074 (MO);
- Prov. Abancay, Cconoc, alt. 2000 m, 26 Apr 1975, R.Chávez 3234 (MO);
- Prov. Abancay, Cconco, alt. 2000 m, 26 Apr 1975, R.Chávez 3240 (MO);
- Prov. Andahuaylas, alt. 2040 m, 26 Mar 1977, H.Ellenberg 7068 (US);
- Prov. Abancay, cerca del puenta de Cunyac, alt. 2100 m, 5 Aug 1954, R.Ferreyra 9832 (MO);
- dry slopes overlooking Río Chalhuanca, 15 km NW of Chalhuanca, alt. 2650-2800 m, 24 Jun 1978, A.Gentry, M.Dillon, P.Berry & J.Aronson 23338A (MO, NY);
- dry slopes along Río Chalhuanca, 15-17 km NW of Chalhuanca, alt. 2600-2650 m, 25 Jun 1978, A.Gentry, M.Dillon, P.Berry & J.Aronson 23359 (MO, NY);
- Prov. Abancay, Valle Matara, alt. 2300 m, 29 Aug 1983, L.v.d.Hoogte & C.Roersch 1452 (MO):
- Prov. Aymaraes, Chalhuanca, alt. 2500 m, 14 Nov 1981, L.v.d.Hoogte & C.Roersch 2427 (F);
- Prov. Abancay, steep hills above Río Pachachaca, 1-2 km below Casinchihua, alt. ca 2300-2500 m, 21 Dec 1962, H.H.Iltis, C.M.Iltis, D.Ugent & V.Ugent 813 (NY, WIS);
Ayacucho:
- Huanta, alt. 2800 m, 19-21 May 1929, E.P.Killip & A.C.Smith 23338 (NY, US);
Cajamarca:
- Prov. Jaén, entre Puente 24 de Julio y Chamaya, alt. 380-400m, 27 Jun 1959, R.Ferreyra 13681 (F, MO);
- Prov. Jaén, cerca a Bellavista, Río Marañón, alt. 400-500 m, 14 Jul 1982, R.Ferreyra & S.Sánchez 19602 (US) 0.5 km E of Pucara, alt. 860 m, 11 Jun 1978, A.Gentry, M.Dillon, J.Aronson & C.Díaz 22698 (G, MO);
- 9-16 km E of Pucara on road to Bagua, alt. 800-835 m, 11 Jun 1978, A.Gentry, M.Dillon, J.Aronson & C.Díaz 22726 (F, G, MO);
- 73 km NE of Pucara on rd. to Bagua, 9 km E of Chamaya, 11 Jun 1978, A.Gentry, M.Dillon, J.Aronson & C.Díaz 22768 (F, G, MO);
- cerca la Puerto Cuyca, 1 May 1987, S.Llatas Quiroz 2195 (F);
- Prov. Celendín, Chachapoyas-Cajamarca rd., alt. 1300-1500 m, 28 May 1984, D.N.Smith & J.Cabanillas 7240 (EIU, MO);
- Condebamba valley, Cajabamba-Cajamarca road, alt. 2100-2600 m, 15 Feb 1983, D.N.Smith & R.Vasquez M. 3402 (ISC);
Cuzco:
- Prov. Convención, Pawac Portage, below Hacienda Pomobamba, Río Apurimac, alt. 1500 m, 7 July 1981, E.W.Davis, N.Marshall, J.Tichenor & M.Lessner 1267 (F, NY);
- Prov. Anta, Río Apurimac, below Limatamba, alt. 2400-2500 m, 10 Jan 1984, A.Gentry, I.Peralta V. & A.Arestegui P. 44109 (EIU, MO);
- Prov. Urubamba, alt. 2820 m, 14 Mar 1950, C.Vargas C. 9291 (MO);
Huánuco:
- 10 km N of Huánuco on rd. to Tingo Maria, alt. 1900 m, 13 Apr 1977, A.Gentry, J.Revilla, D.Daly & D.Alfaro C. 19243 (F, MO, NY);
La Libertad:
- Pacasmayo, 26 Jun 1914, J.N.Rose 18525 (G, NY, US);
Lambayeque:
- Prov. Lambayeque, Pampa de Olmos, alt. 100-120 m, 18 Sep 1975, R.Ferreyra 18560 (MO);
Paita:
- Talara, 1925, O.Haught 56 (US);
Tumbes:
- Prov. Zarumilla, Dtto. Matapalo, cerca de Campo Verde, alt. 600-800 m, 23 Dec 1967, D.R.Simpson & J.Schunke V. 465 (F);