Solanum furcatum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 750. 1814

Saerkinen, Tiina, Poczai, Peter, Barboza, Gloria E., Weerden, Gerard M. van der, Baden, Maria & Knapp, Sandra, 2018, A revision of the Old World Black Nightshades (Morelloid clade of Solanum L., Solanaceae), PhytoKeys 106, pp. 1-223 : 1

4. Solanum furcatum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 750. 1814 Figures 13 , 14

Solanum deltoideum Colla, Herb. Pedem. 4: 273. 1835.

Solanum furcatum Type. Cultivated in Italy at "h. Ripul:" [Hortus Ripulensis], the seeds originally sent by C. Bertero from Chile ["Chili Quillota"] (no specimens cited; lectotype, designated here: TO [herb. Colla]).

Solanum furcatum Dunal var. glabrum G.Don, Gen. Hist. 4: 412. 1837.

Solanum furcatum Type. "In Peruvia" (no specimens cited; no original material located).

Solanum furcatum Dunal var. pilosum G.Don, Gen. Hist. 4: 412. 1837.

Solanum furcatum Type. "In Peruvia" (no specimens cited; no original material located).

Solanum furcatum Dunal var. acutidentatum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843, as " acutedentatum ".

Solanum furcatum Type. "Chile ad Valparaiso, Februario; Peruvia in planitie circa Tacoram, alt. 14000-17000', Aprili" both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located).

Solanum furcatum Dunal var. obtusidentatum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843, as " obtusedentatum ".

Solanum furcatum Type. "Chile. Prov. de San Fernando in Llano del Rio Tinguiririca, 3000' alf, martio"; Peruvia ad Arequipam, Aprili" both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located).

Solanum furcatum Dunal var. subintegerrimum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843.

Solanum furcatum Type. "Chile: Copiapó, Aprili; Peruvia: circa Tacoram, Aprili" both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located).

Witheringia furcata (Dunal) J. Rémy, Fl. Chil. [Gay] 5: 67. 1849.

Solanum furcatum Type. Based on Solanum furcatum Dunal

Solanum pterocaulum Dunal var. dichotimiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, as ' Solanum pterocaulon '.

Solanum furcatum Type. Cultivated in France at Montpellier "Solanum speciosum hort. botan" (no specimens cited, described from living plants "v.v. hort. Monsp."; neotype, designated here: MPU [MPU310703]).

Solanum crenatodentatum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852.

Solanum furcatum Type. Chile. Région VI ( O’Higgins): Colchagua, San Fernando, "in selibus chilensibus San Fernando", Mar 1831, C. Gay 2 (lectotype, designated by D’Arcy 1974a, pg. 738: P [P00337274]).

Solanum rancaguense Dunal, Prodr. [A. P. de Candolle] 13(1): 150. 1852.

Solanum furcatum Type. Chile. Région VI ( O’Higgins): Rancagua, May-Oct 1828, C. Bertero 633 (lectotype, designated by Edmonds 1972, pg. 107 [as holotype], second step designated here: P [P00384088]; isolectotypes: BM [BM000617677], G [G00144259], M [M-0171928], MO [MO-503700], NY [00743695], P [P00384089], P [P00384090], P [P00384091], P [P00384092], P [P00482266], W [1889-0283789]).

Solanum bridgesii Phil., Linnaea 33: 203. 1864.

Solanum furcatum Type. Chile. Región V ( Valparaíso): Panquegue, R.A. Philippi s.n. (lectotype, designated here: SGO [SGO000004549]).

Solanum coxii Phil., Linnaea 33: 200. 1864.

Solanum furcatum Type. Chile. Región X (Los Lagos): Todos los Santos, 1862, G. Cox 38 (lectotype, designated here: SGO [SGO000004555]; isolectotype: W [W1903-0010246]).

Solanum rancaguinum Phil., Anales Univ. Chile 43: 523. 1873.

Solanum furcatum Type. Chile. Región VI ( O’Higgins): Rancagua, Mar 1828, C. Bertero s.n. (lectotype, designated here: SGO [SGO000004594]).

Solanum caudiculatum Phil., Anales Univ. Chile 91: 12. 1895.

Solanum furcatum Type. Chile. Región VIII ( Bío-Bío): prov. Ñuble, Coigüeco, F. Puga s.n. (no original material located, not at SGO).

