Solanum aureum Dunal

Knapp, Sandra, 2013, A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae), PhytoKeys 22, pp. 1-432 : 66-71

7. Solanum aureum Dunal , Solan. Syn. 16. 1816. Figure 20

Solanum loxense Dunal, Solan. Syn. 16. 1816. Type: Ecuador. Loja: Loja, A. Humboldt & A. Bonpland s.n. (holotype: P [P00136336]).

Solanum clematideum Bitter, Repert. Spec. Nov. Regni Veg. 16: 86. 1919. Type: Ecuador. Azuay: Cuenca, Pindilic [Pindilig], 2600-3000 m, 11 Oct 1888, F. Lehmann 4948 (holotype: B [F neg. 2654], destroyed; lectotype, designated here: K [K000438685]).

Solanum aureum Dunal var. riobambense Werd., Notizbl. Bot. Gart. Berlin-Dahlem 12: 379. 1935. Type: Ecuador. Chimborazo: Ríobamba, N of Sanancajas, 3300 m, 28 Jul 1933, L. Diels 355 (holotype: B, destroyed; no duplicates traced).

Type.

Ecuador. Base of Chimborazo, June 1802, A. Humboldt & A. Bonpland 3142 (holotype: P [P00136318, Morton neg. 8151]; isotypes: B-W [B-W-4318, IDC microfiche 271-315.297:5, F neg. 2890], P [P00136319]).

Description.

Woody vines or lax shrubs to 5 m tall, often in open areas. Stems densely pubescent with stiff, golden dendritic trichomes to 1 mm long, the trichome branches numerous and very short; new growth densely golden-pubescent with congested dendritic trichomes. Bark of older stems pale brownish yellow, glabrescent. Sympodial units plurifoliate, not geminate. Leaves simple, 1.5-7(-11) cm long, 0.9-3(-6) cm wide, elliptic to narrowly elliptic, membranous to slightly chartaceous or fleshy, discolorous, the upper surfaces shiny, sparsely and evenly pubescent with erect dendritic trichomes to 0.5 mm long, these with short and congested branches, occasionally glabrous with only a few trichomes along the midvein, the lower surfaces densely pubescent on veins and lamina with erect, golden, dendritic trichomes to 1 mm long, with short, congested branches, the surface appearing golden; primary veins 7-10 pairs, paler than the lamina, golden beneath; base truncate to acute, occasionally oblique; margins entire, not markedly revolute; apex acute; petioles 0.5-2 (3.5) cm long, densely golden pubescent with dendritic trichomes like those of the stems, occasionally twining to aid in climbing. Inflorescences terminal, occasionally becoming lateral and internodal, 5-10 cm long, globose in outline, many times branched, leafy near the basal branches, with 20-40 (+) flowers, densely pubescent with golden dendritic trichomes like those of the stems; peduncle very short, the transition between stem and inflorescence not always clear; pedicels 0.5-0.6 cm long, stout or appearing so from dense pubescence, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading at anthesis, densely golden-pubescent with stiff dendritic trichomes to 1 mm long, articulated at the base leaving a small sleeve or peg 0.5-1 mm long on the axis; pedicel scars irregularly spaced 1-10 mm apart, obscured by the pubescence. Buds globose to broadly ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 2-2.5 mm long, conical, the lobes 1.5-2 mm long, deltate to narrowly deltate, the apex blunt, densely to sparsely pubescent with golden dendritic trichomes to 1 mm, the pubescence usually sparser than that of the pedicel. Corolla 1.5-1.7 cm in diameter, violet or purple to deep purple, stellate, lobed ca. 2/3 of the way to the base, the lobes 5-7 mm long, 3.5-5.5 mm wide, planar at anthesis, densely pubescent abaxially with a mixture of simple and dendritic trichomes to 0.5 mm long, these weak and tangled, denser on the tips and margins, the interpetalar sinuses glabrous, glabrous adaxially. Filament tube ca. 0.5 mm long, the free portion of the filaments ca. 1 mm long, glabrous; anthers 4-5 mm long, 1.2-1.5 mm wide, glabrous, ellipsoid, loosely connivent, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 7-8 mm long, glabrous or densely and minutely papillate in the basal 1.3; stigma capitate, minutely papillate. Fruit a globose berry, ca. 1 cm in diameter, purplish black and shiny when ripe, glabrous, the pericarp thin; fruiting pedicels 1-1.2 cm long, ca. 1 mm in diameter at the base, woody and spreading, the calyx lobes woody and persistent around the base of the berry. Seeds 20-30 per berry, ca. 3 mm long, 2.5-3 mm wide, flattened-reniform, the margins undulate, reddish brown, the testal surface minutely pitted, the cells pentagonal in outline. Chromosome number: not known.

