Solanum nigrum L., Sp. Pl. 1: 186. 1753

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

publication ID

http://dx.doi.org/10.3897/phytokeys.106.21991

persistent identifier

http://treatment.plazi.org/id/7F85C68D-040C-796B-C79B-E2A60284F951

treatment provided by

PhytoKeys by Pensoft

scientific name

Solanum nigrum L., Sp. Pl. 1: 186. 1753
status

 

7. Solanum nigrum L., Sp. Pl. 1: 186. 1753  Figures 22, 23

Solanum nigrum L. var. vulgare  L., Sp. Pl. 186. 1753.

Type. "Solanum 3 α”; cultivated in George Clifford’s garden in Hartekamp, The Netherlands, Anon. s.n. (lectotype, designated here: BM [BM000558026]).

Solanum nigrum L. var. judaicum  L., Sp. Pl. 186. 1753.

Type. Unknown (no specimens or illustrations cited).

Solanum vulgatum  Baumg., Fl. Lips. 120. 1790, nom. illeg. superfl.

Type. Based on Solanum nigrum  L. [ S. nigrum  L. from Syst. Veg. ed. 14: 224. 1784 cited in synonymy]

Solanum humile  Salisb., Prodr. Stirp. Chap. Allerton 134. 1796, nom. illeg. superfl.

Type. Based on Solanum nigrum  L. (cited in synonymy)

Solanum judaicum  Besser, Prim. Fl. Galiciae Austriac. 1: 183. 1809.

Type. No localities of specimens cited; probably from what is now Ukraine (no specimens cited; no original material located).

Solanum morella  Desv., Pl. d’Angers 113. 1818, nom. illegit. superfl.

Type. Based on Solanum nigrum  L. (cited in synonymy)

Solanum parviflorum  Moretti ex Badarò, Giorn. Fis. Chim. Storia Nat. Med. Arti Dec. 2, 7: 364. 1824, nom. illeg, not Solanum parviflorum  Nocca (1793)

Type. Italy. [Liguria]" in olivetis Liguriae occid[ose]", 1824, G.B. Badarò s.n. (no specimens cited; lectotype, designated by D’Arcy 1974a, pg. 735 [as type]: G-DC [G00144311]).

Solanum cestrifolium  Jacq. ex Spreng., Syst. Veg., ed. 16 [Sprengel] 1: 680. 1825.

Type. “Patria?” [origins unknown, although probably from cultivation?] (no specimens cited; no original material found).

Solanum rhinozerothis  Blume, Bijdr. Fl. Ned. Ind. 13: 695. 1826.

Type. Indonesia [no locality cited in protologue], C.L. Blume s.n. (no specimens cited; neotype, designated here: L [L2883159]).

Solanum vulgatum  (L.) Spenn., Fl. Friburg. 2: 427. 1826.

Type. Based on Solanum nigrum L. var. vulgare  L.

Solanum vulgatum (L.) Spenn. var. nigrum  (L.) Spenn., Fl. Friburg. 2: 427. 1826.

Type. Based on Solanum nigrum  L.

Solanum vulgatum (L.) Spenn. var. chlorocarpum  Spenn., Fl. Friburg. 3: 1074. 1829.

Type. Switzerland. Fribourg: Sin. loc. (no specimens cited; no original material found).

Solanum moschatum  C.Presl, Delic. Prag. 77. 1832.

Type. Italy. Sicily: Palermo ("in cultis ruderatis Panormi Siciliae", Anon. s.n. (no specimens cited; original material at PR?, PRC?, not found).

Solanum nigrum L. var. perennans  Bertol., Fl. Ital. [Bertoloni] 2: 634. 1836.

Type. Based on Solanum moschatum  C.Presl.; Solanum parviflorum  Moretti ex Badarò (no specimens cited; no original material found).

Solanum nigrum L. var. atriplicifolium  G.Mey., Chloris Han. 265. 1836.

Type. France. Pays de la Loire: "St. Germain de Calberte, basse Lozère”/” Mans [Le Mans]"[annotation "Solanum atriplicifolium" in Desportes hand], 1806, [illegible] s.n. (lectotype, designated here: G-DC [G00144334]).

Solanum nigrum L. var. atriplicifolium  Desp. ex G.Don, Gen. Hist. 4: 412. 1838, nom. illeg. (isonym), not Solanum nigrum L. var. atriplicifolium  G.Mey. (1836)

Type. Based on same original material as Solanum nigrum L. var. atriplicifolium  G.Mey.

Solanum vulgare  Hegetschw., Fl. Schweiz 219. Dec 1838-Jan 1839, nom. illeg. superfl.

Type. Based on " Solanum nigrum  Willd." (= S. nigrum  L.)

Solanum chenopodium  Raf., Autik. Bot. 107. 1840.

