Solanum elaeagnifolium Cav., Icon. 3: 22, tab. 243. 1795.

Knapp, Sandra, Sagona, Eva, Carbonell, Anna K. Z. & Chiarini, Franco, 2017, A revision of the Solanum elaeagnifolium clade (Elaeagnifolium clade; subgenus Leptostemonum, Solanaceae), PhytoKeys 84, pp. 1-104 : 8-21

1. Solanum elaeagnifolium Cav., Icon. 3: 22, tab. 243. 1795. Figures 2A , 3A, B , 4A , 5

Solanum leprosum Ortega, Nov. Rar. Pl. Hort. Matrit. Dec. 9: 115. 1800. Type. Cultivated in Real Jardin Botánico de Madrid, seeds sent by L. Née, Without collector s.n. (neotype, designated by Knapp 2013b, pg. 59: MA [MA-334600]; possible isoneotypes: MA [MA-334600/2, MA-334600/3]).

Solanum obtusifolium Dunal, Solan. Syn. 26. 1816. Type. Mexico. Hidalgo: "prope Regla et Totonilco [Atotonilco] el Grande, 1200 hex", A. Humboldt & A. Bonpland s.n. (holotype: P-Bonpl. [P001136347]).

Solanum saponaceum Hook., Bot. Mag. 53: tab. 2697. 1826, nom. illeg., non S. saponaceum Dunal, 1813. Type. Argentina. Mendoza: near Río Saladillo, 16 Nov 1820, J. Gillies Solanum -2 (lectotype, designated by Vorontsova and Knapp 2016, pg. 150: K [K000546062]).

Solanum flavidum Torr., Ann. Lyceum Nat. Hist. New York 2: 227. 1828 Type. United States of America. "from Mississippi River to Rocky Mtns.", summer 1820, E. James s.n. (lectotype, designated by Vorontsova and Knapp 2016, pg. 150: NY [NY00759814]).

Solanum dealbatum Lindl., Trans. Hortic. Soc. 7: 52. 1830. Type. Chile. Cumbre "Andium Claustrum", 1826, J. McRae s.n. (lectotype, designated by Vorontsova and Knapp 2016, pg. 151: K [K000546063]).

Solanum texense Engelm. & A.Gray, Boston J. Nat. Hist. 5: 227. 1845. Type. United States of America. Texas: Brazos [River], Jun 1843, F. Lindheimer [66] 135 [fasc. I 1843] (lectotype, designated by Vorontsova and Knapp 2016, p. 151: MO [sheet number MO-3847597]; isolectotypes: BM [BM000514925], GH [GH00217439], LE [2 sheets], MO [sheet number MO-360819]).

Solanum roemerianum Scheele, Linnaea 21: 767. 1848. Type. United States of America. Texas: Travis County, Austin, Apr, F. Roemer s.n. (no herbarium cited, HAL?; possible type material: BM [BM000942973]).

Solanum elaeagnifolium Cav. var. leprosum (Ortega) Dunal, Prodr. [A.P. de Candolle] 13(1): 291. 1852. Type. Based on Solanum leprosum Ortega.

Solanum elaeagnifolium Cav. var. obtusifolium (Dunal) Dunal, Prodr. [A.P. de Candolle] 13(1): 291. 1852. Type. Based on Solanum obtusifolium Dunal.

Solanum elaeagnifolium Cav. var. grandiflorum Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 255. 1879. Type. Argentina. Catamarca: Sin. loc., Nov 1872, P. Lorentz & G. Hieronymus 1243 (holotype: GOET [GOET003503]; isotypes: CORD [CORD00006119], GOET [GOET003504]).

Solanum elaeagnifolium Cav. var. argyrocroton Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 255. 1879. Type. Argentina. Tucumán: between La Oiada and border of provincias Tucumán and Salta, Feb 1873, P. Lorentz & G. Hieronymus 533 (holotype: GOET [GOET003502]; isotype: CORD [CORD00006120]).

Solanum elaeagnifolium Cav. var. albiflorum Cockerell, Bull. Torrey Bot. Club 20: 410. 1893. Type. U.S.A. Texas: El Paso County, El Paso, 1893, T. Cockerell s.n. (lectotype, designated here: NY [NY00759810]).

Solanum elaeagnifolium Cav. var. angustifolium Kuntze, Revis. Gen. Pl. 3(2): 225. 1898. Type. Argentina. Santiago del Estero: San Rafael, O. Kuntze s.n. (lectotype, designated by Vorontsova and Knapp 2016, pg. 151: NY [NY00139141]).

Solanum elaeagnifolium Cav. var. benkei Standl., Rhodora 34: 176. 1932. Type. United States of America. Texas: Cameron County, Brownsville, near mouth of Rio Grande, 20 Mar 1930, H. Benke 5209 (holotype: F [F-642429, F neg. 49341]).

Type.

Cultivated in Madrid from "America calidiore" ["del viaje de los españoles alrededor del mundo, Cult. en el R. J. Bot. 1793"], Anon. s.n. (lectotype, designated by Knapp 2007, pg. 198: MA [MA-476348-2]; isolectotype: MA [MA-476348-1) .

Description.

Herbaceous, woody at base, 20 to 50 cm tall. Stems erect, with running underground stems, sparsely or densely armed, sometimes unarmed; young stems densely pubescent with stellate, sessile to subsessile, porrect lepidote trichomes, the stalk 0.1-0.15 mm long, the rays 10-14(16), 0.1-0.3 mm long, fused at the base, the midpoints 0.1-0.15 mm long, usually absent, unarmed or prickly with the prickles, if present, usually uniform throughout the plant, 2-6 mm long if dense, 0.3-2 mm long if sparse, 0.5-2 mm wide at base, usually straight, orange to brick-red; bark of older stems smooth, brown to grey, the leaf scars small brown stumps. Sympodial units difoliate, not geminate. Leaves simple or shallowly lobed, (1.5)3-6(12) cm long, (0.3)1-2(3) cm wide, (2)3-4(6) times longer than wide, elliptic, sometimes lanceolate or ovate, mostly concolorous, drying yellow-green or silvery green, densely pubescent on both surfaces with sessile to subsessile lepidote trichomes, the rays (9)12-14, 0.1-0.3 mm long, porrect, fused at the base, the midpoint up to 0.15 mm long, sometimes reduced; principal veins 4-7 pairs, raised adaxially, spreading at ca. 30-45° from the midvein, the finer venation mostly not visible to the naked eye; base obtuse to rounded or cuneate, often oblique; margins entire to shallowly lobed, the lobes 4 to 6 on each side, the sinus extending up to ½ of the distance to the midvein, to 2(4) cm long, rounded to deltate; apex obtuse to rounded, rarely acute; petiole 0.2-2(4) cm, 1/2-1/6 of the leaf length, densely stellate-pubescent like the young stems. Inflorescences terminal or lateral, (2)3-5(6) cm long, with (3)4-6(7) flowers, unbranched (simple), densely pubescent with lepidote trichomes like the stems; peduncle 0.8-2.5 cm long; rachis 0.5-3 cm long; pedicels 1-3 cm long, ca. 1 mm in diameter, filiform or apically dilated, densely stellate-lepidote-pubescent like the young stems, usually armed, articulated very near the base; pedicel scars brown, spaced 0.5-1 mm apart, sometimes obscured by the dense pubescence. Buds ellipsoid, the calyx ca. 1/3 of the corolla length prior to anthesis. Flowers 5-merous, usually all perfect, but some distal ones short-styled, functionally staminate and the plants weakly andromonoecious. Calyx tube 2-3 mm long, conical or cup-shaped, the lobes 2- 4 mm long, 1-2 mm wide at base, cuspidate, venation not visible, densely lepidote-pubescent like the leaf blades, sparsely prickly or unarmed, if present the prickles to 2 mm long. Corolla 2.5-3(4) cm in diameter, purple to pale violet or white, stellate, lobed ca. 1/2 of the way to the base, the lobes 0.6-1(1.2) cm, 0.6-1 cm wide, deltate, reflexed to spreading at anthesis, often with a yellow midvein, stellate-pubescent abaxially, mostly glabrous adaxially, the trichomes variously irregular and reduced-multangulate. Stamens equal or slightly unequal in size, with the 2 adaxial anthers very slightly shorter than the 3 abaxial anthers; filament tube 0.8-1.5 mm; free portion of the filaments 1-2 mm; anthers 6-10(12) mm long, 1.5-2 mm wide, tapering, spreading or coherent, the surfaces often papillate, poricidal at the tips, the pores not lengthening with age. Ovary globose to ovoid, usually densely stellate-pubescent like the young stems, sometimes glabrous; style 7-15 mm long, glabrous, slightly dilated toward the apex, in rare short-styled flowers 2-4 mm long; stigma clavate, papillose. Fruit a globose berry, 1-5 per infructescence, 0.6-1.2 cm in diameter, the pericarp thin, smooth, glabrous, yellow to orange at maturity, drying orange to brown and cracking while still on the plant and the seeds released as a mass; fruiting calyx not markedly accrescent, the lobes reflexed or very occasionally covering up to 2/3 of the mature fruit, with prickles ca. 2 mm long; fruiting pedicels 1-2.5 cm long, 0.5-1(2) mm in diameter at the base, 1-2(3)mm in diameter at the apex, somewhat woody, strongly deflexed, usually armed. Seeds 20-60 per berry, 2-3 mm long, 2-3 mm wide, flattened reniform, orange-brown, the surfaces shiny, usually borne in a mass in the berry, the testal cells sinuate (not differing in size between ploidy levels, Nilda Dottori, pers. comm.). Chromosome number: n=12, 24, 36 ( Moscone 1992; Powell and Weedin 2005; Scaldaferro et al. 2012); 2n=24 ( Acosta et al. 2005).

Distribution

(Figure 6 , showing American distribution only). Solanum elaeagnifolium has an amphitropical native distribution, occurring in the deserts and dry zones of the northern hemisphere in the southwestern United States of America and Mexico and in the southern hemisphere in Argentina, Paraguay, Uruguay and Chile, but this species is widespread and invasive in tropical and subtropical regions worldwide (see below). We have seen (non-cultivated) specimens of invasive S. elaeagnifolium from outside its native range from Australia, Cyprus, Egypt, Greece, India, Iraq, Jordan, Kuwait, Malta, Morocco, Pakistan, Saudi Arabia, and South Africa.

Ecology and habitat.

Solanum elaeagnifolium is a plant of disturbed and open places, and often grows in full sun. It occurs at a very wide range of elevations from (-35 m below sea level, Salton Sink, California; Goeden & Ricker s.n.) sea level to 3200m. Solanum elaeagnifolium spreads quickly from underground stems and has become a pest of grazed land in Australia and in the Mediterranean. It is considered a noxious weed in 21 states in the United States (http://www.invasive.org) and in many other countries such as Australia, Canada, Egypt, Greece, India, Syria, Israel, Italy, South Africa and Zimbabwe (Feuerherdt 2010; Uludag et al. 2016; Invasive Species South Africa 2017). It is toxic to livestock and very hard to control, as root stocks less than 1 cm long can regenerate into plants (Cuthertson 1976; Fernández and Brevedan 1972; Stanton et al. 2011).

Common names and uses.

Argentina: capiquí, meloncillo de campo, meloncillo de olor, quillo, suriñado, tutiá chico, granadillo, tomatillo, pocotillo, revienta caballos ( Chiarini 2013). Mexico: trompillo, buena mujer, pera, tomatito de buena mujer (http://www.conabio.gob.mx/malezasdemexico). United States of America: silverleaf nightshade, tomato weed, white horsenettle, trompillo, white nightshade.

In areas where S. elaeagnifolium is invasive it is usually referred to as silverleaf nightshade ( Boyd et al. 1984 Hoffmann et al. 1998; Wu et al. 2016), but in South Africa it is also known as silverleaf bitter apple (English), satansbos, bloubos and silwerblaarbitterappel (Afrikaans) ( Henderson 2011; Invasive Species South Africa 2017).

The berries of S. elaeagnifolium were used in the southwestern United States by Pima, Navajo and Hopi people for making cheese and for tanning leather ( Boyd et al. 1984; Foxx and Hoard 1984) and in local communities in the state of Chihuahua (Mexico) for making filata type asadero cheeses; more recently berries been investigated chemically for use in the food industry due to their milk clotting properties ( Gutiérrez-Méndez et al. 2012). In Argentina, the berries are used as soap due to their high saponin content ( Chiarini 2013).

Preliminary conservation status

( IUCN 2016). LC (Least Concern). EOO = 298,812,730 km2 (LC - Least Concern); AOO = 2,420 km2 (NT - Near Threatened). Solanum elaeagnifolium is an invasive weed (see above) and is actively being eradicated in areas outside its native distribution. The low value for AOO is likely due to the low number of georeferenced collections.

Discussion.

Solanum elaeagnifolium is the most widespread and variable of the species of this group. It can be easily distinguished from the rest of the species by its dense silvery lepidote pubescence, usually orange prickles (if present), mostly perfect flowers that are only slightly zygomorphic in the androecium and multiple berries per infructescence that are exposed at maturity. It is sympatric with S. hindsianum and S. houstonii in Mexico, and with S. homalospermum and S. mortonii in Argentina. Fruit type and pubescence are the most reliable characters for separating the taxa in both North and South America, but in addition, the leaf bases of S. elaeagnifolium are usually more decurrent onto the petiole than any of the other four species.

