Solanum aridum, Morong, Ann.

Wahlert, Gregory A., Chiarini, Franco E. & Bohs, Lynn, 2015, A Revision of Solanum Section Lathyrocarpum (the Carolinense Clade, Solanaceae), Systematic Botany 40 (3), pp. 853-887 : 862-865

1. SOLANUM ARIDUM Morong,

Ann. New York Acad. Sci. 7: 173. 1893.

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TYPE: PARAGUAY. [Presidente Hayes?]: Falls of the Pilcomayo River , 2 May 1888 –1890 (fl), T. Morong 1007 ( lectotype: US – US 00027459 [ US –48029 ]!, designated by C. V. Morton, Revis. Argentine Sp. Solanum : 229. 1976; isolectotypes: BM– BM000087516 p.p. [scan!], E– E00190949 p.p. [scan!], GH– GH00077578 p.p.!, K– K000532483 [scan!], MO–2766698 !, NDG– NDG044996 [scan!], NY– NY00139054 p.p.!, PH – PH 00030383 [ PH – 01029054 ] [scan!], WIS ex WELC–15091 p.p. [scan!]).

Solanum elaeagnifolium var. ovalifolium Kuntze, Revis. Gen. Pl. 3: 225. 1898.—

TYPE: ARGENTINA. Salta: Monte Morro, Nov 1892 (fl, fr), C. E. O. Kuntze s. n. (lectotype, designated here: NY –NY00139143!; isolectotype: US – US 00027563 [ US –701686]!).

Solanum aridum var. pusillum Hassl., Trab. Mus. Farmacol. 21: 106. 1909.—

TYPE: PARAGUAY. Presidente Hayes: Salto Palmares, Jul 1906 (fl, fr), T. Rojas 346 (lectotype, designated here: G–G00306542!).

Solanum conditum C. V. Morton, Revis. Argentine Sp. Solanum 237. 1976.—

TYPE: ARGENTINA. Santiago del Estero: Departamento C. Pellegrini, Estancia El Remate, 500 m, 22 Dec 1927 (fl), S. Venturi 5685 (holotype: US – US 00027522 [ US –1548974]!; isotypes: LIL–LIL88555 [scan!], NY– NY00688148!, SI–SI065951!, SI–SI065952!).

Upright perennial herb up to 0.5 m tall; stems simple or few-branched; roots producing buds and tuber-like ellipsoid swellings ca. 1–3 × 1 cm. Stems sparsely to densely pubescent with sessile stellate hairs 0.15–0.4 mm in diameter, with 4–8 lateral rays, the central ray absent or up to 0.3 mm long, unarmed or very sparsely armed with straight tapered prickles up to 3.5 mm long. Sympodial units 2- to 3-foliate when more than one inflorescence per shoot, but often only one formed and then appearing plurifoliate, the leaves not geminate. Leaves simple, the blades 2–10 × 1–5 cm, ovateelliptic, elliptic, or oblong in outline, discolorous, densely stellate-pubescent abaxially, less densely so adaxially with hairs like those of the stems, unarmed or sparsely armed with prickles up to 3.3(5) mm long on the major veins abaxially and adaxially; base cuneate to subtruncate; margin subentire, sinuate or shallowly lobed with 2–6 lobes per side; apex obtuse to acute; petioles 0.5–5 cm long, densely stellate-pubescent, unarmed or sparsely armed with prickles up to 4 mm long. Inflorescences 1–2.5 cm long, extra-axillary, unbranched or forked, with 1–6 flowers, weakly andromonoecious with the proximal flowers hermaphroditic and the distal flowers staminate, the axes moderately to densely stellate-pubescent, unarmed or very sparsely armed with prickles up to 3 mm long; peduncle up to 2 cm long or absent, with the lowermost flower(s) emerging directly from the node; pedicels 1–2(3) cm long in flower, 2–4 cm long and curved downward in fruit, spaced 0.5–1.5 cm apart, articulated at the base, moderately to densely stellate-pubescent, unarmed or very sparsely armed with prickles up to 1.5 mm long. Calyx 7–10 mm long, the tube 2.5–4 mm long, the lobes 4–7 × 1.8–2.5 mm, linear to lanceolate, often unequal, the apex caudate to acuminate, densely stellate-pubescent abaxially, glabrous adaxially, unarmed or very sparsely armed abaxially with prickles up to 3 mm long; fruiting calyx somewhat accrescent but not completely covering the fruit, 10–20 mm long, the tube 2–4 mm long, the lobes 6.5–15 × 2–5 mm, narrowly triangular, moderately stellate-pubescent, unarmed or very sparsely armed with prickles up to 3.5 mm long. Corollas 1.7–3.5 cm in diameter, 12–16 mm long, stellate to stellate-pentagonal or rotate-stellate, chartaceous, usually white, sometimes pale blue-violet, the tube 2–6 mm long, the lobes 5–9 × 4–8 mm, deltate to triangular, the apex acute, densely stellate-pubescent abaxially, glabrous adaxially. Stamens with filaments 0.6–1.8 × ca. 0.2 mm; anthers 7–8.5 × 1.2–2 mm, narrowly lanceolate, somewhat connivent, yellow, the pores directed distally. Ovary 1.3–2.5 × 1.2–2.5 mm, subglobose-ovoid, glabrous; style 10–12 × 1–1.5 mm, cylindrical, straight, glabrous, exserted in hermaphroditic flowers; stigma capitate. Fruits 1.2–2.2 × 1.2–1.8 cm, ellipsoid to ovoid, the apex obtuse to somewhat pointed, light green with darker green mottling when young, yellow or pale yellow to yellow-orange at maturity, glabrous, the rind tough. Seeds 1.5–2.3 × 1.1–2 mm, flattened-reniform, lenticular, yellow, the surface finely foveolate.