Type. “Europa” (no specimens cited; original material probably lost).

Solanum exaratum  Raf., Autik. Bot. 107. 1840.

Type. “Europa” (no specimens cited; original material probably lost).

Solanum bidentatum  Raf., Autik. Bot. 108. 1840.

Type. "Italia, Sicilia" (no specimens cited; original material probably lost).

Solanum tauschii  Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3: XX. 1843.

Type. Czech Republic. “Prag”, I.F. Tausch s.n. [Herb. Flor. Boehm. 1076] (no herbaria cited; no original material found, perhaps at PR?); “Luzic”, Sadel s.n. (no herbaria cited; no original material found, perhaps at PR?); "Folimanta bei Prag", Tanbler s.n. (no herbaria cited; no original material found, perhaps at PR?); Sin loc., P.M. Opiz 10/8 40 (no herbaria cited; no original material found, perhaps at PR?).

Solanum reineggeri  Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XIX. 1843.

Type. Austria. Niederösterreich; Sin. loc., Reinegger s.n. (no herbaria cited; no original material found, perhaps at PR?).

Solanum decipiens  Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XXIV. 1843.

Type. Czech Republic. “Troja”, P.M. Opiz 10/838; “Ruchelbad”, P.M. Opiz 23/10 835; “Radlic”, P.M. Opiz 4/8 40; “Szaslau”, Janoti 804 (no herbaria cited; no original material found, perhaps at PR?).

Solanum schultesii  Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XXIV. 1843.

Type. Czech Republic. " Im Baumgarten" P.M. Opiz 10/10 35; “Prag”, P.M. Opiz s.n. (no herbaria cited; no original material found, perhaps at PR?).

Solanum nigrum L. forma stenopetalum  A.Braun ex Döll, Rhein. Flor. 412. 1843.

Type. Germany. "Bei Ettlingen", A. Braun s.n. (no herbaria cited; no original material found, not found at KR); Germany. "Bei Carlsruhe in der Nähe des Linkenheimer Thores", A. Braun & J.C. Döll s.n. (no herbaria cited; no original material found, not found at KR).

Solanum nigrum L. forma chlorocarpum  A.Braun ex Döll, Rhein. Flor. 413. 1843.

Type. Germany. "Bei Gerolsau und Lichtenthal unweit Baden", A. Braun & J.C. Döll s.n. (no herbaria cited; no original material found, not found at KR); Germany. "Bei Carlsruhe, Knielingen", A. Braun s.n. (no herbaria cited; no original material found, not found at KR); Germany. "Mannheim, Dossenheim, Oppenheim", J.C. Döll s.n. (no herbaria cited; no original material found, not found at KR).

Solanum guineense (L.) Lam. var. nepalense  Dunal, Prodr. [A. P. de Candolle] 13(1): 49. 1852.

Type. Cultivated in Avignon Botanic Garden, from eastern Nepal, Herb. Requien s.n. (lectotype, designated here: AV [Herb. E. Requien]).

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

Type. Cultivated in France at Montpellier "Solanum deppei. In hortis bot. cultum" (no specimens cited, described from living plants "v.v. Hort. Monsp."; neotype, designated here: MPU [MPU310704]).

Solanum paludosum  Dunal, Prodr. [A. P. de Candolle] 13(1): 57. 1852.

Type. Myanmar "cat. itir. Burm.", 1827, N. Wallich 402 (holotype: G-DC [G00144525]).

Solanum roxburghii  Dunal, Prodr. [A. P. de Candolle] 13(1): 57. 1852.

Type. "In India orientali" (no specimens cited; lectotype, designated here: Wight, Icones plantarum Indiae Orientalis 2: t. 344. 1843) "Peninsula Ind. orientalis", R. Wight s.n. [herb. Wight 2326] (epitype, designated here: E [E00718973]).

Solanum memphiticum Mart. var. repandum  Dunal, Prodr. [A. P. de Candolle] 13(1): 47. 1852.

Type. Germany. Saxony: cultivated in Leipzig [fide JE where original set is stored] “colui”, 1824, W. Gerhard s.n. (holotype: G [G00144263]; isotypes: JE [JE00009838], W [1889-0232903, 1889-0283857]).

Solanum nigrum L. var. atriplicifolium  Desp. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, nom. illeg., Solanum nigrum L. var. atriplicifolium  G.Mey. (1836).

Type. Based on same original material (specimen) as Solanum nigrum L. var. atriplicifolium  G.Mey.

Solanum cechicum  Opiz, Lotos 4: 94. 1854.

Type. Czech Republic. "Um Prag", P.M. Opiz s.n. (no herbaria cited; no original material found, perhaps at PR?).

Solanum nigrum L. forma judaicum  Miq., Fl. Ned. Ind. 2: 637. 1856.

