You selected: Solanum glaucophyllum Desf. Cat. Pl. Hort. Par. 3: 396. 1829. Status: Accepted.

Solanum glaucophyllum Desf., Cat. Pl. Hort. Par. 3: 396. 1829. Type: Cultivated in Botanical Garden at Paris, Desfontaines s.n. (type unknown).

Last edited by Bohs, L. May 2004. Description based on taxon concept by Bohs, L. pages 43-51 in Bohs, L. 2001. Revision of Solanum section Cyphomandropsis (Solanaceae). Syst. Bot. Monog. 61: 1-85.

Habit

Rhizomatous shrubs or slender treelets ca. 0.5-4 m tall. Stems glabrous (rarely moderately to densely puberulent-pubescent with unbranched hairs), usually light-colored and smooth.

Sympodial structure

Sympodial units many-foliate.

Leaves

Leaves simple, the blades 6-18 x 0.6-3.5 (5) cm, 4.5-10 (15) times as long as wide, narrowly elliptic, succulent or fleshy with the midribs and margins often thickened and whitish, surfaces glaucous in fresh material, glabrous adaxially and abaxially (rarely moderately to densely puberulent-pubescent with unbranched hairs); base tapered to decurrent; margin entire; apex acute; the petioles 1.5 cm or less, glabrous, often slightly winged.

Inflorescences

Inflorescences 3.5-9 cm, branched, sometimes highly so, with 20-50 or more flowers, all flowers perfect, the axes glabrous (rarely moderately puberulent-pubescent with unbranched hairs); peduncle 1-3.5 cm; rachis 2-7 cm; pedicels 12-15 mm, ca. 15-20 mm in fruit, spaced 1-15 mm apart, articulated at the base.

Flowers

Flowers with the calyx the radius 2-3 mm, the lobes 0.5-1.5 x 1.5-2.5 mm, deltate, acute at apex, glabrous except for some sparse puberulence at margin. Corollas 2.5-4 cm in diameter, the radius 10-30 mm, rotate-stellate and plicate, chartaceous, whitish to pink or violet, often with a white central star, the the tube 5-8 mm, the lobes 4-10 x 5-10 mm at base, broadly triangular, apiculate at apices, moderately to densely puberulent abaxially, especially on distal parts of lobes and plicae, glabrous adaxially except for a few sparse hairs at tips of lobes. Anthers 5-7 x 2 mm, usually connivent, yellow to orange yellow, ovate, the pores directed distally. Ovary glabrous; style 5-7 x 0.25-0.5 mm in diameter, cylindrical, glabrous; stigma truncate.

Fruits

Fruits 0.75-2 x 0.75-2 cm in diameter, globose, sometimes apiculate at apex when young, obtuse at apex when mature, dark purple or blue-black and glaucous when ripe, glabrous; stone cell aggregates absent.

Seeds

Seeds ca. 4-6 x 3.5-4 mm, angled, smooth or with minute scalloped ridges.

Chromosome number

n = 12 voucher: Bohs 2530 ( Bohs, 2001)

Molecular data

chloroplast ndhF sequence: GenBank U72753 (http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=1644489). nuclear ITS sequence: GenBank AF244714 (http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=7533134). nuclear ITS sequence: GenBank AY523897 (voucher: Bohs 2530, UT). Sequence not yet released (7/04).

Distribution

Bolivia, southern Brazil, Paraguay, northern Argentina, and Uruguay; low, swampy ground at margins of marshes and ponds in seasonally inundated areas; ca. 0-600 m.

Common names and uses

Local names. Argentina: Corcho del agua (Torres 9), duraznillo (Boffa 133; Gibson s.n.; Huidobro 1452; Schulz 2053; Tweedie s.n.), duraznillo blanco (Gibson s.n.; Lefebre s.n.), duraznillo de bañado (Schinini 9033), duraznillo hediondo (Hunziker 310), duraznillo negro (Cordini 61), na’kyet (Arenas 2000), palo hediondo (Boelcke 1396), palo-né or paloné (Boelcke 1396; Irigoyen 221), varilla (Burkart 8727, Hunziker 25304), yaa’tuk or ye:’tuk (Maranta & Arenas 153). Bolivia: Bobo (Murquia 502). Paraguay: sinhejúk (Arenas 714). Uruguay: duraznillo blanco (Gallinal et al. A831; Gibert 140), duraznillo negro (Osten 21883). Brazil: espichadeira (Prance et al. 26173), espixadeiro (Macedo et al. 1209). Uses. The dried stems are used for firewood and for wattling (Burkart 8727; Gibson s.n.). Plants are used medicinally as a purgative (Morton, 1976; Gibert 140; Tweedie s.n.). Foliage is very toxic for cattle and other animals (Prance et al. 26173; Schulz 2053).

Phenology

Flowering and fruiting throughout the year, with a peak in November through March.

Other information

Self-incompatible (Bohs, 2001).

Phylogeny

Solanum glaucophyllum belongs to the Cyphomandra clade of Solanum along with other species traditionally recognized in sections Pachyphylla and Cyphomandropsis (Bohs, in press a). Within the Cyphomandra clade, ITS data ally S. glaucophyllum with S. cylindricum in some trees and analyses, but this relationship does not receive strong parsimony bootstrap support and may be unreliable due to sampling artifacts (Bohs, in press b).

Commentary

Solanum glaucophyllum can distinguished from other species in the section by its usually glabrous, glaucous, lanceolate to elliptic leaves with decurrent bases and short, slightly winged petioles, its rotate-stellate corolla with a relatively long tube and broad lobes, and its globose, purple-black, glaucous fruits. The stems are smooth and usually light yellowish or whitish, and the leaf midribs and margins are often thickened, inrolled, and whitish or cream-colored. Throughout its range, S. glaucophyllum exhibits great morphological variability, especially in leaf shape, which ranges from elliptic to nearly linear, and in corolla and fruit size. Narrow-leaved plants generally have smaller flowers and fruits than those with broader leaves, and narrow-leaved forms seem more common in the northern part of the geographical range of this species.