Distribution and Habitat — Solanum aridum occurs in northern and central Argentina, eastern Bolivia, and western Paraguay ( Fig. 4 ). It typically grows in disturbed areas such as roadsides and waste areas, often in sandy or clay soils. Most specimens examined occur at elevations between 300–1,200 m, but one sheet (C. Peláez 308) reported an elevation of 2,025 m.

Phenology — The species flowers between September and May, and fruits between November and July.

Conservation Status — Solanum aridum is distributed over a large area in Argentina, Bolivia, and Paraguay with a calculated extent of occurrence of ca. 1 × 10 6 km 2 and an area of occupancy of 1,376 km 2. It is common throughout its range and typically grows in disturbed areas, and there is no evidence that the size of its populations or number of locations are declining. Solanum aridum is therefore assigned a preliminary conservation status of “least concern” (LC).

Etymology — The epithet aridum means dry or withered and was applied by Thomas Morong ( Morong and Britton 1893) because of the perceived “parched, dried appearance” of the plant.

Vernacular Names and Uses — Matesevach (2002) and Barboza (2013) report several common names from Argentina: comida de víbora, meloncillo, melon del campo, and pocotillo. Two specimens from Bolivia, G. Bourdy 2082 and R. de Michel et al. 2539, collected in Cordillera Province in Santa Cruz Department, recorded the names tutia and tutiami, respectively. Label information from G. Bourdy 2082 states that the fruit was ground and used to treat scabies. The name tutia very likely refers to the ellipsoid tuber-like swellings on the roots, a name similar to tutía-tutía, which was applied to S. hieronymi , another species with root swellings.

Chromosome Number — Several meiotic and mitotic chromosome counts of vouchered specimens have shown Solanum aridum to be a diploid, with n = 12 (R. Subils et al. 3566; G. Bernardello et al. 487, as S. juvenale ; Di Fulvio 803, as S. juvenale ) and 2 n = 24 (G. E. Barboza et al. 276; G. E. Barboza et al. 782; F. Chiarini 16) ( Moscone 1992; Chiarini and Bernardello 2006; Chiarini 2007).

Notes — In habit and reproductive characters, Solanum aridum resembles S. juvenale , and some intermediate forms can be found where the two species’ ranges overlap ( Morton 1976; Chiarini 2007). However, S. aridum can be separated from S. juvenale by the absence or very sparse density of prickles on the stem, petioles, leaf blades, inflorescence axes, pedicels, and calyces, whereas S. juvenale is typically moderately to densely armed with prickles, particularly on the calyces. The prickles on the leaf blades are usually much shorter in S. aridum (up to 3 mm long), while in S. juvenale they are up to 8 mm long. Solanum aridum also has larger leaf blades (2–10 × 1–5 cm) that are usually sinuate to shallowly lobed, compared to smaller leaf blades in S. juvenale (1.8–5.5 × 1–2.5 cm) that are more deeply lobed.

