Solanum villosum Mill., Gard. Dict. ed. 8, no. 2. 1768

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

19. Solanum villosum Mill., Gard. Dict. ed. 8, no. 2. 1768 Figures 57 , 58

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

Solanum villosum Type. "Solanum annuum hirsutus, baccis luteis", cultivated in England, at James Sherard’s garden in Eltham (Hortus Elthamensis) (lectotype, designated by Henderson 1974, pg. 56: Dillenius, Hortus Elthamensis 2: 366, t. 274, f. 353. 1732).

Solanum luteum Mill., Gard. Dict. ed. 8, no. 3. 1768.

Solanum villosum Type. "Solanumofficinarum lacinis luteis 1735, G.B. 166", cultivated in England at the Chelsea Physic Garden, seeds said to come from America, Herb. Miller s.n. [691 on sheet] (lectotype, designated by Edmonds 1979b, pg. 214: BM [BM000942567]).

Solanum rubrum Mill., Gard. Dict. ed. 8, no. 4. 1768.

Solanum villosum Type. Cultivated in England at the Chelsea Physic Garden, origin said to be the West Indies, Herb. Miller s.n. (lectotype, designated by Edmonds 1979b, pg. 215: BM [BM000942563]).

Solanum nigrum L. subsp. villosum (L.) Ehrh., Hannover. Mag. 1780, 14: 218. 1780.

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

Solanum villosum (L.) Willd., Prodr. Fl. Berol.: 87. 1787, nom. illeg., not Solanum villosum Mill. (1768)

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

Solanum nigrum L. var. rubrum (Mill.) Aiton, Hort. Kew. [W.Aiton] 1: 234. 1789.

Solanum villosum Type. Based on Solanum rubrum Mill.

Solanum aegyptiacum J.F.Gmel., Syst. Nat., ed. 13 [bis] 2: 385. 1791.

Solanum villosum Type. Egypt. Cairo: "circa Cairo", 1762, Herb. P. Forsskål 420 (lectotype, designated here: C [C10013600]).

Solanum villosum Lam., Tabl. Encycl. 2: 18. 1794, nom. illeg., not Solanum villosum Mill. (1768)

Solanum villosum Type. "Ex Gallia australi, &c.", Anon. s.n. (lectotype, designated by Edmonds 1979b, pag. 214 [as holotype]: P-LA [P00357624]).

Solanum villosum Moench, Methodus (Moench) 474. 1794, nom. illeg., not Solanum villosum Mill. (1768).

Solanum villosum Type. Germany. Hessen [?], C. Moench s.n. (no original material located).

Solanum alatum Moench, Methodus (Moench) 474. 1794, nom. cons. prop

Solanum villosum Type. Pakistan. Balochistan: Quetta, 30 km E Gumbaz, 1050 m, 30°02'N, 69°00'E, 17 May 1965, K.H. Rechinger 29684 (type cons. prop. by Knapp et al. 2017, pg. 988: W acc. # 1972-0017910).

Solanum villosum (L.) Willd., Enum. Pl. (Willdenow) 236. 1809, nom. illeg., not Solanum villosum Mill. (1768).

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

Solanum humile Bernh. ex Willd., Enum. Pl. (Willdenow) 236. 1809.

Solanum villosum Type. Cultivated in Berlin Botanic Garden, from "Europa australi" [southern Europe], Anon. s.n. (neotype, designated by Edmonds 2012, pg. 132 [as lectotype]: B [B-W04367 sheet 2]; isoneotype: B [B-W04367 sheet 1]).

Solanum suffruticosum Schousb. ex Willd., Enum. Pl. (Willdenow) 236. 1809.

Solanum villosum Type. Cultivated in Berlin Botanic Garden, said to be originally from “Barbaria” [northern Africa west of Egypt], Herb. Willdenow s.n. (lectotype, designated here: B [B-W04363010]).

Solanum incertum Dunal, Hist. Nat. Solanum 155. 1813.

Solanum villosum Type. “Nelen-Tsjunda” from Malabar [western India] (lectotype, designated here: Rheede von Draakestein, Hortus Malabaricus 10: t. 73. 1690).

Solanum erythraeum Dunal, Hist. Nat. Solanum 238. 1813, nom. illeg. superfl.

Solanum villosum Type. Based on Solanum rubrum L. (as S. rubrum "Murr. Syst. Veg 183, Gmel. Syst. Nat 384")

Solanum kitaibelii Schult., Öster. Fl., ed. 2, 1: 395. 1814.

Solanum villosum Type. Hungary. Sin. loc. “Pannonia”, Without collector [P. Kitaibel] s.n. [349] (lectotype, designated by Edmonds 1979b, pg. 214 [as holotype]: M [M-0166013]).

Solanum flavum Kit. ex Schult., Öster. Fl., ed. 2, 1: 394. 1814.

Solanum villosum Type. Hungary. Sin. loc. [ “Pannonia”], Without collector [P. Kitaibel] s.n. (lectotype, designated by Edmonds 1979b, pg. 214 [as holotype]: B-W [B-W04421010]; probable isolectotypes: BP [herb. Kit. fasc. IX: 105, 3 sheets], M [M0164200]).

Solanum ochroleucum Bastard, J. Bot. (Desvaux) 3: 20. 1814.

Solanum villosum Type. France. Pays de la Loire: Maine-et-Loire, Anjou [Angers], 1813, Without collector [prob. T. Bastard] s.n. (lectotype, designated here: G-DC [G00144474]; isolectotype: ANG [acc. # 2006.213]).

Solanum nigrum L. var. miniatum (Bernh. ex Willd.) Fr., Nov. Flor. Suec.: 111. 1823.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum villosum Lam. var. miniatum (Bernh. ex Willd.) Kostel., Clav. Anal. Fl. Bohem. 34. 1824.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum luteovirescens C.C.Gmel., Fl. Bad. 4: 177. 1826, nom. illegit. superfl.

Solanum villosum Type. Based on S. humile Bernh. ex Willd. (cited in synonymy).

Solanum puniceum C.C.Gmel., Fl. Bad. 4: 176. 1826, nom. illegit. superfl.

Solanum villosum Type. Based on S. miniatum Bernh. ex Willd. (cited in synonymy).

Solanum vulgatum (L.) Spenn. var. luteum Spenn., Fl. Friburg. 2: 427. 1829, nom. illeg. superfl.

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

Solanum nigrum L. subvar. ochroleucum (Bastard) Coss. & Germ., Intr. Fl. Anal. et Descr. Paris, 91. 1842.

Solanum villosum Type. Based on Solanum ochroleucum Bastard

Solanum villosum Lam. subvar. miniatum (Bernh. ex Willd.) Germ. & Wedd., Introd. Fl. Anales Paris 91. 1842.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum villosum Lam. forma miniatum (Bernh. ex Willd.) Döll, Rhein. Flor. 413. 1843.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum nigrum L. forma humile (Bernh. ex Willd.) Döll, Rhein. Flor. 413. 1843.

Solanum villosum Type. Based on Solanum humile Bernh. ex Willd.

Solanum nigrum L. forma villosum (Lam.) Döll, Rhein. Flor. 413. 1843.

Solanum villosum Type. Based on Solanum villosum Lam.