This species grows commonly in flooded or swampy ground, where it forms thickets, known as “varillales” or “duraznillales”, of virgate stems from spreading rhizomes (Cabrera & Zardini, 1978; Okada et al., 1977). It is reportedly deciduous in winter (Okada et al., 1977). The fruits apparently float readily and may be dispersed by water (Nee 37532); they are also eaten by birds (Gibson s.n.).

Solanum glaucophyllum is of economic importance mainly because it causes a disease, “enteque seco” or “espichamento,” of grazing animals (D’Arcy, 1974; Morris, 1977; Okada et al., 1977; Wasserman, 1974). The disease is characterized by calcification of soft tissues, frequently leading to death, and has caused losses of millions of dollars annually to livestock ranchers in Argentina (Cabrera, 1983). The active principle of S. glaucophyllum has been shown to be a vitamin D-like substance that increases calcium and phosphorous absorption (Morris, 1977; Wasserman, 1974). Extracts of S. glaucophyllum are currently being tested for activity as bone growth factors useful in human and veterinary medicine (Morris, 1977; B. Barr, pers. comm.).

Solanum glaucophyllum is most similar to S. confusum of northwestern Argentina and adjacent Bolivia, and to S. matadori of southeastern Brazil. Solanum confusum differs from S. glaucophyllum in being frequently pubescent, and by having leaves with longer petioles, shorter corolla tubes with narrower corolla lobes, and yellow to orange fruits. Solanum matadori is glabrous like most plants of S. glaucophyllum, but it has stellate corollas, green rather than glaucous leaves, and is known only from Santa Catarina, Brazil.

On the basis of morphological characters, Child (1986) considered S. glaucophyllum to be unrelated to section Cyphomandropsis. He removed it to its own section, Solanum section Glaucophyllum Child, included within Solanum subgenus Solanum. Dottori (1995) investigated fruit and seed morphology of this species and concluded that there were significant differences in fruit and seed characters between S. glaucophyllum and the other species of section Cyphomandropsis (S. confusum, S. fusiforme, and S. stuckertii) she investigated. These differences included epicuticular wax covering the fruit exocarp, ventilation cracks rather than stomata in the fruit surface, and seed coat cells with a distinct shape and pattern of wall thickening. While S. glaucophyllum exhibits some unique characters and may represent an isolated clade within the section, its tapered anthers, large angled seeds, and large chromosomes establish it as a member of section Cyphomandropsis. Molecular data from chloroplast and nuclear sequences confirm that S. glaucophyllum belongs to the Cyphomandra clade (Bohs, in press a, b).

A few collections (Arenas 1407, Döbereiner & Tokarnia 794, 797, 800, Hatschbach & Guimarães 21941, Schaller 301) from ca. 20°S latitude in the Río Paraguay drainage are anomalous in being moderately to densely puberulent-pubescent on the axes and leaves. The pubescent plants were recognized as a separate variety by C. V. Morton, but under an unpublished herbarium name. Because they conform to typical S. glaucophyllum in all other respects and occur within the range of the glabrous forms, I do not consider them to be taxonomically distinct.

Solanum glaucophyllum has often been confused with S. amygdalifolium Steud. (also known under the synonyms S. angustifolium Lam., S. persicifolium Mart., S. handelianum Morong, S. brittonianum Morong), which is sympatric and is found in similar habitats. Solanum amygdalifolium, however, is a twining or scandent vine with strongly ridged stems and ovate-lanceolate leaves. The anthers dehisce by broad pores that open into longitudinal slits, and the seeds are much smaller than those of S. glaucophyllum. Solanum amygdalifolium belongs to the dulcamaroid group of Solanum (sects. Dulcamara, Jasminosolanum and relatives; Dulcamaroid clade sensu Bohs, in press a) and, though strikingly convergent in several morphological features, is not closely allied to section Cyphomandropsis.

Desfontaines (1829) published the name S. glaucophyllum in reference to a plant in cultivation in the Paris Botanic Garden. I was unable to locate a type specimen for this name, and a search of the collections at P by Sandra Knapp in 1998 failed to turn up any candidates. D’Arcy (1974) cited a specimen at MPU labelled as “S. glaucophyllum Hort. Paris Cat. Ann. 1829, p. 396” and speculated that it may be a type. Sandra Knapp in 1998 found this specimen at MPU, but with the additional annotation “h.m. Jul 1840,” indicating that it was grown at the Montpellier Botanic Garden in 1840 and thus cannot be a type. Specimens from BR, FI, and G attest to the cultivation of this species in the Paris Botanic Garden in the early nineteenth century. Dunal (1852) indicates that both the names S. glaucophyllum and S. glaucum were in use to refer to this species. It is likely that S. glaucophyllum was described from living material, and that no type specimen exists. Desfontaines original description is adequate to establish the name of this taxon.

Later authors Kuntze (1898), Chodat (1916), and Hassler (1918) proposed a number of infraspecific taxa based on leaf color and size. All of these taxa are subsumed within the range of variation that are encompassed in my concept of the species.

Solanum glaucophyllum was cultivated in French botanical gardens at Angers, Hyeres, and Dijon in the mid-nineteenth and early twentieth century, and specimens are also known from Japan, Nepal, and the United States (Pensacola, Florida). It is not known whether the Asian plants were cultivated or adventive, but probably at least the Nepal collection was a deliberate introduction. D’Arcy (1974) speculates that S. glaucophyllum was introduced to Florida in ships’ ballast. It has not been collected from Florida since 1901.