Solanum elaeagnifolium is extremely variable in its degree of prickliness; this extreme polymorphism also occurs in S. bahamense L. of the Caribbean (Strickland-Constable et al. 2000). Plants are densely prickly (type of S. elaeagnifolium ) to completely unarmed (type of S. leprosum ), and sometimes individual branches on a single plant differ in degree of prickliness (e.g., Bartholomew et al. 2430, MEXU).

Populations of S. elaeagnifolium in North America are diploid, with n=12 ( Powell and Weedin 2005; Chiarini et al. 2016), while those in Argentina are diploid, tetraploid or hexaploid ( Scaldaferro et al. 2012). Morphological differentiation between ploidy levels has not been found (Nilda Dottori, pers. comm.), but hexaploid populations appear to possess a number of differences in plastid haplotypes. Chiarini et al. (2016) suggest this might indicate presence of a cryptic species, but data from the entire genome is needed to test this hypothesis. Tetraploids appear to have arisen multiple times from either diploid or hexaploid populations ( Chiarini et al. 2016). All plants tested outside of the native range (e.g., South Africa, Australia, the Mediterranean) are diploid, and in Australia, genetic variation as measured using AFLP markers suggests that populations in southern Australia are the result of several introductions ( Zhu et al. 2013a).

As discussed above in the section on Habitats and distribution it was long assumed that S. elaeagnifolium was native to North America and was introduced, perhaps by the Spanish or Portuguese, to southern South America ( Goeden 1971; Boyd et al. 1984; Wapshere 1988; OEPP/EPPO 2007). The clustering of all accessions of S. elaeagnifolium , regardless of origin, as sister to the South American S. mortonii + S. homalospermum ( Wahlert et al. 2014) shows this is not the case. Haplotypes based on plastid sequences cluster all diploids in a group with tetraploids in a single cluster and hexaploids in another ( Chiarini et al. 2016).

Introduction of S. elaeagnifolium outside of its native range has resulted in its becoming significant weed of cultivation and pasture (see summaries in Feuerhdert 2010; Uludag et al. 2016). In Australia it was introduced as a contaminant of grain and fodder in the early part of the 20th century ( Parsons and Cuthbertson 2001), but did not become a problem until the 1960s (Stanton et al. 2012). The species became a problem weed in South Africa in the 1970s ( Olckers et al. 1999); our earliest record of it from that country is from 1919 (Carnegie 40). In the Mediterranean it has gone from a few accidental introductions, for example with cotton in Morocco ( Bouhache 2010), to a weed necessitating control in many countries ( Uludag et al. 2016). Plants from botanic gardens such as those used to describe S. elaeagnifolium and S. leprosum appear not to have escaped, and the weedy populations are all thought to be derived from accidental introductions associated with agriculture and subsequent mechanical disturbance of the soil. The Mediterranean populations have been shown to be most genetically similar to those from Texas (Suskiuw 2010). Our records from herbaria indicate that S. elaeagnifolium occurs in India, Iraq, Jordan, Kuwait and Saudi Arabia (see http://dx.doi.org/10.5519/0007624 for complete specimen citations) in addition to in those areas such as Australia, South Africa and the Mediterranean where it is already a listed pest. A worldwide genetic similarity study will be necessary to document and track introduction and invasion of S. elaeagnifolium ; studies on diversity of insect pests as indicators of origins (e.g., Goedin 1971) will need to be tested using new genetic tools. It will also be important to take into account new phylogenetic results (e.g., Wahlert et al. 2014) in determining where invasive populations have originated.

Measures for the control of S. elaeagnifolium have involved both chemical and biological agents. Herbicide control has been used in Australia and in the Mediterranean (Feuerherdt 2010; Uludag et al. 2016; Wu et al. 2016), while biological control using herbivorous insects from the native range in North America ( Goeden 1971) has been successful in South Africa ( Hoffmann et al. 1998; Olckers et al. 1999). In South Africa, the chrysomelid beetle Leptinotarsa texana Schaeffer, a relative of the Colorado potato beetle ( Leptinotarsa decemlineata Say), strips S. elaeagnifolium plants and seriously compromises reproductive potential ( Hoffmann et al. 1998); this species is being considered as a biological control agent in Australia ( Wu et al. 2016). Other insects released to help control S. elaeagnifolium remained at low densities, with no impact; one of these biological control agents has been recorded to have impacted native Solanum species in South Africa (Olckers et al. 1998). Because S. elaeagnifolium reproduces both from seed and vegetatively, the importance of control of both seed and root stocks has been emphasized ( Wu et al. 2016). The great morphological ( Zhu et al. 2013a) and genetic ( Zhu et al. 2013b, c) variability of S. elaeagnifolium surely contributes to its success as an invasive weed. Combining results from phylogenetics and biogeography with those from control measures will surely be important for future control and management of this species as it is introduced elsewhere.

Lectotypes for most of the synonyms of S. elaeagnifolium were designated by Vorontsova and Knapp (2016). At that time, we had not found any original material relating to S. elaeagnifolium forma albiflorum , but a tiny scrap of plant in NY (NY00759810) is annotated as "f. albiflorum " and bears the correct locality. We designate this as the lectotype here.