Solanum nigrum L. forma angulosum Döll, Rhein. Flor. 413. 1843.

Solanum villosum Type. Germany. "Bei Carlsruhe beim Linkenheimer Thore, A. Braun " (no specimens cited; no original material located, not found at KR); Germany. "Bei Philippsburg, Stud. Bauer " (no specimens cited; no original material located, not found at KR): Germany. "Bei Mannheim, Oppenheim, J.C. Döll " (no specimens cited; no original material located, not found at KR).

Solanum nigrum L. subsp. miniatum (Bernh. ex Willd.) Hartm., Sv. Norsk Exc.-Fl.: 35. 1846.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum nigrum L. subsp. villosum (Lam.) Hartm., Sv. Norsk Esc.-Fl. 34. 1846.

Solanum villosum Type. Based on Solanum villosum Lam.

Solanum nigrum L. subsp. humile (Bernh. ex Willd.) Hartm., Sv. Norsk Exc.-Fl. 34. 1846.

Solanum villosum Type. Based on Solanum humile Bernh. ex Willd.

Solanum sinaicum Boiss., Diagn. Pl. Orient. ser. 1, 11: 135. 1849.

Solanum villosum Type. Egypt. [Sinai] "Arabia petraea, Wadi Sheikh" [protologue "in fissuris rupium jugi Sianaitici in valle Wadi Scheick"], Mar 1846, E. Boissier s.n. (lectotype, designated here: G [G00343377] isolectotypes: DD, K [K000759386]).

Solanum plebeium A.Rich., Tent. Fl. Abyss. 2: 100. 1850.

Solanum villosum Type. Ethiopia? [ “Abyssinia”], Prov. Chiré [Shire?], R.Q. Dillon & A. Petit s.n. (lectotype, designated by Lester 1997, pg. 289: P [P00341983]; isolectotype: P [P00341984]).

Solanum nigrum L. subsp. luteovirescens Kirschl., Fl. Alsace 1: 532. 1852.

Solanum villosum Type. Based on Solanum luteovirescens C.C.Gmel. (nom. illeg.)

Solanum nigrum L. subsp. luteum (Mill.) Kirschl., Fl. Alsace 1: 532. 1852.

Solanum villosum Type. Based on Solanum luteum Mill.

Solanum nigrum L. subsp. puniceum Kirschl., Fl. Alsace 1: 532. 1852.

Solanum villosum Type. Based on Solanum puniceum C.C.Gmel. (nom. illeg.)

Solanum ochroleucum Bastard var. flavum Dunal, Prodr. [A. P. de Candolle] 13(1): 56. 1852.

Solanum villosum Type. Greece. Peloponnese: "in vineis agri Argolidis" [Argolis], J. Sartori 86 (lectotype, designated by Edmonds 1979b, pg. 216: G-DC [G00144471]).

Solanum miniatum Bernh. ex Willd. var. stenopetalum Dunal, Prodr. [A. P. de Candolle] 13(1): 56. 1852.

Solanum villosum Type. France. Auvergne-Rhône-Alpes: Isère, "circa Montaud prope Salon", J.L. Castagne s.n. (lectotype, designated here: G-DC [G00144520]; isolectotype: MPU not seen).

Solanum miniatum Bernh. ex Willd. var. villosissimum Dunal, Prodr. [A. P. de Candolle] 13(1): 56. 1852.

Solanum villosum Type. France. Provence-Alpes-Côte d’Azur: Var, Miraval, a l’entre de la grotte, 8 May 1807, A.P. de Candolle s.n. (lectotype, designated here: G-DC [G00144520]).

Solanum miniatum Bernh. ex Willd. var. subserratum Dunal, Prodr. [A. P. de Candolle] 13(1): 57. 1852.

Solanum villosum Type. “Arabie”, Without collector[No. 26] (lectotype, designated here: MPU [MPU091575]).

Solanum minutiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 58. 1852.

Solanum villosum Type. France. Hyères, A. Richard s.n. (lectotype, designated here: G-DC [G00144587]; isolectotype: P [P00582961]).

Solanum nilagiricum Schltdl., Linnaea 26: 484. 1855.

Solanum villosum Type. Cultivated in Germany at Halle [from Nilgiri Hills, India?] (no specimens cited; no specimens located [HAL? not seen]).

Solanum suffruticosum Schousb. ex Lange, Linnaea 28: 361. 1856, nom. illeg., not Solanum suffruticosum Schousb. ex Willd. (1809)

Solanum villosum Type. Cultivated in Denmark at Copenhagen, seeds from "Pontevedra Galleciae Aug 1852 [Galicia, Spain], 1855, Anon. s.n. (lectotype, designated here: C [C10019275]).

Solanum nigrum L. var. croceum Neilr., Fl. Nied.-Oesterr.: 535. 1858, nom. illeg. superfl.

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

Solanum nigrum L. var. miniatum (Bernh. ex Willd.) Neilr., Fl. Nied.-Oesterr. 535. 1858, nom. illeg., not Solanum nigrum L. var. miniatum (Bernh. ex Willd.) Fr. (1823)

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

S. nigrum L. var. luteum Neilr., Fl. Nied.-Oesterr. 535. 1858.

Solanum villosum Type. Based on Solanum flavum Kit. ex Schult.

Solanum nigrum L. var. suffruticosum (Schousb. ex Willd.) Moris, Fl. Sardoa 3: 148. 1858-1859.

Solanum villosum Type. Based on Solanum suffruticosum Schousb. ex Willd. (cites Solanum suffruticosum Schousb. concept of Dunal)

Solanum nigrum L. var. luteum Döll, Fl. Baden [ Döll] 767. 1859, nom. illeg., not Solanum nigrum L. var. luteum Neilr. (1858).

Solanum villosum Type. Germany. "in der Carlsruher Gegend bei Dachslanden und Mühlburg, bei Schwetzingen, Mannheim und Dossenheim" (no specimens cited; no original material located at KR).

Solanum canescens Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg., not Solanum canescens Blume (1826).

Solanum villosum Type. Hungary. "E Cottu Thurócz. v Barodiensi" [protologue "in Cottu Thurociensi et Borsodiensi", P. Kitaibel s.n. (lectotype, designated here: BP [herb. Kit. fasc. IX: 105, numbered sheet]; isolectotypes: BP [unlabelled sheets in fasc. IX: 105]).

Solanum flavum Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg, not Solanum flavum Kit. ex Schult.(1814)

Solanum villosum Type. Hungary. "E Cottu Thurócz. v Barodiensi" [protologue "in Cottu Thurociensi et Borsodiensi", P. Kitaibel s.n. (lectotype, designated here: BP [BP [herb. Kit. fasc. IX: 105, numbered sheet]; isolectotypes: BP [unlabelled sheets in fasc. IX: 105]).

Solanum incanum Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg. superfl., also not S. incanum L. (1753)

Solanum villosum Type. Based on Solanum kitaibelii Kit. ex Schult. (cited in synonymy).

Solanum olivaceum Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg. superfl.

Solanum villosum Type. Based on Solanum humile Bernh. ex Willd. (cited in synonymy).

Solanum pseudovillosum Schur, Enum. Pl. Transsilv. 477. 1866.

