PlantaeSolanalesSolanaceaeKnappSandraBarbozaGloria E.BohsLynnSärkinenTiinaA revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the CaribbeanPhytoKeys3052019123114410.3897/phytokeys.123.31738 Solanum furcatum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 750. 1814Figures 18, 19 Solanum deltoideum Colla, Herb. Pedem. 4: 273. 1835. 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 by Särkinen et al. 2018, pg. 73: TO [herb. Colla]). Solanum furcatum Dunal var. glabrum G.Don, Gen. Hist. 4: 412. 1837. Type. “In Peruvia” (no specimens cited; no original material located). Solanum furcatum Dunal var. pilosum G.Don, Gen. Hist. 4: 412. 1837. 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”. Type. “Chile ad Valparaiso, Februario; Peruvia in planitie circa Tacoram, alt. 14,000–17,000’, 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”. Type. “Chile. Prov. de San Fernando in Llano del Rio Tinguiririca, 3,000’ alt., 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. 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. Type. Based on Solanumfurcatum Dunal Solanum pterocaulum Dunal var. dichotimiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, as ‘pterocaulon’. Type. Cultivated in France at Montpellier “Solanum speciosum hort. botan” (no specimens cited, described from living plants “v.v. hort. Monsp.”; neotype, designated by Särkinen et al. 2018, pg. 73: MPU [MPU310703]). Solanum crenatodentatum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852. 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. Type. Chile. Región VI (O’Higgins): Rancagua, May-Oct 1828, C. Bertero 633 (lectotype, designated by Edmonds 1972, pg. 107 [as holotype], second step designated by Särkinen et al. 2018, pg. 73: P [P00384088]; isolectotypes: BM [BM000617677], G [G00144259], M [M-0171928], MO [MO-503700], NY [NY00743695], P [P00384089], P [P00384090], P [P00384091], P [P00384092], P [P00482266], W [acc. # 1889-0283789]). Solanum bridgesii Phil., Linnaea 33: 203. 1864. Type. Chile. Región V (Valparaíso): Panquegue, R.A. Philippi s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004549]). Solanum coxii Phil., Linnaea 33: 200. 1864. Type. Chile. Región X (Los Lagos): Todos los Santos, 1862, G. Cox 38 (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004555]; isolectotype: W [acc. # 1903-0010246]). Solanum rancaguinum Phil., Anales Univ. Chile 43: 523. 1873. Type. Chile. Región VI (O’Higgins): Rancagua, Mar 1828, C. Bertero s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004594]). Solanum caudiculatum Phil., Anales Univ. Chile 91: 12. 1895. 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 Solanumsubandinum F.Meigen (1893). Type. Chile. Región XIII (Metropolitana): Santiago, Las Condes, R.A. Philippi s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004600, F neg. 2745]). Solanum ocellatum Phil., Anales Univ. Chile 91: 14. 1895. Type. Chile. Región XIII (Metropolitana): Prope Colina, F. Philippi s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004582]; isotypes: SGO [SGO000004581], W [acc. # 1903-0010230]). Solanum nigrum L. var. crentatodentatum (Dunal) O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909. Type. Based on Solanumcrenatodentatum Dunal Solanum bridgesii Phil. var. ocellatum (Phil.) Witasek ex Reiche, Anales Univ. Chile 124: 460. 1909. Type. Based on Solanumocellatum Phil. Solanum andinum Reiche, Fl. Chile 5: 346. 1910. Type. Based on (replacement name for) Solanumsubandinum Phil. Solanum tredecimgranum Bitter, Repert. Spec. Nov. Regni Veg. 11: 6. 1912. Type. Chile. Región V (Valparaíso): Valparaíso, 17 Aug 1895, O. Buchtien s.n. (lectotype, designated by Barboza et al. 2013, pg. 246: US [US00432692, acc. # 139293]; isolectotypes: HBG [HBG511497], US [US00681745, acc. # 139294]). Solanum robinsonianum Bitter, Repert. Spec. Nov. Regni Veg. 11: 7. 1912. Type. Chile. Región V (Valparaíso): Juan Fernández Island, R.A. Philippi 742 (holotype: B, destroyed, F neg. 2743; lectotype, designated by Särkinen et al. 2018, pg. 74: W [acc. # 0001347]). Solanum masafueranum Bitter & Skottsb., Nat. Hist. Juan Fernandez & Easter Island 2: 167, pl. 14. 1922. 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 by Särkinen et al. 2018, pg. 74: 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. Type. Argentina. Río Negro: Bariloche, 19 Mar 1939, A.L. Cabrera 5024 (holotype: GH [GH00077764]; isotypes F [v0073411F, acc. # 1007493], LP [LP006791]). Type.