Selected specimens examined

(American range only). Argentina. Buenos Aires: Rivadavia, F.C.C.A, 3 Mar 1945, Álvarez 573 (W); Hurlingham, F.C.S, 16 Mar 1945, Álvarez 631 (NY, SI); Tornquist, Sierra de La Ventana, desvío por camino de tierra rumbo a La Hoya (unos 70 m del Camping de Sierra de la Ventana, bordeando el río, 22 Jan 2010, Barboza et al. 2308 (CORD, CTES, SI); Bahía Blanca, Bahía Blanca, en los alrededores de la ciudad, en baldíos sobre Avda. Colón al 2000, 23 Jan 2010, Barboza et al. 2319 (CORD, CTES, SI); La Plata, Dec 1944, Boffa s.n. (NY, SI); San Miguel, Bella Vista, Feb 1946, Castellanos 806 (CORD, LIL); Campana, 27 Nov 1938, Eyerdam & Beetle 23093 (BH, K); Coronel Dorrego, Sauce Chico Río, 120 km east of Bahía Blanca on road to Tres Arroyos, 13 Dec 1938, Eyerdam et al. 23735 (K, SI); Buenos Aires en la calle Nazca, 9 Feb 1943, Hunziker 3252 (CORD); Tigre, cerca del Río las Conchas, 25 Jan 1945, Hunziker 280 (CORD); Patagones, Carmen de Patagones, 13 Apr 1945, Hunziker 347 (CORD); Tigre, 24 Dec 1943, Lanfranchi 35 (SI); Martín Coronado, 12 Jan 1939, Nicora, E.G. 2101 (SI); Guaminí, Isla Chica, Laguna del Monte, 19 Jan 1946, Nicora 4162 (SI); Rincón Viedma, Dec 1933, Ringuelet, E.J. 255 (SI); Pergamino, 4 Dec 1944, Schulz 5590 (W); Bahía Blanca, camino a Cuatreros (km 700), San Fernando, Jan 1902, Stuckert 11590 (CORD); barrancas al sur de Buenos Aires, 12 Mar 1902, Venturi 63 (CORD, SI). Catamarca: Santa Maria, Santa María, 23 Nov 1949, Araque M. & Barkley 19Ar - 341 (K); La Paz, La Guardia, 10 Feb 2012, Barboza et al. 3435 (BM, CORD); La Paz, km 981 F. C, 4 Apr 1950, Brizuela 1159 (CORD, LIL); Andalgalá, 4 km N of Andalgalá, on roadside, 15 Dec 1972, Cantino 505 (CORD, SI); Capayán, Ruta 60 (km 1044), entre El Médano y San Martín, 11 Feb 1988, Hunziker 25228 (CORD); Tinogasta, El Cordoncito, 26 Feb 1950, Hunziker & Caso 4097 (BAB, CORD); Santa María, El Medanito, 19 Feb 1948, Reales 952 (CORD, LIL); Santa María, El Rasguñado, 3 Feb 1949, Reales 1615 (CORD, LIL); Santa María, Famatanca, 15 Feb 1949, Reales, A. 1678 (CORD, LIL); Andalgalá, Fuerte de Andalgalá, Oct 1875, Schickendantz 82 (CORD); Poman, Dec 1909, Spegazzini 28174 (SI); La Paz, Casa de Piedra: km 1103, 28 Feb 1986, Subils 3881 (CORD); Del Alto, Balcogna, 18 Jan 1928, Venturi 7132 (CAS); Santa Maria, La Puntilla, 12 Apr 1947, Villafañe 1265 (W); Santa María, Fuerte Quemado, 28 Apr 1947, Villafañe 1271 (E). Chaco: Resistencia, Margarita Belén, 13 Oct 1946, Aguilar 912 (W); General Güemes, 5 km N Castelli (Colonia Fortuny), 24 Aug 1973, Bordón s.n. (CTES); 1 de Mayo, al costado de la RN 11, entre Resistencia y Colonia Benítez, 4 Mar 2012, Chiarini & Wahlert 879 (CORD); Independencia, Baldecito, 15 Jan 1907, Kurtz 14206 (CORD); General Güemes, Fortín Lavalle, 24 Jan 2006, Martínez, G.J. 486 (CORD); General Güemes, Río Bermejo, 27 Jan 2006, Martínez 524 (CORD); Vientecinco de Mayo, Machagay, 8 May 1945, Meyer 9863 (K). Chubut: Biedma, Puerto Madryn, saliendo de la ciudad rumbo a Trelew, a 4 km de P. Madryn sobre Ruta 3, 24 Jan 2010, Barboza et al. 2334 (CORD, CTES, SI); Rawson, km 54, en las proximidades de Trelew, 24 Jan 2010, Barboza et al. 2340 (CORD, CTES, SI). Córdoba: Colón, Estancia Santo Domingo, por el camino de tierra paralelo al camino de la entrada a la estancia, antes de llegar a los maizales, 14 Mar 2005, Ariza Espinar et al. 3587 (CORD); San Alberto, desde San Carlos Minas, rumbo a Córdoba, por RP 15, 1 Feb 2008, Barboza et al. 2010 (CORD); Tulumba, Salinas Grandes, ca. 50 km al norte de Quilino, antes de llegar al límite con Catamarca, 10 Feb 2012, Barboza et al. 3434 (BM, CORD); Minas, La Playa, 26 Feb 2014, Barboza & Palchetti 4120 (CORD); San Justo, Laguna del Plata, 16 Feb 2015, Barboza et al. 4337 (CORD); Gutenberg, 24 Mar 1943, Bartlett 19871 (SI); Tulumba, Cerro Colorado, 4 Mar 1984, Bernardello 449 (CORD); Sobremonte, San Francisco del Chañar, unos 6 km al oeste de la plaza del pueblo, yendo por el camino hacia L.V. Mansilla, 13 Jan 1986, Bernardello 534 (CORD); Colón, Caroya, 15 Jan 1950, Borsini 1201 (CORD); Altautina, 29 Dec 2009, Cantero & Núñez 6183 (CORD); Calamuchita, Valle de Los Reartes, 8 Feb 1919, Castellanos 90 (SI); Colón, La Calera, 28 Jan 1999, Chiarini 40 (CORD); Capital, Ciudad de Córdoba, calle Ovidio Lagos esquina Libertad, 13 May 2001, Chiarini 462 (CORD); Tulumba, sobre Ruta 60 rumbo a Totoralejos, 28 Feb 2002, Chiarini et al. 559 (CORD); Pocho, Los Crestones, cerca de la estancia Las Gramillas, por el camino que va desde Taninga a Chancaní, a ca. 5 km de Taninga, 15 Mar 2012, Chiarini et al. 928 (CORD); San Justo, La Francia, RN 19 y calle Cagnolo, 16 Nov 2013, Chiarini 1033 (CORD); San Javier, Yacanto, 12 Mar 1994, Cosa 162 (CORD); Cruz del Eje, Serrezuela, Punta de Sierra, 7 Jun 1945, Cuezzo 898 (BM, K); San Justo, entre Marull y La Para, frente a la Laguna Mar Chiquita, 18 Apr 1987, Di Fulvio 831 (CORD); Pocho, camino de Taninga a Las Palmas, Feb 2007, Giorgis 693 (CORD); Salinas Grandes, entre el límite con Catamarca y Totoralejos, unos 5 km antes de Totoralejos (al E de las vías férreas), 26 Apr 1964, Hunziker 17347 (COL); Cruz del Eje, saliendo de Soto, rumbo al oeste, hacia Paso Viejo, 12 Mar 1990, Hunziker et al. 25385 (CORD); Próximo a Mar Chiquita, 22 Mar 1988, Juliani 34 (CORD); Río Primero, Arroyo de La Parra y Los Molles, 25 Feb 1887, Kurtz 4744 (CORD); Capital, Quebrada de las Rosas, 20 Mar 1956, Lanfranchi 1315 (SI); Capital, Estancia Germania pr [ope] Cordoba, Jun 1874, Lorentz 9 (BM, G, W); Cruz del Eje, Cruz del Eje a 10 km de la ciudad, 29 Jan 2001, Matesevach s.n. (CORD); Calamuchita, Falda del Sauce (Prop. Marcellino), 19 Apr 1985, Moscone 111 (CORD); San Justo, Ruta nacional n° 19, inmediaciones de Jean Marie, hacia La Francia, 8 Feb 2001, Moscone & Hunziker 245 (CORD); San Marcos, 23 Jan 1941, Nicora s.n. (SI); La Oyada, 2 Feb 1908, Puyssegur s.n. (SI); Tulumba, entre Mansilla y km 907, 25 Jan 1947, Ragonese & Piccinini 6390 (BAB, CORD); Punilla, Huerta Grande, 16 Feb 1897, Stuckert 1720 (CORD); Punilla, La Falda, Jan 1898, Stuckert 4300 (CORD); Ischilín, Quilino, 20 Feb 1899, Stuckert 6517 (CORD); Río Cuarto, Río Cuarto, 20 Feb 1900, Stuckert 8592 (CORD); San Alberto, Mina Clavero, 10 Dec 1901, Stuckert 10520 (CORD); Río Primero, Estancia San Teodoro, Mar 1916, Stuckert 23189 (CORD); Río Segundo, Pilar, 18 Feb 1990, Subils s.n. (CORD); Marcos Juárez, Río Carcarañá, a la altura de Los Surgentes, 1 May 1985, Torriglia 327 (CORD); Ischilín, Deán Funes, RN 60, estacion de Servicio YPF, 27 Mar 2012, Urdampilleta et al. 583 (CORD); San Javier, Cumbre de Achala (Falda O), Las Tapias, al ESE de Villa Dolores, 22 Jan 1990, Zygadlo 4 (CORD). Corrientes: Goya, en los alrededores de la ciudad, 26 Nov 1945, Boelcke 1412 (SI); Capital, Ciudad, 26 Mar 1944, Ibarrola 151 (BM); San Roque, San Roque, 2 leguas al sud, 18 Apr 1945, Ibarrola 2955 (W); Capital, a 2 km del acceso a paraje Perichón, 2 Nov 2010, Medina et al. 57 (CTES); San Cosme, near Santa Ana, 13 Feb 1961, Pedersen 5789 (C, CORD, K, NY); San Cosme, near Santa Ana [de los Guácaras], 12 Feb 1961, Pedersen 5790 (K); Corrientes, Quinta La Eloisa, 6 May 1969, Plowman 2711 (K); Saladas, San Lorenzo, 18 Feb 1950, Schwarz 9758 (K). Entre Ríos: Uruguay, Colonia Caseros, cerca del Palacio San José, a ca. 5 km al oeste de Concepción del Uruguay, 9 Feb 1991, Bernardello & Galetto 746 (CORD); La Soledad, 1904, Britton s.n. (BM); Islas del Ibicuy, Desvío a Médanos, Burkart 23491 (SI); Paraná, Ruta Nacional 18 entre Paraná y Viale, 16 Nov 2013, Chiarini 1034 (CORD); Nogoyá, Sobre ruta provincial 26 en la entrada a Nogoyá, 20 Nov 2013, Chiarini 1039 (CORD); Paraná, Parque Urquiza, 25 Feb 1990, Cosa 115 (CORD); Diamante, Salto, 26 Nov 1946, Huidobro 3514 (W); Paraná, Paraná: entrada al campig Toma Vieja, muy cerca del Túnel Subfluvial, 19 Dec 1985, Hunziker 24860 (CORD); Concepción del Uruguay, Concepcion del Uruguay, Feb 1877, Lorentz 905 (BM, CORD, GOET, K); Nogoyá, Estancia Las Aguadas, 3 Apr 1967, Pedersen 8235 (K); Paraná, Paraná, 27 Dec 1928, Serrano 128 (SI). Formosa: Matacos, Ingeniero G.N. Juárez. Barrio Obrero, orillas del pueblo, 27 Feb 1983, Arenas 2360 (BACP, CORD); Patiño, 5 km antes de Las Lomitas, cerca del aeropuerto, 18 Dec 1984, Bernardello 503 (CORD); Laishí, RP 5, a un par de km del empalme con la RN 11, 5 Mar 2012, Chiarini & Wahlert 890 (CORD); Patiño, RN 81, entre Ibarreta y Las Lomitas, 5 Mar 2012, Chiarini & Wahlert 892 (CORD); Patiño, RN 81, a ca. 6 km de Pozo del Tigre viniendo desde Las Lomitas, 6 Mar 2012, Chiarini & Wahlert 899 (CORD). La Pampa: Utracán, General Acha, 18 Feb 1948, Burkart 15989 (SI); Utracán, Ataliva Roca, 16 Jan 1990, Cocucci 439 (CORD); Ruta 154, km 100-101. ca. 40 km al norte de Río Colorado, 17 Jan 1990, Cocucci 440 (CORD); cruce de ruta prov 26 con RP 19, 31 Oct 2012, Cocucci & Sérsic 5041 (CORD); Caleu-Caleu, desde Hucal, 41 km antes de La Aldea por Ruta Nacional 154 (km 100), 1 Apr 2003, Di Fulvio 1126 (CORD); Catriló, La Gloria F.C.O, 2 Apr 1944, Fortuna 22 (BM, K, SI); entre Chacharamardi y La Reforma, 16 km antes de La Reforma, sobre la izquierda de la ruta, 28 Apr 1990, Galetto 237 (CORD); Jacinto Arauz, 13 Dec 1951, Solbrig 140 (SI); Lihuel Calel, Sierras de Lihuel Calel, 6 Mar 1976, Steibel & Troiani 4092 (CORD). La Rioja: General Belgrano, Las Chacritas, a ca. 35 km al oeste de Jagüe hacia la Laguna Brava, Alrededores del campamento de la Empresa Benito Roggio, 7 Feb 2001, Barboza et al. 220 (CORD); Vinchina, desde Laguna Brava rumbo Jagüé, en el Campamento de B. Roggio, 16 Dec 2011, Barboza et al. 3131 (CORD); Chilecito, Cuesta de Miranda, 17 Dec 2011, Barboza et al. 3159 (CORD); Famatina, alrededores de la Hostería de Famatina, 17 Dec 2011, Barboza et al. 3160 (CORD); Capital, desde Chumbicha rumbo a Mazán por RN 60, km 148-149, 11 Feb 2012, Barboza et al. 3449 (BM, CORD); Chamical, Ruta nacional 79, entre Chamical y Olta a 2-3 km de la primera, 7 Mar 1988, Biurrun & Pagliari 2103 (CORD); General San Martín, E of Chepes, c. 4 km W of crossroad RN 79 and RN 141, 13 Jan 2009, Chiapella & Vitek 2175 (CORD, W); Castro Barros, Anillaco, 10 Nov 2011, Chiarini 781 (CORD); Famatina, Chilecito Cueva de las Brujas, 17 Nov 2015, Cocucci 5760 (CORD); Independencia, Ruta 74, entre Cueva del Chacho y Patquía, 1 May 1997, Di Fulvio 949 (CORD); General San Martín, Por Ruta prov. 32, 5 km antes del empalme con Ruta nac. 141 desde Tello a Chepes (36 km antes de Chepes), 12 Feb 2008, Filippa et al. 79 (CORD); Arauco, Villa Mazán, 20 Dec 1989, Galetto et al. 232 (CORD); Vinchina, Vinchina, 21 Feb 1879, Hieronymus & Niederlein 288 (CORD, E); General Belgrano, Ruta Nacional 38 (km 217-218), entre el límite con Prov. Córdoba y Castro Barros, 12 Mar 1990, Hunziker, A.T.et al. 25395 (CORD); Ruta nacional 38 (km 230/232), entre Castro Barros y Chañar, 12 Mar 1990, Hunziker 25936 (COL); Vinchina, Jagüé, 10 Feb 1949, Krapovickas & Hunziker 5950 (BAB, CORD); Famatina, Las Gredas, 21 Feb 1907, Kurtz 14406 (CORD); Aracico, Aimogasta, 12 Dec 1948, Marín 150 (BH, NY, W); Rosario Vera Peñaloza, Ciudad de Chepes, 29 Jan 2001, Matesevach s.n. (CORD); General Ortíz de Ocampo, El Milagro, 29 Jan 2001, Matesevach s.n. (CORD); Chilecito, 10 Jan 1942, Meyer 3506 (SI); General Lamadrid, Villa Castelli, 19 Jan 1942, Meyer 4067 (SI); General Belgrano, por Ruta prov. 79, entre Chañar y Olta a 700 m del primero, en campo Experimental del Instituto Castro Barros, 24 Oct 1996, Riedel 54 (CORD); La Diana, Apr 1900, Stuckert 9337 (CORD). Mendoza: Las Heras, Uspallata, Ruta Nac. 7, 1 km al W de la villa de Uspallata, 28 Feb 1985, Ambrosetti & Moscone 1480 (CORD, MERL); San Rafael, Salina del Diamante, en los alrededores más cercanos a la propia salina, 26 Nov 2015, Barboza et al. 4450 (CORD); San Felipe de Aconcagua, Mar 1899, Bridges s.n. (W); Luján de Cuyo, El Carrizal, entre Ugarteche y El Carrizal de Arriba, 27 Feb 1985, Del Vitto & Moscone 850 (CORD, MERL); Las Heras, Reserva Pcial. Villa Vicencio, 38, 8 km de Las Heras, camino a Villa Vicencio por ruta 52, 29 Jan 2010, Ferrucci et al. 2953 (CORD, CTES); Tupungato, Ruta 89, trayecto Tupungato-Potrerillos, 6, 5 km de San José, sentido NW, río Las Carreras, 31 Jan 2010, Ferrucci et al. 2979 (CORD, CTES); Luján, Lunlunta, 8 Jan 1948, Garcia 413 (W); Capital, Parque General San Martín, CRICYT, 21 Nov 1991, Illanes & Pacheco 104 (CORD); Las Heras, Quebrada del Camino, above Uspallata on road to Las Cuevas and Chilean frontier, 5 Feb 2013, Knapp et al. 10470 (BM, CORD, MERL); Las Catitas, 5 Dec 1939, la Barrera 20 (SI); Junín, Finca San José, Calle Cervalaán, 4 Feb 1945, Lourteig 1019 (W); General Alvear, Ruta 188, 8 Feb 1987, Naranjo et al. 937 (SI); Capital, Mendoza (Parque), 19 Jan 1944, O’Donell 224 (CORD); Las Heras, Laguna La Seca, Ruta Nac. 40, 23 Feb 2008, Pensiero et al. 7374 (SF); San Rafael, Cuadro Benega, 12 Nov 1949, Reales 2015 (CORD); Luján, Luján de Cuyo, 4 Mar 1947, Villafañe 937 (W); Luján, Chacras de Coria, 3 Mar 1947, Villafañe 903 (CAS); Maipú, 7 Sep 1947, Villafañe 993 (DS); San Rafael, Valle del Río Atuel, Jan 1897, Wilczek 239 (CAS). Neuquén: Confluencia, Plaza Huincul, 13 Jan 1960, Articó 244 (CORD); Collón Curá, Bajada colorada: entre Piedra del Águila y Neuquén, 2 Feb 1960, Articó 250 (CORD); Confluencia, en los alrededores del aeropuerto de Cutral Có, 29 Jan 1990, Barboza 76 (CORD); Chos Malal, Chos Malal, 20 Jan 1964, Boelcke et al. 11691 (BAA, BAB, SI); Zapala, San Antonio, a 20 km de Zapala rumbo a Neuquén por RN º 22, 14 Feb 2007, Chiapella et al. 1804 (CORD, CTES, SI); inter Zapala et Cutral Co, prope diversorium ruta 22 cum ruta 34, 11 Dec 1988, Fernández Casas 11162 (MA); Confluencia, sobre un baldío cerca de la Universidad, 2 May 1990, Galetto 239 (CORD). Río Negro: vicinity of General Roca, Sep 1914, Fischer 238 (BM, K, NY, SI); General Roca, Allen, Feb 1939, Hunziker 20 (CORD); Pichi Mahuida, Pichi-Mahuida, 23 Nov 1944, O’Donell 1712 (W); Avellaneda, Choele-Choel, 27 Nov 1944, O’Donell 1815 (W); General Roca, Cipolleti, 4 Dec 1944, O’Donell 1910 (W); Pichi Mahuida, Río Colorado, barrancas del Río, en cruce ruta 3, 18 Feb 2008, Pensiero et al. 7209 (SF). Salta: Cachi, San José de Cachi, 28 Mar 1979, Cabrera et al. 30771 (SI); General Güemes, por Ruta 34, entre General Güemes y Metán, 5 km antes del Río Juramento, 17 Dec 1989, Galetto et al. 214 (CORD); Rosario de la Frontera, Rosario de la Frontera, 19 Feb 1947, de la Sota 132 (W); Capital, Cerro San Bernardo, 4 Feb 1949, Legname 221 (CORD, LIL); Anta, El Dorado, 5 May 1948, Luna 1021 (DS); Coronel Moldes, 2 Jan 1945, Mintzer s.n. (SI); Coronel Moldes, 2 Jan 1945, Mintzer s.n. (SI); Candelaria, El Datil, 7 Feb 1949, Montenegro 380 (W); Capital, Ciudad de Salta, vías del FFCC, playa de maniobras entre la estación y molino Batule, 5 Dec 1988, Novara 8248 (CORD); La Viña, Coronel Moldes, entrada da cidade, 25 Jan 2007, Paula-Souza et al. 7856 (ESA, K, SI); Rivadavia, Ruta 81, km 1517, 16 Nov 2009, Schinini & Flaschland 37059 (CTES); Rosario de la Frontera, Pasando Rosario de la Frontera, 13 Nov 1984, Subils 3589b (CORD); Guachipas, Alemania, 16 Nov 1929, Venturi 9882 (BM, NY); Chicoana, Ruta Prov. 33, de Pulares a San Fernando de Escoipe, 15 Feb 2007, Zuloaga et al. 9373 (SI). San Juan: Calingasta, Barreal, cerca de la Gendarmería, 1 Nov 1969, Ariza Espinar 2393 (CORD); Caucete, Rumbo a la Difunta Correa, desvío de 2 km hacia la derecha, en el km 195 de la RN 141, 21 Dec 2007, Barboza 1944 (CORD); Sarmiento, Media Agua, 28 Jan 1950, Borsini 1333 (CORD, LIL); Ullum, Hualilán, 8 Dec 1979, Cabrera et al. 31065 (SI); Ullum, road RN 40 between San Juan city and Talacasto, 14 Jan 2009, Chiapella & Vitek 2186 (CORD, NY, W); Ullum, entre ciudad de San Juan y Ullún, cerca de la empresa Valle de la Luna S.A, 4 Apr 1998, Cocucci et al. 11 (CORD); Trinidad, 20 Nov 1945, Cuezzo 1302 (BM, K); Jáchal, Entre El Balde y Tucunuca, unos 7 km después de Ing. Matías G. Sanchez (ruta 40), 10 Feb 2002, Di Fulvio 1112 (CORD); Iglesia, R 149, alrededores de Iglesia, 7 Mar 2011, Fortunato et al. 9942 (BAB, CORD); Sarmiento, Puesto Bachongo, 25 Jan 1986, Guaglianone et al. 1555 (SI); Rawson, Dique Bello, 26 Jan 1986, Guaglianone et al. 1571 (SI); Caucete, Marayes, 8 Nov 1986, Haene 61 (SI); Valle Fértil, Puesto de las Chaves, cerca de Mareyes Viejo, 16 Feb 1921, Hosseus 2638 (CORD); Caucete, Ruta 20, km 1158, entre Caucete y Vallecito (=Difunta Correa), 13 Jan 1979, Hunziker et al. 23318 (CORD); Jáchal, Camino a Mogna, 9 Nov 1980, Kiesling 3006 (K, SI); Ullum, Talacasto a Alto de Colorado, 17 Nov 1982, Kiesling & Sáenz 4241 (SI); Angaco, Sierra de Pie de Palo, ruta cerca de Angaco, 23 Nov 1984, Kiesling et al. 4827 (SI); Calingasta, 5 km N of Villa Nueva (measured from the bridge over the Río de Castaño Viejo), on Ruta 412 towards Iglesia, 11 Jan 1995, Leuenberger et al. 4459 (B, CORD, Z); Chimbas, Finca de la familia Matesevach, calle Oro 4730, 1 Jan 2001, Matesevach s.n. (CORD); Iglesia, Reserva de San Guillermo, Baños Termales de San Crispín, 10 Jan 1984, Meglioli 33 (SI); km 24, camino de Mendoza a San Juan, 31 Jan 1946, Nicora 4282 (SI); Jáchal, Entre Jáchal Estación Adan Quiroga, 19 Jan 1950, Palacios et al. 1695 (CORD, LIL); Reserva de San Guillermo. Baños de San Crispín camino de deyección sobre las vegas termales, 24 Mar 1983, Pujalte 210 (SI); Iglesia, curso superior del Río Valle del Cura, Llanos de Conconta, Agua de Alutiaca, 25 Jan 1981, Subils 2909 (CORD); Capital, Estación Ferrocarril, 22 Jan 1972, Volponi 241 (SI). San Luis: Ayacucho, a orillas del Río Luján, 19 Dec 2007, Barboza et al. 1923 (CORD); Capital, Chosmes, 25 Jan 1950, Borsini 1013 (W); Chacabuco, Concarán, alrededores del Establecimiento El Sauce, 4 Feb 2012, Chiarini et al. 824 (CORD); Junín, Ruta nacional 146: entre Quines y Villa Dolores, cerca del cruce hacia La Unión, Jan 1988, Cocucci 220 (CORD); La Capital, entre el Chorrillo y Cruz de Piedra, por la ruta 20, 2 Dec 1988, Del Vitto & Petenatti 3558 (CORD); Ayacucho, Ruta 20, km 390, La Chañarienta, 9 Feb 2002, Di Fulvio 1110 (CORD); Chacabuco, Estanzuela, 4 Mar 1882, Galander s.n. (CORD); La Capital, a la salida de San Luis, sobre la rotonda que separa las rutas que van a Potrero de Funes y Trapiche, 14 Nov 1988, Galetto 24 (CORD); Buena Esperanza, 24 Nov 1962, Hunziker 16035 (COL); Las Pampas cerca de San Luis, 6 Mar 1863, Isern 8096 (SI); Junín, Merlo, frente al aeródromo, 10 Apr 1988, Juliani 37 (CORD); San Luis del Palmar, in Bahnhofsgelande von Mercedes, ca. 90 km ESE San Luis, 9 Feb 1993, Starlinger 22 -93 (W); Desaguadero ( límite entre San Luis y Mendoza), 26 Jan 1939, Troncoso 6030 (SI); Ayacucho, Luján, 2 Dec 1944, Varela 713 (BM, K). Santa Fé: San Justo, Soledad, 3 Dec 1946, Álvarez 989 (W); General López, Southern District, Venado Tuerto, Dec 1975, Ardiar s.n. (K); General Obligado, Cerca de Berna, Ruta 11, km 759, 25 Jan 1992, Bernardello & Galetto 788 (CORD); 9 de Julio, RN 95, 8 Mar 2012, Chiarini & Wahlert 917 (CORD); San Cristóbal, Saliendo de La Rubia, 17 Dec 1981, D´Angelo 285 (SF); San Cristóbal, entre San Cristóbal y Gobernador Crespo, Ruta 39, 4 Jan 1985, Di Fulvio 799 (CORD); Cuty Lai, 6 Apr 1917, Hosseus 65 (CORD); San Lorenzo, Carcarañá, Kurtz 5117 (CORD); Castellanos, Cañada Las Calaveras. Proximo a Lehmann, 24 Apr 1989, Luchetti 5 (SF); San Cristóbal, Ceres, 17 Jan 1989, Luchetti et al. 6 (SF); Las Colonias, Reserva UNL, 21 Nov 1991, Marino 201 (SF); Vera, 10 km al N de Fortín Olmos, campo de Sr. Sanchez, 11 Nov 2003, Marino 1914 (SF); Vera, Campo del Sr. Sanchez, 10 km al norte de Fortin Olmos, 11 Nov 2003, Marino 1942 (SF); Vera, Escuela EFA, km 50, Ruta 98 S, 13 Nov 2003, Marino 2047 (SF); San Justo, 5 km al O de Naré, Cañada Naré, 28 Dec 1985, Pensiero & Faurie 2424 (SF); Las Colonias, Esperanza, Reserva de la Escuela Granja, 25 Jan 2000, Pensiero & Exner 6063 (SF). Santiago del Estero: Santa Rosa, 5 Jan 1938, Argañaras 51 (SI); entre Brea Pozo y Río Huayco Hondo, 23 Mar 1943, Bartlett 19760 (SI); Río Hondo, Termas de Río Hondo. Ruta 9, 26 Jun 1991, Cosa 124 (CORD); General Taboada, Averias, 23 Jan 1985, Fernandez 156 (CORD); Fortin Tostado, Jun 1906, Goubat s.n. (SI); General Taboada, Tacanitas km 443, 17 Apr 1917, Hosseus 233 (CORD); Atamisqui, Isla Verde, Por ruta 9, Isla Verde, entre El Guanaco y San Gregorio, 11 Feb 1986, Hunziker et al. 24868 (BM); Ojo de Agua, Entre el límite con Córdoba y Ojo de Agua, unos 10 km al sur de Ojo de Agua, 13 Sep 1987, Hunziker 25089 (CORD); Copo, Ruta 16, 15 km NW de Los Tigres, 28 Jan 2007, Paula-Souza et al. 8046 (ESA, SI); Robles, Colonia Jaime, 9 Nov 1948, Ruiz 53398 (BA, CORD); Moreno, Quimili, 24 Feb 1947, Schulz 1363 (W); Añatuya, 27 Jan 1944, Soriano 548 (BAB, SI); Choya, Ruta Nacional 158: alrededores de Frías, camino a San Antonio, 1 Jun 1990, Subils & Guzmán 4414 (CORD); Quebracho, Sumampa, 6 Jan 1948, Terribile 808 (W). Tucumán: alrededores de Tucumán, 8 Dec 1907, Dinelli 654 (BA, SI, SI); Graneros, La Madrid, 26 Jan 1886, Kurtz 4222 (CORD); Graneros, La Madrid, 26 Jan 1886, Kurtz 4224 (CORD); Trancas, Tapia, 28 Nov 1920, Venturi 1091 (SI); Burruyacú, El Puestito, 2 Dec 1928, Venturi 7678 (BM, CAS, K, SI).