Solanum subandinum Phil., Anales Univ. Chile 91: 13. 1895, nom. illeg., not Solanum subandinum F.Meigen (1893).

Solanum furcatum Type. Chile. Región XIII (Metropolitana): Santiago, Las Condes, R.A. Philippi s.n. (lectotype, designated here: SGO [SGO000004600, F neg. 2745]).

Solanum ocellatum Phil., Anales Univ. Chile 91: 14. 1895.

Solanum furcatum Type. Chile. Región XIII (Metropolitana): Prope Colina, F. Philippi s.n. (lectotype, designated here: SGO [SGO000004582]; isotypes: SGO [SGO000004581], W [1903-0010230]).

Solanum nigrum L. var. crentatodentatum (Dunal) O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909.

Solanum furcatum Type. Based on Solanum crenatodentatum Dunal

Solanum bridgesii Phil. var. ocellatum (Phil.) Witasek ex Reiche, Anales Univ. Chile 124: 460. 1909.

Solanum furcatum Type. Based on Solanum ocellatum Phil.

Solanum andinum Reiche, Fl. Chile 5: 346. 1910.

Solanum furcatum Type. Based on (replacement name for) Solanum subandinum Phil.

Solanum tredecimgranum Bitter, Repert. Spec. Nov. Regni Veg. 11: 6. 1912.

Solanum furcatum Type. Chile. Región V ( Valparaíso): Valparaíso, 17 Aug 1895, O. Buchtien s.n. (lectotype, designated by Barboza et al. 2013, pg. 246: US [00432692, acc. # 1399293]; isolectotypes: HBG [HBG511497], US [00681745, acc. # 1399294]).

Solanum robinsonianum Bitter, Repert. Spec. Nov. Regni Veg. 11: 7. 1912.

Solanum furcatum Type. Chile. Región V ( Valparaíso): Juan Fernández Island, R.A. Philippi 742 (holotype: B, destroyed, F neg. 2743; lectotype, designated here: W [0001347]).

Solanum masafueranum Bitter & Skottsb., Nat. Hist. Juan Fernandez & Easter Island 2: 167, pl. 14. 1922.

Solanum furcatum Type. Chile. Región V ( Valparaíso): Juan Fernández Islands, Masafuera [Isla Alejandro Selkirk], Las Chozas, 715 m, 3 Mar 1917 [20 Feb 1917 on label], C. Skottsberg & I. Skottsberg 363 (lectotype, designated here: S [acc. # 04-2947]; isolectotypes: BM [BM000617676], LD [1643307], K [K000585692], NY [00172084], GOET [GOET003548], GB [GB0048742], P [P00337092], UPS [acc. # 104031]).

Solanum spretum C.V.Morton & L.B.Sm., Revis. Argentine Sp. Solanum 132. 1976.

Solanum furcatum Type. Argentina. Río Negro: Bariloche, 19 Mar 1939, A.L. Cabrera 5024 (holotype: GH [GH00077764]; isotypes F [V0073411F, acc. # 1007493], LP [LP006791]).

Type.

Peru?. "Cette plante croît au Perou", J. Dombey [343] (lectotype, first step designated by Edmonds 1972, pg. 107 [as holotype], second step designated by Barboza et al. 2013, pg. 246: P [P00335357]; isolectotypes: CORD [CORD00006928], F [V0043232F, acc. # 976846], G [G00359946], G-DC [G00144483], P [P00335358]) .

Description.