Type. Indonesia. Java: [West Java] Buitenzorg [Bogor], C.L. Blume s.n. (lectotype, designated here: L [L2880952]).

Solanum nigrum L. forma paludosum  (Dunal) Miq., Fl. Ned. Ind. 2: 637. 1856.

Type. Based on Solanum paludosum  Dunal

Solanum hirsutum  Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg., not Solanum hirsutum  (Vahl) Dunal (1813).

Type. Slovakia. "ex itinere Arvensi ”[Árva], P. Kitaibel s.n. (lectotype, designated here: BP [Herb. Kit. fasc. IX No. 103]).

Solanum acutifolium  Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg., not Solanum acutifolium  Ruiz & Pav. (1799).

Type. Slovakia. "ex itinere Arvensi ”[Árva], P. Kitaibel s.n. (lectotype, designated here: BP [Herb. Kit. fasc. IX No. 103]).

Solanum nigrum L. var. macrocarpum  Schur, Enum. Pl. Transsilv. 478. 1866.

Type. Romania. Sibiu: Sibiu "In ruderalis prope Cibinium Transilv." [protologue "Schur sert. n. 1994, auf unbebauten Orten bei Hermannstadt, Aug."], Oct, F. Schur s.n. (neotype, designated here: LW [LW00210121]; no duplicates found at BRNU).

Solanum nigrum L. var. chlorocarpum  (Spenn.) Schur, Enum. Pl. Transsilv. 478. 1866.

Type. Based on Solanum vulgatum Spenn. var. chlorocarpum  Spenn.

Solanum nigrum L. var. glabrum  Lowe, Man. Fl. Madeira 2: 73. 1872.

Type. Portugal. Madeira: Mr. Gordon’s kitchen garden, the Mt., 8 Dec 1932, R.T. Lowe 16 [a] (lectotype, designated by Edmonds 2012, pg. 127: BM [BM000943025]).

Solanum nigrum L. var. hebecaulon  Lowe, Man. Fl. Madeira 2: 73. 1872.

Type. Portugal. Madeira: ["Levada de Sta. Luzia, above Funchal, Feb", protologue], R.T. Lowe [?] s.n. (no specimens corresponding to this locality and date located).

Solanum morella Desv. subsp. nigrum  (L.) Rouy, Fl. France 10: 364. 1908.

Type. Based on Solanum nigrum  L.

Solanum ganchouenense  H. Lév., Repert. Spec. Nov. Regni Veg. 11: 295. 1912.

Type. China. Guizhou: Gan Chouen, Aug 1910, J. Cavalérie 3815 (holotype: E [E00284474]; isotypes: E [E00284475], K [K001080605], P [P00055234]).

Solanum chenopodiifolium  H. Lév., Repert. Spec. Nov. Regni Veg. 12: 531. 1913.

Type. China. Yunnan: Tong-Tchouan plaine, Sep 1912, E.E. Maire s.n. (holotype: E [E00284477]).

Solanum nigrum L. subvar. atriplicifolium  (Desp. ex Dunal) Schinz & Thell., Fl. Schweiz, ed. 3, 2: 295. 1914.

Type. Based on Solanum nigrum L. var. atriplicifolium  Desp. ex Dunal (= Solanum nigrum L. var. atriplicifolium  G.Mey.)

Solanum peregrinum  E.P.Bicknell, Bull. Torrey Bot. Club 42: 332. 1915.

Type. United States of America. Massachusetts: Nantucket County, Nantucket street, E.P. Bricknell 7719 (holotype: NY [00138955]; isotype: NY [00073847]).

Solanum probstianum  Polg., Mitteil. Naturfor. Gesellsch. Solothurn 12: 30. 1938.

Type. Cultivated in Hungary at Györ, from seeds sent by R. Probst from Switzerland (Solothurn: Derendingen [Kammgarnfabrik Derendingen] in 1932), 23 Aug 1933, S. Polgár s.n. [Herb. Polg. 4051] (no specimens cited; lectotype, designated here: BP [BP-272406]).

Solanum nigrum L. var. schultesii  (Opiz) Rouy, Acta Horti Gothob. 10: 201. 1935.

Type. Based on Solanum schultesii  Opiz

Solanum pseudoflavum  Pojark., Bot. Mater. Gerb. Inst. Komarova Akad. Nauk S.S.S.R. 17: 338. 1955.

Type. Kazakhstan. Between Vernoy Alma-Ata, Vernenskiy area, between town of Vtrnym and the station Karasukskaya, the village of Dimitrivka, V.S. Titov 2220 (holotype: LE).

Solanum nigrum L. forma pallidum  Wessely, Repert. Spec. Nov. Regni Veg. 63: 311. 1960, as "Solanum nigrum subsp. nigrum var. atriplicifolium forma pallidum".

