Solanum crispum Ruiz & Pav., 1799

Knapp, Sandra, 2013, A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae), PhytoKeys 22, pp. 1-432 : 82-86

11. Solanum crispum Ruiz & Pav. , Fl. Peruv. 2: 31, t. 158a. 1799 Figure 28

Solanum ligustrinum Lodd., Bot. Cab. 1963. 1833. Type: Chile. Sin. loc., seeds sent by H. Cuming in 1831 (lectotype, designated here: Loddiges, Bot. Cab. No. 1963. 1833).

Solanum concavum Lindl., Edwards’s Bot. Reg. 28 (Misc.): 57. 1842. Type: Chile. sin. loc., H. Cuming 263 (lectotype, designated here: CGE; isolectotypes: BM [BM000935822], E [E00057570], GH [GH00077603], K [K000585719]).

Solanum laetum Kunze, Linnaea 16: 352. 1842. Type: Germany. Cultivated in Leipzig, 1837, Anon. s.n. (no specimens cited or found; synonymy ex descr.).

Solanum syringaefolium Kunth & C.D. Bouché, Ind. Sem. Hort. Berol. 10. 1845. Type: Chile. sin. loc., T.C. Bridges s.n. (holotype: B, destroyed [F neg. 2746]).

Witheringia berteroana J. Rémy, in Gay, Fl. Chil. 5: 65. 1849, as " berterianum " Type: Chile. Tagua-Tagua, C.G.L. Bertero s.n. (lectotype, designated by Knapp 1989, pg. 74: P [P00324728]).

Witheringia crispa (Ruiz & Pav.) J. Rémy, in Gay, Fl. Chil. 5: 63. 1849. Type: Based on Solanum crispum Ruiz & Pav.

Witheringia gayana J. Rémy, in Gay, Fl. Chil. 5: 67. 1849. Type: Chile. sin. loc., C. Gay s.n. (lectotype, designated by Knapp 1989, pg. 74: P [P00335390]; isolectotype: K [K000585718]).

Witheringia tomatillo J. Rémy, in Gay, Fl. Chil. 5: 64. 1849. Type: Chile. sin. loc., C. Gay s.n. (lectotype, designated by Knapp 1989, pg. 74: P [P00325721]; isolectotype: K [K000585722]).

Solanum angustifolium Lam. var. brevifolium Dunal, Prodr. [A.P. de Candolle] 13(1): 90. 1852. Type: Chile. Sin. loc., 1828, E. Poeppig 74 [diar. n. 34] (holotype: G-DC [G00144942, Morton neg. 8402, IDC microfiche 800-61.2068:II.7]; isotypes: BM [BM000935826], LE, P [P00319643, Morton neg. 8145]).

Solanum congestiflorum Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852. Type: Chile. Aviluco, C.G.L. Bertero 634 (lectotype, designated by Knapp 1989, pg. 74: P [P00325722]; isolectotype: MO [MO-3461608]).

Solanum congestiflorum Dunal var. longifolium Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852. Type: Chile. Región × (Los Lagos): Valdivia: "in Chile australis provincia Valdivia", C. Gay herb. 3e envoi 211 (lectotype, designated by Knapp 1989, pg. 74: P [P00325724]).

Solanum crispum Ruiz & Pav. var. elaeagnifolium Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852. Type: Chile. Región VI (Liberador): Prov. O’Higgins: circa Rancagua, 1833, C.G.L. Bertero 640 (lectotype, designated by Knapp 1989, pg. 74: G-DC [G00144948, Morton neg. 8406]; isolectotype: P [P00325727, Morton neg. 8175]).

Solanum crispum Ruiz & Pav. var. ligu strinum (Lodd.) Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852. Type: Based on Solanum ligustrinum Lodd.

Solanum pyrrhocarpum Phil., Anales Univ. Chile 21(2): 383. 1862. Type: Chile. Región VIII ( Bío-Bío): Prov. Ñuble: Chillan, sin. coll. (lectotype, designated by Knapp 1989, pg. 74: SGO [SGO-55460, barcode SGO000004551]).

