Solanum hindsianum Benth., Bot. Voy. Sulphur 39. 1844.

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 : 21-26

2. Solanum hindsianum Benth., Bot. Voy. Sulphur 39. 1844. Figures 2C , 7

Type.

Mexico. Baja California: Bay of Magdalena [ Bahía Magdalena], [1841], R. Hinds s.n. (holotype: K [K000063726]) .

Description.

Erect shrub, 0.5-3 m tall. Stems armed or more often unarmed; young stems pale yellowish green, densely stellate-pubescent with sessile to short-stalked porrect-stellate trichomes with 8-12 rays (0.03-)0.1-0.2 mm long, the midpoints up to 0.07(0.1) mm long, often reduced; prickles if present 0.2-1.5 cm long, 0.2-2 mm wide at base, usually straight, reddish brown, bark of older stems smooth, brown or greyish or reddish brown, glabrescent. Sympodial units difoliate, not markedly geminate. Leaves simple, (1-)2-3 cm long, (0.5-)0.8-2.3(-6) cm wide, ovate to elliptic (unusually narrowly elliptic), usually concolorous, drying yellowish green to pale green, densely pubescent on both sides and the blade surface not visible, the pubescence usually more dense beneath, the trichomes porrect-stellate, sessile or short-stalked, translucent, the rays 8-14, 0.05-0.2(0.3) mm long, straight, slightly fused near the midpoint base, the midpoint to 0.1 mm, often absent; primary veins 4-7 pairs, raised adaxially, flat abaxially, spreading at ca. 45° to the midvein, the tertiary venation mostly obscured by the dense stellate pubescence; base rounded to truncate, usually oblique; margins entire, rarely shallowly lobed, if so the lobes 2-3 on each side, the sinus extending only to 1/8 of the distance to the midvein, up to 3 mm long, rounded; apex obtuse or rounded; petiole 0.5-1.5 cm long, 1/3-1/4 of the leaf length, densely stellate-pubescent like the young stem. Inflorescences terminal or lateral, 3-4 cm long, unbranched, with 3-4 flowers; peduncle 1-2 cm long, rachis 0.5-1 cm long; pedicels 0.4-0.5 cm long, ca. 1 mm in diameter, filiform or apically dilated, densely stellate-pubescent like the leaf blade, unarmed, even in plants with prickly stems, articulated near the base; pedicel scars spaced 1-2 mm apart, forming prominent brown stumps. Buds ellipsoid, the calyx ca. ½ of the corolla length prior to anthesis. Flowers 5-merous, heteromorphic and the plants andromonoecious, the lower ones (1-3) hermaphroditic and long-styled and the more distal staminate and short-styled. Calyx tube 6-7 mm long, conical or cup-shaped, strongly keeled at the midvein, the lobes 2-3 mm long, 2-3 mm wide at base, long-triangular from a deltate base, densely stellate-pubescent abaxially, unarmed. Corolla 2.5-4(-5) cm in diameter, apparently expanding with age (slightly smaller in short-styled flowers), violet, drying yellowish-light brown, lobed for 2/3-1/2 of its length, pentagonal with abundant interpetalar tissue, the lobes 0.8-1.2 cm long, 1-1.2 cm wide, broadly deltate with an abruptly acuminate tip, spreading or slightly reflexed, densely stellate-pubescent abaxially where exposed in bud, the interpetalar tissue glabrous abaxially. Stamens equal or very slightly unequal, with the 2 adaxial anthers shorter than the 3 abaxial anthers; filament tube 1-1.5 mm long; free portion of the filaments 2-3 mm long, glabrous; anthers 6-10 mm long, free, slightly unequal, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated, the anther surface smooth to finely papillose. Ovary globose, 2-3 mm in diameter, glabrous; style in long-styled flowers 13-18 mm long, glabrous, strongly curved, in short-styled flowers 5-6 mm long, straight or only slightly curved; stigma capitate in long-styled flowers, the surface minutely papillose. Fruit a globose berry, 1(-3) per infructescence, 1-1.5 cm in diameter when dry, the pericarp thin, smooth, glabrous, light green, sometimes with dark stripes or a marbled pattern when young, drying dark brown or brown-reddish, drying and cracking open to release the seeds when mature; fruiting pedicels 1-2 cm long, 1-2 mm in diameter at the base, 2-3 mm at the apex, woody, erect, usually unarmed; fruiting calyx not accrescent, up to 1.5 cm long, usually unarmed. Seeds ca. 10-30 per berry. 2-3 mm long, 2-3 mm wide, flattened reniform, golden to dark reddish brown, the surface minutely pitted, the margins thickened, the testal cells sinuate, more rectangular in shape near the margins. Chromosome number: n=12 ( Averett and Powell 1972).