Solanum villosum Type. Transylvania [Romania], Hermannstadt [Sibiu] and Kronstadt [Brașov], Aug-Sep, P.J.F. Schur s.n. (no herbaria cited; no original material located; LW?, BRNU?).

Solanum nigrum L. subsp. alatum (Moench) Čelak., Prodr. Fl. Böhmen [2:?] 309. 1871.

Solanum villosum Type. Based on Solanum alatum Moench

Solanum nigrum L. subsp. villosum (Lam.) Čelak., Prodr. Fl. Böhmen 309. 1871.

Solanum villosum Type. Based on Solanum villosum Lam.

Solanum patens Lowe, Man. Fl. Madeira 2: 74. 1872.

Solanum villosum Type. Portugal. Madeira: Ribeiria do Sta. Luzia, 1 Jul 1833, R.T. Lowe 547 [a] (lectotype, designated by Edmonds 2012, pg. 133: BM [BM000083209]; other duplicates of Lowe 547 with different collecting dates are not type material).

Solanum villosum Lam. var. velutinum Lowe, Man. Fl. Madeira 2: 76. 1872, as " velutina "

Solanum villosum Type. Portugal. Madeira: Valle [garden], 24 Aug 1831, R.T. Lowe [722] (lectotype, designated here: BM [BM000056199]).

Solanum villosum Lam. var. laevigatum Lowe, Man. Fl. Madeira 2: 78. 1872, as " laevigata "

Solanum villosum Type. Portugal. "Lisbon, court of the Museum [Jan 1865]", R.T. Lowe s.n. (lectotype, designated by Edmonds 2012, pg. 133: BM [BM000847650]; isolectotype: K [K000414066]).

Solanum hildebrandtii A.Braun & C.D. Bouché, Ind. Sem. Hort. Berol. 18. 1874.

Solanum villosum Type. Somalia. Saanag: Yafir, Ahlgebirge [Karin Yafir, Ahl Mountains, N Somalia], J.M. Hildebrandt 865 (neotype, designated by Edmonds 2012, pg. 133 [as lectotype]: BM [BM000907531]; isoneotype: L [L.2879554]).

Solanum nigrum L. var. humile (Bernh. ex Willd.) Boiss., Fl. Orient. 4: 284. 1879.

Solanum villosum Type. Based on Solanum humile Bernh. ex Willd.

Solanum nigrum L. var. induratum Boiss., Fl. Orient. 4: 284. 1879, nom. illeg. superfl.

Solanum villosum Type. Based on Solanum nigrum L. var. suffruticosum (Schousb. ex Willd.) Moris (cited in synonymy)

Solanum villosum Lam. subsp. ochroleucum (Bastard) Nyman, Consp. Fl. Eur. 3: 526. 1881.

Solanum villosum Type. Based on Solanum ochroleucum Bastard

Solanum nigrum L. subsp. miniatum (Bernh. ex Willd.) Arcang., Comp. Fl. Ital. (Arcangeli): 497. 1882.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum morella Desv. subsp. luteum (Mill.) Rouy, Fl. France 10: 366. 1908.

Solanum villosum Type. Based on Solanum luteum Mill.

Solanum plebeium A.Rich. var. brachysepalum Bitter, Bot. Jahrb. Syst. 49: 564. 1913.

Solanum villosum Type. Ethiopia. Oromia: "Gallahochland, Ego", 2000 m, H. Ellenbeck 358 (holotype: B, destroyed; no duplicates located).

Solanum plebeium A.Rich. var. subtile Bitter, Bot. Jahrb. Syst. 49: 564. 1913.

Solanum villosum Type. Ethiopia. Harari: "Harar, Hararmaja-See", H. Ellenbeck 462 (holotype: B, destroyed; no duplicates located).

Solanum nigrum L. forma glabrum Chiov., Res. Sci. Somalia Ital. 128. 1916.

Solanum villosum Type. Somalia. Baay: Berdale [Berdaale], 12 Oct 1913, G. Paoli 935 (holotype: FT [FT003070]).

Solanum nigrum L. forma humile (Bernh. ex Willd.) Lindm., Svensk Fan. Fl. 480. 1918.

Solanum villosum Type. Based on Solanum humile Bernh. ex Willd.

Solanum nigrum L. var. sancti-marini Pamp., Boll. Mus. Republ. San Marino 4(3-4): 116. 1920.

Solanum villosum Type. San Marino. Sin. loc., Without collector s.n. (no specimens cited; possibly at FI?; no original material located).

Solanum nigrum L. var. alatum (Moench) Fiori, Nuov. Fl. Italia 2: 311. 1926.

Solanum villosum Type. Based on Solanum alatum Moench

Solanum nigrum L., var. aurantium Maire, Bull. Soc. Sci. Nat. Maroc 8(4-6): 139. 1928.

Solanum villosum Type. Morocco. Souss-Massa: Anti-Atlas Mountains "Djebel Inter à Talaïnt” [Djebel Inter à Talguit (=Tafraoute?)], 600 m, 4 Apr 1922. R. Maire s.n. (lectotype, designated here: RAB [RAB-44657]; isolectotypes: MPU [MPU001957], P [P00489756]).

Solanum villosum Lam. var. miniatibaccatum Blom, Act. Hort. Gothoburg. 10: 207. 1935.

Solanum villosum Type. Sweden. Göteborgstrakten: Göteborg, som ogräs i Botaniska Trädgården, 1935, C. Blom s.n. (lectotype, designated here: GB; isolectotypes: LD, S, UPS).

Solanum luteum Mill. subsp. alatum (Moench) Dostál, Květena Č.S.R. 1270. 1949.

Solanum villosum Type. Based on Solanum alatum Moench

Solanum transcaucasicum Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 332. 1955.

Solanum villosum Type. Azerbaijan. Lankaran: Talysh, near Tatuni "Talysh bei Tatuni", R.F. Hohenacker s.n. (holotype: LE).

Solanum olgae Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 333. 1955.

Solanum villosum Type. Uzbekistan. Samarkandaya Oblast’, [Tadzkikistan, Hissar, near Kschtut in protologue] Kschtut, on the road to Sherbut, Fedotov & Gol’bek s.n. (holotype: LE; isotype: LE).

Solanum zelenetzkii Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 336. 1955.

Solanum villosum Type. Ukraine: Crimea, "Tauria Australis", Jalta [Yalta in protologue], Oreanda, 21 Aug 1906, K. Golde s.n. (holotype: LE).

Solanum woronowii Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 337. 1955.

Solanum villosum Type. Georgia. Abkhazia AR: " Abchazia [Abkhazia], Gagry", 30 Oct 1954, A. Pojarkova 6 (holotype: LE).

Solanum luteum Mill. var. calvum Wessely, Feddes Repert. Spec. Nov. Regni. Veg. 63: 315. 1960.

Solanum villosum Type. Cultivated in Germany at Greifswald, seeds from Hort. Bot. Marseille, I. Wessely 214 (holotype: GFW).

Solanum luteum Mill. var. procerum Wessely, Feddes Repert. Spec. Nov. Regni. Veg. 63: 315. 1960.

Solanum villosum Type. Cultivated in Germany at Greifswald, seeds from München Botanical Garden, 1957, I. Wessely 320 (holotype: GFW).