Bahamas. Eleuthera: Hatchet Bay Farm, 24 Jul 1969, Proctor 30960 (BM); edge of cultivated field 2 miles south of Gregory Town , 2 Jun 1979, Sauleda 2649 (MO, NY) .

British Virgin Islands. Anegada: around The Settlement , 1 Aug 1970, D’Arcy 4861 (MO) ; Tortola: East End , vacant lot, 15 Apr 1965, D’Arcy 44A (BM) .

Chile. Región II (Antofagasta): El Loa, Patio del Hostal Puri, San Pedro de Atacama, 16 Feb 2001, Ackermann 218 (BM); El Loa, Rio Grande , San Pedro de Atacama, 1 Dec 2008, Baines et al. 255 (BM, E); El Loa, Calama, beside Ensueno land development area, part of the Stadium in Calama , 12 Mar 1988, Dunnett 13 (K); Hostería San Pedro de Atacama, 5 Feb 1967, Martin 272 (SI). Región III (Atacama): Caldera, prov. Atacama, 21 Feb 1939, Beetle 26132 (BH, K); Paipote, 25 Jan 1989, Cocucci 369 (CORD); Copiapó, 30 km S of Copiapo on road to Los Loros, 17 Sep 1987, Hannington 23 (K); Huasco, Yerba Buena Chica, 25 miles east of Carrizal Bajo , Vallenar, 25 Dec 1871, King s.n. (E); Copiapó, Dorf Tierra Amarilla am Weg zur Mine Las Bateas, 12 Nov 1987, Rechinger & Rechinger 63645 (W); Vallenar, Mine Algarrobo, Nov 1925, Werdermann 142 (BM, CAS, E, SI); Huasco, all along road from Vallenar to San Felix, 24 Oct 1938, Worth & Morrison 16215 (K). Región IV (Coquimbo): Elqui, Vicuña, 6 Oct 1927, Elliot 62 (E, K); Elqui, Paiguano, 7 Dec 1997, Gardner & Matthews 58 (E); Vicuña, a 62 km al este de La Serena, en las inmediaciones de la bodega pisquera Capel, 22 Feb 1996, Hunziker 25554 (CORD); Elqui, NE of Monte Grande , Paiguano , 2 Mar 1996, Instituto de Investigaciones Ecologicas Chiloe 502 (E). Región IX ( Araucanía): El Loa, San Pedro de Atacama, 28 Feb 1988, Wickens & Dunnett 828 (K). Región V ( Valparaíso): San Felipe de Aconcagua , 1868, King s.n. (E); Los Andes, 18 Jan 1904, Scott-Elliot 442 (BM, E). Región VIII ( Bío-Bío): Ñuble, Chillán, Gillies 669 (K) .

Colombia. Cundinamarca: prés Bogotá, Chaquiero , Jul 1908, Apollinaire s.n. (E) [possibly cultivated?] .

Cuba. Cienfuegos: Cienfuegos , 5 Jun 1923, Ekman 13942 (NY) . La Habana: Carmelo Hill, Vedado , 13 Jun 1919, Fr. León 8782 (NY) .

Ecuador. “Andes”, Jameson s.n. (E) [possibly Peru, from Lima?].

Haiti. Sud: Massif de la Hotte, aprox. 4 km al E de Les Anglais en el camino costero a Port-a-Piment y Les Cayes, 25 Jan 1985, Zanoni et al. 33168 (JBSD) .

Jamaica. Saint Catherine: Boldes Research Station , ca. 2 miles W of Old Harbour, beside the piggery, 5 Aug 1987, Collins s.n. (MO, NY) .