Distribution

(Figure 8 ). Solanum hindsianum is endemic to the Sonoran Desert region of southern Arizona (Organ Pipe Cactus National Monument) and northern Mexico (States of Baja California, Baja California Sur, Sonora and Sinaloa); it occurs in matorral xerófilo, bosque tropical caducifolio and bosque espinoso (classification of Rzedowski 1978), from sea level to 400 m.

Ecology and habitat.

In the Sonoran Desert biome ( Pinkava et al. 1992), S. hindsianum is most often recorded as growing in rocky outcrops and scrubby areas, coastal scrub (matorral) in rocky areas and dunes near the shore.

Common names and uses.

Mexico. Baja California: mariola (many collections); Baja California Sur: mariola (many collections), trompillo (MEXU-1203067), malva (Ramirez 33); Sonora: mariola ( López 85). United States of America. Arizona: Hinds’ nightshade, Baja nightshade (in cultivation).

Preliminary conservation status

( IUCN 2016). LC (Least Concern). EOO 315,043 km2 (LC - Least Concern); AOO 544 km2 (VU- Vulnerable). Solanum hindsianum is widespread throughout Baja California and along the coast of the Gulf of California, and although it is habitat restricted, it appears to be common where it occurs.

Discussion.

Solanum hindsianum is similar to the widespread S. elaeagnifolium , but has larger, more pentagonal flowers with a curved style and somewhat dimorphic anthers (although not as dimorphic as those in S. houstonii ). Anderson et al. (2007) do not record the anthers of S. hindsianum as dimorphic, nor the flowers as zygomorphic. Solanum hindsianum is sister to S. houstonii (sometimes identified as S. tridynamum ) in all molecular analyses to date ( Anderson et al. 2007; Stern et al. 2011; Wahlert et al. 2014). Solanum hindsianum can be distinguished from S. houstonii by its less strongly zygomorphic flowers, its usually rounder, smaller leaves and its berries that are not completely enclosed in the accrescent calyx. In areas of sympatry in northwestern Mexico, populations of S. houstonii often have anthers that are purple or purple-tinged, while those of S. hindsianum have not been recorded as other than bright yellow, and populations of S. houstonii in these areas of sympatry often have deeply pinnatifid or lobed leaves (e.g., type of S. azureum ).

Solanum hindsianum has entered the garden plant trade in the southwestern United States (e.g., http://www.desert-tropicals.com/Plants/Solanaceae/Solanum_hindsianum.html) where it is used in xeric landscaping. As a wild species in the United States it is only known from extreme southern Arizona in Organ Pipe National Monument (see Specimens examined).

Selected specimens examined.