Solanum luteum Mill. forma miniatibaccatum (Blom) Wessely, Feddes Repert. Spec. Nov. Regni. Veg. 63: 315. 1960.

Solanum villosum Type. Based on Solanum villosum Lam. var. miniatibaccatum Blom

Solanum purpureilineatum Sabnis & Bhatt, Bull. Bot. Surv. India 12(1-4) 258. 1972 [ “1970”].

Solanum villosum Type. India. Gujarat: Baroda District, Baroda, L.V. Palace compound, 2 Oct 1960, S.D. Sabnis 2762, 2763 (holotype: BARO [two sheet holotype, single gathering).

Solanum carmanicum Schönb.-Tem., Fl. Iranica 100: 8. 1972.

Solanum villosum Type. Iran. Kerman: Bam, ca. 900 m, 7 May 1948, K.H. Rechinger & F. Rechinger 3630 (holotype: W [1958-0003176]; isotype: E [E00251353], G [G00343074], US [00992058, acc. # 2316288]).

Solanum villosum Mill. subsp. alatum (Moench) Edmonds, Bot. J. Linn. Soc. 75: 171. 1977.

Solanum villosum Type. Based on Solanum alatum Moench

Solanum nigrum L. var. humile (Bernh. ex Willd.) C.Y. Wu & S.C.Huang, Acta Phytotax. Sin. 16(2): 72. 1978, nom. illeg., not Solanum nigrum L. var. humile (Bernh. ex Willd.) Boiss. (1879).

Solanum villosum Type. Based on Solanum humile Bernh. ex Willd.

Solanum villosum Mill. subsp. puniceum (Kirschl.) Edmonds, Bot. J. Linn. Soc. 78: 215. 1979.

Solanum villosum Type. Based on Solanum nigrum L. subsp. puniceum Kirschl.

Solanum villosum Mill. subsp. miniatum (Bernh. ex Willd.) Edmonds, Bot. J. Linn. Soc. 89: 166. 1984.

Solanum villosum Type. Based on Solanum miniatum Bernh. ex Willd.

Solanum villosum Mill. subsp. alatum (Moench) Dostál, Folia Mus. Rerum Nat. Bohemiae Occid., Bot. 21: 11. 1984, nom. illeg., not Solanum villosum Mill. subsp. alatum (Moench) Edmonds (1977)

Solanum villosum Type. Based on Solanum alatum Moench

Solanum luteum Mill. var. miniatibaccatum (Blom) Hyl., Uppsala Univ. Årsskr. 7: 279. 1945.

Solanum villosum Type. Based on Solanum villosum Lam. var. miniatibaccatum Blom

Type.

Cultivated in Chelsea Physic Garden from "Barbadoes where it is supposed to grow naturally", Herb. Miller s.n. (lectotype, designated by Henderson 1974, pg. 54: BM [BM000942575]) .

Description.

Annual to short lived erect or weakly scrambling perennial herbs up to 0.5 m tall, subwoody at base and much branched. Stems spreading to decumbent, terete to ridged, green to purple, not hollow; new growth densely pubescent with simple, spreading, uniseriate, translucent, eglandular and/or glandular trichomes, these 3-10-celled, 0.2-2.0 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5-5.0(-10.0) cm long, 0.7-2.5(-6.5) cm wide, broadly to narrowly ovate to elliptic, membranous, green on both sides, without smell or somewhat pleasant smelling; adaxial surfaces sparsely to densely pubescent with spreading, simple, uniseriate eglandular and/or glandular trichomes like those on stem evenly along veins and lamina; abaxial surfaces more densely pubescent on veins and lamina; major veins 4-6 pairs; base acute to truncate, short-attenuate, often asymmetric; margins sinuate-dentate to rarely entire; apex acute; petioles 0.5-3.0(-4.5) cm long, pubescent with simple uniseriate glandular and/or eglandular trichomes like those on stems. Inflorescences 0.4-2.0 cm long, internodal, simple, the flowers spaced along the rhachis, with (2-)3-5(-8) flowers clustered at the tip or more commonly spaced along the rhachis, pubescent with spreading simple glandular and/or eglandular uniseriate trichomes like those of the stems; peduncle 0.4-1.5 cm long, straight; pedicels 4-7 mm long, 0.2-0.3 mm in diameter at the base and 0.4-0.5 mm at apex, spreading, articulated at the base; pedicel scars spaced 0-1.0 mm apart. Buds globose, the corolla exserted ca. 1/5 from the calyx before anthesis. Flowers 5-merous, all perfect. Calyx tube 1.2-1.5 mm long, conical, the lobes 0.8-1.5 mm long, 0.5-0.8 mm wide, elliptic to triangular with obtuse thickened apices and paler (almost scarious) sinuses, pubescent with spreading simple uniseriate eglandular and/or glandular trichomes like those on stem. Corolla 8-15(-20) mm in diameter, white with a yellow-green central portion near the base and occasionally with purple stripes along lobe midveins abaxially, stellate, lobed 1/2 way to the base, the lobes 2.5-4.5 mm long, 2.0-3.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate-pubescent abaxially with simple uniseriate eglandular trichomes. Stamens equal; filament tube minute, pubescent with spreading uniseriate simple eglandular trichomes adaxially; free portion of the filaments 1.0-1.3 mm long, pubescent like the tube; anthers 1.8-2.2(-2.4) mm long, 0.5-0.7 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.8-3.5(-4.0) mm long, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower half, exserted 0-1 mm beyond anther cone; stigma capitate, the surface minutely papillate, green in live plants. Fruit an ellipsoid berry, usually somewhat longer than broad, 8.5-10 mm long, 8.0-9.5 mm wide, (red-)orange to yellow at maturity, the pericarp thin, shiny and translucent; fruiting pedicels 8-14 mm long, 0.4-0.5 mm in diameter at the base, 0.7-1.5 mm at apex, strongly reflexed, becoming woody, spaced 1.0-2.0 mm apart not falling with the fruit, remaining on the plant and always persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.0-1.5 mm long, the lobes 2.0-3.0 mm long, strongly reflexed in fruit. Seeds 20-40 per berry, 1.8-2.2 mm long, 1.5-1.7 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent, but occasionally 1-2 found in North African and Arabian material, ca. 0.5 mm in diameter. Chromosome number: 2n =4x=48 ( Jørgensen 1928; Tokunaga 1933; Stebbins and Paddock 1949; Soria and Heiser 1961 [as S. roxburghii and S. villosum ]; Heiser et al. 1965 [as S. roxburghii ]; Gerasimenko and Reznikova 1968; Venkateswarlu and Rao 1972; Henderson 1974; Tandon 1974; Randell and Symon 1976; Edmonds 1977, 1982, 1983, 1984a; Ganapathi and Rao 1986a; Bukenya 1996; Ojiewo et al. 2006; Olet et al. 2015).

Distribution

(Figure 59 ). Native to the northern Hemisphere, from Europe, especially around the Mediterranean basin, Arabian Peninsula to dry sub-tropical Asia and eastern Africa (where it is cultivated for its fruit); introduced globally. The native distribution of S. villosum is restricted to the Old World, but cultivated and/or introduced populations that never seem to persist are occasionally found in North America and the Caribbean ( Stebbins and Paddock 1949).