Annual or perennial erect to sprawling herbs or small shrubs to 1.0 m tall, subwoody and branching at base. Stems spreading to decumbent, terete or ridged, green to purple tinged, older stems pale yellowish-brown, not markedly hollow; new growth sparsely pubescent with simple, spreading, uniseriate, eglandular trichomes, these 1-5-celled, 0.1-0.5 mm long; older stems sparsely pubescent to glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, (1.5-)4.0-8.0(-12.0) cm long, (0.6-)2.2-4.6(-6.5) cm wide, ovate to rhomboidal, membranous, slightly paler beneath, without smell; adaxial surface sparsely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along lamina and veins; abaxial surface more densely pubescent; major veins 4-6 pairs; base cuneate to acute, slightly oblique, decurrent on the petiole; margins sinuate-dentate or entire; apex acute; petioles 1.0-3.5 cm long, sparsely pubescent with simple uniseriate trichomes like those on stem. Inflorescences (1.0-)1.5-3.0(-4.0) cm long, internodal, furcate or more rarely unbranched, sub-umbelliform or the flowers evenly spaced along the rhachis, with 6-14 flowers, sparsely pubescent with simple uniseriate trichomes like those on stems; peduncle (1.0-)1.5-2.0 cm long, straight; pedicels 4.0-7.5 mm long, 0.2-0.3 mm in diameter at the base and 0.3-0.4 mm at apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0.2-2.5 mm apart. Buds subglobose, the corolla exserted 1/3-1/2 from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 0.5-1.5 mm long, conical, the lobes 0.8-1.5 mm long, 0.6-1.0 mm wide, rectangular to narrowly obovate, tips obtuse to shortly acute, pubescent with simple uniseriate trichomes like those on stem but shorter. Corolla 12-20 mm in diameter, white to lilac with a green or yellow-green central portion near the base, this sometimes purplish near the lobe midvein, stellate, lobed 1/3-1/2 of the way to the base, the lobes 5.5-7.0 mm long, 2.8-5.5 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially with 1-4-celled simple uniseriate trichomes, especially along the margins and apex, these shorter than the trichomes of the stems and leaves. Stamens equal; filament tube minute; free portion of the filaments 0.9-1.6(-2) mm long, adaxially pubescent with tangled uniseriate 4-6-celled simple trichomes; anthers 2.3-3.3(-3.6) mm long, 0.8-1.0 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 6.0-6.5 mm long, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower 1/2-2/3 where included in the anther cone, exserted 2-3 mm beyond the anther cone and somewhat curved; stigma capitate, minutely papillate, yellow or green in live plants. Fruit a globose berry, 6-9 mm in diameter, dull green to purple at maturity, the pericarp thin and opaque; fruiting pedicels 7-12 mm long, 0.2-0.4 mm in diameter at the base and 0.5-1.0 mm at apex, strongly reflexed, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1.5-2.5 mm long, appressed against the berry. Seeds 30-40 per berry, 1.8-2.0 mm long, 1.4-1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow-brown, the surface minutely pitted, the testal cells pentagonal in outline. Stone cells 2-13 per berry, largest 0.8-1.0 mm in diameter but varying in size. Chromosome number: 2n =6x=72 ( Stebbins and Paddock 1949; Edmonds 1982, 1983), but see 2n =2x=24 in Randell and Symon (1976; possible voucher identification issue).

Distribution

(Figure 15 ). Native to Pacific coastal South America and locally introduced and naturalised in the states of Victoria and Tasmania in Australia and in the western United States of America (California and Oregon, see Bohs in press); Although the disjunct distribution of the species appears peculiar, other taxa native to the Pacific coast of South America have become established in similar areas with Mediterranean climates (e.g. Nicotiana acuminata (Graham) Hook.).

Ecology.

Open places and disturbed areas, along roadsides and field margins; between sea level and 3,000 m elevation in its native range, in Australia collected in coastal habitats along the foreshore or along creeks near sea level.

Common names.

Australia: broad nightshade ( Walsh and Entwisle 1999).

Uses.

None recorded for Old World collections.

Preliminary conservation status

( IUCN 2016). Solanum furcatum , while only rarely encountered in the Old World, is common in its native range in Chile ( Barboza et al. 2013; Särkinen et al. 2015b) and can be assessed as LC (Least Concern; Table 7 ) on a worldwide basis. In its native range in South America, it has an EOO of 1,809,315 km2 and is not of conservation concern.

Discussion.

Solanum furcatum can be distinguished from all other Old World species based on its branched inflorescences, anthers 2.3-3.3 mm long and flowers where styles are exserted up to 3.0 mm beyond anthers. It has been sparingly introduced into areas of Mediterranean habitat (e.g. Australia and California) but appears not to spread. It is potentially confusable with S. nigrum , but is distinguished from it by the characters mentioned above and by its 2-13 stone cells per berry. Its subglobose buds are also distinct from the more ellipsoid buds of S. nigrum and S. villosum , but this character can be hard to see in herbarium specimens.

The name S. douglasii Dunal has been misapplied to some specimens of S. furcatum in Australia (e.g. Symon 1981; Walsh and Entwisle 1999). Careful assessment of the specimens cited in these publications and material labelled as S. douglasii confirms that the specimens from Victoria represent S. furcatum . Solanum douglasii is native to Mexico and the south-western United States of America and is known only from cultivation in the Old World.