Mexico. Baja California: Ensenada, Cataviña, 10 km N of Cataviña, 26 Feb 1991, Breedlove & Burns 71604 (CAS, MEXU); Ensenada, San Agustin, 12 miles S, 29 Jul 1955, Chambers 720 (DS, MEXU); Ensenada, Socorro, about 12 miles NE of Socorro, Distrito del Norte, 6 Feb 1947, Constance 3116 (DS, K, MEXU); Punta Prieta, 15 miles SW of Punta Prieta, 10 Feb 1935, Epling & Robison s.n. (DS, K, NY); Mulegé, Santa Rosalia, 9 miles from Santa Rosalia on the San Ignacio road, 10 Mar 1934, Ferris 8634 (DS); Bahia de Los Angeles, S of inner road, 13 Mar 1992, Fritsch et al. 1325 (MEXU); Ensenada, afuera de la cueva de las Pinturas Rupestres de Cataviña, 6 May 2008, García-Mendoza et al. 9016 (MEXU); Sierra Calavario, in eastern bajada of Sierra, systema de Sierra Viscaino (Desierto Viscaino Region), 10 Mar 1947, Gentry 7502 (DS); Calmalli, 8 miles W of Calmalli, 8 Feb 1935, Haines & Stewart s.n. (DS, K); Mesquital, about 3 miles inland, 29 miles N of Mesquital, 27 Sep 1941, Hammerly 75 (CAS, DS); Ensenada, 7.1 miles by road east of Rancho Laguna Chapala, 13 Oct 1963, Hastings & Turner 63- 182 (DS); Turtle Cove, Magdalena Bay, type region, 10 Aug 1932, Howell 10643 (CAS, DS, NY); San Fernando, 1 Mar 1958, Huey s.n. (W); Ensenada, Socorro, 2.4 miles N of Socorro, 21 Apr 1935, Ingram & Chisaki 650 (DS, NY); Mexicali, Isla Angel de la Guarda, opposite Pond Island, 30 Jun 1921, Johnston 4201 (CAS, K); Ensenada, 0.5 miles E of Hwy 1, 20 miles S of San Quintin, 24 Dec 1973, Kipping 308 (CAS); Ensenada, Isla San Lorenzo Sur, cañon grande que da al S, lado SW de la isla, 6 May 1985, Lott & Atkinson 2469 (CAS, MEXU); 5 mi S of San Quintin on Hwy 1, 17 Apr 1992, Merello & Brunner 250 (MEXU); Manuela, ca. 6.4 miles S of Rancho de Mesquital, ca. 5.3 miles N of site of Manuela, 13 May 1983, Michener et al. 4253 (CAS); San Juan Mine, Sierra San Borja, 24 Mar 1960, Moran 8030 (DS); Arroyo de la Escopeta, head of arroyo, S slope, 3 Jun 1975, Moran 22371 (CAS, MEXU); Aguascalientes, 2 Oct 1925, Orcutt 2793 (DS); Sierra la Asamblea, SW foot of the range near the W edge of Mesa Yubay and the SSW side of Mesa Cuerno de Borrego, ca. 7 road miles NNE of the abandoned site of El Desengaño., 3 May 1993, Ross et al. 7032 (BM, CAS); Isla Mellizas, norte, Bahía de Guaymas, 26 Sep 2006, Suárez Gracida 2006- 25 (MEXU); Ensenada, San Simone, N of San Quintin, 22 Mar 1949, Thomas 111 (DS); Arroyo just W of Highway 1 near gravel quarry, ca. 6.7 miles S of Socorro Wash (Arroyo Hondo), 23 Apr 1984, Thorne et al. 58047 (MEXU); Ensenada, El Marmol, 6 Mar 1930, Wiggins 4366 (CAS, DS, K, NY); Mexicali, San Felipe, 28 miles S of San Felipe, 4 Mar 1962, Wiggins 16972 (DS); Arroyo Grande, side canyon of Arroyo Grande E of El Rosario, near end of road N of San Juan de Dios, 2 May 1993, Wisura et al. 4843 (MEXU). Baja California Sur: Los Cabos, rancho Boca del Salado carretera costera Buena Vista-Los Cabos, 31 Oct 1983, Agúndez 297 (MEXU); La Paz, Isla Espiritu Santo, Gulf of California, 15 May 1939, Berry, E.O. 19 (CAS); Comondú, Bahia San Juanico, 8 Oct 1939, Berry 34 (CAS, DS); Mulegé, northern Sierra La Giganta, S of Mulegé, 11.2 mi. W of Hwy 1 on road from Rosarito to San Isidro, 5 Apr 1991, Boyd & Ross 5956 (MEXU); Mulegé, Puerto Nuevo, Sierra de Placeros, 10 Mar 1991, Breedlove & Burns 71795 (CAS, MEXU); La Paz, a 2 km del entronque del a Colonia Tabachines, sobre la carretera de terraceria a la Colonia Oriental, por la carretera federal La Paz a Los Cabos, 23 Feb 2008, Calzada 25037 (K); Mulegé, Arroyo San