Additional specimens examined

BRAZIL. Mato Grosso: Mun. Paconé, Fazenda Cassange, 11 Jul 1969 (fr), Döbereiner & Tokarnia 699 (US); Transpantaneira Highway, Pocone, Fazenda Santa Rosa, km 40, 11 Sep 1978 (st), Macedo et al. 1209 (NY); Transpantaneira Highway, Fazendo Jofre, 56°30’W, 17°10-17’S, 13 Jun 1979 (fl, fr), Prance et al. 26173 (NY); Transpantanal Highway, Fazenda Jofre, 2 Nov 1978 (fl), Schaller 258 (NY). Mato Grosso do Sul: Mun. Corumbá, Fazenda São Sebastião, 3 Jul 1970 (fl), Döbereiner & Tokarnia 717 (L, US); Mun. Aquidauana, Fazenda Tabaco, 28 Aug 1971 (fl, fr), Döbereiner & Tokarnia 794 (F, US); Mun. Corumbá, Fazenda Brasil Fronteira, 2 Sep 1971 (fl), Döbereiner & Tokarnia 797 (US); Mun. Aquidauana, Fazenda Tupacireta, 5 Sep 1971 (fl, fr), Döbereiner & Tokarnia 800 (F, US); Mun. Aquidauana, Fazenda Santa Cruz, 17 Jul 1969 (fr), Hatschbach & Guimarães 21941 (F); Mun. Miranda, Rio Miranda, 13 Jun 1976 (fl, fr), Hatschbach 38640 (F, US); Porto Murtinho, 18 Mar 1985 (fl. fr), Hatschbach & da Silva 49222 (NY, US); Pantanal, Fazenda Miranda, 5 Apr 1979 (fl, fr), Schaller 301 (NY); Mun. de Miranda, Marimbondo, 60 km N de Guaicurus, 80-200 m, 5 Jun 1973 (fl, fr), Silva 80 (SP). Rio Grande do Sul: I.A.S, Pelotas, margem do Canal do I.A.S., São Gonçalo, 4 Dec 1957 (fl), Sacco 816 (F, NY). BOLIVIA. Beni: Prov. Cercado, 7 km SW of Trinidad, vic. Puerto Almacen, along the Río Ibare, 14°52’S, 64°57’W, ca. 200 m, 7 Jan 1989 (fl, fr), Nee 37532 (G, LPB, NY, USZ). Chuquisaca: Prov. Luis Calvo, margenes del lugar “Arbol Solo,” 7 Sept 1986 (fl, fr), Murquia 502 (LPB); Prov. Luis Calvo, El Salvador, Laguna Seca, Campo Quemado, a orillas de represa, 9 Dec 1992 (fl, fr), Penseiro & Marino 4423 (SI). Santa Cruz: Prov. Velasco, San Miguelito, ca. 500 m, Apr 1974 (fl, fr), Arnold s.n. (K, US); Prov. Andrés Ibañez, Jardín Botánico de Santa Cruz, 12 km E of center of Santa Cruz on road to Cotoca, 17°47’S, 63°04’W, 375 m, 5 June 1998 (fl), Bohs & Nee 2767 (USZ, UT); Prov. Andrés Ibañez, en Tarope, cantón Cotoca, 20 Dec 1989 (fl), Coimbra 1006 (JBSC); Prov. A. Ibañez, Sur de Campanero, camino entre Cotoca y Payla, 375 m, 24 June 1992 (fl, fr), Coimbra 2320 (JBSC); Prov. Nuflo de Chávez, Estancia San Miguelito, 200 km NE de Santa Cruz, Puesto Corrales, 4 km NW del puesto, 17°09’S, 61°43’W, 270 m, 30 Mar 1995 (fl, fr), Fuentes 586 (USZ); Curuyuqui, trail along Río Parapetí ca. 8 km SW of airstrip, 18°50’S, 62°20’W, 350 m, 27 Oct 1991 (fl, fr), Gentry & Foster 75300 (NY, USZ); Bañados de Izozog, 3 Apr 1990 (fl), Navarro et al. 19 (LPB); Prov. Cordillera, Bañados de Izozog, Monte Alto, bosque inundado del Río Parapetí, 400 m, 12 Mar 1991 (fl, fr), Navarro & Vargas 346 (BOLV, LPB); Prov. Andrés Ibañez, Jardin Botánico de Santa Cruz, 12 km E of center of Santa Cruz on road to Cotoca, 17°47’S, 63°04’W, 375 m, 15 July 1987 (fl), Nee 35186 (JBSC, LPB, NY); Prov. Andrés Ibañez, 12 km SE of Comunidad Don Lorenzo, Estancia Caracore, 0.5 km W of the Río Grande (= Río Guapaí), 17°51’S, 62°47’W, 300 m, 17 Nov 1990 (fl, fr), Nee & Coimbra 39996 (JBSC, LPB); Prov. Andrés Ibañez, 7 km SE of comunidad Don Lorenzo, 17°50’S, 62°49’W, 310 m, 5 Dec 1990 (fl, fr), Nee 40174 (LPB, USZ); Prov. Andrés Ibañez, 10 km S de Cotoca, camino a la localidad de Paurito, 17°49’0”S, 62°57’05”W, 450 m, 26 Apr 1990 (fl, fr), I. Vargas 518 (USZ); Prov. Cordillera, Bañados de Izozog, 500 m al N de las instalaciones de Estancia Cachari, 18°55’S, 62°25’W, ca. 350 m, 5-15 Jan 1993 (fl, fr), I. Vargas et al. 1935 (USZ); Missiones Guarayos - Santa Cruz de a Sierra, ca. 250-300 m, Oct 1926 (fl), Werdermann 2625 (K, LPB, MO, S). PARAGUAY. Alto Paraguay: Puerto Diana, 5 km de Bahía Negra, Jan 1976 (fl, fr), Arenas 1407 (CTES); Estancia