Type. Germany. Rhineland-Palatinate: Neuwied, Wirtgen s.n. (holotype: W [not seen]).

Solanum nigrum L. forma luridum  Wessely, Repert. Spec. Nov. Regni Veg. 63: 311. 1960, as "Solanum nigrum subsp. schultesii forma luridum".

Type. Germany. Saxony: Dresden, Trümmerstellen am Postplatz, 22 Sep 1957, I. Wessely 0.23 (holotype: GFW).

Solanum nigrum L. subsp. schultesii  (Opiz) Wessely, Feddes Repert. Spec. Nov. Regni Veg. 63: 311. 1960.

Type. Based on Solanum schultesii  Opiz

Solanum nigrum L. var. incisum  Täckh. & Boulos, Publ. Cairo Univ. Cairo Herb. 5: 101. 1974 [ “1972”].

Type. Egypt. Faiyum, Sinnuris, L. Boulos s.n. (holotype: CAI [CAI000175]).

Type.

"Habitat in Orbis totius cultis" [sheet marked with Θ, meaning central part of Asia = Middle East], Without collector (lectotype, designated by Henderson 1974, pg. 19: LINN [LINN 248-18]).

Description.

Annual or short-lived erect to sprawling perennial herbs to 1.0 m tall, subwoody and branching at base. Stems spreading to decumbent, terete to sharply angled and ridged, green, the ridges often spinescent, older stems not appearing spinescent, not markedly hollow; new growth pubescent with simple, spreading, uniseriate, eglandular or glandular trichomes, these 1-6-celled, 0.5-0.6 mm long; older stems glabrescent, the trichome bases persisting as pseudospines. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.8-7.2(-14.5) cm long, 2.5-5.0(-9.5) cm wide, broadly ovate, membranous, green, concolorous, without smell or smell somewhat foetid; adaxial surface sparsely pubescent with spreading, simple, uniseriate trichomes like those on stem evenly scattered along veins and lamina; abaxial surface more densely pubescent along veins and sparsely along lamina with eglandular and/or glandular trichomes like those of the stems; major veins 5-7 pairs; base obtuse to truncate, somewhat attenuate; margins sinuate-dentate, especially in the lower 2/3, to occasionally entire or deeply toothed; apex acute; petioles 0.5-3.0 cm long, pubescent with simple uniseriate glandular and eglandular trichomes like those of the stems. Inflorescences 0.8-2.0 cm long, internodal, simple to occasionally furcate, the flowers spaced along the rhachis, with (3-)4-10 flowers, pubescent with spreading simple uniseriate trichomes like those on stem; peduncle 0.5-1.5 cm long, straight; pedicels 3-5 mm long, 0.2-0.3 mm in diameter at the base and 0.2-0.3 mm at the apex, spreading, articulated at the base; pedicel scars spaced 0.3-0.7 mm apart. Buds subglobose, the corolla approximately halfway exserted from the calyx before anthesis. Flowers 5-merous, all perfect. Calyx tube 0.8-1.0 mm long, conical, the lobes 0.5-0.8 mm long, 0.6-0.8 mm wide, triangular with acute or somewhat rounded apices, pubescent with spreading simple uniseriate eglandular and glandular trichomes like those of the pedicels. Corolla 10-12 mm in diameter, white with a yellow-green central portion near the base, stellate, lobed 1/2 to 2/3 of the way to the base, the lobes 4.0-5.0 mm long, 2.0-2.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate-pubescent abaxially with simple uniseriate eglandular trichomes. Stamens equal; filament tube very short to minute; free portion of the filaments 0.5-0.7 mm long, adaxially pubescent with spreading uniseriate simple trichomes; anthers 1.8-2.5 mm long, 0.8-1.0 mm wide, ellipsoid, very slightly wider at base, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5-3.5 mm long, densely pubescent with tangled 2-3-celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted 0-1 mm beyond anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 6-10 mm in diameter, purple-black or green to yellowish-green at maturity, the pericarp dull or slightly shiny; fruiting pedicels 10-12 mm long, 0.4-0.5 mm in diameter at the base and 1.0-1.1 mm at apex, generally spreading to occasionally recurved, spaced 1.0-2.0 mm apart, dropping with mature fruits, not persistent but occasionally remaining on the inflorescence; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1.0-2.0 mm long, spreading to reflexed in fruit. Seeds (15-)20-40 per berry, 1.8-2.0 mm long, 1.5-1.6 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent (North America and Europe) but usually 2(-8) per berry in other areas (Asia), ca. 0.5 mm in diameter, brown. Chromosome number: 2n=6x=72 ( Jørgensen 1928; Tokunaga 1933; Ellison 1936; Nakamura 1937; Stebbins and Paddock 1949; Baylis 1958; Heiser et al. 1965; Saarisalo-Taubert 1967; Venkateswarlu and Rao 1972 [as S. nigrum  S8, S11, S19]; Henderson 1974; Tandon 1974; Randell and Symon 1976; Edmonds 1977, 1981, 1982, 1983, 1984a; Ganapathi and Rao 1986a; Symon 1981; Bukenya 1996).