José de Castro, Desierto Vizcaino, 26 Jun 1983, Cancino 20 (MEXU); La Paz, Ejido El Centenario, 16 km por ONO de La Paz por la carretera transpeninsular, 11 Jun 2001, Carrillo-Reyes & Cabrera 1974 (MEXU); Mulegé, Tres Virgenes, at base of southernmost of Tres Virgenes, 25 miles W of Santa Rosalia on W side of pass, 3 Aug 1955, Chambers 771 (DS, MEXU); along road between Hwy 1 and La Purisima, 9.5 km SW of Hwy 1, 11 Dec 1994, Daniel & Butterwick 6784 (CAS, MEXU); La Paz, carretera por Pichilingue, 1 km por carretera a Puerto Carralvo desde la entrada en el estero Enfermeria, norte de La Paz, 12 Oct 2001, Domínguez Cadena 2420 (MEXU); Mulegé, Rancho El Cuarenta, 29 km al SE de Laguna San Ignacio, 18 Oct 2009, Domínguez Cadena 3063 (MEXU); Santa Rosalia, 0 Apr 1986, Elizondo 319 (MEXU); Los Cabos, Rancho Cabo El Sol, 10 km al noroeste de Cabo San Lucas, 2 Aug 1984, Encarnación & Agúndez 32 (MEXU); Isla Espiritu Santo, 1 km al O de la bahia, en frente del Islote del Gallo, 11 Jan 1987, Flores-Franco 400 (MEXU); Isla Espiritu Santo, 12 May 1975, Gaviño de la T. s.n. (MEXU); La Paz, Isla Cerralvo, 10 May 1975, Gaviño de la T. s.n. (MEXU); Isla Monserrat, 19 May 1975, Gaviño 13 (MEXU); Los Cabos, El Arco, 14 miles NE, 18 Nov 1938, Gentry 4037 (DS,K); Desierto Vizcaino, Las Tinajas and vicinity in cerros east of Los Picachos de Santa Clara, 21 Mar 1947, Gentry 7542 (DS); Mulegé, Arroyo de Tecolote, near lava flows, 19 Nov 1947, Gentry 7863 (DS); Los Cabos, Cabo San Lucas, 6 Aug 1932, Howell 10610 (CAS, DS); La Paz, Pescadero, S of Todos Santos, 4 Apr 1930, Johansen 551 (CAS, DS); Los Cabos, San Juan del Cabo, 18 Jan 1923, Jones 24004 (CAS,DS); Sierra Cacachilas, base del Cerro del Puerto de Los Soldados, 6 Feb 2003, León de la Luz 10161 (MEXU); Sierra la Giganta, Llanos de Kakiwi, Rancho El Choyal, 19 Nov 2004, León de la Luz 10591 (MEXU); a 30 km al NW de Ciudad Constitución, 10 Feb 1975, López-Forment 292 (MEXU); La Paz, 12.2 mi NW of junction carretera Las Garzas on Hwy 1, 31 Aug 1985, Luckow et al. 2837 (MEXU); Mulegé, 81 km entre Guerrero Negro y Santa Rosalia, 28 Nov 2004, Martínez 6618 (MEXU); 5 mi N of San Ignacio on Mexican Highway 1, 30 Apr 1992, Miller et al. 7291 (MEXU); San Juanico, 27 Mar 1952, Moran 3518 (BH, BM, DS); Isla Cerralvo, south end, Gulf of California, 3 Apr 1952, Moran 3603 (DS); Isla San Francisco, 11 Apr 1952, Moran 3740 (DS); Isla San Jose, south end of Amortajada Bay, 11 Apr 1952, Moran 3779 (DS); Aguaje de San Esteban, 5 Oct 1905, Nelson & Goldman 7210 (BM); about 10 miles north of Loreto, 3 Feb 1977, Reeder & Reeder 6772 (MEXU); Vizcaino Desert, mountain pass through the Sierra El Placer, 11.45 road miles S of the main Vizcaino-Bahia Tortugas road, along the road toward Cerro El Elefante and Bahia Asuncion, 1 May 1993, Ross et al. 6995 (CAS); Isla San José, costa suroeste de la isla, a 20 m de la playa, 4 Nov 1986, Sousa-Peña 79 (MEXU, NY); Isla Santa Catalina, lado SW de la isla, en una pequeña ladera a unos 500 m del mar, 6 Nov 1986, Sousa-Peña, M. 102 (MEXU); Isla del Carmen, lado W de la isla, Puerto Balandra, a 100 m de la playa, 7 Nov 1986, Sousa Peña 133 (CAS, MEXU); Isla Cerralvo, playa SW, Los Viejitos, cerca del Faro, 8 Nov 1986, Sousa-Peña 175 (MEXU); Santa Rosalia, 31 km al SE de Bahía de Tortugas, brecha a Viscaino, 21 Apr 1987, Tenorio L. & Romero de T. 1306 (MEXU); San Antonio, 4 km al SE de San Pedro, carretera a Cabo San Lucas, 13 Apr 1987, Tenorio L. & Romero de T. 12824 (MEXU); Mulegé, El Arco, 4-5.5 miles