Sorpresa, 28 July 2002 (fr), González Parini et al. 1076 (FCQ); Palmar de las Islas, Estancia San José, 7 Jan 1998 (fl), Insua 14 (FCQ); Estero Doña Cynthia, 28 Sept 1988 (fl), Keel 1626 (FCQ); desembocadura del Río Negro en el Paraguay, 20°09’15.6”S, 58°18’12.4”W, 4 July 1997 (fl), Mereles 6779 (FCQ). Boquerón: camino de Filadelfia a Mcal. Estigarribia, 10 Oct 1987 (fl), Basualdo & Zardini 1262 (FCQ); cerca y al S de Mariscal Estigarribia, 23 Oct 1980 (fl, fr), Casas & Molero 4434 (MO, NY); 9.6 km al S del Cruce de Picadas Petroleras, 25 May 1994 (fr), Degen & Mereles 3201 (FCQ); Zinfunké, orillas del Río Pilcomayo, 30 Oct 1985 (fl), Mereles 704 (CTES, FCQ); Area Linea 12/3, 2 Sept 1992 (fl), Mereles & Degen 4671 (FCQ); Picada a Mistolar, 4.2 km S de linea 10, 21 May 1994 (fl, fr), Mereles & Degen 5648 (FCQ); Tte. Ochoa, 11 Dec 1993 (fl, fr), Mereles & Degen 5514 (FCQ); 11 km de la Pista 4ta. División de la Infantería, S/Linea 10, 24 May 1994 (fr), Mereles & Degen 5685 (FCQ); Estancia Catán, 23°27’47”S, 60°48’54”W, 26 Oct 1994 (fl), Mereles & Degen 5847 (FCQ); Puerto Casado and vicinity, 26 Oct 1956 (fl, fr), Pedersen 4198 (BR, C). Capital: Asunción, 27 Nov 1888 (fl, fr), Morong 181 (BM, F, GH, K, MO, NY, US, WIS); Puerto Itá Enramada, 6 Sept 1976 (fl, fr), Schinini & Bordas 13344 (CTES, G, MO); Fortín Infante Rivarola, 21°42’S, 62°26’W, 250-300 m, 12 Oct 1987 (fl), Spichiger et al. 2247 (FCQ). Central: Piquete-cué (Jukyty), 18 May 1985 (fl, fr), Bordas 4061 (CTES); Salado, Laguna Ypacarai, Jul 1914 (fl), Chodat 33 (G); Laguna Ypoa, Sep 1914 (fl, fr), Chodat 35 (G); Ruta Nva. Italia – Carapeguá, arroyo Caañabé, 12 Apr 1989 (fl), Degen 1441 (FCQ); Laguna Ipacarary, Nov 1902-Mar 1903 (fl), Fiebrig 463 (E, F, G, GH); Areguá, orillas del Lago Ypacaraí, Dec 1982 (fl, fr), Mereles 163 (CTES, FCQ, NY); Areguá, costa del Lago Ypacaraí, 5 jan 1982 (fl, fr), Mereles 1017 (G); Luque, Compañia, Tarumandy, Aug 1991 (fl), Mereles 4042 (FCQ); Luque, Tarumandy, Propiedad Nueva Asunción, 9 Feb 1992 (fl, fr), Mereles 4420 (FCQ); Ipacaraí, orillas Lago Ipacaraí, 14 Aug 1985 (fl), Ortíz 510 (FCQ, G); Nueva Italia, Compañia Jukyty, cercanías del arroyo ka’a ñave, 14 Mar 1985 (fl, fr), Pérez et al. 496 (CTES); Simpio a Emboscada, cerca del Cruce del Salado, July 1971 (fl), Schinini 3919 (G); Ypacaraí, Dec 1985 (fl), Soria 1044 (FCQ); Aregua, orillas del Lago Ypacarai, 25°20’S, 57°20’W, 19 Mar 1990 (fr), Soria 4181 (FCQ, NY, TEX); Trinidad, Bahia P. Caballero, 19 Mar 1990 (fl, fr), Sparre & Vervoorst 67 (S). Chaco: Laquerenza, cauce seco del Río Zimane, Mar 1989 (fl, fr), Mereles & Ramella 2692 (CTES, FCQ); Palmar de las Islas, Laguna Palmar, 6 Mar 1989 (fl), Mereles 2893 (FCQ); Mayor Pedro Lagerenza, cauce seco del Río Timane, 60°45’W, 20°0’S, 4 Apr 1978 (fl, fr), Schinini & Bordas 14871 (CTES); Cerro León, 60°15’W, 20°26’S, en Laguna Nueva Misión, 2 Oct 1979 (fl), Schinini & Bordas 17881 (CTES, F, G, MO, NY). Concepción: Chaco y prope Concepción, Oct 1914 (fl), Chodat 34 (G); near Concepción, shores of Río Paraguay, Sep 1901 (fl, fr), Hassler 7483 (BM, G, GH, K, LIL, MO, NY, S, W). Cordillera: San Bernardino, Ciervo-cuá, July 1988 (fl), Mereles 1282 (FCQ); Altos, Cnia. Bernal Cué, orillas del arroyo Itagaza, 18 June 1973 (fl), Schinini 6721 (CTES); Río Salado, 57°22’W, 25°13’S, 27 Dec 1985 (fl, fr), Schinini & Mereles 24560 (CTES, G). Ñeembucú: Curupayty, Humaitá, 11 Nov 1978 (fl), Bernardi 18517 (BM, F, MO, NY); Dist. Cambacu, Albera, Mar 1975 (fl, fr), Walter 93 (BM). Nueva Asunción: Tte. Montanía, km 8 – Estancia Nva. Asunción km 20 N, 17 Oct 1992 (fl, fr), Mereles & Degen 4769 (FCQ); camino Mcal. Estigarribia a Tte. Pico, 19 Feb 1987 (fl), Soria 1334 (FCQ). Presidente Hayes: Estancia Loma Pyta, 23°40’S, 59°35’W, 12 Feb 1978 (fl, fr), Arenas 714 (NY); Colonia Menno, Paratodo, 30 Jan 1976 (fl, fr), Arenas 1490 (CTES); Ruta Trans-Chaco, about 1 km after crossing from Asuncíon over the Río Paraguay on Puente Remanso en route to Villa Hayes, 25°09’32.7”S, 57°33’36.1”W, ca. 65 m, 29 Nov 2003 (fl), Bohs et al. 3201 (FCQ, NY, PY, UT); Ruta a General Bruguez, Ninfa, 24°44’16.4”S, 58°23’16.2”W, 24 Oct 1995 (fl), Degen & Mereles 2414 (FCQ); Estancia Maroma, Arroyo Conchita, a 3 km del casco de la estancia, camino a Arroyo Conchita, 23°33’20”S, 57°54’33”W, 23 Mar 1995 (fl, fr), Degen & Mereles 3263 (FCQ); 5 km sobre ruta a Ftín. Cadete Pando, a 60 km de Ninfa, 24°26’01.9”S, 58°54’09”W, 31 Oct 1995 (fl), Degen 3428 (FCQ); Colonia Falcón, lado del Río Pilcomayo, 25°14’S, 57°44’W, 30 Oct 1987 (fl), Keel et al. 1082 (FCQ); Villa Hayes, Estancia La Golondrina, 24°55’S, 57°40’W, 9 Sep 1982 (fl), Hahn 671 (BH, NY); Zona Confuso, 28 Oct 1988 (fl), Mereles 1566 (FCQ); km 120, Ruta Trans Chaco, 21 Feb 1989 (fl), Mereles 2592 (CTES, FCQ); Tinfunke, Estancia Brusquetti, 20 Dec 1989 (fl, fr), Mereles & Geissler 3327 (FCQ); 30 km NE, Estancia Palo Santo, 17 June 1993 (fl, fr), Mereles & Degen 5189 (FCQ); Estancia Cunu’ú, 23°54’S, 59°39’19”W, 22 Sept 1993 (fl), Mereles & Degen 5393 (FCQ); Estancia Quebracho, Estero, 23°46’03”S, 59°52’19”W, 23 Sept 1993 (fl), Mereles & Degen 5396 (FCQ); Estancia Maroma, 90 km E de Pozo Colorado, 23°33’20”S, 57°54’33”W, 22 Mar 1995 (fl), Mereles & Degen 5903 (FCQ); Estancia Lapacho, 23°16’30”S, 58°20’53.4”W, 18 July 1995 (fl, fr), Mereles & Degen 6050 (FCQ); Tte. Esteban Martínez y alrededores, 28 Nov 2000 (fl, fr), Mereles et al. 8292 (FCQ); Benjamín Acerval, a costado de la Ruta trans-chaco, 7 Sept 1988 (fl), Rueslede 50 (FCQ); Chacoí, campo inundable del Río Negro, 25°12’S, 57°38’W, 5 Dec 1989 (fl, fr), Schinini 26770 (CTES, NY); Ruta Trans Chaco, 8 May 1985 (fl, fr), Schmeda 706 (FCQ, US); Estancia Tinfunque, Puesto Clorinda-í, 23°45’S, 60°08’W, 100-150 m, 20 Sept 1987 (fl), Spichiger et al. 2024 (FCQ). San Pedro: Puerto Antequara, Dec 1992 (fl, fr), Soria 5539, 5544 (FCQ); Puerto Rosario, island in Río Paraguay, 27 Jan 1958 (fl, fr), Woolston 937 (C, K, S, SP, U, US). Without Department: Linea 1, 60 km E de Agua Dulce, Tyto. Bahía Negra, 19°59’04”S, 60°03’16.8”W, 13 Apr 1997 (fl), Mereles 6564 (FCQ); Mcal. Estigarribia, camino a Santa Teresita, 1 June 1980 (fl, fr), Schmeda 192 (CTES, FCQ). ARGENTINA. Buenos Aires: Pdo. Maipu, Ruta 74 a 7 km de Las Armas, 6 Dec 1967 (fl, fr), Boelcke et al. BAA 5888 (CTES); Pdo. Magdalena, Punta Indio, 21 Feb 1973 (fl, fr), Boelcke et al. BAA 12613 (CTES); Depto. La Plata, Berisso, Feb 1945 (fl, fr), Boffa 133 (F); alrededores de La Plata, Los Talas, 26 Mar 1933 (fl), Cabrera 2795 (NY); Tigre, 13 Dec 1936 (fl), Castro BAB 52328 (K); Laguna de Monte, 13 Jan 1940 (fl, fr), Cordini 6 (SI); Laguna La Adela, 6 Apr 1939 (fl), Dawson 663 (NY); Depto. Campana, Campana, 10 m, 27 Nov 1938 (fl, fr), Eyerdam & Beetle 23064 (BH, G, GH, MO); Los Yngleses, Ajó, near Cape San Antonio, mouth of the Río de La Plata, 20 Oct 1915 (st), Gibson s.n. (BM); Depto. La Plata, Punta Lara-La Plata, 4 Apr 1968 (fl, fr), Gómez Sosa 61 (CTES); Quilmes, no date (fl), Hicken 948 (NY); Azul, 15 Dec 1944 (fl), Huidobro 1452 (NY, S); Partido de Campana, Otamendi, 3 Jan 1940 (fl), Hunziker 310 (NY); La Plata, 1890 (fl), Lefebre s.n. (BR); vicinity of La Plata, 12 Dec 1986 (fl, fr), Passarelli 1 (LP); same locality and date (fl), Passarelli 2 (LP); Miramar, Partido Gral. Alvarado, 1 Apr 1951 (fl, fr), Pedersen 1045 (C, BR, S, US); Quilmes, Plátanos, 9 Dec 1912 (fl), Rodríguez 111 (A, SI); near Buenos Aires, 1884 (fl, fr), Saint-Yves 