Mexico. Aguascalientes: 35 miles N of Aguascalientes, 24 Aug 1953, Manning & Manning 531244 (MEXU); Rincón de Romos, Colonia 16 de Septiembre, 17 Sep 1989, Marmolejo P. s.n. (MEXU). Baja California: Mexicali, 26 May 1933, Harvey 585 (US); La Hechicera, 2 km S of La Hechicera on highway, Sierra Juarez, 7 Jul 1979, Moran 27703 (CAS); Mexicali, Mexicali, about 14 miles W along road to Tecate, 30 Jun 1962, Wiggins & Thomas 444 (DS, MEXU). Baja California Sur: La Paz, Carretera entronque al aeropuerto, km 3 carretera a Ciudad Constitución, 15 Jun 1994, Domínguez C. 1237 (MEXU). Chihuahua: Villa Ahumada, ca. 10 miles S of Villa Ahumada along Highway 45 between Ciudad Juárez and Chihuahua, 1 Aug 1975, Davidse & Davidse 9252 (MEXU); 9.4 miles W of Hwy 45 along Hwy 10, ca. 15 miles SW of Ciudad Juárez, 18 Aug 1971, Henrickson 5708 (MEXU); Parral, 10 km al este de Parral, 31 Jul 1982, Hernández Magaña 8352 (CAS, MEXU); Ciudad Juárez, 35 km al S de Ciudad Juárez, 5 Aug 1982, Hernández Magaña et al. 8415 (MEXU); Juárez, Samalayuca, south of Juárez, 7 Jul 1957, Knobloch 70 (BM); along Hwy 45, km 1615-1616, S of Chihuahua, 20 Jun 1965, Mertz 15 (MEXU); Aldama, km 68 carretera Chihuahua-Ojinaga, 25 Aug 1978, Molinar et al. 47 (MEXU); Ahumada, Laguna Santa María, near Lake Santa Maria, 7 Sep 1899, Nelson 6429 (K); Chihuahua, plains and mesas near Chihuahua, 16 Aug 1887, Pringle 1566 (BM, K, MEXU); between Cuidad Juárez and Cuidad Chihuahua on Hwy 45, 33 miles from US border, 7 Aug 1984, Randolph 173 (MEXU); entronque a Santa Clara, km 91 Chihuahua-Ciudad Juárez, 12 Jun 1980, Siqueiros & Baray 515 (MEXU); Saucillo, Naica, 14 Sep 2011, Tejero Diez & Canek 6517 (MEXU); Villa Ahumada, 15 km al N de Villa Ahumada, km 102 carretera Chihuahua-Ciudad Juárez, 13 Sep 1978, Valdés et al. 15 -1978 (MEXU); Jiménez, Escalón, 26 Jul 1939, White 2052 (CAS); Janos, border of Chihuahua and Sonora, 26 Aug 1939, White 2526 (MEXU). Coahuila: Saltillo, 12 miles E on Mexican Highway 40, 4 Apr 1964, Canedo et al. 9075 (DUKE); Torreón, Valle Verde, 21 Sep 1996, Cano R. & López C. 21 (MEXU); along road to E of Hwy 57, 18 mile SE of Saltillo, 15 Jul 1979, Dunn et al. 23177 (MEXU); General Cepeda, ca.de 0.5 km del entroque a Hipolito, 24 Mar 2009, Guzman 3013 (K); ca. 444 (air) miles W of Cuatro Cienagas, 3 miles NE of Est. Zacatosa in Bolson de Mapimi region, 21 Sep 1972, Henrickson 7917 (MEXU); Parras, 0.5 km al sur de San Rafael de los Milagros, poblado que esta a mitad de camino de la cettera Saltillo-Torreón, 16 Oct 2000, Montero Castro & Torres C. 136 (MEXU); San Pedro, along hwy. Mex. 30, 108 km by road NE of San Pedro de Las Colonias, 27 Jul 1982, Nee & Diggs 25319 (CORD, F, XAL); Ramos Arizpe, Paredón, 1 Oct 1924, Orcutt 1270 (DS); San Lorenzo de Laguna, Feb 1880, Palmer 935 (K); Saltillo, Fraile, 59 km S of Saltillo, 10 Jul 1941, Stanford et al. 273 (DS, MEXU); 18 m W of Concepción de Oro, on mountain, 24 Jul 1941, Stanford et al. 586 (DS, NY); Torreón, Ejido El Tajito, 10 km al E de Torreón, 21 May 1965, Vázquez s.n. (MEXU); along Mex. 40 30 miles W of Monterrey, ca. 25 miles NW of Saltillo, 29 Jun 1966, Ward 5742 (DUKE); Monclova, 5 Jul 1939, White 1712 (DS); Ramos Arizpe, between Hipolito and Sacramento, in El Desierto de Payla, 15 Jun 1936, Wynd & Mueller 76 (K, NY); Torreón, near Horizonte, 24 Aug 1937, Wynd 780 (K, NY). Colima: Tecomán, 2 km de Calera rumbo a Madrid, 28 May 1990, Román Miranda 1337 (MEXU). Distrito Federal: City of Mexico, Nov 1884, Carruthers s.n. (BM); Ticomán Cuauhtepec, Jul 1952, Gallegos Harking 81 (MEXU); Rosario, 1 Sep 1936, MacDaniels 703 (BH); Azcapozalco, 1 Apr 1950, Matuda 19545 (MEXU); Xacateno, 1 km al N de la Unidad Profesional de Zacateno, 17 Aug 1969, Pérez H. 86 (DS); E. de Agricultura, Valle de México, 14 Apr 1882, Urbina s.n. (MEXU). Durango: Ceballos, 1 Aug 1979, Aguilar C. et al. 164 (MEXU); Durango, La Ferreria, Río El Tunal, Jul 1992, Aguirre T. 45 (MEXU); Durango, P.P. Morelos, Oct 1994, Aguirre T. 151 (MEXU); Mapimí, 45 km al E de la Ciudad de Ceballos [26.29-52 DMS N/103.32-58 DMS W], 26 Jun 1981, Alba 45 (MEXU); Santiago Papasquiaro, km 3 de la carretera Santiago Papasquiaro-Los Altares, 30 Jul 1990, Benítez P. 1814 (MEXU); Durango, km 27 carretera Durango-Torreón, Jul 1994, Domingo & Aceval A. 524 (MEXU); Durango, 7 km al NE de Durango, 7 Jul 1982, Hernández Magaña & Tenorio L. 7714 (MEXU); Mezquital, km 20 carretera Durango-Mezquital, Sep 1992, Martínez Marín 416 (MEXU); Durango, camino a Malaga, km 1, Jun 1993, Martínez Marín 464 (MEXU); Mapimí, 49 km al NE de Ceballos, Rancho San Ignacio (Ojo de Agua), 25 Oct 1975, Martínez O. et al. 413 (K); Poanas, Rancho 4 Hermanos, Jun 2000, Martínez 683 (MEXU); Durango, city of Durango and vicinity, Apr 1896, Palmer 233 (BM, E, G, MEXU, NY); Pénuco de Coronado, km 38 carretera Durango-Torreón, May 1994, Rangel Ramírez 130 (MEXU); San Juan de Guadelupe, Ejido Acacio, 12 Jul 1990, Rendón s.n. (MEXU); Mezquital, Durango, 50 miles S of Durango, 10 Aug 1953, Selander 4-53 (DS); Cuencamé, Velardeña, 23 km al NNE del centro Cuencamé, 22 Jul 2010, Tejero-Díaz & Torres 6256 (MEXU); Tepehuanes, Presidios, 12 km al S de Tepehuanes, 19 Jul 1982, Tenorio L. & Romero de T. 1122 (MEXU); Mapimí, Puente de Hojuelas, 8 km SE de Mpio de Mapimí, 8 Sep 1983, Torrecillas 208 (MEXU); Conejos, 25 miles S of Ceballos, ca. 50 miles N of Gomez Palacio, 3 Jul 1966, Ward 5771 (DUKE). Guanajuato: carretera Irapuato a Mungía, 20 May 1973, Boege 2793 (MEXU); Apaseo el Grande, Apaseo, 1938, Figueroa s.n. (MEXU); San Felipe, 10 km al SW de San Felipe, 25 Aug 1990, Galván & Galván 3614 (MEXU); San Miguel de Allende, highway 51, 21 Jun 1971, Genelle & Fleming 819 (DUKE, MEXU); Guanajuato City to San Luis de la Paz, 14 Sep 1946, Hernández X. et al. X-2362 (MEXU); San Miguel Allende, el sendero que continua al S de Calle Recreo, 19 Sep 1977, Kishler, J. 145 (MEXU); Irapuato, 1.66 km al OSO de Lo de Juárez, 7 Sep 2007, Martínez S. & Ramos 39643 (MEXU); Predio El Cortijo, a 16 km al NE de la ciudad Dolores Hidalgo sobre la carretera a San Luis de la Paz, 4 May 1996, Ocampo 39 (MEXU); León, Lagunillas, 28 Jun 1996, Pérez C. & Zamudio 3359 (MEXU); Irapuato, 10 May 1901, Pringle 9370 (BH, K); Dolores Hidalgo, Las Yerbas, 17 May 1996, Rojas Villegas s.n. (MEXU); Estación Río Laja, 19 Jul 1973, Valdés et al. 55- 2 (MEXU); San Luis de la Paz, Rancho La Misión, 8 km al NE de San Luis de la Paz, 11 Oct 1988, Ventura & López 6112 (MEXU); San Luis de la Paz, Potosino Dos, 24 May 1990, Ventura & López 8022 (MEXU); San Luis de la Paz, Cerro Prieto, 19 Jul 1990, Ventura & López 8329 (MEXU). Hidalgo: Actopan, El Arenal, Mar 1936, Bravo H. s.n. (MEXU); Zimapán, Zimapán, Coulter 1246 (K); Ajacuba, Cerro El Crestón, ca. 2 km antes de llegar a la desviación al centro de Emiliano Zapata, rumbo a Ajacaba, Ejido Tecomatlán (de E a W), 24 Aug 1988, Díaz Vilchis 124 (MEXU); Ajacuba, Rincón del Gato, barranca al N del poblado Emiliano Zapata, Sierra de Chicavasco, ejido Emiliano Zapata, 28 May 1989, Díaz Vilchis 438 (MEXU); Tecozautla, 4 km al oeste en el desvío a Pañhé, 11 Jul 1980, Hernández Magaña 4642 (CAS, MEXU); Pachuca, San Bartolo, 2 km al W de Pachuca, 22 Jun 1977, Medina C. 2036 (MEXU); El Arenal, valley of Actopan between El Arenal and Jiadi, 26 Oct 1946, Moore 1640 (BH); Cerro Gordo, 5 km al W de Pachuca, 18 Aug 1972, Rzedowski 29179 (CAS, NY); Santiago de Anaya, 500 m al O de Santiago de Anaya, 5 Jun 1992, Soriano M. 144 (MEXU); Tepeapulco, 2 Jun 1976, Ventura A. 1509 (MEXU). Jalisco: 21.1-24.1 km de la salida de San Juan de los Lagos por la carretera a Lagos de Moreno, 28 Jul 1978, Guzmán et al. 999 (MEXU); Ojuelos, 3 km N, 17 Sep 1946, Hernández X. et al. X- 2493 (MEXU); Villa Hidalgo, Villa Hidalgo, May 1977, la Torre 47 (MEXU); Amacueca, cerca de Cofradía del Rosario, 1 Aug 1993, Novoa L. & Villa M. 919 (MEXU); 5 km antes de llegar a Lagos de Moreno en la desv. hacia la Mesa, 31 Jul 1987, Román Miranda, C-742 (MEXU); Ojuelos, 3 km al E de Ojuelos, 25 Sep 1962, Rzedowski 16116 (MEXU). México: Cerro de San Cristobal, 2 Jul 1950, Matuda 18953 (MEXU); Icatepec, [Ecatepec] a Peñon, 28 Jun 1953, Matuda 28754 (MEXU); Huehuetoca, Tajo de Nochistongo, 22 Jun 1981, Romero & Rojas 1405 (MEXU); Huehuetoca, Huehuetoca, vía de ferrocarril, 22 Jun 1981, Romero & Rojas 1406 (MEXU); Venta de Carpio, 16 Jul 1930, St Pierre 2601 (K); Otumba, Ahuatepec, 27 Jun 1976, Ventura A. 1361 (MEXU); Otumba, Ahuatepec, 15 Jul 1976, Ventura A. 1595 (CORD, ENC, MEXU). Michoacán: Pátzcuaro, a 300 m de la salida a Uruapan, sobre la vía del FF.CC, 29 May 1986, Escobedo 928 (MEXU); Morelia, La Huerta, Estación del Tren, 22 Aug 1988, Escobedo 1594 (MEXU); Coahuayana, Boca de Apiza, 16 Jul 1985, Soto N. 9508 (MEXU). Morelos: Zacatepec, [Zacatepec de Hidalgo], 5 Apr 1969, Vázquez 2164 (MEXU). Nuevo León: Monterrey, ½ West of ITESM Campus, 29 Jul 1970, Burge 25 (EIU); 7.8 km E of turnoff to Los Herreras on Mex 40 near Presa of San Juan River drainage, 18 Sep 2001, Bye et al. 28334 (MEXU); Dr. Arroyo, Tanquecillos, Aug 1842, Karwinski 583 (LE); near Montemorelos, 19 Apr 1947, Kelley 21 (BM, MEXU, NY); Los Ramones, 2 kilo. west of Ramones, 70 kilo. east of Monterrey, 10 Jul 1971, Parker 379 (EIU); La Escondida, N of Monterrey, 1946, Roybal 904 (MEXU); Montemorelos, Ojo de Agua, 24 Apr 1988, Tirado 51 (MEXU); Montemorelos, 4.1 miles S on Highway 85, 4 Jul 1969, Weaver 2044 (DUKE, MEXU); Monterrey, Cerro de la Silla, foot of La Silla, 4 Sep 1937, White & Chatters 183 (MEXU). Oaxaca: Salina Cruz, Salina Cruz, dry dock area, 18 Sep 1978, D’Arcy 12027 (MEXU); Salina Cruz, La Ventosa, al S de Salina Cruz, 31 Aug 1988, Martínez 1796 (MEXU); Playa Abierta at Salina Cruz about 15 miles S of Tehuantepec, 1 Aug 1984, Wilbur 36096 (BM). Puebla: Zinacatepec, 1 km from Zinacatepec, on the federal Tehuacán-Teotitlan de Flores Magón-Oaxaca road, 17 Jul 2004, Calzada 24391 (K); Coxcatlán, San Rafael, campos de cultivo al oriente de San Rafael, 24 May 2009, Cervantes M. 58 (K); Chietla, carretera Axochiapan-Izucár de Matamoros, entre Ahuehuetzingo y Atencingo, 19 Jun 1998, Guízar N. & Herrera 4031 (MEXU); Matamoros, 22 Mar 1941, Miranda 1418 (MEXU); San José Miahuatlán, Axusco, 4-6 km al S, 18 Oct 1992, Salinas T. 7104 (CAS). Querétaro: cerca acueducto, 4 May 1975, Argüelles 23 (MEXU); calle cerca de Constituyentes, 31 May 1986, Argüelles 2468 (MEXU); Ciudad de Querétaro, Barrio Carretas, 19 Jun 2016, Chiarini 1270 (CORD); Peñon de Bernal, 22 Jul 1984, Espejo 1023 (MEXU); Cadereyta, Jardín Botánico Regional de Cadereyta Ing. Manuel González de Cosío, 8 Apr 1992, Hernández Magaña et al. 9752 (MEXU); El Marqués, La Cañada, 12 Jun 2007, Martínez 6827 (MEXU); Querétaro cerca de las paredes, 20 Apr 1886, Urbino s.n. (MEXU); Peñamiller, ladera norte y noreste de Cerro Picacho, 29 Jul 1977, Zamudio 2305 (MEXU). San Luis Potosí: 4 miles NE of San Luis Potosi, 29 Aug 1947, Barkley et al. 783 (MEXU); Villa de Arriaga, a 22 km de Villa de Arriaga sobre la autopista, San Luis Potosi a Villa de Arriaga, a Lagos de Moreno por el km 54-55, 21 Aug 2009, Calzada 25305 (K); km 124 carretera Querétaro-San Luis Potosí, 24 Sep 1979, Equipo 5 (MEXU); km 30 carretera San Luis Potosí-Zacatecas, 7 Oct 1972, García M. et al. 322 (MEXU); Matehuala, Matehuala, frente del Motel de las Palmas, orilla de la carretera que va hacia Saltillo, entre el motel y la carretera, 24 Sep 1978, García P. 670 (CAS, MEXU); San Luis Potosí, Colonia Del Valle, 14 Jun 1973, Gómez Lorence 3 (MEXU); Vanegas, 35.8km de Vanegas a El Salado, 8 Nov 2008, Guzman 2947 (K); Charcas, Charcas, Jul 1934, Lundell 5145 (DS, K, MEXU); San Luis Potosí, carretera a la presa de San Jose, 20 Oct 1973, Moncada 4741 (MEXU); Central Mexico, chiefly in the region of San Luis Potosi, 1878, Parry & Palmer 636 (BM, E); San Luis Potosí, Aug 1876, Schaffner 695 (K, MEXU); 2 km del camino Rioverde-Tablas, 7 Aug 1960, Takaki 326 (MEXU); Guadalcázar, 1 km N del crucero Guadalcázar sobre la autopista a Matehuala, 27 Jun 2000, Torres C. 15684 (MEXU); Guadalcázar, Charco Blanco, 1 km del crucero San Luis Potosí-Matehuala, hacia Guadalcázar (bajos y lomerios entrando por un arroyo), 22 Sep 2012, Torres C. et al. 17481 (MEXU); Charcas, Jul 1934, Whiting905 (MEXU). Sonora: Plutarco Elios Calles, Sonoyta, NW side of town ca. 0.5 km S of river, 12 Oct 1986, Felger & Joseph 86-400 (MEXU); Puerto Peñasco, Puerto Peñasco, along railroad tracks, middle of town, 25 Jun 1985, Felger 85-795 (MEXU); Empalme, end of causeway at Empalme, along Mexican Hwy 15, 14 May 1973, Hansen et al. 1392 (MEXU); Bocum, Ejido Javier Mina, Block 409, Lote 2, 9 Apr 2009, López Valencia s.n. (MEXU); Naco, 10 km al SE de Naco, 24 May 1987, Tenorio L. & Romero de T. 13676 (MEXU); Fronteras, Bacoachic, 23.5 miles NE on road to Esqueda, 10 May 1948, Wiggins 11750 (DS). Tamaulipas: Matamoros, Mezquital, 61 km al E del carretera a Matamoros, 3 Oct 1994, Baro et al. 495 (MEXU); Burgos, El Mulato, vicinity, 16 Aug 1930, Bartlett 10991 (DS); east of Interamerican Hwy 85 23.7 miles S of Nuevo Laredo, 1 Jul 1969, Broome 313 (DUKE); Nuevo León, Nuevo Laredo, 14 km (9 miles) S of Nuevo Laredo on road to Monterrey, 17 Apr 1939, Frye & Frye 2383 (DS); Salinas Victoria, Transito Pesados Naciones Unidas, 1 Jun 1995, Galván 618 (MEXU); Nuevo Laredo, 50 miles southeast of Nuevo Laredo, 28 Mar 1964, García & García 40 (CAS, DUKE); Rancho el Mezquite, 35 km al SE de Santa Teresa, Mar 1963, González-Medrano 115 (MEXU); Jaumave, Ejido San Antonio, 17 Nov 1985, Jiménez 426 (MEXU); Nuevo Laredo, 20 km al oeste de [Nueva] Ciudad Guerrero, Oct 1973, López F. & Hernández Magaña 6335 (MEXU); Soto de la Marina, Poblado La Pesca, 24 Jul 2008, Martínez S. & Ibarra 40416 (MEXU); Miquihauana, 8 Aug 1941, Stanford et al. 802 (DS); Soto la Marina, due E of Cuidad Victoria about 100 miles along hwy 180 that parallels the coast, across from gas station in town, 8 Jul 1983, Taylor 1898 (DUKE); 30 miles S of Matamoros towards San Fernando, 14 Jul 1984, Wilbur 35155 (DUKE). Veracruz: Tampico Alto, Río Panuco, side road across from Tampico, ca. 2 mi. E of Pueblo Viejo, 11 Jun 1973, Hansen et al. 1791 (BH, MEXU); Villa Cuauhtemoc, Pueblo Viejo, at ferry landing across from Tampico, 11 Jul 1980, Nee & Hansen 18972 (MEXU); Puerto Viejo, vicinity of Pueblo Viejo, 2 km south of Tampico, 10 Feb 1910, Palmer 388 (BM, K). Zacatecas: Villa de Cos, El Capirote, 3 Jul 1997, Arteaga Saucedo 840 (MEXU); Pánuco, Valle Hermoso, 21 Jul 1986, Brigada Zacatecas 86 (MEXU); Calera, Calera, junction of Hwys 45 & 49, 16 Aug 1975, Gillett & Delgado 17013 (MEXU); Río Grande, Rancho El Carrizal, potrero Las Remudas, 7 km al E carretera Zacatecas-Torreó, entrando por km 41 desde el entroque Torreón-Durango, 31 Aug 1979, González E. & Cano s.n. (CAS); Campo experimental Noris de Guadelupe CNIZA, 35 km S de Concepción de Oro, 12 Jul 1975, González Medrano et al. 8003 (MEXU); 50 miles from Aguas Calientes on road to Zacatecas, ca. km 695, 7 km S of Ojo Caliente, 9 Sep 1958, Hawkes et al. 1463 (K); 70 (air) miles NE of Zacatecas, 66 (rd) miles NE of Hwy 41 along Hwy 54, 13 Sep 1971, Henrickson 6673 a (MEXU); Francisco Villa, 1 km S (on map Estación Symon), 28 Mar 1973, Johnston et al. 10443 (CAS, MEXU); Guadelupe, Lo de Vega, 13 Oct 1987, Medina 241 (MEXU); Fresnillo, km 97-98 carretera Mex. 45 entre Fresnillo y Sombrerete, justo en el crucero a Río La Medina, 5 Sep 2000, Rodríguez C. et al. 6100 (MEXU); Teul de González Ortega, La Haciendita, 16 Sep 1985, Saucedo Q. & Aceves s.n. (MEXU); Ojocaliente, Predio La Vibora, 28 Sep 1987, Silva R. 184 (MEXU); Ojocaliente, Predio La Lomas, 2 Oct 1987, Silva R, L. 187 (MEXU).