Distribution

(Figure 24). Widespread species native to Eurasia (western Europe to Japan), northern Africa and Australia, sporadically introduced in South Africa and naturalised locally in temperate North America.

Ecology.

Weed of cultivated land, open spaces in forests and roadsides; found in disturbed areas between 0-2,200 (3,500) m elevation.

Common names.

Australia: blackberry, black berry nightshade ( Symon 1981); China: paak fa tsai ts’o [Hainan], long kui ( Zhang et al. 1994); Denmark: kirtel-natskygge, sort natskygge ( Hartvig 2015), sort natskygge ( Frederiksen et al. 2006); Egypt: anabadib [Berber], aneb el dib, enab athib; Estonia: must maavits ( Kukk and Kull 2005; Finland: mustakoiso, karvamustakoiso ( Hämet-Ahti et al. 1998); France: morelle noire ( Jauzein and Nawrot 2011); Germany: schwarzer Nachtschatten; India: makoi [Hindi], kakamachi [Bengali, Sanskrit], kakahva [Sanskrit]; Iraq: dubbais, habbat seda; Italy: morella  commune, erba morella  , pomidorella, ballerina ( Pignatti 1982), solano nero; Japan: inuhozuki, inn-hodzuki, kanzashi-inuhoozuki; Libya: aneb el dib/aneb el dhib; Lithuania: juodoji kiauliauogė ( Natkevičaitė-Ivanauskienė 1976); Malaysia: beliwan; Nepal: kamai; New Zealand: black nightshade ( Webb et al. 1988); Norway: svartsøtvier ( Lid et al. 2005); Pakistan: tolangur; Philippines: amti, amtih (Ifugao language), moti; Poland: psianka czarna ( Pawłowskiego 1963); Portugal: herva moira morelle commune, pera de Santa Maria; Saint Helena: Tristan blackberry; Slovenia: lulok čierny ( Bertová and Goliašová 1993); Spain: yerba mora, hierba mora; Sweden: nattskatta, hårig nattskatta ( Mossberg et al. 2003); United Kingdom: black nightshade

Uses.

Leaves used as vegetable in India, China, southeast Asia and in Europe (in local areas); thought to be poisonous by association with the deadly nightshade, Atropa belladonna  (see Uses in introductory section). Berries sometimes used for jam ( Viljoen 2011).

Preliminary conservation status

( IUCN 2016). Solanum nigrum  is a widely distributed amphitropical species across temperate and subtropical areas in the Old World; it can be assigned a status of LC (Least Concern; Table 7).

Discussion.

Solanum nigrum  , the type species of the genus Solanum  , is a widespread weed with much morphological variation recognised at various infraspecific levels by many different authors (e.g. Linnaeus 1753; Opiz 1843). There are almost 100 names associated with the taxon and the status of some of these remains uncertain due to difficulty in finding types but we include them here based on their descriptions (but see Doubtful species). Here, we adopt an inclusive and broad concept of the species and recognise all infraspecific taxa under a single, more widespread and morphologically variable species. Two of the most commonly recognised infraspecific units include S. nigrum subsp. schultesii  (Opiz) Wessely (villous, with glandular hairs) and S. nigrum subsp. nigrum  (eglandular). Previous studies have found no support for the recognition of these or any other infraspecific taxa within S. nigrum  based on morphological ( Henderson 1974), seed protein ( Edmonds and Glidewell 1977) or molecular marker data ( Dehmer 2001; Dehmer and Hammer 2004; Olet 2004; Manoko 2007).

We do not include material identified as S. nigrum  in recent African regional floras ( Edmonds 2006a, 2006b, 2012) in our circumscription of this species. We follow Olet (2004) and Olet et al. (2011) and consider this material to represent the wild form of S. scabrum.  In Africa, we consider only material from northern Africa around the Mediterranean and a few specimens (probably introduced from Europe) from South Africa to be true S. nigrum  . Both S. nigrum  and S. scabrum  are hexaploids and morphologically similar, but can be distinguished on the following suite of morphological characteristics; Solanum nigrum  has leaves with rather indistinct petioles, inflorescences with the flowers spaced along the rhachis, acute calyx lobes that are more or less appressed to the berry in fruit, and berries that not markedly shiny and often have stone cells (particularly in Asia). Solanum scabrum  has distinctly petiolate leaves, the flowers are usually tightly congested at the tips of inflorescences or inflorescence branches, the calyx lobes are rounded, irregular and strongly reflexed in fruit and the berries are shiny with thick pericarp and lack stone cells. In addition, the fruiting pedicels of S. nigrum  are spreading to somewhat recurved, while those of S. scabrum  are erect and strongly spreading.