Distribution

( Figure 21 ). In Andean Ecuador and Peru, from 2500 to 3700 m elevation, most commonly collected around 3000 m.

Ecology.

Most commonly collected in montane forests, páramos and páramo margins ("ceja de selva" or “jalca”).

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 aureum is widespread in Ecuador and has been considered to include several species here recognised as distinct: Solanum sanchez-vegae of northern Peru, Solanum dichroandrum of northern Colombia and Venezuela and Solanum aspersum of Ecuador and Colombia, with which Solanum aureum is sympatric. Solanum aureum is almost always described as a vine, but occasionally as shrubby, a habit variously common in the Dulcamaroid clade. Solanum sanchez-vegae and Solanum dichroandrum differ from Solanum aureum in their looser, less dense leaf pubescence and somewhat larger flowers. All three taxa have dark purplish black berries, but those of Solanum aureum are somewhat smaller. Solanum aureum and Solanum sanchez-vegae may overlap in distribution in northern Peru in the Huancabamba depression; Solanum dichroandrum is only known Colombia and Venezuela. Specimens now recognised as Solanum aspersum had long been considered as conspecific with Solanum aureum , but differ from it in their simple trichomes from bullate bases, elliptic buds and flowers with more lanceolate corolla lobes. These two species are sympatric in northern Ecuador. Solanum cutervanum is also sympatric with Solanum aureum through much of its range and is easily confused with it. The most obvious difference in the two taxa is habit, with Solanum aureum being a vine and Solanum cutervanum a shrub, but also the leaf and stem trichomes of Solanum cutervanum are beige, more elongate and the branches shorter than those of Solanum aureum , in which the trichomes are golden and more classically dendritic.

In Prov. Azuay, Ecuador, Solanum aureum has markedly shiny upper leaf surfaces and is superficially very like Solanum aspersum , but the trichomes from these plants are always dendritic with short branches and the flowers have deltate corolla lobes. The type of Solanum clematideum comes from among these populations.

In describing both Solanum aureum and Solanum loxense , Dunal cited specimens in the Humboldt and Bonpland herbarium ("v.s.h. H. et B."); no sheets of either are presently in P-Bonpl., but material collected by Humboldt and Bonpland is housed in the general collection and is certainly that used by Dunal in describing these taxa. Many specimens from the original Humboldt and Bonpland collection were apparently re-integrated into the general collections after use in the 19th century rather than being returned to the special collection (see Knapp and Spooner 1999; Knapp 2007b).

Specimens examined.

Colombia. Cauca: Pitayó, Prov. of Popayán, Anonymous s.n. (K); Putumayo: road from Sibundoy to Pasto, between La Maria and Páramo de San Antonio, 2900 m, 1 Jun 1946, Schultes & Villarreal 7536 (GH, K, US); Valle del Cauca: Cuchilla de Barragán, Cordillera Central, vertiente occidental, hoya del Río Bugalagrande, 3250 m, 12 Apr 1946, Cuatrecasas 20605 (F, US).