Solanum sadae Phil., Linnaea 33: 203. 1864 -1865. Type: Chile. Región VI ( O’Higgins): Colchagua, sin. coll. (lectotype, designated by Knapp 1989, pg. 74: SGO [SGO-55450, barcode SGO000004595]; isotype: E).

Solanum landbeckii Phil., Linnaea 33: 204. 1864 -1865. Type: Chile. Región VI ( O’Higgins): Colchagua, C.L. Landbeck s.n. (lectotype, designated by Knapp 1989, pg. 74: SGO).

Solanum izquierdii Phil., Anales Univ. Chile 43: 522. 1873. as " izquierdi " Type: Chile. Región Metropolitana: Prov. Santiago, Aculeo (ca. 33°50'S, 70°55'W), V. Izquierdo s.n. (lectotype, designated by Knapp 1989, pg. 74: SGO [SGO-55455, barcode SGO000004573]; isolectotype: K [K000585727]).

Solanum gayanum (J. Rémy) F.Phil., Cat. Pl. Vasc. Chil. 228. 1881. Type: Based on Witheringia gayana J. Rémy

Solanum tomatillo (J. Rémy) F.Phil., Cat. Pl. Vasc. Chil. 229. 1881. Type: Based on Witheringia tomatillo J. Rémy

Solanum pugae Phil., Anales Univ. Chile 91: 7. 1895. Type: Chile. Región VIII ( Bío-Bío): Prov. Ñuble, Cerro Centinela, F. Puga s.n. (holotype: SGO, specimens not located).

Solanum pannosum Phil., Anales Univ. Chile 91: 9. 1895. Type: Chile. Región VII (Maule): Prov. Curicó, Los Maquis, ca. 35°43'S, 70°47'W, M. Vidal s.n. (holotype: SGO, specimens not located; isotype: W [W-1908-10271]).

Solanum berteroanum (J. Rémy) Phil., Anales Univ. Chile 91: 8. 1896. Type: Based on Witheringia berteroana J. Rémy

Solanum tagua Kuntze, Revis. Gen. Pl. 3(2): 226. 1898. Type: Based on Witheringia berteroana J. Rémy

Solanum congestiflorum Dunal var. syringaefolium (Kunth & Bouché) Reiche, Anales Univ. Chile 123: 721. 1908. Type: Based on Solanum syringaefolium Kunth & C.D. Bouché

Solanum congestiflorum var. pannosum (Phil.) Reiche, Anales Univ. Chile 123: 722. 1908. Type: Based on Solanum pannosum Phil.

Type.

Chile. Región VIII ( Bío-Bío): Concepción: "in Chile ruderatis copiosé in Conceptionis urbis sepibus, et ad Carcamo et Palomares tractus", H. Ruiz & J. Pavón s.n. (lectotype, designated by Knapp 2008c, pg. 312: MA [MA-747012]; isolectotype: MA [MA-747101]).

Description.