Peru? [more likely Chile]. “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. # 976864], G [G00359946], G-DC [G00144483], P [P00335358]).

Description.

Annual or perennial herbs to 1.0 m tall, erect to lax, subwoody at base, sprawling to ca. 2 m across. Stems terete or ridged, green to purple tinged, not markedly hollow sparsely pubescent with simple, uniseriate 1–5-celled eglandular trichomes 0.1–0.5 mm long; new growth sparsely to densely pubescent with similar simple, uniseriate 1–5-celled eglandular trichomes; older stems sparsely pubescent to glabrescent, pale yellowish brown. Sympodial units difoliate, the leaves not geminate. Leaves simple, (1.5–)4.0–8.0(–2.0) cm long, (0.6–2.2–4.6(–6.5) cm wide, ovate to rhomboidal, green above, slightly paler beneath; 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, the two sides slightly unequal, 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, lateral, internodal, forked or more rarely unbranched, with 6–14 flowers clustered at the tips (sub-umbelliform) or evenly spaced along the rhachis, sparsely pubescent with simple uniseriate trichomes like those on stem; peduncle (1.0–)1.5–2.0 cm long; pedicels 4.0–7.5 mm long, 0.2–0.3 mm in diameter at the base and 0.3–0.4 mm in diameter at the 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 2–3 mm long, conical, the lobes 0.8–1.5 mm long, 0.6–1.0 mm wide, rectangular to narrowly obovate with obtuse to shortly acute apices, 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, 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, opaque, the surface of the pericarp matte; fruiting pedicels 7–12 mm long, 0.2–0.4 mm in diameter at the base, 0.5–1.0 mm in diameter at the apex, strongly reflexed, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube 1.0–2.0 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 6–14 per berry, 0.8–1.0 mm in diameter. Chromosome number: 2n=6×=72 (Stebbins and Paddock 1949; Edmonds 1982, 1983; Chiarini et al. 2017).

Solanumfurcatum Dunal A Habit B detail of adaxial leaf surface C detail of abaxial leaf surface D Bud E dissected flower F detail of infructescence G fully mature fruit (A–GSärkinen et al. 4095). Drawing by R. Wise (previously published in “PhytoKeys 106”).

Solanumfurcatum Dunal A Flowering branch B inflorescence with flowers at full anthesis C developing fruits D mature fruits (A–DKnapp s.n. Golden Gate Park). Photos by S. Knapp.

Distribution.

(Figure 20) Solanumfurcatum is native to Chile (incl. the Juan Fernández Islands) and adjacent Andean Argentina. It is probably locally introduced and naturalised along the west coast of the United States of America, Australia and New Zealand. Only a few specimens have been seen from California and Oregon, but in those areas the species is clearly naturalised. Wiggins (1980) recorded S.furcatum from Baja California (Mexico), with no specimen citations; all specimens we have seen identified as S.furcatum from Baja California are plants of S.douglasii.

Distribution of Solanumfurcatum Dunal.

Ecology.

In western North America S.furcatum is a plant of disturbed areas in winter-wet areas along sea cliffs and bluffs between sea level and 100 m elevation.

Common names.

United States of America. Forked nightshade (USDA Plants 2017).

Uses.

None recorded in the region.

Preliminary conservation status (<xref ref-type="bibr" rid="B115">IUCN 2017</xref>).

Least Concern (LC). Solanumfurcatum is introduced into the western United States, where it is not common, but its worldwide range is very large. For EOO see Table 6.

Discussion.

Solanumfurcatum can be distinguished from the similar and sympatric S.douglasii in its usually forked inflorescences, globose buds from which the style is often exserted, ellipsoid anthers on distinct filaments, and berries with usually more than 10 stone cells. Solanumfurcatum is not sympatric with S. nigrescens but differs from it in the same set of characters and in its style that is exserted for as long as the anther cone (e.g., exserted portion of the style equal to the length of the anther cone).

Solanumfurcatum was considered to be an introduction from Chile by Stebbins and Paddock (1949) and is well-established but not common in coastal areas from Oregon to central California. Other Chilean species have similarly become established along the west coast of the United States [such as Nicotianaacuminata (Graham) Hook., see Goodspeed 1954; Knapp in press].

Details of typification for the synonyms of S.furcatum can be found in Särkinen et al. (2018).

Specimens examined.

See Suppl. materials 1 and 3.