Colla (1835) described S. deltoideum from material grown in Italy from seeds sent by C. Bertero from Chile as "Solanum scabrum". Of the two specimens from Colla’s herbarium in TO, only one is labelled as originally identified as "S. scabrum" and this is the sheet we select as the lectotype of S. deltoideum .

Solanum furcatum was lectotypifed by Edmonds’ (1972) citation of "Type: Peru, Dombey s.n. (P, holotype!)"; Barboza et al. (2013) selected one of the two sheets in P (P00335357) as the second step lectotype.

The infraspecific taxa of S. furcatum described by Nees van Esenbeck (1843) were all based on specimens collected by F. Meyen (fide protologue) during his 1830-1832 voyage on the Princess Louise to Chile and Peru; we have found no duplicates of these and so have chosen not to lectotypify these names until further in-depth searches have been conducted. These may belong to S. arequipense Bitter described from Arequipa which was previously considered as synonym of S. furcatum but is distinct based on morphology, ecology and molecular evidence.

Solanum rancaguense was lectotypified by Edmonds (1972) by citation of a P specimen of Bertero 633 as “holotype”, but because there are several sheets of this collection number preserved in P, this is correctable to lectotype and we select a particular sheet (P00384088) of the five there as a second step lectotype.

We have selected lectotypes in SGO for the taxa described by Philippi here considered synonyms of S. furcatum following the advice of Smith and Figueiredo (2011). None of the protologues specifies herbaria and, although Philippi worked in Chile, we have followed McNeill (2014) in lectotypifying these names.

Selected specimens examined.

Australia. Tasmania: Circular Head, B 22 between Irishtown and Edith Creek, 2 Nov 2004, Baker 981 (HO); Circular Head, Grooms Cross rd (B22), between Irishtown and Edith Creek, 2 Nov 2004, Baker 1010 (HO); Circular Head, Edith Creek, 2 Nov 2004, Baker 1019 (HO); Circular Head, Copper Creek nr Smithton, May 1947, Curtis s.n. (HO); Circular Head, Copper Creek , May 1948, Without collector s.n. (HO) ; Victoria: Greater Geelong, Edwards Point State Faunal Reserve, 19 May 1984, Albrecht 483 (MEL); Along Billy Creek in proposed extension to Morwell National Park , 15 Jul 1988, Harris 4 (K); South Gippsland , Boolarra to Mirboo North rd, 22 Nov 1987, Harris s.n. (MEL); East Gippsland , Gauging Station on the Genoa River nr the Wangarabell rd, 10 Mar 2011, Jeanes 2662 (MEL); Yarra Ranges, Tremont, 30 Jul 2006, Stajsic 4292 (HO); Yarra Ranges, Tremont, 30 Jul 2006, Stajsic 4292 (MEL)Greater Geelong, St Leonards, Bellarine Peninsula, 9 Nov 1947, Willis s.n. (MEL); Greater Geelong, St Leonards, Bellarine Peninsula, 9 Nov 1947, Willis s.n. (MEL); Mornington Peninsula, Main Creek , 24 Mar 1984, Willis s.n. (MEL); McCrae, between Dromana & Rosebud, 24 Feb 1963, Willis s.n. (MEL); McCrae, between Dromana & Rosebud, 24 Feb 1963, Willis s.n. (MEL) .

New Zealand. North Island: Manawatu-Wanganui, Vinegar Hill, Rangitikei River near Rowa , 18 Nov 1964, Healy 64/425 (AK) .

Figures

Figure 13. Solanum furcatum Dunal A Habit B Detail flower (A-B Anonymous s. n., grown from seed sent by J. Edmonds, originally from California [ADW 42421]). Drawing by M. L. Szent-Ivany, first published in Symon (1981), courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Figure 14. Solanum furcatum Dunal A Habit B Flowering branch C Infrutescence with developing fruits. D Fully mature fruit dropping off with pedicel (A-D Knapp s. n., United States of America, California, San Francisco, Golden Gate Park, in front of California Academy of Sciences, 11 June 2013). Photos by S. Knapp.

Figure 15. Distribution of Solanum furcatum Dunal in its non-native range in the Old World.