Shrubs or small trees, often lax and scrambling, 0.4-5 m tall. Stems glabrous or pubescent with tiny dendritic trichomes; leaf scars somewhat prominent; new growth glabrous to densely pubescent with fine, dendritic trichomes. Bark of older stems pale brownish-yellow, glabrous, and shiny. Sympodial units plurifoliate. Leaves simple, 2.7-7.5 (10) cm long, 1-3 (7) cm wide, ovate to narrowly ovate, occasionally somewhat elliptic, larger and broader in plants growing in shade and in juvenile plants (see discussion), the adaxial surfaces glabrous or with a few dendritic trichomes along the main veins, the abaxial surfaces glabrous or puberulent with dendritic trichomes, these usually denser along the veins; primary veins 6-12 pairs, pubescent below; base truncate or somewhat cordate, not winged on to the petiole; margins entire, undulate or crispate; apex acute to acuminate; petiole 0.5-1 (2.2) cm long. Inflorescences terminal, later appearing lateral from overtopping of shoots, 2-10 cm long, flat-topped or pyramidal, branching 5-7 times, with 10-20 flowers, glabrous or sparsely pubescent with dendritic trichomes like those of the stems and leaves; peduncle 1-5 cm long; pedicels 1-1.3 cm long, tapering from a basal diameter of ca. 0.5 mm to an apical diameter of ca. 1 mm, nodding at anthesis, glabrous or with a few scattered dendritic trichomes, articulated at the base and inserted in a sleeve ca. 0.5 mm long; pedicel scars closely spaced in clusters. Buds globose when young, later elliptic, the corolla strongly exserted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube 1-1.5 (2) cm long, conical, the lobes 0.5-1 mm long, deltate to long-triangular, glabrous or with a few scattered dendritic trichomes abaxially, adaxially glabrous. Corolla 1.2-2.5 cm in diameter, violet or occasionally white, lobed 3/4 to 7/8 of the way to the base, the lobes 5-9 mm long, 3-5 mm wide, planar or somewhat reflexed at anthesis, densely pubescent with simple (in glabrous plants) or dendritic (in pubescent plants) trichomes abaxially, the trichomes denser at the tips of the lobes, glabrous adaxially. Filament tube less than 0.5 mm long; free portion of the filaments 1-1.5 mm long, glabrous; anthers 3.5-5 mm long, 1-2 mm wide, loosely connivent, poridical at the tips, the pores becoming slit-like with age. Ovary glabrous or with a few dendritic and simple trichomes at the apex, especially in otherwise pubescent plants; style 0.6-1 cm long, pubescent with dendritic or simple trichomes along the entire length; stigma clavate or capitate, the surface minutely papillose. Fruit a globose or ellipsoid berry, 0.8-1 cm in diameter, bright red when ripe, changing from green to yellow or orange during ripening, with thin pericarp; fruiting pedicels 1.2-1.6 cm long, ca. 1 mm in diameter at the base, woody, deflexed. Seeds ca. 11 per berry, 2-3 mm long, 1.5-2 mm wide, flattened lenticular, reddish-brown, the surfaces minutely pitted. Chromosome number: n = 12 (vouchers: Knapp 8632, Knapp 8633 cultivated material).

Distribution

( Figure 29 ). Chile from Quillota south to the island of Chiloé, from 10-2500 m elevation. Solanum crispum is also known from scattered collections in Argentina along the border with Chile; in the Neuquén area these plants are usually found in association with villages and it is suspected that rather than being native, they have been brought from Chile and cultivated for their medicinal properties (C. Ezcurra, pers. comm.).

Ecology.

Solanum crispum grows in Nothofagus ( Nothofagaceae ) forest, often in second growth, and in a wide variety of moist microsites in otherwise dry habitats.

Common names.

Chile: tomatillo, natre, natrien, natri, tomatilla (see Knapp 1989).

Conservation status.

Least Concern (LC); EOO>50,000 km2 (LC) and AOO>5,000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum crispum is one of the most variable and is the most southerly species of the Solanum nitidum species group, occurring to 43 degrees S latitude. It also has a huge elevational range, occurring from sea level to nearly 3000 m in a wide variety of habitats. Two pubescence forms occur throughout the range of Solanum crispum : glabrous plants were traditionally called Solanum crispum and pubescent ones Solanum congestiflorum (see Knapp 1989). Specimens of intermediate pubescence are rare, but the new growth of glabrous plants is always dendritic-pubescent. In a cladistic analysis ( Knapp 1989) the two forms were treated as separate, but were strongly resolved as sister taxa. Pubescence may be related to habitat, but this effect has not been studied in any detail; polymorphism in pubescence is extremely common in members of the Dulcamaroid clade and elsewhere in Solanum . The pubescent form often has larger, more repand leaves than does the glabrous form. This raises the intriguing possibility that the pubescent form is paedomorphic, retaining the shape and indument of juvenile leaves.

Medicinal uses of Solanum crispum have been recorded for over two centuries, beginning with Ruiz and Pavón in Flora Peruviana (1797), where the plant was reported to be used as a febrifuge. It has been reported as used against the fevers called ‘congo’ and ‘chavalongo’.