NETHERLANDS ANTILLES. Curaçao: sin. loc., Oct 1968, Arnoldo-Broeders 3621 (DUKE) .

PARAGUAY. Boquerón: Filadelfia, 8 Jun 1983, Hahn 1420 (BH); Río Verde, 26 May 1993, Mereles 5109 (CTES, FCQ); Fuerte Olimpo, 30 Oct 1946, Rojas 13678 (W); Colonia Fernheim, Estancia Laguna Porá, 1 Mar 1991, Vanni et al. 2620 (CTES); Central: Estero del Ypoa : Puerto Guyratí on Paraguay River, 22 Jun 1993, Zardini 36391 (AS, MO, SI) . Presidente Hayes: Ruta Transchaco km 82, 8 Mar 1991, Mereles 4010 (CAS, NY); km 65, Estancia Santa Maria del Doce Ret. San Juan , 1 Dec 2003, Mereles et al. 9095 (FCQ); Estancia Santa Asunción, 12 Oct 2003, Peña-Chocarro et al. 1483 (BM) ;

Peru. Cusco: Urubamba, Machu Picchu , 25 Jan 1975, Schwabe s.n. (B) ; Lima: Lima, Cuming 1090 (BM, E, K); Lima , La Molina , Dec 1986, Dreyfus s.n. (USM); Lima, 1825, Durville s.n. (W); Lima, La Molina, 8 Nov 1986, Vilcapoma 520 (USM) .

Puerto Rico. Ensenada, 28 Jun 1963, Liogier 9732 (DUKE, NY); Barrio Llanos Costa, Rancho Cabassa, on S. side of Sierra Bermeja , 16 Nov 1994, Proctor & Colón 49664 (MO) ; Sabana Grande, 4 Feb 1935, Sargent 513 (MO) ; Lajas, along road 324 several miles east of its intersection with road 304, 28 Jun 1965, Stimson 1403 (DUKE, MO); Punta Jorobado, rte 235, south of Ensenada , 17 Jul 1968, Wagner 1559 (DUKE) ; Ponce, 7 Sep 1979, Woodbury & Del Llano s.n. (MO, NY) ; Guanica, Ensenada , 19 Jun 1969, Wagner 1880 (BM, DUKE) ;