Throughout its range in Europe and into Eurasia as far as western China, S. nigrum  is sympatric with S. villosum  . The simplest distinguishing character is mature berry colour; S. villosum  has red, orange or yellow berries, while those of S. nigrum  are black or green. Many old collections, however, do not state berry colour on the label, so identification can be difficult. Calyx lobes are useful for distinguishing these taxa; S. nigrum  calyx lobes are usually deltate and acute, with sharp triangular sinuses, while those of S. villosum  are longer, usually rounded at the tip and the sinuses are broad and quite transparent (see description of S. villosum  ), leaving a paler “window” just below the sinus in flower buds and early flowers. The shiny, translucent berries of S. villosum  (mostly slightly ellipsoid) usually dry blackish-brown, but are distinct from the matte, more opaque berries of S. nigrum  . Neither species has stone cells in Europe, but in Asia, S. nigrum  usually has 2 (or occasionally more) stone cells in the berries. Both species retain pedicels after fruits drop, but S. nigrum  is not as extreme in this regard and often plants are found with old inflorescences with no remaining pedicels.

Solanum nigrum  has been considered native only to Europe, but our study of populations of this widespread weed across its range has shown that the largest morphological variation can be observed in Asia. Populations of S. nigrum  from Asia have more stone cells in the fruit and the plants have a more delicate look overall with longer peduncles and often fewer flowers per inflorescence. Material from Asia has been described as different taxa (e.g. S. guanchounense  , S. chenopodiifolium  ) but the variation is continuous across the range, with European populations being more invariant (except in leaf shape and indumentum). Populations in Europe represent a relatively monomorphic set of populations compared to material from China and eastern Asia. We therefore consider that the species is native to the entire Eurasian area, with limited introductions to North America. North American material we have seen appears most similar to European plants based on morphology, suggesting the introduction to North America came from European populations, but this has not been tested genetically.

Australian populations of S. nigrum  could have originated either from Europe or from Asia. In general, S. nigrum  in Australia does not seem to have stone cells in general and plants fall into two continuously variable groups: one is similar to classic European S. nigrum  and the other is more similar to plants from eastern and south-eastern Asia.

Solanum nigrum  is an autoalloploid species, now thought to have originated from a tetraploid S. villosum  and a diploid S. americanum  by spontaneous amphiploidy ( Edmonds 1979a; Ganapathi and Rao 1986b, 1986c; Poczai and Hyvönen 2011). Cytogenetic and crossing studies show high fertility and regular pairing of bivalents at meiosis in crosses of S. villosum  , S. retroflexum  and S. americanum  with S. nigrum  ( Edmonds 1979a; Ganapathi and Rao 1986b, 1986c, 1986d). Of the two tetraploid species, S. villosum  and S. nigrum  are sympatric at least across parts of their native ranges which would make it more likely that S. nigrum  is an autoalloploid derived from the autopolyploid S. villosum  and the diploid S. americanum  . Molecular analyses have now shown that the two hexaploids S. nigrum  and S. scabrum  share a parental species with the tetraploid S. villosum  ( Poczai and Hyvönen 2011). Artificially produced hexaploids from tetraploid crosses of S. villosum  and S. americanum  have been shown to be fertile and represent the glandular variant of S. nigrum  (sometimes recognised as subsp. schultesii  (Opiz) Wessely; Edmonds 1979a). It is hence becoming more clear that the tetraploid parent of S. nigrum  is S. villosum  and not S. retroflexum  . Artificial hexaploids have been produced using S. villosum  and S. americanum  by several authors and the artificial hexaploids resemble S. nigrum  morphologically ( Tandon and Rao 1964; Venkateswarlu and Rao 1969, 1972; Soria and Heiser 1959; Edmonds 1979a). Reciprocal backcrossing of the artificial hexaploids with S. nigrum  results in a high fruit set ( Venkateswarlu and Rao 1969, 1972; Edmonds 1979a), supporting the hypothesis that S. villosum  and S. americanum  are the parent species of S. nigrum  . The autoallopolyploid origin of S. nigrum  has been suggested by previous authors ( Magoon et al. 1962; Rao 1971). In the light of the current molecular evidence combined with the accumulating evidence from cytological and crossing studies, the alternative hypotheses on the origin of S. nigrum  as an autohexaploid ( Jørgensen 1928; Nakamura 1935, 1937) and an allopolyploid derived from three distinct genomes (e.g. Tandon and Rao 1964, 1966a; Henderson 1974; Edmonds 1979a) are becoming less likely.