The ploidy level consistently differs between the two species, with Solanum aridum being 2 n = 24 and S. juvenale being 2 n = 48. Experimental studies in the greenhouse by Chiarini (2007) showed that crosses between S. aridum and S. juvenale produced triploid plants (2 n = 36) and that ca. 90% of reciprocal crosses between the two species resulted in offspring that produced fruit with an expected amount of seeds, yet only ca. 7% of seeds were viable. Solanum aridum was also shown to be self-incompatible.

Of the 11 sheets of T. Morong 1007 we examined, five are mixed collections of Solanum aridum and S. multispinum (the WIS ex WELC sheet has a specimen of Turnera sp.) and one sheet is composed only of S. multispinum . Nee et al. (2006) outlined the mixed elements of the T. Morong 1007 collection, but here we explicitly identify the elements of each sheet. The sheets at K, MO, NDG, and PH consist solely of S. aridum . The BM sheet contains seven elements; six are Solanum aridum , but the densely prickly plant outlined in pencil with the round fruit is S. multispinum . The sheet at E contains three elements. The left-hand one is S. multispinum and the two right-hand ones are S. aridum ; these have been annotated by M. Nee as “a.” and “b.” for S. multispinum and S. aridum , respectively. The GH sheet consists of five elements; the four densely prickly specimens, each with a round fruit, are S. multispinum , and the central element with few prickles and buds only is S. aridum . The sheet at NY has five elements; the four flowering ones are S. aridum , and the densely prickly one with the round fruit is S. multispinum . The WIS ex WELC sheet contains six elements; five are S. aridum (four with fruit), and the other was determined by M. Nee as Turnera sp. and is indicated in pen on the sheet.

Finally, of the two sheets of T. Morong 1007 at US, one sheet ( US – US 01014263 [ US –1324449]) is composed only of S. multispinum while the other ( US – US 00027459 [ US –48029]) contains only S. aridum . When selecting one of the two US sheets as the lectotype, Morton (1976) was unambiguous: “One of these plants [ US –1324449] is the very spiny plant with stipitate-glandular stems, peduncles, and pedicels, and leaf-blades cordate at [the] base that was described a little later as S. multispinum N. E. Brown ; the other [ US –48029] is a nearly spineless plant (the few spines present being small and inconspicuous), lacking stipitate-glands, and with cuneatebased leaf-blades. I choose the latter as lectotype ...” The sheet is also annotated as “ lectotypus ” in Morton’ s handwriting. Nee (in Nee et al. 2006) incorrectly states that the lectotype sheet is a mixture of S. aridum and S. multispinum when in fact it is composed exclusively of S. aridum .

In the protologue of Solanum elaeagnifolium var. ovalifolium, Kuntze (1898) cited his collection (C. E. O. Kuntze s. n.) from Salta Province, Argentina. Two specimens of this collection were located at NY and US. Kuntze did not indicate the location of a holotype, so we have chosen the specimen C. E. O. Kuntze s. n. (NY–NY00139143) as the lectotype. Both NY and US specimens are from Kuntze’ s herbarium and are annotated in his handwriting, but we chose the NY specimen as lectotype because it is the more complete of the two and contains several additional flowering and fruiting elements. Similarly, the collection T. Rojas 346 from Presidente Hayes Department, Paraguay was cited in the protologue of S. aridum var. pusillum , but the herbarium where the collection was deposited was not indicated. Although the specimen now conserved at G constitutes original material, we cannot be completely certain that it was the only one used for the description, and thus we have designated the specimen T. Rojas 346 (G–G00306542) as lectotype.