Ecology.

Grows in open and disturbed areas in a wide variety of habitat types, including in cities; between sea level and 3,000 (-3,600) m elevation.

Common names.

Austria: Rotbeeren-Nachtschatten, Gelbbeeren-Nachtschatten, Gelbrotbeeren-Nachtschatten ( Fischer et al. 2005), Flügel-Nachtschatten; Chad: abriki, waatenni, waha tinni; China: hong guo long kui ( Zhang et al. 1994); Denmark: højrød natskygge ( Hartvig 2015); Egypt: 'enneyb, aneb el dib/aneb el dik; ilshsmdhut, ribai hauwei; Ethiopia: auwt; Greece: styphnos; India: gammie, kambe, tutguna (Kalahandia language); Iraq: anab dheeb; Italy: Solanum morella rossa ( Pignatti 1982); Kenya: isoiyot/isoyot, itulu, ksoya, kusuyo, manangwe, managu, munavu, ndolu, ol’ momoit/ ol’ momoye, ormomoi; New Zealand: red-berried nightshade ( Webb et al. 1988); Nigeria: gautan kaji; Oman: mjaj; Pakistan: tol angur; Palestine: inab ed dib; Poland: psianka niska, psianka kosmata, psianka skrzydlata ( Pawłowskiego 1963); Portugal: erva de Santa Maria; Saudi Arabia: aah’al, khur ma; Slovenia: lulok žltý, lulok krídlatý ( Bertová and Goliašová 1993); Somalia: biyeche El, karire; Spain: modalicou, yerba mora; Sweden: Gul nattskatta, röd nattskatta ( Mossberg et al. 2003); Tanzania: emunauui, en’songwe, manakk, mnafu/mnavu, osuga, soku; Uganda: eswiga, ocuga; United Arab Emirates: black nightshade; United Kingdom: red nightshade ( Stace 2010).

Uses.

Leaves used as spinach (usually boiled, often in milk) and as a pot-herb in Europe, Africa and the Middle East; berries eaten raw (especially by children, see Uses above) and cooked ( Pojarkova 1955a, b). Medicinal uses include treatment of eye conditions and swellings (see Yang and Ojiewo 2013).

Preliminary conservation status

( IUCN 2016). Solanum villosum is a very widespread species and can be assigned a preliminary status of LC (Least Concern).

Discussion.

This common European species was long known as S. luteum (e.g. Hawkes and Edmonds 1972) and this name was in wide use in botanical gardens as the name for this red-fruited taxon. The name S. villosum Lam. was recognised as a synonym for S. luteum . The name S. villosum Mill. was not in use in Europe until resurrected by Edmonds (1979b), but had been used by Stebbins and Paddock (1949) in their treatment of the morelloids of California. Edmonds (1979b), following what is now Article 11.5 of the Code ( McNeill et al. 2012), changed the name to S. villosum Mill. citing the explicit synonymisation of S. luteum with S. villosum Mill. by Stebbins and Paddock (1949). Dostál (1949), only a month later than Stebbins and Paddock (1949), synonymised S. villosum with S. luteum , but unfortunately used the name in the Lamarckian sense, although in a letter to Edmonds (quoted in Edmonds 1979b) equated S. villosum Mill. and S. villosum Lam. Many specimens in European herbaria are still identified as S. luteum or as S. alatum (see Knapp et al. 2017 for discussion of S. alatum ).

Variation in pubescence and stem characteristics within S. villosum has been recognised at many ranks from subspecies, varieties and different species ( Hawkes and Edmonds 1972; Edmonds 1979b, 1984b; Henderson 1974; Symon 1981; Edmonds and Chweya 1997; Olet 2004; Dehmer and Hammer 2004). The various synonyms and infraspecific ranks reflect the wide morphological variation observed within the species, where indumentum can range from nearly absent (the plants glabrous) to densely pubescent with eglandular and/or eglandular trichomes. The recognition of the glandular variants at infraspecific rank is not supported by AFLP data ( Manoko 2007; Olet et al. 2011). In fact, studies have shown that pubescence varies in individuals according to age, where seedlings can exhibit glandular hairs which are lost during growth ( Henderson 1974; Seithe 1979; Edmonds 1982). Here we have an inclusive, broad concept of S. villosum that includes this variability but does not recognise it at any rank.

Solanum villosum can be easily recognised from similar species by its orange-red berries (ranging from yellow to bright red) that are slightly elongate and ellipsoid. The only other species present in the Old World with pale yellow fruits is S. palitans , but this taxon differs in being a prostrate plant with thinly membranous 3-lobed leaves and is easy to tell apart. The rest of the morelloids in the Old World have rounded or subglobose black or green berries. Solanum villosum is much more common in southern Europe than is S. nigrum , while S. nigrum becomes more common towards northern and central Europe. These latter two species are difficult to tell apart based on herbarium material alone, especially when mature fruit colour is not available. Calyx lobes of S. villosum are generally longer than those of S. nigrum (relative to width), strongly reflexed at fruit maturity and the sinus is rounded (as opposed to acute in S. nigrum ) and, in flower, the calyx tube has pale patches that look like small windows or white/translucent areas below the sinus itself. This is particularly easy to see in herbarium specimens. The leaves of S. villosum are similar to those of S. retroflexum in being usually more or less rhomboid with dentate margins, but leaves are more narrowly rhomboid than those of S. retroflexum (i.e. narrower relative to length). Specimens from the southern part of the range of S. villosum and the northern range of S. retroflexum (e.g. Malawi and Mozambique) can be difficult to distinguish in the absence of information on fruit colour ( S. retroflexum has dull purple-black berries, also with strongly reflexed calyx lobes in fruit).

Solanum villosum has generally distinctly longer filaments relative to anther length than other species (except S. opacum and S. retroflexum , see descriptions of these species), although large variation can be observed within and between individuals especially in herbarium specimens. We have observed in the field that filaments elongate during flower maturation, where younger flowers in the same inflorescence can be seen with shorter filaments and older flowers with fully extended and distinctly longer filaments (SK photo 101532). Solanum villosum has a large capitate stigma that does not generally extend much beyond the anthers, in contrast to S. tarderemotum with which it often grows in sympatry in Africa, where the style is long-exserted and often bent and the stigma is smaller.

A set of herbarium and seed bank accessions of S. villosum from Kenya have narrower leaves with entire margins and larger, branched inflorescences with fruits in which calyx lobes do not recurve as strongly as in more typical populations, but that lack stone cells in the berries. These accessions are somewhat morphologically intermediate between S. villosum and S. tarderemotum and could present man-made or natural introgression between the two species (see discussion of S. tarderemotum ).