United States of America. Arizona: Yavapai County, Ashfork, 10 km W on road to Seligman, 27 Sep 1985, Bartholomew et al. 2430 (CAS, MEXU); Cochise County, Paradise, Chiracahua Mountains, 27 Sep 1907, Blumer 1732 (DS, K, NY, W); Mohave County, Kingman, 3 miles SE of Kingman in foothills of Hualapai Mountains, 7 Sep 1961, Breedlove 1150 (DS); Pima County, Twin Buttes, Sierrita Mountains, NW of Placer Peak, 25 Jun 1981, Butterwick & Hillyard 7782 (CAS); Pinal County, Bloody Tanks Wash, near Miami, 11 Jun 1982, Butterwick & Mittleman 8217 (CAS); Cochise County, Warren, Country Club S of Warren, Sulphur Springs Valley, 7 Jun 1915, Carlson s.n. (CAS); Yavapai County, Verde River, ca. 0.5 mile NE of Clarkadale [Clarkdale], 14 Apr 1960, Crosswhite 720 (K); Santa Cruz County, route 82 toward Patagonia. 5 miles past N of Nogales, 12 Oct 1979, Davis 1000 (BM, CAS, MEXU); Apache County, Springerville, Hwy 666, 3 miles S of P.O, 11 Jul 1963, Deaver 6439 (CAS); Santa Cruz County, Huachaca Military Reservation, inside west gate, 21 Jul 1969, Dreyer s.n. (CAS); Maricopa County, Wadell, ca. 1/ 4 mile N of Wadell, or 14 miles NW of Peoria, 17 Apr 1960, Dullas 135 (K); Gila County, Roosevelt Dam, Apache trail and adjacent regions, 22 May 1929, Eastwood 17097 (CAS); Yavapai County, Camp Verde, 4 miles ESE on main road, 24 Jun 1979, Ertter & Strachan 2939 (CAS); Adamana, 1954, Ewan 15 (BM); Pima County, below White House Canyon, Santa Rita Mountains, 16 Apr 1928, Graham 3619 (DS); Apache County, Canyon de Chelly National Monument, Antelope House Ruins, Canyon del Muerto, 29 Jul 1971, Halse 568 (ARIZ); Coconino County, Grand Canyon National Park, Havasu Canyon, 27 May 1950, Howell 26563 (CAS); Monroe, 22 Jul 1921, Jones s.n. (UC); Cochise County, San Pedro Riparian National Conservation Area, Upper San Pedro River floodplain, Escalante Crossing, near northern border of SPRNCA, 15 Jul 2002, Makings 1102 (MEXU); Yuma County, Gila Mountains, Mt. Ord, at base of Mt. Ord Road (FSR 636) in paved area where road widens to allow parking, 22 Oct 2005, Price 338 (MEXU); Pima County, Fort Lowell, 16 May 1894, Price s.n. (DS); Maricopa County, Tempe, corner of 8th Street and Van Ness, parking next to street, 19 Apr 1963, Sprankle 96 (DUKE); Pinal County, Sacaton, 13 Jul 1928, Thackery 226 (DS); 10 mi W on rt. 80 from Douglas, 12 Jun 1974, Tilton 374 (CORD, MO); Navajo County, Holbrook, 16 Jun 1901, Ward s.n. (K, NY); Santa Cruz County, Coronado Forest, 0.6 km S of Yellow Jacket Road junction, 29 Jun 1995, Way et al. WSSB- 3 (K); Maricopa County, Glendale, 1 mile W of Glendale, 21 Apr 1960, White 6 (K); Pima County, Tucson, 3 miles SW of Tucson, 4 Jul 1928, Wolf 2485 (CAS, DS); Cochise County, Douglas, 42 miles NE on road to Rodeo, New Mexico, 7 Jul 1928, Wolf 2558 (CAS, DS). Arkansas: Monroe County, Brinkley, 28 Aug 1934, Demaree 10857 (DS); Cleveland County, New Edinburg, 4 Jul 1938, Demaree 17916 (CAS, DS). California: Los Angeles County, San Fernando Valley, near Cahuenga Pass, 17 Jul 1917, Abrams 6607 (DS, NY); Yuba County, Honcut, Glen Clark Farm, 25 Aug 1969, Ahart s.n. (CAS); Tulare County, Sultana, at Santa Fe railway crossing on outskirts of town, 29 Jul 1941, Bacigalupi et al. 2663 (DS); Sonoma County, Joe Kohuke Place at mouth of Adobe Canyon, 24 Oct 1945, Baker 11244 (CAS); Contra Costa County, Pittsburgh, hiway S of Pittsburgh, 3 Sep 1946, Baker 11574 (CAS); Sacramento County, Sacramento valley and south, 24 Sep 1931, Ball s.n. (CAS, DS); Sutter County, Yuba City, 5 miles W along road, 13 Aug 1965, Bennett 8486 (DS); Kern County, just N of the jct between Hwy 58 and Bealville Road, on road to Caliente, between Bakersfield and Tehachapi, 17 Jun 1989, Charlton 3259 (MEXU); Yolo County, Davis, 3 miles N along Southern Pacific Railroad, 2 Aug 1964, Crampton 7136 (CAS); Solano County, railroad track between Suisern and Van Den, 28 Aug 1920, Eastwood 1040 (CAS); San Bernardino County, Needles, 23 Jun 1916, Eastwood 5958 (CAS); Los Angeles County, West Los Angeles, 25 Aug 1932, Ewan 680 (BM, CAS, DS, DUKE, K, NY); Amador County, Jackson, Don Favre property, Previtali Road, Sierra Nevada, 15 Jul 1968, Farnham s.n. (CAS); Glenn County, Feenstra Farm, N side of Newville Road between Rd. FF and Rd. E, ca. 2 miles W of Orland, 27 Jul 1987, Feenstra s.n. (CAS); Riverside County, E side of Hwy 95, W bank of Colorado River, 2.1 miles N of Palo Verde Dam Road, 22.4 miles W of San Bernardino County line, 3 Oct 1962, Fuller 9706 (CAS); Imperial County, Bard, 3.1 miles S of Bard, W of San Pasqual School, 17 Oct 1962, Fuller 9796 (CAS); Calaveras County, Wallace, W limits, N side of Hwy 12, Sierra Nevada, 23 Jun 1970, Fuller 19758 (CAS); Imperial County, Salton Sink, Brawley, 29 Jun 1965, Goeden & Ricker s.n. (CAS); San Luis Obispo County, El Pomar, 7 miles E of Templeton, Rienitz Ranch, 8 Aug 1957, Hardham 2804 (CAS); San Francisco County, San Francisco, Embarcadero, 1956, Howell s.n. (CAS); Tehama County, Red Bluff, 6 miles S of Red Bluff, 17 Jun 1934, Howell 12238 (CAS); Monterey County, King City, 21 Jun 1963, Howell 39457 (CAS); Mariposa County, White Rock Road, 6 miles SW of Westfall Road, 22 Sep 1971, Johnson & Keffer s.n. (CAS); Inyo County, Haiwee, 17 May 1935, Kerr s.n. (CAS); Merced County, Ingomar, 30 Jul 1948, Mason & Smith 8256 (DS); Contra Costa County, Pittsburgh, 2 miles W of Pittsburgh, 3 Sep 1946, Mason & Grant 13097 (DS); Imperial County, El Centro, 2 miles E of El Centro, 30 May 1917, McGregor 857 (DS); Mendocino County, Boonville, 9 Jul 1934, Ornbaum s.n. (CAS); Santa Barbara County, Santa Barbara, Southern Pacific Railroad W of town, 13 Sep 1953, Pollard s.n. (CAS); Ventura County, Foster Park, W side of Ventura Avenue (freeway) about 0.5 mile S of Foster Park, 9 Jul 1971, Pollard s.n. (CAS); Santa Barbara County, Santa Barbara, South Pacific Railway S of Samarkand Hotel, 24 Jul 1957, Pollard s.n. (W); Yolo County, Merritt Island, 3.5 miles S of Clarksburg near junction of county roads 140 & 142, 18 Aug 1969, Quick 69-15 (CAS); San Bernardino County, Cajon Pass, just below Cajon Campground, 17 Sep 1961, Raven 16690 (CAS); Mariposa County, Coulterville, 4.4 miles W on N side of Hwy 132, 22 Sep 1971, Roberson & Ferlatte s.n. (CAS); San Bernardino County, Yucaipa, between Ave E and F, east from Wilson Creek channel to top of slope, 4 Oct 1996, Sanders 19612 (CAS); Riverside County, The Badlands, west fork of Laborde Canyon near old rocket test facility, 26 Apr 2002, Sanders et al. 25026 (CAS); Tulare County, Prixley Wildlife Sanctuary, 15 Oct 1970, Shevock 502 (CAS); Kern County, along California Highway 178 just W of Kern River Canyon entrance, 9 Jul 1994, Shevock 12209 (CAS); San Mateo County, East Palo Alto, 779 Bell Street, 1 Sep 1962, Thomas 9978 a (DS); Tulare County, Angiola, 0.25 mile N of Angiola, 13 Jul 1955, Twisselmann 2248 (CAS); Kern County, Devil’s Den region, Twisselmann Road 1.5 miles W of Highway 33, Western San Joaquin Valley, 10 Nov 1963, Twisselmann 9088 (CAS); Kern County, Jerry Slough, at the Tracy Ranch ( US Public Health study area), San Joaquin Valley (E side of Kern County), 22 Aug 1968, Twisselmann 14887 (CAS); Monterey County, Spence, south of Salinas, 19 Oct 1943, Wheeler 5859 (BM); Riverside County, Hemet, at NW corner of Stetson Ave and Stanford intersection, 22 Sep 1996, White 4526 (CAS); Tehama County, Red Bluff, Jun 1917, Wickes s.n. (CAS); Riverside County, Blythe, field at edge of town, Palo Verde Valley, 28 Apr 1932, Wolf 3086 (DS); Monterey County, Lewis Creek, 4.5 miles into Lewis Creek, 13 Sep 1984, Yadon s.n. (CAS). Colorado: Mesa County, public right-of-way, along a fence-line just S of Patterson Road near 27 1/2 Road, 22 Aug 1998, Austin 277 (MESA); El Paso County, E of Fountain in a valley on old RR grade, 12 Jul 1935, Christ 991 (CS); Baca County, Comanche National Grassland, at junction of County Road M and County Road 13, ca 16 mi S of Pritchett, 4 Oct 2003, Elliot 11929c (RM); Fremont County, E edge of Florence, 4 Jun 1955, Harrington 796 (CS); Huerfano County, Wet Mountains, Wet Mountain Valley, Sangre de Cristo Range, and vicinity; County Road 330, 1.5 road mi S of Walsenburg (where road and railroad parallel), 28 Jul 1999, Hartman 65234 (RM); Las Animas County, Comanche National Grassland, along Purgatoire River Valley near Rourke Ranch, ca 28 air mi SSW of La Junta, 13 Jul 2007, Kuhn et al. 2605 (RM); Bent County, near Las Animas, 4 Aug 1954, Zonitch s.n. (CS). Florida: Escambia County, Pensacola, 30 Jun 1892, Curtiss 5913 (K, NY); Monroe County, Newport, Key Largo, 26 Mar 1898, Pollard et al. s.n. (BM); Monroe County, Key West, 30 Nov 1913, Small & Small 4841 (K, NY, W). Illinois: Chicago [cultivated?], 3 Aug 1897, Umbach s.n. (CAS). Indiana: Lawrence County, Anderson Farm belonging to Purdue University, 5 miles west of Bedford, 6 Jul 1939, Kriebel 8208 (DUKE). Kansas: Meade County, Meade, 13 Jul 1950, Horr 3521 (DUKE); Sumner County, South Haven, 3 miles E of town, 19 May 1967, Stephens 10953 (DS). Louisiana: Richland Parish, beside La. 183, 1.1 miles N of Holly Ridge and US 80, 24 Jul 1980, Dixon 3636 (MEXU); Calcasisu Parish, Lake Charles, 18 May 1915, Palmer 7678 (CAS, K); Boissier Parish, at edge of Bodcau Wildlife Management Area southeast of La. 160 and Ivan, Sec. 4, T20N, R11W, 29 May 1986, Thomas & Dorris 96112 (BM). Mississippi: De Soto County, on Red Banks Rd 2.8 mi S of the junction of this road and US Hwy 78, 2 Jul 1969, Ferrari 313 (MMNS); Washington County, 3 mi ESE of Leland, farm field by Bogue Philia Creek, 30 Aug 1994, MacDonald 7551 (MMNS); Rankin County, on Reese’s Farm in Old Byram community, field surrounding Stumps Lake, 24 May 1975, Snow s.n. (MMNS); Lafayette County, ca. 2 mi N of a point off College Hill Rd, ca. 2 mi NW of College Hill, 19 Jun 1967, Temple 5513 (MISS, MMNS); Wilkinson County, along county blacktop 2.8 mi. E of a point 9.4 mi. S of Adams-Wilkinson county line on U.S. Hwy. 61, 8 Aug 1969, Temple 12151 (MMNS). Missouri: Barry County, sin. loc., 25 Jul 1893, Bush 266 (K); Howard County, along valley of Missoury [i] River, north of quarry, 2 1/2-2 3/ 4 mi. north of Lisbon, 9 Oct 1956, Steyermark 83071 (BM). Nevada: Nye County, Ash Meadows, SE Ash Meadows, US Atomic Energy Commission’s Nevada Test Site, Amargosa drainage basin, 17 Jun 1969, Beatley 9028 (DS); Clark County, Overton, 6 May 1941, Clokey 5937 (CAS, DS); Clark County, Las Vegas, NW limits of Las Vegas, Hwy 95 1 mile SE of Decatur Lane, 10 May 1962, Fuller 8567 (CAS); U. S. Highway 95, 7 mi north of Searchlight, 7 Jul 1952, Gullion 383 (CORD); Lincoln County, Dray Lake Valley, ca. 2 mi by air SW of intersection with Bennet Pass Road on Black Canyon power line road, 15 Jun 2013, Gust 2206 (ENLC); Clark County, Moapa, Warm Springs, 24 May 2012, Johnson 12-029 (BRY); Clark County, Kyle Canyon, leaving canyon, Spring Mountains, 8.2 miles from Hwy 95, 30 Jul 1979, Williams 79-176 1 (CAS). New Mexico: Luna County, mesa at base of Little Florida Mountains, 24 Jul 1919, Abrams s.n. (DS); Eddy County, White’s City, 5 miles N of city, 17 Jun 1965, Barbour 119 (DUKE); Otero County, along Rt. near Rio Tularosa at west base of Sacramento Mt. front, 8 miles N.E. from Tularosa, 4 Aug 1965, Bennet 8659 (BM); San Miguel County, Las Vegas, vicinity, Oct 1919, Brother Anect 115 (CAS); Eddy County, Hope, Jun 1907, Campbell s.n. (CAS); San Miguel County, Soham, Foothills Ranch, 30 Jun 1958, Clear s.n. (CAS); Grant County, Silver City, 8 May 1919, Eastwood s.n. (CAS); Hidalgo County, Lordsburg, 15 May 1919, Eastwood 8550 (CAS); Santa Fe County, Glorieta, 12 Oct 1928, Eastwood 15419 (CAS); Bernalillo County, Sandia Mountains, on road from Santa Fe to Albuquerque, 16 Oct 1928, Eastwood 15646 (CAS); Socorro County, San Antonio, 21 Jun 1921, Ferris & Duncan 2302 (CAS, DS); Dona Ana County, Mesquite, 28 Jul 1930, Fosberg S-3399 (CAS, W); valley of Rio Grande, two miles north of Socorro, 3 Jul 1927, Goddard 836 (BM, K); Dona Ana County, Las Cruces, bank of Rio Grande River S of Las Cruces, 17 Jun 1930, Goodman & Hitchcock 1142 (CAS, DS); San Miguel County, Las Vegas Hot Springs, 30 May 1903, Grant 5550 (DS); Lea County, Paduca Habitat Evaluation Area, 31.7 air miles SW of Hobbs, 21 air miles WSW of Eunice, at old oil well drilling pad, 14 Aug 2009, Hansen 4421 (W); Doña Ana County, banks of Rio Grande near Doña Ana, 1 Aug 1958, Hawkes et al. 1166 (K); Santa Fe County, Canoncito, 18 Jun 1897, Heller & Heller 3733 (BM, DS, E, G, K, NY); Sandia County, Albuquerque, 3 Sep 1884, Jones 4119 (BM, CAS, G); Dona Ana County, Jornada del Muerto, base of Mt. Summerford, 8 Jun 1983, Lajtha 123 (DUKE); Colfax County, Vermejo Park Ranch, fields along south side of Ponil Creek at Cimarron Headquarters on northeast side of Cimarron, 15 Aug 2008, Legler 10779 (RM, UNM); Grant County, Dog Spring, 27 May 1892, Mearns 122 (DS, NY); Luna County, Carrizalillo Mountains, 15 Apr 1892, Mearns 135 (DS); Grant County, Mangas Springs, Aug 1901, Metcalfe s.n. (DS); Grant County, Santa Rita, 8 Aug 1895, Mulford 684 (K); Grant County, Lordsburg, 15 miles E of Lordsburg, 4 Sep 1938, Rollins & Chambers 2749 (DS); San Miguel County, Pecos Indian Ruins, Santa Fe National Forest, 21 Jul 1937, Sagalyn 69 (DUKE); Lincoln County, Gray, 1 Jun 1898, Skehan 16 (K, W); Guadelupe County, Santa Rosa, 20 miles W at junction of US 84 and 66, 20 Jul 1958, Smith 1030 (CAS); Grant County, Pinos Altos, mountains near Pinos Altos, 26 Jun 1936, Stewart s.n. (CAS); Union County, 7 mi. SW of Logan, 7 Jul 1949, Stiteler s.n. (BM); San Miguel County, 75 mi W of Roswell,, 11 Jun 1974, Tilton 321 (CORD, MO); San Miguel County, 3 mi W of Alamogordo, NM on rt. 82 toward Las Crucas, 11 Jun 1974, Tilton, D. 330 (CORD, MO); Dona Ana County, Mesilla, 28 Jun 1897, Wooton 59 (DS, E, G, K, NY); Dona Ana County, Mesilla Valley, 10 Aug 1907, Wooton & Standley 3127 (DS). North Carolina: Dare County, Nag’s Head, along US 158 business, northern part of Nag’s Head, 30 Jul 1970, Leonard & Radford 3394 (MEXU, NY). Oklahoma: Cleveland County, Norman, 2 Aug 1924, Bruner s.n. (DS); Creek County, Sapulpa, "I. Ten." (Plants of Indian Territory), 19 Jun 1894, Bush 392 (K); Beckham County, north side of North fork of Red River about 6.5 miles N of Texola, 9 Jun 1940, Clausen 4597 (K, NY); Comanche County, Fort Sill, 14 Jun 1916, Eastwood 11763 (CAS); Cleveland County, Norman, Dec 1944, Hopkins et al. 897 (DS); Custer County, 1 mile north of Weatherford, Jul 1938, Mericle 500 (BM, CAS, DS, DUKE, K); Alfalfa County, Aline, 8 Jun 1913, Stevens 802 (DS, K); Dewey County, Putnam, 11 Jun 1913, Stevens 889 (SI); near Oklahoma City, 16 Sep 1938, White 1180 (MEXU). Oregon: Umatilla County, Hermiston, Jul 1945, Hachler s.n. (OSC). South Carolina: Aiken County, corner of Ridgecrest Avenue and Martintown Road in: North Augusta, 9 Jun 1962, Ahles & Baird 56834 (BM, E, NY). Texas: Reeves County, Balmorhea, 4 mi S of Balmorhea on Texas 17, 4 Sep 1966, Anderson & Laskowski 3504 (DUKE); Bexar County, Bejar [Presidio San Antonio de Bexar], Berlandier s.n. (K); Jim Hogg County, Hebronville, 26 miles SW on Farm Road 496, 27 Mar 1965, Botello & Ayala 14 (DS); La Salle County, Encinal, US Hiway 81, 7 miles N of Encinal, 27 Oct 1962, Bruno 4 (DUKE); Nolan County, Sweetwater, 5 May 1925, Burkill 432 (K); Brazoria County, Columbia, [West Columbia], 20 Oct 1900, Bush 1588 (K); Grimes County, College Station, 13 miles E of city on Hwy 30, 3 May 1974, Calhoun 129 (CAS); El Paso County, Fort Bliss, 30 Apr 1915, Carlson s.n. (CAS); Edwards County, Substation No. 14, 5 Oct 1945, Cory 52468 (DS, NY); Zapata County, Zapata, 2 miles SE of Zapata, 25 Apr 1965, Cuesta 57 (DS); Travis County, Austin, junction Texas 270, Texas 271, 5 Sep 1978, D’Arcy 11689 (MEXU); Lubbock County, Lubbock, Texas Tech. campus, 13 May 1930, Demaree 7674 (DS); Jeff Davis County, Fort Davis, 19 Sep 1920, Eggleston 7424 (BM); Harris County, Houston, 21 Jul 1917, Fisher 5089 (CAS); San Patricio County, on N side of Nueces Bay, west of Portland, near junction of county rds. 66 and 67, 22 Apr 1998, Fryxell 5131 (MEXU); Zapata County, Mecom Ranch, hiway 83, 6 miles S of San Ignacio, 15 Mar 1964, García 23 (DUKE); Grayson County, Denison, edge of town, 20 Jun 1949, Gentry 51-393 (MEXU); Palo Pinto County, Mineral Wells, 8 Jun 1931, Gillespie 5222 (DS); Zapata County, Laredo, 22 miles S on US Highway 83, 4 Apr 1965, Guerra et al. 601 (DS); Waller County, Hempstead, 10 Jun 1872, Hall 498 (BM, G, K); Nueces County, Corpus Christi, 23 Mar 1894, Heller 1511 (E, G, K); Bowie County, near Texarkana, 31 Aug 1898, Heller & Heller 4185 (E, G); Presidio County, Chinati Mountains State Natural Area, along jeep trail W of Cienaga, SW of Sierra Parda, 4 Aug 2004, Lott et al. 5121 (CAS); Presidio County, Chinati Mountains State Natural Area, NE- to SW-trending tributary to San Antonio Canyon, below old San Antonio Mine, 21 Aug 2005, Lott et al. 5573 (CAS); Willacy County, San Perlita, 8 May 1940, Lundell & Lundell 8784 (DS); Bexar County, San Antonio, Our Lady of the Lake College campus, 29 Oct 1939, Metz 3164 (CAS); Webb County, Laredo, 29 May 1974, Nee 11787 (K, MEXU); Travis County, Austin, west side of Austin, along Red Bud Road just W of Colorado River, 30 May 1974, Nee & Whalen 11815 (MEXU); Hidalgo County, near entrance to Bentsen-Rio Grande State Park, 4 mi SW of Mission, 18 Dec 1981, Nee 24064 (CORD, F); Parker County, 8.3 km NW of center of Weatherford, Shady Grove Road, 1.3 km N of Old Garner Road, 22 May 2010, Nee 57070 (MEXU, NY); Tarrant County, along Long Ave. at Braswell Drive, 1/ 4 km W of I-35 W, 6 km NNE of center of Fort Worth, 26 May 2010, Nee 57083 (NY, SI); Terrell County, Sanderson, 26 Jun 1924, Orcutt 680 (DS); Taylor County, Abilene, Aug 1883, Parish s.n. (DS); Dallas County, Dallas, Jun 1880, Reverchon s.n. (G); Frio County, Dilley, US Highway 81, 2 miles N of Dilley, 22 Mar 1964, Rodríguez 43 (DUKE); Victoria County, Victoria, 16 Sep 1908, Schallert 540 (DUKE); Washington County, Millerick Settlement, 1856, Schlottmann 269 (LE); Atascosa County, 'southern Atascosa County’, 20 Apr 1920, Schulz 97 (CAS); Maverick County, 5 miles N of Normandy, 10 Sep 1962, Scora 2285 (MEXU); Jeff Davis County, Valentine, 0.5 miles S of Valentine, 18 Aug 1953, Selander 21 (DS); Kinney County, Brackettville, 4 miles NE of town, 15 May 1965, Strother 258 (DS); Hidalgo County, about 5 miles W of Mission, in railroad cut, 30 Apr 1949, Tharp & York 51- 261 (MEXU); Wichita County, 10.6 mi outside Wichita Falls on route 82 W to Lubbock, 9 Jun 1974, Tilton 218 (CORD, MO); Baylor County, 21 mi W of Seymour on rt. 82 (roadside has been moved very recently from Seymour to Vera), 9 Jun 1974, Tilton 237 (CORD, MO); Taylor County, Camp Barkeley, 1 Nov 1942, Tolstead 5859 (W); Galveston County, Galveston Island, 23 Sep 1901, Tracy 7576 (BM, CAS, W); Chambers County, Wallisville, ca. 1/ 4 mile S of Tex hwy 73, ca. 1/ 8 mile N of Trinity River, along road toward river, S side of road, 14 Jul 1958, Traverse 802 (MEXU); Kaufman County, Mabank, on highway 175, 1/ 4 mile E of the second bridge over the reservoir, 5 Jun 1996, Whitson & Whitson 803 (DUKE); Kimble County, on West I-10 at the Fort McKavett overpass, 41 miles E of Sonora, 16 Jun 1996, Whitson & Whitson 817 (DUKE); Cameron County, Brownsville, near the Western Union Station on St. Charles Street, 14 Jul 1984, Wilbur 35189 (DUKE); Hale County, about 3 miles N of Abernathy, near small lake, 25 Sep 1948, York & Cowan 52-300 (MEXU); Lubbock County, Lubbock, 2 May 1934, Zobel s.n. (CAS). Utah: San Juan County, 2 mi E of Bluff, 21 May 1985, Atwood 11023 (NY); Washington County, near Zion National Park, Jun 2006, Bohs s.n. (UT); Kane County, Kaiparowits Plateau, south side of US Highway 89 ca. 1 mile east of The Cockscomb, 15.5 miles west of Big Water, 7 Jun 2006, Fertig 22568 (NY); Wshington County, St. George, 12 Nov 1999, Higgins 20971 (NY); San Juan County, Glen Canyon NRA, San Juan River mile 44, Honaker Trail, 16 Jun 2003, Hill 225 (BRY); Tooele County, Open Reventment Area, 3 Aug 2000, Long 1100 (BRY, RENO). Washington: Asotin County, Snake River Canyon, 5 miles upstream (SE) from intersection (in Asotin) w/ Hwy 3 at east end of town, and ca ½ SSE of confl[uence] of Snake River and Tenmile Cr. along road from Asotin to Heller Bar, 21 Aug 1983, Henderson & Cholewa 6848 (NY).