Additional Specimens Examined — ARGENTINA. Catamarca: Dpto. La Paz, km. 981, 4 Apr 1950 (fr), J. Brizuela 1164 (C); Dpto. Santa Rosa, Bañado de Obanta, 12 Mar 1944 (fl, fr), S. A. Pierotti s. n. (NY); El Alto, Balcozna, 19 Jan 1928 (fl), S. Venturi 7124 (SI). Chaco: Dpto. Resistencia, Margarita Belén, 2 Feb 1948 (fl, fr), R. M. Aguilar 1235 (NY); Dpto. Primero de Mayo, a 6 km de Cnia. Benítez, rumbo al Río Paraná, 15 Dec 2002 (fl), G. E. Barboza et al. 527 (CORD, MO); Dpto. Primero de Mayo, al costado de la RN11, entre Resistencia y Colonia Benítez, 27°23′57.9″S, 58°59′49.8″W, 4 Mar 2012 (fl), F. Chiarini & G. A. Wahlert 878 (CORD, UT); Dpto. Bermejo, al costado del camino, cerca de General Vedia, 27°00′47.1″S, 58°42′08.6″W, 5 Mar 2012 (fl), F. Chiarini & G. A. Wahlert 887 (CORD, UT); Dpto. Maipú, RP9, cerca de Tres Isletas, 26°20′20.3″S, 60°18′06.7″W, 6 Mar 2012 (fl), F. Chiarini & G. A. Wahlert 905 (CORD, UT); Dpto. Presidencia de La Plaza, RP7, Parque Nacional Chaco, 26°43′48.1″S, 59°39′30.8″W, 7 Mar 2012 (fl), F. Chiarini & G. A. Wahlert 907 (CORD, UT); Presidencia de la Plaza, RP 7, 26°53′47.0″S, 59°49′07.8″W, 07 Mar 2012 (fl, fr), F. Chiarini & G. A. Wahlert 912 (CORD, UT); San Fernando, RN 11, 10 km saliendo de Resistencia hacia Formosa, 5 Jan 1985 (fl), T. E. Di Fulvio 803 (CORD); Cmte. Fernández, Col. Rivadavia, 27 Mar 1977 (fl), Pérez 257 (CTES). Córdoba: Capital, Barrio General Paz, 1 Dec 1998 (fl), F. Chiarini 16 (CORD); Capital, Barrio Observatorio, 20 Nov 1950 (fl), A. T. Hunziker 8566 (CORD). Formosa: Pilcomayo, alrededores de Laguna Blanca, 25°07′50″S, 58°15′57″W, 14 Dec 2002 (fl), G. E. Barboza et al. 513 (CORD); Dpto. Patiño, RN81, a ± 3km de Las Lomitas, 24°42′38.5″S, 60°34′44.5″W, 5 Mar 2012 (fl, fr), F. Chiarini & G. A. Wahlert 894 (CORD, UT); Laishi, San Francisco de Laishi, Reserva Ecológica El Bagual, 20 Dec 1998 (fr), A. Di Giacomo 378 p.p. (CTES);