Ecuador. Azuay: Cantón Cuenca, Yanasacha, Parroquia Baños, 2925 m, 26 Dec 1976, Boeke 595 (GH, MO); Cruz Pamba region above Baños, ca. 15 km southwest of Cuenca, 2743 m, 29 Jun 1945, Camp E-3940 (MO); Campamento Molón, road in construction Sigsig-Gualaquiza, 2900 m, 11 Apr 1968, Harling et al. 8218 (MO x2); Cumbe, 24 Sep 1918, Rose et al. 22963 (GH, US); Bulán, Padrehurco-Amapala, en las riberas del río Chanín, 2776 m, 18 Oct 2000, Verdugo et al. 285 (HA); Cantón Girón, Girón, El Chorro, 2700 m, 22 Jun 2001, Verdugo et al. 353 (HA); Bolívar: Río Tataguazó, carretera Santiago-San Vicente-Alizo "San Agustin", 2500 m, 24 Feb 1987, Zak 1741 (F); Carchi: Páramo del Ángel, 3300 m, 19 Jul 1945, Acosta Solís 10571 (F); El Ángel, 1 Jan 1931, Benoist 3630 (P); La Rincónada, ranch between Ibarra and Tulcán, 3000 m, 10 Aug 1923, Hitchcock 20947 (GH, US); ca. 2 km along the road El Angel-Tulcán, 3150 m, 14 May 1973, Holm-Nielsen et al. 5207 (F, MO); Road Julio Andrade-Palestina, 3300 m, 27 Dec 1980, Holm-Nielsen et al. 29707 (BM); near El Angel on old road to Tulcán, 3060 m, 23 Feb 1984, Juncosa 2335 (MO); Cantón Espejo, Reserva Ecológica El Ángel, Libertad-Morán, El Salado, 3300 m, 30 Oct 1993, Palacios 11547 (MO); Quebrada del Río Angel, 3 km NE of San Isidro on road to El Angel, 28 Jan 1967, Sparre 14280 (S); Cañar: north of Biblian, 2900 m, 6 Apr 1974, Harling & Andersson 13239 (MO); carretera Alausi-Cañar, desvio a Oyashig-Hacda. El Carmen al N de Cañar, 3270 m, 12 Aug 1987, Zak 2400 (F, MO); carretera Alausi-Cañar, desvio a Oyashig-Hacda. El Carmen al N de Cañar, 3270 m, 12 Aug 1987, Zak 2403 (F, S); Chimborazo: Cubillín, Cordillera Oriental, 3300 m, 1 Mar 1944, Acosta-Solís 7575 A (F); Cantón Riobamba, Parque Nacional Sangay, páramo de Pinlilligue, entre Alao y La Tranca, 3300 m, 18 Aug 1990, Cerón et al. 11842 (MO); 3 km E of Alao, above Río Alao, on road to Huamboya (Morona-Santiago), 2 Dec 1988, Dorr & Barnett 6207 (MO); Cantón Riobamba, Riobamba, 3300 m, 22 Mar 1934, Schimpff 896 (G); Bayushig, carretera Riobamba-Penipe-Bayushig, 3400 m, 18 Feb 1986, Zak 1635 (F); Cantón Riobamba, Parque Nacional Sangay, carretera Chimborazo-Chambo-Pungalá-Alao, 3150 m, 9 Aug 1987, Zak 2360 (F, F, MO); Cotopaxi: Pilalo, 2400 m, 9 Jul 1968, Holm-Nielsen, & Jeppesen 1541 (S); Road Pilaló-Zumbagua, 10 km above Pilaló, 3150 m, 28 Jul 1980, Holm-Nielsen & Quintana 24654 (BM); Imbabura: Shanshipamba-La Esperanza, 2950 m, 16 Nov 1949, Acosta-Solís 14422 (F); Laguna Cuicocha, crater lake 30 W of Ibarra, steep slopes around lake and on the smaller island in the lake, 3100 m, 24 May 1973, Holm-Nielsen et al. 6321 (S); Cantón Otavalo, Otovalo, Páramo de Mojanda, 3200 m, Lehmann K-222 (K); Lake