southwestward (by road) from El Arco on road to La Banderita, 22 Oct 1959, Thomas 8285 (CAS, DS, MEXU); Comondú, Cadaje, 1.5 mi S of Cadaje, 19 Dec 1959, Wiggins & Ernst 596 (DS, MEXU); La Paz, 11 miles E of La Paz on road to Las Cruces, 27 Nov 1959, Wiggins 15648 (CAS, DS, MEXU). Sinaloa: Mazatlán, 20 Nov 1926, Jones 22514 (MEXU). Sonora: San Luis Río Colorado, Sonoyta, 11.4 miles S on Mexico 2, 26 Jan 1962, Breedlove 1455 (DS); Isla Pájaros, Bahía de Guaymas, 7 Oct 2006, Búrquez 2006- 89 (MEXU); El Himalaya, peña Blanca, faja costera entre San Carlos y Tastiota, 21 Mar 1991, Búrquez 91-316 (MEXU); Guaymas, ca. 5 mi (8 km) N of San Carlos towards Playa del Mar, 17 Mar 1992, Eggli et al. 1958 (MEXU); al E de Pinacate, 16 Apr 1981, Equihua et al. s.n. (MEXU); Cañón del Nacapule, ca. 6 km N of Bahía San Carlos, 13 Dec 1992, Felger & Búrquez 92-1031 (MEXU); Isla San Esteban, 5 Apr 1963, Felger 7071 (MEXU); Puerto Peñasco, pinacate region, ca. 4 km SSE of Tinajas de los Papagos, 26 Sep 1964, Felger 10542 (MEXU); Guaymas, Ensenada Grande, (= Bahía San Pedro), bajada at N end of bay, 13 Nov 1964, Felger et al. 11613 (MEXU); Isla Turners, (=Isla Datil), small island off shore of Isla Tiburon, NW side of island, 20 Dec 1966, Felger & Cooper Miller 15319 (CAS); Hermosillo, Isla Tiburon, Tecomate, 24 Mar 1968, Gold 365 (DS, MEXU); Pitiquito, Desemboque, 6 miles N (by road) of Desemboque, near Río San Ignacio, 29 Apr 1964, Hastings, J.R. & Turner, R.M., 64- 51 (DS); Hermosillo, Tastiota, 19 Oct 1965, Hastings & Turner 65-165 (DS); Hermosillo, Isla San Esteban, 19 Apr 1921, Johnston 3178 (CAS); Pitiquito, Bahia Tepoca, 25 Apr 1921, Johnston 3302 (CAS); Altar, Quitovac, 11.6 miles N of Quitovac, N of summit of Paso San Emeterio (just S of Sierra de Cubabi), ca. 17 miles S of Sonoyta, 13 Mar 1936, Keck 4142 (CAS, K); Puerto Peñasco, Highway 2, 11 miles (17.7 km) by road S of Sonoyta, near where ridge of Sierra Cubabi approaches the road, 13 Apr 1992, Levin 2159 (MEXU); Hermosillo, antes de llegar a Punta Sargento, al NO de Punta Chueca, 27 Mar 1987, López 85 (MEXU); Empalme, despues de La Palma, ejido Santa Maria Morelos (La Atravezada) a 13 km por la carretera 85 del km 115 de la carretera 15, 23 Jul 1995, Saucedo et al. 338 (MEXU); Bahia San Carlos, inland from bay, Guaymas, Cerro Otatal, 5 Apr 1964, Sherwin 226 (DUKE); Guaymas, 27.4 km south of Restuarant Los Arrieros, 4.4 km north of Pemex El Caballo (toll road turnoff north of Guaymas) km 143 on Mex 15, 12 Jan 1998, Van Devender et al. 98- 29 (MEXU); Sierra Pinacate, Campo Rojo (Palo Verde), E of Pinacate Peak, 21 Mar 1980, Webster 24269 (MEXU).

United States of America. Arizona: Pima County, Organ Pipe Cactus National Monument, Puerto Blanco Mountains , ca. 18 km S of Pinkley Peak and 1 km NW of Red Tanks Wall, 18 Oct 1987, Baker et al. 7568 (ASU); Pima County , Organ Pipe Cactus National Monument , Pinkley Peak , 28 Jan 1986, Lechner s.n. (ARIZ) .

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 7. Solanum hindsianum Benth. A Flowering branch B Fruiting branch showing stellate trichomes C Buds D Corolla E Anthers F Gynoecium G Fruiting branch of prickly individual with broad based stem prickles H Fruiting branch with mature berries I Mature berry with irregular dehiscence J Seed. Drawn by Bobbi Angell. A, D-F from Halse s. n. (NY 00854332) B, C from Sousa-Pena 79 (NY 00751522); G from Orcutt 1339 (NY 00751531); H, J from Daniel 1856 (NY 00751517).

Figure 8. Distribution of S. hindsianum and S. homalospermum.