In parts of eastern England, S. x procurrens  A.C.Leslie ( Leslie 1978), a sterile tetraploid hybrid between S. nitidibaccatum  and S. nigrum  , is locally established where the two species grown together ( Stace 2010: 531; Stace et al. 2015). The hybrid has also been recorded in New Zealand ( Webb et al. 1988). These plants are intermediate between the two species, with glandular pubescence and black berries with somewhat accrescent calyx lobes. A good description and distribution map for this local hybrid can be found in Stace et al. (2015). We have not seen convincing material from elsewhere in Europe of this hybrid, but it potentially occurs wherever S. nitidibaccatum  and S. nigrum  co-occur. As it is sterile, however, it does not spread or persist.

Linnaeus (1753) recognised six varieties of his S. nigrum  , of which four are now considered distinct species (var. patulum  = S. americanum  ; var. villosum  = S. villosum  ; var. guineense  = S. scabrum  ; var. virginicum  = S. emulans  Raf.). Henderson (1974) lectotypified S. nigrum  with the single sheet in the Linnean herbarium (LINN 248.28) corresponding to this species. In the protologue, Linnaeus (1753) cited several other elements and it is from these that we select the lectotype for var. vulgare  while, for var. judiacum  , he cites no elements. It is possible that a specimen from his herbarium was the basis for this infraspecific epithet and that the sheet in LINN is that referred to by this name.

In describing S. judaicum  , von Besser (1809) did not specifically cite Linnaeus’s varietal name, so we have assumed he was coining a new name, rather than making a combination.

The name S. cestrifolium  has a complex history; we can find no place of publication for "Solanum cestrifolium Jacq." as cited by Sprengel (1825) and Weinmann (1824:100; in synonymy with his nomen nudum S. besserianum  , see Doubtful names under S. besseri  Weinm. ex Roem. & Schult.). The name S. cestrifolium  has also been attributed to Willdenow by Gussone (1825, as a nomen nudum) and subsequently validated by Colla (1835); this epithet refers to plants now considered part of the species S. bahamense  L. ( Strickland-Constable et al. 2008). We have not lectotypified S. cestrifolium  Jacq. ex Spreng. in the hope that material amongst the cultivated holdings at W will reveal potential original material.

Blume (1826) cited no specimens for any of the names in his Bijdragen tot de flora van Nederlandsch Indië nor do many of the descriptions have specific localities. We have found a specimen labelled S. rhinozerothis  in Blume’s hand in L (L.2883159) that we here select as the neotype for that species, but have not found any material directly attributable to Blume that we could associate with S. uliginosum  (see under Doubtful names).

The varietal epithet " atriplicifolium  " was used by many authors to refer to plants of S. nigrum  with dentate leaf margins. It appears to have been used in reference to a specimen that was annotated "Solanum atriplicifolium" by Narcisse Desportes, probably seen in Paris or Geneva. Georg Meyer of Hannover (1818) was the first botanist to effectively and validly publish this epithet and did so at the varietal rank. Dunal (1852) cited the same material and said that the collector was “Prost”, though the G herbarium catalogue states the collector as Desportes. We select this G-DC (G00144334) sheet as the lectotype for S. nigrum var. atriplicifolium  because it appears to be original material by way of locality and annotation. The combinations made by Don (1837) and Dunal (1852) are isonyms and therefore have no standing, but we include them here because they have been widely used and cited and other combinations have been made based upon them.

The various names coined by the amateur Czech botanist Philipp Maximilian Opiz and here recognised as synonyms of S. nigrum  were all included by him in the “superspecies” S. nigrum  and, from descriptions and key, represent leaf shape, pubescence and inflorescence size variations of that species. Opiz’s enormous herbarium is housed mostly in PR ( Kirschner et al. 2007) with duplicates in PRC and other major European herbaria. We have been unable to visit Prague to examine the no doubt extensive gatherings of S. nigrum  and have found few unambiguous duplicates of the collections cited in the protologues. We therefore leave the typification of these infraspecific names until further study of Opiz’s herbarium can be undertaken. The same is true for the infraspecific names coined by Johann Döll (1843), whose protologues cited some presumed collections, but no herbaria. Searches at KR reveal no original material and the A. Braun collections cited were probably at B and were destroyed. Again, these infraspecies refer to the highly variable populations of S. nigrum  in central Europe. Döll (1843) attributed the infraspecific epithets var. chlorocarpum  and stenopetalum  to Braun; he was thus coining a new name “chlorocarpum” and not citing Spenner’s earlier use of this epithet.

Solanum chenopodium  , S. exarmatum  and S. bidentatum  were all coined by Rafinesque (1840) for European plants. He cited "S. nigrum var. undatum", a name never published, in the protologue of S. chenopodium  and suggested that S. bidentatum  was the same as " S. patulum  of India". We here place these three in the synonymy of S. nigrum  based on their rather meagre descriptions; Rafinesque’s herbarium was destroyed, but occasionally fragments he sent to European herbaria survive (e.g. material relating to S. emulans  Raf.)