Dpto. Matacos, 13 km N de Ing. G. Juárez, ruta prov. 39, 23°47′S, 61°49′W, 23 Nov 2000 (fl, fr), R. H. Fortunato et al. 6729 (BAB-n.v., MO, NY); Dpto. Pilagá, Pilagá, 3 Oct 1945 (fl), S. A. Pierotti 4257 (BM); Patiño, Las Lomitas, 14 Nov 1984 (fr), A. Schinini & G. Bernardello 24108 (SI). Jujuy: San Pedro de Jujuy, 16 May 1945 (fl, fr), J. Herrera 455 (BM); Dpto. San Pedro, Ruta Nac. 34, El Quemado, 9 Nov 1974 (fl, fr), A. Krapovickas et al. 26695 (CTES, MO, WIS); Ledesma, entre Fraile Pintado y Ledesma, 17 Dec 1968, P. R. Legname & A. R. Cuezzo 5975 (LIL); Dpto. San Pedro, 2 km al SW de La Mendieta, 24°19.87′S, 64°59.02′W, 800 m, 13 Apr 2000 (fl, fr), M. H. Nee et al. 50760 (CORD, NY); Dpto. El Carmen, Perico, 936 m, 15 Feb 1937 (fl), J. West 8355 (MO). Salta: Dpto. Chicoana, El Carril, en el pueblo, al costado del camino, rumbo a Cachi, 30 Sep 2001 (fl, fr), G. E. Barboza et al. 276 (CORD, NY); Dpto. Capital, alrededores de la Plaza de Barrio Ceferino Velarde, 19 Jan 2002 (fr), G. E. Barboza 331 (CORD, NY); Dpto. Anta, por ruta prov. 5, 15 km antes de Las Lajitas, 24°49′29″S, 64°19′29″W, 21 Mar 2005 (fl, fr), G. E. Barboza et al. 1262 (CORD); Dpto. Capital, vías del ferrocarril, 20 Mar 1989 (fl), A. Del Castillo 1138 (MCNS); Salta, pr. oppidum, 24 Sep 1901 (fl, fr), R. E. Fries 579 (S); Dpto. Capital, Parque San Martín, 1200 m, 25 May 1978 (fl), G. Jaime 46 (MCNS); Dpto. Anta, San Severo a San Javier, 29 Mar 1985 (fl), F. Juárez & A. Del Castillo 1195 (MCNS); Dpto. San Martín, pasando Río Bermejo, 30 Nov 1988 (fl, fr), F. Juárez 1737 (MCNS); Dpto. Capital, ampliación Barrio El Tribunal, calle Peñaloza, 1200 m, 19 Mar 1989 (fl, fr), F. Juárez 1775 (MCNS); Dpto. Capital, camino a la Isla, 1200 m, 19 Mar 1989 (fl, fr), F. Juárez 1791 & 1792 (MCNS); Dpto. Metán, El Tunal, 9 Nov 1974 (fl, fr), A. Krapovickas et al. 26711 (MO); Orán, 5 km antes de Embarcación, 2 Jun 1963 (fl), P. R. Legname & A. R. Cuezzo 4064 (LIL); Hwy 34 from Jujuy to Tartagal, 20.5 km SW of turnoff to San Ramón de la Nueva Orán, 23°25.95′S, 64°21.24′W, 335 m, 10 May 2005 (fl, fr), M. H. Nee 52964 (NY); along Hwy 24 from Jujuy to Tartagal, 9.4 km SW of turnoff to General Mosconi and 15.4 km SW of Tartagal, 22°40.71′S, 63°48.83′W, 430 m, 10 May 2005 (fl), M. H. Nee 52965 (NY); Dpto. Capital, ciudad de Salta, Parque San