156 (G); Campos de Hudson SE of the capital, 14 Dec 1939 (fl), Santesson 75 (S); 2 km SW of Ensenada on Ruta Provincial 10, 7 Dec 1978 (fl, fr), Solomon & Solomon 4026 (CTES, MO); outskirts of Buenos Aires, no date, Tweedie s.n. (K); Buenos Aires, Barracas al Sud, 3 May 1902 (fl), Venturi 76 (K, S). Chaco: Depto. Resistencia, Margarita Belén, 17 Jan 1948 (fl, fr), Aguilar 1212 (BR, MO); Makallé, 7 Dec 1939 (fl, fr), Birabén 63 (LP); Depto. Primero de Mayo, Antequera Cerrito, ca. 55 m, 4 Nov 1986 (fl, fr), Charpin & Eskuche 20145 (G); Las Palmas, Nov 1917 (fl, fr), Jörgensen 2204 (GH, MO, US); Depto. Gral. Güemes, Zaparinqui, 13 km S de J. J. Castelli, 26°04’S, 60°33’W, 25 Jan 1980 (fl, fr), Piccinini & Hilfer 4230 (CTES); Colonia Benítez, 5 May 1955 (fl, fr), Schinini 9033 (CTES); Depto. Donovan, 13 km N de La Verde, Estancia Dos Tranqueras, 21 Sept 1982 (fl, fr), Schinini 22746 (CTES); Colonia Benítez, 18 Mar 1937 (fl, fr), Schulz 2053 (CTES). Córdoba: Depto. Marcos Juárez, unos 8 km al SE de Camilo Aldao, a casi 20 km al W del límite con Santa Fé, 1 Dec 1966 (fl), Hunziker 18957 (UT). Corrientes: Depto. Bella Vista, Ruta 117, 7 km E de Ruta 27, 11 Mar 1976 (fl, fr), Ahumada 309 (CTES, MO); Depto. Itatí, Ruta Nac. 12, 60 km E de Itatí, Arrocera Rzepecki, 28 Feb 1977 (fl), Ahumada et al. 858 (CTES, F); Depto. Esquina, 47 km W de Sauce, Ruta 126, 23 Oct 1977 (fl), Ahumada et al. 1449 (CTES, LP); Depto. San Cosme, 28 km E de Corrientes, 26 Feb 1978 (fl), Ahumada 1653 (CTES); Depto. San Miguel, 12 km. S de Caa-Catí, Ruta 5, 13 Mar 1978 (fl), Ahumada 1762 (CTES, MO, WIS); Depto. General Paz, 29 km S de Caa-Catí, Ruta 13, 17 Mar 1978 (fr), Ahumada 2021 (CTES); Depto. General Paz, 26 km W de Caa-Catí, Ruta 5, 18 Mar 1978 (fl), Ahumada 2105 (CTES); Depto. Esquina, sobre Ruta 25, 59°15’W, 29°48’S, 2 Mar 1980 (fl), Ahumada et al. 3458 (CTES); Depto. San Cosme, 4 km E de Paso de la Patria (desvío a Pto. González), 20 Mar 1975 (fl), Arbo et al. 721 (CTES, MO, WIS); Depto. San Roque, Estancia Caaguazú, 11 km NE de Chavarría, camino a Tacuaritas, approx. 4 km al S del casco, orrilas del Río Corriente, 22 Oct 1996 (fl), Arbo et al. 6675 (LPB); Dept. Mercedes, Macrosistema Ibera, nacimiento del Río Corriente, 28°43’S, 58°07’W, 27 Aug 1998 (fr), Arbo et al. 7968 (FCQ); Goya, 26 Nov 1945 (fl), Boelcke 1396 (SI); Depto. San Roque, San Roque, bajos del Santa Lucía, 6 Dec 1975 (fl, fr), Cristóbal et al. 1437 (AAU, G, CTES); Paso de La Patria, 11 Jan 1975 (fl, fr), Cuezzo et al. 11271 (LIL); Depto. Itatí, Pueblo Cercanía, 28 Sep 1944 (fl), Ibarrola 890 (NY, S); Depto. Empedrado, 4 leguas al E de El Pollo, 30 Aug 1945 (fl), Ibarrola 3205 (F, L); Depto. Bella Vista, camino de Bella Vista a San Roque, 6 km del empalme a Bella Vista, 28 Sep 1975 (fl), Irigoyen 221 (AAU, CTES, G, WIS); Depto. San Cosme, Paso de La Patria, orillas del Río Paraná, 7 May 1967 (fl, fr), Krapovickas & Cristóbal 12964 (CTES); Depto. Bella Vista, 7 km S de Bella Vista, 28 Jan 1971 (fl, fr), Krapovickas & Irigoyen 17844 (CTES); Depto. Empedrado, Río Empedrado, Ruta Nacional 12, 26 Sep 1971 (fl), Krapovickas et al. 19897 (CTES); Depto. Esquina, Esquina, costa del Río Corriente, 30 Nov 1974 (fl, fr), Krapovickas et al. 26742 (CTES, G, MO); Depto. Esquina, islas frente a Esquina, 30 Aug 1974 (fl, fr), Krapovickas et al. 26861 (CTES, G, MO); Depto. Goya, Ruta 12, 40 km S de Goya, 11 Mar 1975 (fl), Krapovickas et al. 27317 (CTES, G, MO); Depto. Esquina, 26 km SE de Libertador, Estancia La Blanca, 59°20’W, 30°20’S, 12 Mar 1975 (fr), Krapovickas et al. 27503 (CTES); Depto. Esquina, 26 km SE de Libertador, Estancia La Blanca, 59°20’W, 30°20’S, 12 Mar 1975 (fl, fr), Krapovickas et al. 27517 (CTES); Depto. Bella Vista, Ruta 