Dunal (1852) described several varieties of his complex taxon S. pterocaulum  (in 1852 written as pterocaulon  ); most of these correspond to S. scabrum  . His var. deppei  was described from living material "v.v. Hort. Monsp."; a sheet in MPU (MPU310704) labelled "Solanum deppei. In hortis bot. cultum" is here selected as neotype for this name.

A single collection of a plant from Nepal cultivated in Avignon was cited in the protologue of S. guineense var. nepalense  ( Dunal 1852), housed in "h. Requien". A sheet at AV clearly labelled as "S. guineense β nepalense Prodr." is most likely original material for the name and we designate this as the lectotype.

Solanum roxburghii  was coined by Dunal (1852) as a replacement for "S. rubrum" as used by Roxburgh (1824) in his Flora Indica. Dunal cited the publications of Roxburgh (1824, but with the incorrect page number of 216 rather than 246) and Nees van Esenbeck (1837) and an illustration by Wight, but cited no herbarium material. We consider the illustration in Wight (1843) as the only unambiguous original material for this name and designate it as the lectotype for S. roxburghii  . Both the citations of "Solanum rubrum" are based on the epithet of Miller and/or Linnaeus and are not at all clear. Differentiating S. nigrum  from S. villosum  (both which occur in India) can be difficult without ripe berries and, since berry colour in the original hand-coloured illustration at E is green, we are certain the Wight illustration depicts a plant of S. nigrum  , with flowers spaced along the inflorescence and sepals that are not reflexed in fruit. We therefore designate an epitype from amongst the sheets collected by Wight (E00718973) to fix the application of this name. Early authors, such as Roxburgh and Nees van Esenbeck, included both red/orange and black-fruited plants in their circumscriptions of both S. nigrum  and S. rubrum  . The sheets in the East India Company herbarium at K (see de Candolle and Radcliffe-Smith 1981; Noltie 2005) are a complex and confusing mixture of S. americanum  and S. villosum  ; none of these specimens matches the illustration as well as does the E sheet we select here as the epitype. This again illustrates the difficulty that early authors had with these very similar plants and the dangers of assuming that collections are duplicates.

Miquel (1857) recognised several forms of S. nigrum  in his broad species circumscription. His forma judiacum  was specifically coined to encompass S. judaicum  Besser in the sense of Blume (1826), but excluding Besser’s material. We therefore consider this the coining of a new name and not a new combination based on Besser’s S. judaicum  ; the lectotype selected is a sheet in L (L2880952) with the locality "in uliginosis circa Buitenzorg" as cited by Blume and in Blume’s hand.

The names coined by the Hungarian botanist Pál Kitaibel were published posthumously by Kanitz (1863) and are largely illegitimate (some were used in earlier publications by Schultes, see discussion under S. villosum  ). Solanum hirsutum  and S. acutifolium  were published as alternative names "Solanum hirsutum vel acutifolium" and therefore have the same type; we have selected the sheet in the Kitaibel herbarium at BP with the locality matching that in the protologue ("ex itinere Arvensi", BP [Herb. Kit. fasc. IX: 103]).

We have selected a specimen collected by Schur (LW00210121), but with a different date of collection and exact locality, as the neotype for S. nigrum var. macrocarpum  ; searches in the other relevant herbaria revealed no original material.

Edmonds (2012) only lectotypified one of the varieties of S. nigrum  from Lowe’s Manual Flora of Madeira (1872). Lowe’s var. hebecaulon  cited a single locality and date "Levada de Sta. Luzia, above Funchal, Feb"; no specimens corresponding exactly to that are found in BM or at K.

Solanum probstianum  was described from material cultivated in Hungary from seeds sent by Rudolf Probst to Sandor Polgár in 1932. Although no specific specimens were cited in Polgár’s protologue (in Probst 1938), there are many specimens annotated "Solanum probstianum" in Polgar’s hand at BP that are clearly cultivated and appear to be from this original seed collection. We have selected one of these with both flowers and fruits as the lectotype of S. probstianum  [BP-272406], it bears a collecting number (4051) and was collected in July 1933. We do not consider the other sheets with this number as necessarily duplicates; the sheets do not have consistent collection numbers and may represent individual plants from the cultivated population collected on different dates in the same or different years.

Selected specimens examined.

A total of 1,537 specimens were examined from 82 countries during the study across Africa, Asia, Australia, Eurasia and the Pacific. Adventive specimens from the New World were also studied in order to understand the full range of morphology within the species. All specimens examined can be seen in Appendix 2 (csv format) and Appendix 3 (traditional Specimens Examined list in pdf format).