Preliminary conservation assessments for morelloid species from the Caribbean and North and Central America. For details see Materials and Methods and individual species treatments. Preliminary assessments are based on EOO only (see Materials and Methods) and have been calculated for worldwide ranges for each species. The EOO and conservation status of species known to be solely cultivated, introduced or adventive in the region has been assessed in Särkinen et al. (2018).

SpeciesPreliminary conservation assessment (IUCN 2017)EOO (km2) [worldwide range]
Solanumamericanum Mill. LC 444,094,992
Solanumchenopodioides Lam. LC 77,207,558
Solanumcorymbosum Jacq. LC 1,621,244 (all); 148,300 (Mexico and Central America only)
Solanumdouglasii Dunal LC 6,419,607
Solanumemulans Raf. LC 5,394,300
Solanumfurcatum Dunal LC 209,035,647 (North America only 4,169 – EN)
Solanuminterius Rydb. LC 4,506,327
Solanummacrotonum Bitter LC 3,804,650
Solanumnigrescens M.Martens & Galeotti LC 15,340,166
Solanumnigrum L. LC 78,076,619
Solanumnitidibaccatum Bitter LC See Särkinen et al. 2018
Solanumpruinosum Bitter LC 294,305
Solanumpseudogracile Heiser LC 1,048,309
Solanumretroflexum Dunal LC See Särkinen et al. 2018
Solanumsarrachoides Sendtn. LC 100,440,077
Solanumscabrum Mill. LC See Särkinen et al. 2018
Solanumtriflorum Nutt. LC 91,711,478
Solanumvillosum Mill. LC See Särkinen et al. 2018

Specimens cited in pdf format (traditional format; only specimens from the region cited)

Data type: PDF file

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Sandra Knapp, Gloria E. Barboza, Lynn Bohs, Tiina Särkinen

Searchable CSV file of all specimens examined for this treatment, including Old World and South America

Data type: CSV file

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Sandra Knapp, Gloria E. Barboza, Lynn Bohs, Tiina Särkinen
SärkinenTPoczaiPBarbozaGEvan der WeerdenGMBadenMKnappS (2018) A revision of the Old World black nightshades (Morelloid clade of Solanum L., Solanaceae).PhytoKeys106: 1223. https://doi.org/10.3897/phytokeys.106.21991D’ArcyWG (1974a) [1973]) Solanaceae, Flora of Panama. Part 9.Annals of the Missouri Botanical Garden60: 573780. https://doi.org/10.2307/2395139EdmondsJM (1972) A synopsis of the taxonomy of Solanumsect.Solanum (Maurella) in South America.Kew Bulletin27(1): 95114. https://doi.org/10.2307/4117874BarbozaGEKnappSSärkinenT (2013) Grupo VII. Moreloide. In Anton AM, Zuloaga FO (Eds), Barboza GE (coord.) Flora Argentina vol. 13, Solanaceae. IBODA-IMBIV, CONICET: Buenos Aires & Córdoba, Argentina, 231–264.StebbinsGLPaddockEF (1949) The Solanumnigrum complex in Pacific North America.Madrono10: 7081.EdmondsJM (1982) Epidermal hair morphology in SolanumL.sectionSolanum. Botanical Journal of the Linnean Society 85(3): 153–167. https://doi.org/10.1111/j.1095-8339.1982.tb02583.xEdmondsJM (1983) Seed coat structure and development in SolanumL.sectionSolanum (Solanaceae).Botanical Journal of the Linnean Society87(3): 229246. https://doi.org/10.1111/j.1095-8339.1983.tb00992.xChiariniFELipariDBarbozaGEKnappS (2017) Solanaceae. In: MarholdKKucěraJ (Eds) IAPT/IOPB chromosome data 25.Taxon66: 12461247. https://doi.org/10.12705/665.29WigginsIL (1980) Flora of Baja California. Stanford University Press, Stanford.USDA Plants (2017) USDA- NCRS The PLANTS Database National Plant Data Team, Greensboro [http://plants.usda.gov, accessed 26 November 2017].IUCN (2017) Guidelines for Using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. http://www.iucnredlist.org/documents/RedListGuidelines.pdfGoodspeedTH (1954) The genus Nicotiana. Chronica Botanica 45. Chronica Botanica Company, Waltham MS.KnappS (in press) Nicotiana. In: Flora of North America Editorial Committee (Eds) Flora of North America north of Mexico. Volume 14: Magnoliophyta: Gentianaceae to Hydroleaceae. Oxford University Press, New York, in press.