The origin of the tetraploid S. villosum has been the focus of previous crossing and cytological studies (e.g. Edmonds 1979a). Molecular ( Poczai and Hyvönen 2011), experimental breeding ( Tandon and Rao 1966a, 1966b; Venkateswarlu and Rao 1972; Rao 1978) and chromosome binding data ( Bhiravamurty and Rethy 1984; Sultana and Alam 2007) suggest that the species is an ancient autopolyploid with regular bivalent formation derived from S. americanum . Edmonds (1979a) suggested that S. villosum is derived from the diploid parents S. americanum and S. sarrachoides , because these two diploids are interfertile. The hypothesis in which a native of the New World, S. sarrachoides , would have given rise to an Old World endemic is not particularly convincing, but current distributions of progenitor taxa do not necessarily have to overlap (e.g. Nicotiana tabacum L., see Chase et al. 2003). The autopolyploid origin of S. villosum and its contribution to the autoalloploid S. nigrum derived from S. villosum and S. americanum , is more likely, considering their sympatry across the entire range of S. villosum . Furthermore, S. sarrachoides is reproductively isolated from all other diploids studied thus far ( Edmonds 1977), making it less likely that the species has been involved in allopolyploidy. A possible diploid progenitor could be S. chenopodioides , however, as suggested by Edmonds (1979a), with which S. nigrum and S. villosum share some of their distributions. It is much more likely, however, based on the current molecular, morphological, artificial crossing and cytological studies that S. nigrum is an autoallopolyploid derived from a genome duplication of the tetraploid S. villosum that comprises genomes of two distinct populations of S. americanum . Morphological characters support this, in that S. americanum and S. villosum both drop their fruits without pedicels.

Edmonds (1979b) superfluously lectotypified S. nigrum var. villosum with a specimen in the Linnean herbarium (LINN 248.19), apparently not realising the name had already been lectotypified completely in accordance with the protologue by Henderson (1974), using the illustration from Hortus Elthamensis ( Dillenius 1732; see Jarvis 2007).

Although Miller (1768) used Linnaean names extensively in his Gardener’s dictionary, it is unlikely that he had seen Linnaeus’ (1767) 12th edition of Systema naturae where S. rubrum was first published (J.E. Dandy pers. comm. to J.M. Edmonds, quoted in Edmonds 1979b), so S. rubrum Mill. is considered a new name, rather than a citation of S. rubrum L. (nom. utique rej. prop. see Doubtful names).

Edmonds (2007) typified the names in morelloid solanums coined by J.F. Gmelin (1791) for plants collected by Pehr Forsskål, but for some reason did not designate a lectotype for S. aegyptiacum . We here select the specimen in the Forsskål herbarium (C10013600) that corresponds to Gmelin’s description and is labelled "aegyptiacum a) fructu rubro". Edmonds (2007) considered " S. aegyptiacum Forssk." to be not validly published because it lacked a description or diagnosis for the species, although she included one of its varieties in the synonymy of S. memphiticum (as " S. aegypticum b) fructu nigro; foliis integris, villosissimus "). We agree that Forsskål did not validly publish the name S. aegyptiacum , but it was validly published by J.F. Gmelin.

Lamarck (1794) coined his name S. villosum without reference to Miller and, because the more commonly used name for this species was S. luteum until Edmonds (1979b) changed the name (see above), many of the new combinations and infraspecific taxa attributed S. villosum to Lamarck rather than Miller.

The name S. alatum Moench has been commonly used for densely pubescent eglandular populations of S. villosum ( Edmonds 1979b). There was considerable confusion over the identity of the type of S. alatum ; Edmonds (1979b) typified it with an element she, in the same publication, excluded from her concept of S. alatum . The name has been proposed for conservation with a conserved type ( Knapp et al. 2017) to overcome this disruptive typification; the conserved type proposed by Knapp et al. (2017) is cited here.

Solanum suffruticosum apparently was a “herbarium” name of Peter Schousboe, a Danish botanist who worked in Tangiers in the early part of the 19th century. The epithet was used and attributed to Schousboe by Willdenow (1809) who referred to plants grown in Berlin that originated from northern Africa (seeds sent by Schousboe?); we have selected the sheet in the Willdenow herbarium with annotations of the name "Solanum suffruticosum" and "Hort. bot. Berol. W." as the lectotype for this name. Almost 50 years later, Johan Lange, working in Copenhagen, also validated S. suffruticosum , but with a “?” as to the name ("d) Solanum suffruticosum Schousb.?" and citing a different collection from Spain (C10019275), that we here designate as the lectotype. Lange’s name is a later homonym and illegitimate.

Dunal (1813) cited a single element for his S. incertum , an illustration in Rheede von Draakestein’s Hortus Malabaricus (1689) with the common name “Nelen-Tsundja”; we here designate this plate as the lectotype for S. incertum . The accompanying description states the berries are orange.

Solanum kitaibelii and S. flavum are names coined by Schultes (1814) based on specimens of Pál Kitaibel; S. kitaibelii as a replacement for Kitaibel’s "Solanum incanum" that Schultes realised was pre-occupied ("Hr. Prof. Kitaibel nannte diese Art S. incanum ; da aber Flora peruviana schon ein S. incanum hat, so nannte ich es nach meinem hochverdienten Freunde, der es zuerts in Ungern fand" - Professor Kitaibel named this species S. incanum , but there is already one in the Flora Peruviana, so I name it after my esteemed friend, who found it first in Hungary). Edmonds (1979b) cited specimens as holotypes for these names, effectively lectotypifying them. She cited a specimen in the Willdenow herbarium (B-W04421010) as the “holotype” of S. flavum ; it is more likely however, that Schultes used specimens now held in M ( Schultes’s herbarium at Landshuth, where he worked at the time of description of these two taxa and this is now part of M). The lectotype for S. kitaibelii has the name in Schultes’s hand (M-0166013), but the isolectotype for S. flavum does not (M-0164200).

We have designated a specimen with the exact locality as the lectotype of S. ochroleucum ; the G-DC duplicate is better preserved (G00144474).

Döll’s various infraspecific taxa described in his Rheinische Flora (1843) as “Formen” (here taken as the rank of forma) are mostly new combinations of previously published epithets, but his forma Solanum nigrum angulosum is based on several cited localities in Germany. We have not been able to locate specimens for these collections and defer typification until more in-depth searches are done. It may be that these names are based on field observations and not herbarium material.

Solanum sinaicum was described by Boissier (1849) using a collection of his own from Arabia; he did not cite a herbarium, so we are lectotypifying this name with a specimen in G (G00343377). Although the protologue mentions "baccis nigricantibus", the plants are unmistakably S. villosum ; we suspect the fruit colour was cited from the dry specimen.

The spelling of Richard’s (1850) name S. plebeium has sometimes been corrected to “plebejum” (e.g. Edmonds 2012). This is not necessary and incorrect, "plebeium is the nominative neuter singular of “plebeius”, meaning “commoner” or "of the common people".

Dunal (1852) described three varieties of S. miniatum ; var. Solanum miniatum villosissimum and var. Solanum nigrum stenopetalum based on European specimens and var. Solanum subserratum subserratum based on a specimen from “Arabia”. Edmonds (1979b) discussed the identities of these varieties, but did not typify them. Dunal cited the same collection Castagne s.n. from "Montaud prope Salon" for both var. Solanum nigrum stenopetalum and var. Solanum miniatum villosissimum ; we have lectotypifed var. Solanum miniatum villosissimum with the other specimen cited that does not overlap with that used for var. Solanum nigrum stenopetalum (G00144520). Material in Dunal’s herbarium at MPU (MPU091757), labelled as “Arabie” and cited by Edmonds (1979b), is labelled as var. Solanum miniatum villosissimum and numbered "No. 26" and is undoubtedly original material of var. Solanum subserratum subserratum . This sheet is here designated the lectotype for this name, despite its having an annotation label with the incorrect infraspecific name. In describing S. minutiflorum , he ( Dunal 1852) cited two collections "Pannonia circa Bannat - (Terenbinki l.c.)" and a specimen from Richard. We have chosen the latter as the lectotype because it has two duplicates; neither Edmonds (1979b) nor we were able to locate the other syntype.