23 y Río Santa Lucía, 17 Nov 1973 (fl), Lourteig et al. 2678 (CTES); Depto. Capital, Capital, 24-28 Jan 1944 (fl, fr), Meyer 6458 (A); Corrientes, 12 Jan 1855 (fl), Palmer s.n. (US); Depto. Mburucuyá, Estancia Santa María, 1 Feb 1954 (fl, fr), Pedersen 2595 (BR, C, G, GH, K, MO, S, US); same locality, 4 Jun 1977 (fl, fr), Pedersen s.n. (AAU, MO, NY); Saladas, road to Mburucuya, 7 May 1969 (fl, fr), Plowman 2724 (F, GH, L, S, US); Depto. San Luis del Palmar, Arroyo Riachuelito, 25 km E de San Luis de Palmar, 29 Jan 1972 (fl), Quarín et al. 383 (CTES, US, WIS); Depto. Mercedes, Cnia. Pellegrini, 6 Mar 1973 (fl, fr), Quarín & Schinini 1044 (CTES, WIS); Depto. San Luis del Palmar, San Luis del Palmar, Cañada Grande, Ruta 8, 26 Sep 1973 (fl), Quarín & Tressens 1436 (CTES); Esquina, canal de entrada al puerto, Dec 1936 (fl), Rodrigo 994 (NY); Dept. Bella Vista, 10 km S de Bella vista, cauce seco de Ayo. Toropí, 13 Sept 1972 (fl), Schinini 5350 (CTES); Depto. Capital, Riachuelo, 10 km S de Corrientes, 8 Aug 1973 (fl), Schinini et al. 6915 (CTES); Depto. Capital, Corrientes, 3 oct 1974 (fl, fr), Schinini et al. 9659 (CTES, G, MO); Depto. Bella Vista, Ruta 27, 10 km S de Bella Vista, Toropí, orilla del Río Paraná, 13 Oct 1974 (fl), Schinini & Cristóbal 9870 (CTES); Depto. Saladas, Saladas, 30 Mar 1975 (fl, fr), Schinini et al. 11059 (CTES); Depto. Mercedes, 75 km N de Mercedes, Laguna Trin, Estancia Culantrillar, sobre la orilla inundable de la Laguna Trin, Ea. Culantrillar, 17-24 Oct 1975 (fl), Schinini et al. 11829 (CTES); Dept. Mercedes, 75 km N de Mercedes, Laguna Trin, en la isla El Disparo, Ea. Culantrillar, 17-24 Oct 1975 (fl), Schinini et al. 12048 (AAU, CTES, G); Dept. Bella Vista, Ruta 117, 7 km E de Ruta 27, 11 Mar 1976 (fl, fr), Schinini 12611 (CTES, MO); Depto. San Roque, San Roque, orillas de la ciudad, Río Santa Lucía, 11 Mar 1976 (fl), Schinini & Ahumada 12666 (CTES); Depto. Curuzú Cuatiá, 13 km NW de Perugorría, Ruta 12, 8 Jan 1977 (fl, fr), Schinini 13956 (CTES); Depto. San Martín, 6 km SW de La Cruz, sobre el Río Uruguay, 8 Feb 1979 (fl, fr), Schinini et al. 16868 (CTES, MO); Depto. Goya, 36 km S de Goya por Ruta 12, 22 Nov 1979 (fl), Schinini et al. 18929 (CTES); Depto. Curuzú Cuatía, Ruta 12, 8 km E de Paso López, 29-31 Oct 1974 (fl), Tressens et al. 531 (CTES, G); Depto. Mercedes, Laguna Iberá, Paso Picada, Reserva Natural Provincial de Iberá, 24-28 Feb 1989 (fl, fr), Tressens et al. 3580 (CTES). Entre Ríos: Islas de Victoria, 18 Dec (fl, fr), Burkart 8727 (F); Depto. Paraná, Paraná, un poco al S de la ciudad, a orillas del Río Paraná, Bajada Grande, 5 Jan 1989 (fl, fr), Hunziker 25304 (UT); Isla Puentes (frente a Paraná), 5 Nov 1946 (fl, fr), Meyer 10094 (B); Paraná, 15 Dec 1944 (fr), Schulz 62 (W); Constanza, Brazo largo del Río Paraná, 3 Nov 1965 (fl), Walter & Walter 216 (B). Formosa: Depto. Patiño, Pozo Navagán, Reducción de Indígenas Pilagás, 24°15’S, 60°0’W, 20 Jan 1982 (fl, fr), Arenas 2000 (CTES); camino de Pilcomayo a la Esmeralda, 9 Nov 1941 (fl), Cordini 61 (SI); Depto. Formosa, Formosa, 600 m, 11 Nov 1938 (fl, fr), Eyerdam & Beetle 22974 (BH, G, GH, K); Depto. Patiño, 3 km al N de Las Lomitas, 28 Apr 1992 (fl), Filipov 110 (SI); Depto. Pilcomayo, Parque Nacional Pilcomayo, alrededores de Laguna Blanca, 25°10’N, 58°06’W, 7 Nov 1991 (fl, fr), Fortunato et al. 2041 (SI); Depto. Laishi, Ruta 11, 10 km N de Puerto Velaz, 22 Sep 1967 (fl), Krapovickas & Cristóbal 13196 (CTES, MO, US, WIS); Depto. Pirané, 3 km W of Pirané, 17 Nov 1945 (fl, fr), Morel 305 (L, MO); Depto. Pirané, Los Matacos, 18 Dec 1945 (fl, fr), Morel 655 (G); Depto. Pironé, Cosco-Cué, 9 Feb 1946 (fl, fr), Morel 898 (B, W); Depto. Pilcomayo, Ruta 11, km. 6, 19 Sep 1946 (fr), Morel 