Cuicocha, island in lake, 3100 m, 29 May 1939, Penland & Summers 761 (F, GH, US); Cantón Cotacachi, Reserva Ecológica Cotacachi-Cayapas, islote Teodoro Wolff, laguna de Cuicocha, 3100 m, 30 Aug 1991, Peñafiel et al. 313 (BM, MO); Carretera Pimampiro-Chuga-Palmar Chico; entre Chuga y Palmar Chico, 2800 m, 16 Jan 1988, Zak & Jaramillo 3353 (BM, F x2, GH, MO, MO); Loja: between Loja and San Lucas, 2100 m, 6 Sep 1923, Hitchcock 21497 (GH, US); Cerro de Celica. Celica-Guachanamá, Km 14-18, 2700 m, 13 Apr 1994, Jørgensen et al. 157 (BM); El Cisne- Zaruma, unfinished road km 2.9, 2340 m, 12 Dec 1994, Jørgensen et al. 1397 (MO); Catamayo-Catacocha, km 25, turnoff at Las Chinchas towards Piñas, km 2.3, 2250 m, 13 Dec 1994, Jørgensen et al. 1463 (MO); Pichincha: Cantón Cayambe, Oyacachi, Cordillera Oriental, 3200 m, 26 Sep 1945, Acosta-Solís 11148 (F); entre Oyacachi y Comenia, Cordillera Oriental, 3000 m, 27 Sep 1945, Acosta-Solís 11204 (F); El Llalo, 3201 m, 29 Dec 1938, Balls B-5826 (BM, E, K, US); base du Pichincha, 23 Mar 1930, Benoist 2211 (P); Paluguillo, 19 Feb 1931, Benoist 3908 (P); Cantón Cayambe, Reserva Cayambe-Coca, zona de amortiguamiento, 3200 m, 2 Jan 2000, Cuamacás et al. 579 (MO); Cantón Quito, Pifo, Hacienda Los Andes, 3500 m, 24 Jan 1991, Palacios 6906 (MO); Lloa, Sodiro s.n. (P); Hac. Paluguillo, on road Pifo-Papallacta, 3200 m, 19 Jul 1967, Sparre 17727 (S); carretera Quito-Guantopugro-Yanacocha, 3400 m, 22 Mar 1987, Zak 1845 (F, MO); Cantón Mejía, Parroquia El Chaupi, faldas del Volcán El Corazón, 3300 m, 23 May 1988, Zak & Jaramillo 3698 (BM, MO); Tungurahua: entre Leito y La Cima, Cordillera Oriental, 2700 m, 15 Nov 1944, Acosta-Solís 8994 (F); Cantón Mocha, Mocha, 6 Jul 1896, André 3922 (K); road between El Triunfo and Patate, summit of road that descends to Patate, 3140 m, 26 Feb 2003, Bohs et al. 3132 (UT); Cantón Mocha, Mocha, base of Chimborazo, 2743 m, Jameson 46 (K); Cantón Ambato, Ambato, 3048 m, Jul 1939, Sandeman 37 (BM, K); Cantón Mocha, Mocha, alrededores del pueblo, 2900 m, 30 Sep 1995, Villacres 373 (F).

Peru. [perhaps Ayabaca?], 1909, Weberbauer 6411 (F); Cajamarca: San Ignacio, San José de Lourdes. Base del Cerro Picorana, 25 Aug 1999, Díaz et al. 10743 (MO); Piura: Huancabamba, carretera entre Canchaque y Huancabamba, 2800 m, 14 Jan 1988, Díaz et al. 2719 (MO); Huancabamba, Cruz Blanca-Turmalina, 2600 m, 15 Sep 1981, López M. et al. 8926 (BM, MO); Huancabamba, Huancabamba, 2591 m, Aug 1943, Sandeman 4262 (K).

Figures

Figure 20. Solanum aureum Dunal. (A drawn from Juncosa 2335, NY B drawn from Zak 1845 C-G drawn from Rose 22963 H drawn from Ceron et al. 11842). Illustration by Bobbi Angell.

Figure 21. Distribution of Solanum aureum Dunal.