Martín, 1187 m, 28 Jul 1977 (fr), L. J. Novara 416 (CORD, MCNS); Dpto. Capital, FFCC a Quijano, altura de la calle Leguizamón, 1200 m, 6 Nov 1977 (fl, fr), L. J. Novara 539 (CORD, MCNS); Dpto. San Martín, Ruta 81, km. 14, entre Senda Hachada y Hickmann Fca. Yaveré, 1 Oct 1983 (fl), L. J. Novara 3790 (MCNS, MO); Dpto. Capital, La Lagunilla, 5–6 km E del Cerro San Bernardo, 1,000 m, 23 Oct 1988 (fl, fr), L. J. Novara 8150 (MCNS); Dpto. Captital, ciudad de Salta, FF.CC., 1187 m, 3 Apr 1999 (fl), L. J. Novara 11272 (MCNS); Dpto. Capital, ciudad de Salta, 1187 m, 4 Mar 2000 (fl), L. J. Novara 11397 (MCNS); Dpto. Metán, Metán, 30 Mar 1945 (fl), C. A. O’ Donell 2468 (NY); Dpto. Campo Santo, Güemes, 3 Apr 1945 (fl, fr), C. A. O’ Donell 2647 (NY); Dpto. Capital, ciudad de Salta, Parque San Martín, 1187 m, 21 Apr 1983 (fl, fr), M. del C. Otero 3481 (MCNS); San Javier, 19 km al S de Joaquín V. González, 16 Dec 1987 (fl), C. Saravia Toledo 1517 (CTES); Dpto. Capital, San Bernardo, 1800 m, 3 Dec 1946 (fl), B. Sparre 1117 (S); Dpto. Capital, ciudad de Salta, 1187 m, 28 Aug 1978 (fl), C. Suárez 20 (MCNS); Dpto. Capital, Cerro San Bernardo, 5 Mar 1991 (fl, fr), J. Tolaba 98 (MCNS); Dpto. Capital, ciudad de Salta, Cementerio de la Santa Cruz, 23 Dec 1998 (fl), J. Tolaba 1294 (MCNS); Dpto. Capital, Finca La Lagunilla, camino a Mojotoro, 4 km N de Ruta Nac. 9, km. 1585, 12 km E de la ciudad de Salta, 1100–1200 m, 29 Jan 2000 (fl), J. Tolaba 2340 (MCNS); Dpto. General Güemes, Torzalito, Ruta 9, 9.5 km S del Río Mojotoro, 750 m, 24 Oct 1986 (fl, fr), F. Varela 1509 (MCNS); Dpto. Capital, Parque San Martín, 1190 m, 9 Oct 1964 (fl), L. Vargas 1005 (MCNS); same locality, 3 Mar 1965 (fl), L. Vargas 1036 (MCNS); Dpto. Capital, ciudad de Salta, 1200 m, 25 Oct 1976 (fl), S. Zapata 80 (MCNS). Santa Fe: without precise locality, 28 Oct 1964 (fl), Alonso et al. 550 (CORD). Santiago del Estero: Robles, Colonia Jaime, 12 Nov 1948 (fr), F. E. Luna 1367 (CORD); Dpto. Banda, Dique Los Quiroga, 23 Dec 1978 (fl), R. Subils & G. Bernardello 23155 (CORD). Tucumán: Colombres, Camino a San Andrés, 4 Feb 1945, A. R. Cuezzo 14 (LIL); without precise locality, 6 Mar 1900, T. J. V. Stuckert 8723 (CORD); Capital, Río Salí, 4 Apr 1922 (fl), S. Venturi 1777 (SI); Dpto. Capital, Río Sali, 450 m, Jan 1926 (fl, fr), S. Venturi 4093 (NY).