Uruguay. Colonia: Colonia , 6 Jan 1902, Berro 7942 (G) . Río Negro: Fray Bentos , 5 Feb 1879, Felippone 5092 (K) . Soriano: Villa Soriano , 24 Jan 1908, Berro 4411 (G) .

Figures

Figure 2. Flowers of members of the Elaeagnifolium clade. A Solanum elaeagnifolium, with divergent anthers of approximately equal size and shape (Argentina, Mendoza, Knapp et al. 10470) B Solanum homalospermum (Argentina, Catamarca, Chiarini et al. 505) C Solanum hindsianum (cultivated in Arizona) D Solanum houstonii, hermaphroditic flower and strongly curved buds (Mexico, Yucatan, Pena-Chocarro et al. 407) E Solanum houstonii, staminate flower (Mexico, Queretaro, Ochoterena et al. 976) F Solanum mortonii (cultivated in Cordoba; from Barboza et al. 644). Photographs A, D, E by S. Knapp; B, F by F. Chiarini; C by Eugene Sturla.

Figure 3. Fruits of members of the Elaeagnifolium clade. A Solanum elaeagnifolium, immature berries with green mottling (Argentina, Cordoba, Barboza et al. 3434) B Solanum elaeagnifolium, mature fruit with sticky tan seeds (Argentina, Cordoba, Barboza et al. 3435) C Solanum mortonii, immature fruit exerted from the calyx (Argentina, Catamarca, Barboza et al. 3439) D Solanum mortonii, mature fruit exerted from the calyx (Argentina, Catamarca, Barboza et al. 3438) E Solanum houstonii, immature fruit enclosed in accrescent calyx (Mexico, Queretero, Ochoterena et al. 976) F Solanum houstonii, mature fruit enclosed in accrescent calyx with black seeds (Mexico, Queretero, Ochoterena et al. 976). Photographs by S. Knapp.

Figure 4. Habitats of members of the Elaeagnifolium clade. A Solanum elaeagnifolium, old stems with persistent berries and young new stems from underground rhizomes (Argentina, Mendoza, Knapp et al. 10470) B Solanum houstonii on rocky slope (Mexico, Queretaro, Ochoterena et al. 976) C Solanum mortonii, plant dug up showing extensive underground stems (Argentina, Catamarca, Barboza et al. 3437) D Solanum mortonii, large population on rocky slopes (Argentina, Catamarca, Barboza et al. 3437). Photographs by S. Knapp

Figure 5. Solanum elaeagnifolium Cav. A Fruiting branch B Flower C Spread corolla D Dorsal view of stamen E Ventral view of stamen F Gynoecium G Glandular trichome from gynoecium H Fruit I Lepidote trichome from calyx, seen from above J Lepidote trichome from calyx, seen from the side K Seed L Transverse section of seed M Embryo. Drawn by Leonor Sanchez. Reproduced with permission from Flora Argentina (Chiarini 2013).

Figure 6. Distribution of S. elaeagnifolium.