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 above). Solanum flavum and S. canescens were published as alternative names "Solanum flavum vel canescens" and therefore have the same type; we have selected the sheet in the Kitaibel herbarium at BP with the locality on the label matching that in the protologue ("E Cottu Thurócz. v. Borsodiensi", BP [Herb. Kit. fasc. IX: 105]).

In his Manual flora of Madeira, ( Lowe 1872) coined several names we here consider synonyms of S. villosum , both at the specific ( S. patens ) and infraspecific level ( S. villosum var. velutinum , S. villosum var. laevigatum ). Edmonds (1979b: 215) lectotypified var. Solanum villosum velutinum citing the type as the "same as the holotype of Solanum villosum Lam.". Lamarck (1794) was cited along with many other works, including Linnaeus’ (1753) S. nigrum var. villosum , so arguably var. Solanum villosum velutinum is an illegitimate superfluous name because S. nigrum var. villosum was cited in synonymy and the epithet “villosum” should have been taken up. We here re-lectotypify var. Solanum villosum velutinum with a specimen cited by Lowe and represented in BM (BM000056199). Edmonds (1979b) lectotypified S. patens with "Lowe 547 BM!", but we exclude all but the sheet labelled by J.M. Edmonds as “lectotype” (BM000083209) as type material; other specimens at BM labelled "Lowe 547" have different collecting dates and localities.

Edmonds (2012) designated wild-collected material as the lectotype of S. hildebrandtii stating that, in Bitter’s re-description of this species in Solana africana (1913a: 467-468), he had designated a holotype. However, Braun and Bouché clearly stated that the material they used was from cultivation and Bitter (1913a) did not re-typify S. hildebrandtii with wild-collected material, he merely cited it as something he had seen. This typification is corrected here to neotype from lectotype.

Two un-numbered collections were cited in the protologue of S. nigrum var. aurantium ( Maire 1928); Maire s.n. (coll. 4 Apr 1922) and Maire s.n. (coll. 26 Mar 1923). We have selected the duplicate in Morocco (RAB-44657) of the former as the lectotype because it has numerous duplicates. We have only been able to find a single duplicate (MPU001957) of the collection made in March 1923.

Blom (1935) cited two collections (H. Christoffersson s.n., coll. 1930; C. Blom s.n., coll. 1935) and several herbaria in his description of S. villosum var. miniatibaccatum ; we have selected the GB duplicate of his un-numbered collection from 1935 as the lectotype.

The names coined by Pojarkova (1955b) and here regarded as synonyms of S. villosum were first published in Russian only ( Pojarkova 1955a), but later in the same year validated with Latin descriptions and type specimen designations.

Solanum purpureilineatum ( Sabnis and Bhatt 1972) was described using two specimens with two different collecting numbers from a single gathering. Rather than lectotypifying this name, we are treating this as a two sheet holotype (see McNeill et al. 2012: 14-15, Art. 8.3 Ex. 4).

Selected specimens examined.

A total of 1,287 specimens were examined from 86 countries during the study across Africa, Asia, Australia, Eurasia and the Pacific. Specimens from the entire range (including adventives) were studied in order to understand the full range of morphology within the species. All specimens examined from the Old World can be seen in Appendix 2 (csv format) and Appendix 3 (traditional Specimens Examined list in pdf format).

Doubtful names

Solanum cestrifolium Jacq., mentioned by Weinmann (1824), as well as by Sprengel (1825), but no original publication of this epithet by Jacquin has been found (see discussion under S. nigrum ).

Solanum foetidum Rottb., Acta Lit. Univ. Hafn. 1: 287. 1778. Type. “Surinam”, C.F. Rottbøll s.n. (no original material located); this name has sometimes been referred to the morelloids, but is likely not to even be a Solanum , it might refer to a plant of Datura .

Solanum foliosum Link, Phys. Beschr. Canar. Ins. [Buch] 144. 1825. Type. Spain. Islas Canarias: Tenerife, Puerto Orotava, L.C. Buch s.n. (no specimens cited; original material seen by Link in B, destroyed). Link’s description suggests this plant is like, not the same as, S. nigrum , but then goes on to describe a plant with pinnatifid leaves and flowers larger than those of S. nigrum . Possibly S. nava Webb & Berthel. (1845) or perhaps the cultivated S. laxum Spreng. (1824)

Solanum nigrum L. var. aspergilliflorum Sendtn., Fl. Bras. (Martius) 10: 16. 1846. Type. Sin. loc. [Brazil?] (no specimens or collectors cited). Might be S. chenopodioides Lam. based on Sendtner’s notes " S. chenopodioides Lam.?" beneath the description in the original publication, but the description mentions stellate hairs on the leaf undersides, a character that does not occur in the Morelloid clade.

Solanum nigrum L. var. humile Macloskie, Rep. Princeton Univ. Exped. Patagonia 8: 707. 1905. The “description” consists solely of a reference to " S. humile Salisb." an illegitimate superfluous name citing S. nigrum L. in synonymy. Solanum nigrum does not occur in Patagonia, so this name cannot be considered homotypic; identity uncertain, but certainly a species native to the New World.

Solanum nigrum L. var. stylosum Witasek ex Reiche, Anales Univ. Chile 124: 456. 1909. Type. Chile. "Provincias Centrales " (no specimens cited; no original material found). The description is very short and no specimens annotated as this from Chile have been found at W or WU; it could be S. alphonsei Dunal of the Dulcamaroid clade; the description of the peduncle as twice as long as the branches is indicative of that species, rather than a morelloid.

Solanum rubrum L., Syst. Nat. ed. 12: 173. 1767, nom utique rej. prop. No original type material traced, Edmonds (1979b) treated this as a nomen dubium and it has been proposed for rejection ( Knapp et al. 2017).

Solanum uliginosum Blume, Bijdr. Fl. Ned. Ind. 13: 695. 1826. No specimens or localities cited, not treated by Nees van Esenbeck (1837), description merely copied from Blume (1826) by Dunal (1852) could be S. americanum , S. nigrum or S. scabrum. We have found no specimens at L that could be unequivocally associated with this name; perhaps material is held in Bogor.

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

Type. Based on Solanum uliginosum Blume

Names (designations) not validly published

Designations are listed in alphabetic order; Code Articles are numbered as in McNeill et al. (2012) - Melbourne Code.

Solanum series Luteum Pojark., Flora SSSR 22: 32. 1955, as " Lutea ", not validly published; in Russian, no diagnosis or description in Latin (Art. 39.1). [only included species S. luteum Mill. = S. villosum Mill.]

Solanum aegyptiacum Forssk., Fl. Aegypt.-Arab. 46. 1775, nomen nudum (see Edmonds 2007) = S. memphiticum J.F.Gmel. and S. villosum Mill.