BOLIVIA. Chuquisaca: Prov. Luis Calvo, Ñancaroinza, just E of railroad tracks of railroad from Santa Cruz to Yacuiba, 20°42′20″S, 63°17′16″W, 740 m, 19 Mar 2007 (fl, fr), M. H. Nee & R. Flores S. 54781 (NY); Prov. Luis Calvo, El Salvador, 7 Dec 1992 (fl, fr), J. Pensiero & D. Marino 4334 (MO); El Salvador, 18 Dec 1991 (fl), C. Saravia Toledo & J. Joaquín 10643 (CORD). Cochabamba: Prov. Mizque, on the road Vila Vila to Station Cruz, 18°00′S, 65°30′W, 619 m, 21 Dec 1949 (fl), W. M. A. Brooke 5901 (BM, NY); Mizque, entre la plaza y el río, saliendo hacia el sur (a dos cuadras de la plaza) 17°33′49″S, 65°12′09″W, 2025 m, 2 May 2008 (fl, fr), C. Peláez 308 (CORD). Santa Cruz: Prov. Cordillera, around highway and railroad bridges over Río Seco on N side of settlement of Río Seco, along new highway from Santa Cruz to Abapó, 18°39′S, 63°15′W, 550 m, 24 May 1998 (fl), L. Bohs & M. Nee 2733 (UT); Prov. Cordillera, bañados del Río Parapetí, Izozo [Izozog], 16 km al N de al comunidad Cuarirenda, 18 Sep 1998 (fl), G. Bourdy 2082 (NY); Prov. Cordillera, Izozog, Comunidad guaraní de Ibasiriri, a 3 km de “La Brecha”, en la ribera del Río Parapetí, 300 m, 18 May 1999 (fl), R. de Michel et al. 2539 (NY); Prov. Cordillera, 20 km N de San Antonio Parapatí, 19°50′S, 63°10′W, 700 m, 20 Jun 1992 (fr), T. Killeen et al. 4237a (AAU-n.v., LPB, MA-n.v., MO, NY); Prov. Andrés Ibáñez, between Puerto Pailas and Monte Bello, 28 Apr 2000 (fl, fr), N. Kuroiwa & A. Maeda 1158 (NY); Prov. Andrés Ibáñez, W side of Santa Cruz, between second and third Anillos, 17°47′S, 63°12′W, 420 m, 16 Jan 1987 (fl, fr), M. H. Nee 33575 (CORD, LPB, NY); Prov. Andrés Ibáñez, Boyuibe, 20°26′S, 63°17′W, 810 m, 23 Jul 1987 (fr), M. H. Nee 35329 (CORD, NY, P); Prov. Andrés Ibáñez, on E side of city of Santa Cruz, 17°47′S, 63°10′W, 420 m, 21 Nov 1988 (fl, fr), M. H. Nee 36907 (LPB, NY); Prov. Andrés Ibáñez, 2 km NW of Puerto Pailas on road to Montero Hoyos, 17°39′S, 62°48′W, 290 m, 17 Dec 1991 (fl, fr), M. H. Nee 42132 (LPB, MO, NY); Prov. Andrés Ibáñez, Puerto Pailas, north side of railroad bridge and edge of floodplain of the Río Grande, 17°40′S, 62°47′W, 280 m, 17 Dec 1991 (fl), M. H. Nee 42146 (LPB, MO, NY); Prov. Andrés Ibáñez, 3 km W of Puerto Pailas, along old dirt road to Santa Cruz and railroad, 17°40′S, 62°49′W, 290 m, 11 Dec 1994 (fl), M. H. Nee 45887 (NY); Prov. Andrés Ibáñez, old dirt road highway and railroad from Cotoca to Puerto Pailas, 2.5 km W of Río Grande bridge at Puerto Pailas, 17°40′S, 62°48′30″W, 290 m, 13 Jan 1998 (fr), M. H. Nee 47938 (CORD, NY); Prov. Andrés Ibáñez, 3.5 km NE of Montero Hoyos on dirt road to San Antonio, 17°37′S, 62°49′W, 280 m, 24 Feb 1998 (fl), M. H. Nee 48430 (NY); Prov. Cordillera, between new Hwy from Santa Cruz to Abapó and the railroad at turnoff to Paliza, 45 km S of turnoff from Santa Cruz –Samaipata Hwy, 18°17′S, 63°12′W, 525 m, 27 Feb 1998 (fl), M. H. Nee 48501 (MO, NY); Prov. Cordillera, along new Hwy from Santa Cruz to Abapó, N side of settlement of Río Seco, around Hwy and bridges over Río Seco, 18°39′S, 63°15′W, 550 m, 19 Apr 1998 (fl, fr), M. H. Nee 49091 (NY); Prov. Andrés Ibáñez, along railroad tracks in Basilio, 18°06.9′S, 63°11.7′W, 480 m, 8 Jul 2004 (fl, fr), M. H. Nee 52657 (NY); Prov. Cordillera, 3 km NW of center of Boyuibe, along Hwy to Camiri, 20°24.99′S, 63°18.27′W, 840 m, 8 Feb 2006 (fl, fr), M. H. Nee & I. I. Linneo F. 53978 (NY); Prov. Florida: along road between Pampa Grande and the Mairana–Mataral Hwy, 18°05′S, 64°07.25′W, 1285 m, 31 Mar 2002 (fl, fr), M. H. Nee et al. 52086 (NY [2 sheets]); Prov. Cordillera, El Curiche, 30 Jan 1946 (fl), Peredo s. n. (LIL); Cabezas, 420 m, 19 Jan 1945 (fl, fr), Peredo 7 (LIL, NY); Prov. Cordillera, 10 km by road E of Gutiérez, Laguna Caucaya, 875 m, 24 Apr 1984 (fl), G. Schmitt & D. Schmitt 98 (LPB, MO, NY). Tarija: Prov. Gran Chaco, 15 km hacia el N de Yacuiba, 630 m, 23 Sep 1985 (fl), S. G. Beck et al. 11524 (LPB, MO, NY); 14 km E de Villa Montes, 5 May 1983 (fl), A. Krapovickas & A. Schinini 39193 (LPB); Yacuiba, 1977 (fl, fr), R. Lara s. n. (LPB); Yacuiba, pueblo El Palmar, 21°53′S, 63°37′W, 612 m, 21 Sep 2001 (fl), M. Mendoza 126 (MO); Prov. Gran Chaco, 20 km E of Villamontes, 21°18.20′S, 63°18.26′W, 355 m, 13 May 2005 (fl, fr), M. H. Nee & J. M. Mendoza 53004 (NY); Prov. Gran Chaco, Villa Montes, 21°15.94′S, 63°28.55′W, 385 m, 12 Feb 2006 (fl, fr), M. H. Nee & I. I. Linneo F. 54112 (NY).