Solanum crispum has been cultivated in the United Kingdom since the early part of the 19th century. Specimens were introduced to Kew Gardens from the island of Chiloé (Chile) by Mr. Anderson (Hooker 1844) and a variety still in cultivation today was developed at the Glasnevin Botanic Gardens in Dublin. Solanum crispum is usually classed as a climber, but this is due to its lanky habit in the British climate. It does not possess the twining stems, petioles or tendrils of a true climber, but it is not a robust, erect plant.

Many monographers in Solanum have stated that holotypes or lectotypes for Ruiz and Pavón names were in the Madrid herbarium (MA), but without specifying a particular sheet. In a few cases, only one sheet exists, thus making lectotypification relatively straightforward, but in others multiple sheets exist in the Ruiz and Pavón herbarium at MA, meaning that previous type designations are not sufficiently precise. It is unlikely that any of these specimens are actually holotypes; the dispersal of specimens at the time of the expedition and subsequently through sale and loss means lectotypification is essential even if only a single sheet is present at MA. Knapp (1989) lectotypified Solanum crispum citing only a sheet in MA; this was rectified in 2008 by citation of the particular sheet ( Knapp 2008c) as the lectotype.

Knapp (1989) erroneously lectotypified Solanum ligustrinum using one of the specimens cited by Dunal (1852) when making the new combination rather than the plate from Loddiges’ (1833) original description (in which seeds from an un-numbered collection sent by H. Cuming are cited). The epithet ligustrinum is correctly lectotypified here using Loddiges’ plate ( Figure 30 ), as it is the only extant element unambiguously related to the protologue.

Specimens examined.

Argentina. Mendoza: Santa Rosa de los Andes to Uspallato Pass, Moseley s.n. (BM, LE); Neuquén: Minas, a 18 km de las Ovejas camino a las lagunas Epu-Lauquén, mallin de la Culebra, 1450 m, 14 Jan 1964, Boelcke et al. 10786 (SI).