1346 (F); Depto. Pilcomayo, Ruta 11, Clorinda a Pilcomayo, 13 Nov 1946 (fl, fr), Morel 1624 (C); Depto. Pilcomayo, Ruta 11, k. 7, Cda. a L. Blanca, 14 Mar 1947 (fl), Morel 2286 (MO); Depto. Pilcomayo, Puente Ceibo, 22 Sep 1947 (fl), Morel 3592 (B); Depto. Pilcomayo, Ruta 11, km. 112, 25 Sep 1947 (fl, fr), Morel 3650 (C); Depto. Pilcomayo, Isla Pé, 29 Sep 1947 (fl), Morel 3739 (BR, MO); Depto. Pilcomayo, 2 km NW of Riacho Negro, 25 Oct 1947 (fl, fr), Morel 4072 (B); Depto. Pilcomayo, Puente Sastre, 15 Mar 1948 (fl), Morel 5087 (C); Depto. Pilcomayo, Ruta 86 al km. 50, 20 Mar 1948 (fl, fr), Morel 5174 (C); Depto. Pilcomayo, Ruta 86 al km. 75, 9 Nov 1948 (fl), Morel 6566 (BR); Depto. Bermejo, Las Lomitas, 4 Sep 1945 (fl), Pierotti 4085 (F, K); Depto. Pirané, Chacras, 30 Sep 1945 (fl), Pierotti 4157 (BR, G); 5 km. E of Capitan J. Page, 20 Nov 1978 (fl, fr), Renvoize 3546 (C, K, MO, NY, S, US); Ing. Juárez, 10 Nov 1969 (fl), Torres 9 (CTES). Misiones: Depto. Candelaria, Puerto Santa Ana, 29 Jan 1945 (fl), Schwarz 599 (F, NY). Salta: Orán, Laguna del Palmar, 26 Feb 1873 (fl), Hieronymus s.n. (Z); Depto. San Martín, 12 km de Hickmann, 2 Apr 1977 (fl), Krapovickas & Schinini 30853 (CTES); Depto. Rivadavia, J. Solá (Morillo), Barrio Mataco, 23°28’S, 62°53’W, 9 Jan 1983 (fl, fr), Maranta & Arenas 153 (CTES, NY); Depto. San Martín, Fca. “Marianito de los llanos” entre Ruta 81 y el Río Bermejo, unos 10 km al E de Hickmann, antes de llegar a Dragones, 220-240 m, 5 Oct 1977 (fl), Novara 500 (MCNS); Depto. Orán, km 58, Vespucio a Hickmann, 16 Feb 1940 (fl), Schreiter 11214 (F, GH, SI); Depto. Orán, Embarcación, orilla del Río Bermejo, 450 m, 21 Dec 1926 (fl, fr), Venturi 5158 (A, SI, US). Santa Fe: Depto. Gral. López, Chapuy, 9 Mar 1983 (fl, fr), Franceschi 762 (CTES); Depto. Castellanos, Rafaela, 15 Nov 1946 (fl, fr), Huidobro 3419 (G); Depto. San Jeronimo, autopista Rosario-Santa Fé, km 115, cerca del desvío a Coronda, 19 Dec 1985 (fr), Hunziker et al. 24857 (UT); Depto. San Lorenzo, entre La Salada e Ibarlucea, Ruta 34, a 1 km al E de la ruta, 18 Dec 1985 (fl), Moscone & Barboza 113 (UT); Depto. Gral. Obligado, Mocoví, 22 Dec 1972 (fl, fr), Quarín 590 (CTES, WIS); Depto. General Obligado, La Reserva, Arroyo Las Garzas, 30 Mar 1975 (fl), Quarín 3058 (CTES); Depto. La Capital, camino a Guadalupe, Jan 1936 (fl, fr), Ragonese 2139 (US); Depto. Gral. Obligado, entre San Juan y Adelaida Vieja, 14 Feb 1938 (fl, fr), Ragonese 3255 (US); Depto. Garay, Colonia Mascias, 20-30 Nov 1942 (fl, fr), Spegazzini BAB 64608 (K); Depto. Capital, El Puerto, 20 Dec 1946 (fl), Terribile 563 (L); Mocoví, 22 Nov 1905 (fl), Venturi 312 (SI). URUGUAY. Canelones: Barra Arroyo, 11 Feb 1930 (fl, fr), Osten 21883 (GH, S). Cerro Largo: Rio Negro, estancia Palleros, Dec 1935 (fl), Gallinal et al. A831 (US); Sierra del Aceguá, Jan 1926 (fl, fr), Herter 18512 (K). Colonia: Aerodromo, 25 May 1947 (fl, fr), Herter 659a (US, Z). Montevideo: Montevideo, 1880 (fl), Arechaveleta 140 (Z); Montevideo, Jul 1868 (fl), Gibert 140 (K, W); Montevideo, 18 Nov 1899 (fl), Hieronymus 3057 (Z); Carrasco, 28 Nov 1901 (fl), Osten 5281 (GOET, US). San José: Barra, 10 m, Dec 1926 (fl, fr), Herter 659 (B, F, G, GH, HBG, MO, NY, S, SI, U, US, WIS, Z). Introduced plants: UNITED STATES. Florida: Pensacola, 6 Aug 1901 (fl, fr), Curtiss 6862 (BH, G, HBG, NY, WU, Z). JAPAN. Koyasan, Jul 1917 (fl), Wilson & Suzuki s.n. (A). NEPAL. Kathmandu, 27°42’N, 85°19’E, 1350 m, 19 Aug 1970 (fl), Dobremez 616 (BM). FRANCE: Cult Jard. Bot. d’Angere, 28 Sep 1858 (fl, fr), Boreau s.n. (BM); Jard. Bot. de Dijon, Martens 1932 (BR); Hort. Huber, Hyéres, 25 Jun 1867 (fl), no collector (NY).

References

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