PARAGUAY. Alto Paraguay: proposed Biosphere Reserve “Gran Chaco Americano,” Madrejon, 20°30′47″S, 59°52′48″W, 6 Feb 2002 (fr), E. M. Zardini & L. Guerrero 58102 (MO, NY). Boquerón: Nueva Asunción, Ruta Transchaco km 630, 7 May 1988 (fl), A. Charpin & L. Ramella 21372 (CORD); Filadelfia, 22°20′S, 60°05′W, 26 Nov 1982 (fl, fr), W. Hahn 783 (MO, NY, PY); Joel Estigarribia, línea 11, 17 km W del cruce a base Pratt Gill, 16 Oct 1996 (fl), F. Mereles 6468 (FCQ); Tyto Pozo Brillante, 15 Jul 1992 (fl, fr), F. Mereles & R. Degen 4529 (FCQ); Area Línea 12/S, 2 Sep 1992 (fl), F. Mereles & R. Degen 4667 (FCQ); Campo Vía, Tyto a Colonia Neuland, 22°52.05′S, 59°52.08′W, 27 May 1993 (fl, fr), F. Mereles & R. Degen 5126 (FCQ); Picada a Mistolar, 4.2 km S de Línea 10, 21 May 1994 (fl), F. Mereles & R. Degen 5644 (FCQ); Mayor Gardel, cercanías de la Estancia, canal de desvío del Pilcomayo, lado Paraguayo, 22 May 1994 (fl), F. Mereles & R. Degen 5660 (FCQ); 18 km de Línea 10, Picada Estancia “Palmar Quemado”, 25 May 1994 (fl, fr), F. Mereles & R. Degen 5694 (FCQ); Camping “Flor de Chaco” a 15 km al E de Filadelfia, 25 Aug 1985 (fl, fr), L. Molas & J. Facetti 634 (MO, PY); Ruta Trans Chaco, 8 km SE de Mcal. Estigarribia, 15 Dec 1987 (fl, fr), A. Schinini & R. A. Palacios 25774 (FCQ); Fortín Teniente E. Ochoa, 21°43′S, 60°54′W, 21 Jan 1994 (fl, fr), E. M. Zardini & L. Guerrero 38076 (AS-n.v., MO, NY); Fortín Teniente E. Ochoa, 21°43′S, 60°54′W, 21 Jan 1994 (fr), E. M. Zardini & L. Guerrero 38105 (AS-n.v., MO, NY); between La Patria and Fortín Teniente Emilio Ochoa, Route 9, km 635, 21°27′S, 61°22′W, 180 m, 25 May 1994 (fl, fr), E. M. Zardini & L. Guerrero 39281 (AS-n.v., MO, NY); Parque Nacional Teniente Agripino Enciso, 21°12′37″S, 61°39′26″W, 250 m, 14 Dec 1998 (fr), E. M. Zardini & N. Duarte 49876 (AS-n.v., MO, NY). Presidente Hayes: Chacoí, 25°12′S, 57°38′W, 5 Dec 1989 (fl, fr), A. Schinini 26737 (NY); Estancia Brusquetti, 29 Oct 1985 (fl, fr), N. Soria 1116 (FCQ); Filadelfia, 13 Jun 1986 (fl), N. Soria 1261 (FCQ); between Cruce to Loma Plata and Estancia Montiel, 2 km S of Route 9, km 413, 22°40′S, 59°45′W, 26 May 1994 (fl), E. M. Zardini & L. Guerrero 39369 (AS-n.v., MO, NY); between Cruce to Loma Plata and Estancia Montiel, 2 km S of Route 9, km 413, 22°40′S, 59°45′W, 26 May 1994 (fl), E. M. Zardini & L. Guerrero 39389 (AS-n.v., MO, NY); Laguna Capitán, 22°32′21″S, 59°40′31″W, 29 Jul 1997 (fl, fr), E. M. Zardini & L. Guerrero 47077 (AS-n.v., MO, NY); Pozo Colorado–Monte Lindo, 23°32′16″S, 58°46′14″W, 30 Jul 1997 (fl, fr), E. M. Zardini & L. Guerrero 47299 (AS-n.v., MO, NY).

Figures

FIG. 4. Distribution of Solanum aridum and S. juvenale.