Chile. Región IV (Coquimbo): Elqui, sector bordering road to Pachon, 38 km SE of Guard Control Post at Cerro Tololo, 2416 m, 16 Jan 2004, Acosta-Solís & León BB 181 (K); Illapel, Dec 1862, Landbeck s.n. (W); Elqui, Baño de Pangue, 1800 m, 20 Sep 1947, Sparre 2628 (S); Ovalle, Ovalle, ca. 86 km from Ovalle on road Caren to Río Mostazal, 1800 m, 11 Nov 1938, Worth Morrison 16448 (G); Región Metropolitana: Rio Colorado, Paso Uspallata, 6 Jan 1886, Philippi & Borchers s.n. (BM); Santiago, Oct 1933, Grandjot s.n. (GOET); Melipilla, Las Vizcachas, ca. 10 km from La Dormida, 1920 m, 7 Dec 1938, Morrison 16760 (G); Santiago, 1876, Philippi s.n. (G); Santiago, 1862, Philippi s.n. (G); Farallones, 2500 m, 15 Apr 1969, Plowman 2683 (LE);); Santiago, Cerro San Cristobal, Nov 1869, Reed 1869 (BM); Santiago, Prov. Santiago, Cord de Santiago, Rio San Francisco, 2200 m, Dec 1924, Werdermann 439 (BM); Río San Francisco, Cordillera de Santiago, 2200 m, Dec 1927, Werdermann 479 (G); Región V ( Valparaíso): Santa Rosa de los Andes, May 1882, Ball s.n. (LE); Los Señales, Dec 1829, Bertero 1323 (G-DC); Quillota, Aug 1829, Bertero 1327 (G-DC); Llaillai, “Atacama”, 9 Oct 1884, Borchers s.n. (F); Región VI ( O’Higgins): prov. Colchagua, Cerro Echaurrina, San Fernando, 13 Oct 1926, Montero 33 A (F); Colchagua, 1862, Philippi s.n. (G); Región VII (Maule): Talca, Cordillera de Los Andes: 18 km on road to Laguna del Maule from first border post, 4 Feb 1998, Baxter et al. 26 (BM); Cordillera de Curicó, Valle del Toro, 1500 m, 1903, Bürger 48 (GOET); Región VIII ( Bío-Bío): Baños de Chillán, Jan 1878, Anonymous s.n. (W); Concepción, 26 Sep 1939, Bailey 941 (BH); Concepción, Quebrada Honda, entre Liriquén y Tomé, 50 m, 16 Oct 1986, Basualto et al. 37 (MA); Ñuble, Termas de Chillán, Refugio El Aserradero, 1240 m, 13 Nov 1986, Basualto et al. 156 (MA); Talcahuano, 12 Nov 1950, Brooke 6951 (F); Biobío, Antuco, Cordillera de los Andes, Fundo Los Ciervos, passing into El Toro hydroelectric central and crossing the Río Polcura, 860 m, 28 Jan 2004, Brownless et al. DCI-1045 (BM); Ñuble, Chillán, road to Termas Chillán at Puente Torrealba, 1514 m, 25 Dec 2003, Gardner & Knees 6776 (BM); Ñuble, Chillán, 1856, Germains.n. (G); Prov. Ñuble, Chillán, 1856, Germain s.n. (F); Concepción, 1825, Macrae s.n. (G); Coronel, 1864, Oschenius s.n. (GOET); Chillán, Philippi s.n. (K); Ñuble, Termas de Chillán, 1450 m, 11 Dec 1987, Rechinger & Rechinger 64317 (B); Ñuble, Río Ñuble, 40-50 km desde San Fabian hacia la Cordillera, entre rio Nuble y rio Los Sauces, 26 Feb 1968, Zalensky III 81-82 (LE); Región IX ( Araucanía): Malleco, Victoria, Hotel El Bosque on Ruta 5 south-bound from Victoria, 338 m, 24 Jan 2004, Brownless et al. DCI-903 (BM); Cautín, Villarrica, road from Meseta San Judas to western edge of Lago Colico, 500 m, 20 Dec 2003, Gardner & Knees 6734 (BM); Malleco, road from Victoria to Termas de Tolhuaca before Parque Nacional Tolhuaca near to Puente Tacadero, 942 m, 30 Dec 2003, Gardner & Knees 6870 (BM); Cautín, Río Zuerpe, 28 Sep 1905, Middleton s.n. (G x2); Región X (Los Lagos): Island of Chilóe, Anonymous 7966 (BM); Island of Chilóe, Anonymous 7967 (BM); Archipelago de Chiloe, Downton 6 (BM); Región XIV (Los Ríos): Valdivia, 20 Oct 1904, Buchtien s.n. (G); Valdivia, 22 Oct 1905, Buchtien s.n. (G); Valdivia, 28 Sep 1896, Buchtien s.n. (G); Chiloé, Chiloé, Caldeleugh s.n. (G); Valdivia, Valdivia, coastal road from Curiñanco to Niebla, 19 Dec 2003, Gardner & Knees 6723 (BM); Valdivia, Panguipulli, 150 m, Oct 1924, Hollermayer 324 (BM); Cordillera de Ranco, Lechler 827 (GOET); Valdivia, road from La Union to El Mirador, 700 m, 30 Mar 1969, Plowman 2640 (GH); Valdivia, Corral, Amargos, San Carlos, 20 m, 6 Jan 1954, Sparre & Smith 399 (G); Valdivia, Panguipulli, 150 m, Oct 1924, Werdermann 324 (G);

Figures

Figure 28. Solanum crispum Ruiz & Pav. (A drawn from Taylor 10235 B drawn from Nee 54654 C drawn from Landrum 8216 D drawn from Biese 78 E-H drawn from Landrum 4459). Illustration by Bobbi Angell.

Figure 29. Distribution of Solanum crispum Ruiz & Pav.

Figure 30. Lectotype of Solanum ligustrum Lodd. (Loddiges 1833: tab. 1963). Reproduced with permission of the Natural History Museum Botany Library.