Solanum aegyptiacum Forssk., var. b) Fructu nigro; foliis integris villossissimus, Fl. Aegypt.-Arab. 46. 1775, polynomial, not validly published (see Edmonds 2007). = S. memphiticum J.F.Gmel.

Solanum amaranthifolium Gillies ex Rusby, Bull. Torrey Bot. Club 26: 152. 1899, nomen nudum; based on a Gillies manuscript name at Kew; two specimens by Gillies (K001166701, K001166704) are annotated "S. Solanum amaranthifolium Gill." in Gillies’ hand = S. chenopodioides Lam.

Solanum arenarium Schur, Enum. Pl. Transsilv. 477. 1866, pro syn. Solanum humile Bernh. ex Willd. = S. villosum Mill.

Solanum asperum Hornem. ex Walp., Repert. Bot. Syst. (Walpers) 3: 49. 1844, pro syn. Solanum rumphii Dunal = S. americanum Mill.

Solanum atriplicifolium Desp. ex Dunal, Solan. Syn. 12. 1816, herbarium name pro syn. Solanum nigrum L.= S. nigrum L.

Solanum atriplicifolium Desp. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, pro. syn. Solanum nigrum var atriplicifolium Dunal = S. nigrum L.

Solanum besserianum Weinm., Elench. Hort. Pawlowsk.: 100. 1824, nomen nudum, no description or diagnosis; Cult. "hort. Pawl.", said to be "S. cestrifolium Jacq.?" [no such published name found; validated by Sprengel (1825), see also Doubtful names] = S. nigrum L.

Solanum cestrifolium Gussone, Ind. Sem. Hort. Boccadifalco 11. 1825, nomen nudum = S. nigrum L.

Solanum chlorocarpum Schur, Enum. Pl. Transsilv. 478. 1866, pro. syn. Solanum nigrum L. var. chlorocarpum Koch. = S. nigrum L.

Solanum chousboe var. merrillianum (T.N.Liou) C.Y.Wu & S.C.Huang. This citation from Flora of China ( Zhang et al. 1994) that also appears on Tropicos (accessed 12 August 2017) is a misprint and confounding of the attribution to Schousboe (Schousboe ex Willd.) with a specific epithet.

Solanum cuneifolium Hort. Avens ex Dunal, Prodr. [A. P. de Candolle] 13(1): 53. 1852, pro syn. Solanum nigrum L. = S. nigrum L.

Solanum decurrens Wall. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, pro syn. Solanum rhinozerothis Blume = S. nigrum Mill.

Solanum deppei Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, pro syn. Solanum pterocaulon Dunal var. deppei Dunal = Solanum scabrum Mill.

Solanum gracile Otto ex Baxter, Hort. Brit. [Loudon] 2, Suppl.: 639 1850, nomen nudum; also later homonym of S. gracile Sendtn. (1846) = identity uncertain

Solanum heterogonum Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, pro syn. Solanum pterocaulon var. Solanum pterocaulon heterogonum Dunal = S. scabrum Mill.

Solanum judaicum Hack. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 53. 1852, pro syn. Solanum tauschii Opiz. = S. nigrum L.

Solanum kraussianum Buchinger ex Krauss, Flora 27: 831. 1844, pro syn. of polynomial Solanum nigrum L. var. fol. dentato-lobatis puberulis and not intended as a new name, refers to J.F. Drège 7864a = S. retroflexum Dunal

Solanum muricatum Bertero ex Dunal, Prodr. [A. P. de Candolle] 13(1): 150. 1852, pro syn. Solanum rancaguense Dunal = S. furcatum Dunal

Solanum nigrum L. var. aegyptiacum Lam. ex C.C.Gmel., Hort. Carlsruh. 253 1811, nomen nudum, no description or diagnosis; the epithet “aegyptiacum” was not used by Lamarck in the genus Solanum and must be an error. = identity uncertain

Solanum nigrum L. var. Solanum nigrum angustifolium Filov, Kult. Fl. SSSR (Zhukovskii) 10: 382. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. villosum Mill.

Solanum nigrum L. var. atriplicifolium Desp. ex Filov, Kult. Fl. SSSR (Zhukovskii) 10: 370-386. 1958, not validly published, no direct citation of basionym (Art. 39.1); also nom. illeg., later homonym of S. nigrum L. var. atriplicifolium G.Mey. = S. nigrum L.

Solanum nigrum L. subsp. australiense Filov, Kult. Fl. SSSR (Zhukovskii) 10: 380. 1958, not validly published; no diagnosis or description in Latin (Art 39.1) = S. nigrum L.

Solanum nigrum L. var. burbankii (Bitter) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 383. 1958, as " berbankii ", not validly published, no direct citation of basionym (Art. 38.1) = S. nigrum L.

Solanum nigrum L. subsp. chinense Filov, Kult. Fl. SSSR (Zhukovskii) 10: 382. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. americanum Mill.

Solanum nigrum L. var. chlorocarpum Koch, Syn. Fl. Germ. Helv., ed. 2: 600. 1846, not a new name, clear citation of Solanum nigrum var. chlorocarpum Spenn. ("S. nigrum chlorocarpum Spenn. fl. fri.") = S. nigrum L.

Solanum nigrum L. subsp. cultum Filov, Kult. Fl. SSSR (Zhukovskii) 10: 383. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. scabrum Mill.

Solanum nigrum L. var. frutescens Macloskie, Rep. Princeton Univ. Exped. Patagonia 8: 707. 1905, nomen nudum = S. nitidibaccatum , S. triflorum , S. chenopodioides , or S. pygmaeum that all occur in Northern Patagonia

Solanum nigrum L. forma genuinum Döll, Rhein. Flor. 412. 1843, not validly published (Art. 24.3) = S. nigrum L.

Solanum nigrum L. var. genuinum Godr., Fl. Lorraine 2: 133. 1843, not validly published (Art. 24.3) = S. nigrum L.

Solanum nigrum L. var. genuinum Hassl., Trab. Mus. Farm. Fac. Cienc. Med. Buenos Aires 21: 104. 1909, not validly published (Art. 24.3) = S. nigrum L.

Solanum nigrum L. var. griseocarpum Filov, Kult. Fl. SSSR (Zhukovskii) 10: 378. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. nigrum L.

Solanum nigrum L. var. humile F.M.Bailey, Queensland Philos. Soc. 3: 1-4. 1881, not validly published; no diagnosis or description (Art. 38.1) = S. nigrum L.

Solanum nigrum L. var. inerme K.Koch, Linnaea 17: 283. 1843, not intended as a new name (also nomen sub nudum); protologue "Solanum nigrum L. β Solanum inerme . Rami non murieulati. In Transcaucasia abundat" is much less detailed than where in the same work Koch describes new taxa, neither does he attribute the name to himself as he does for other new taxa in this paper.

Solanum nigrum L. subsp. irano-algiricum Filov, Kult. Fl. SSSR (Zhukovskii) 10: 378. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. villosum Mill.

Solanum nigrum L. var. legitimum Neilr., Fl. Nied.-Oesterr. 535. 1859, not validly published (Art. 24.3) = S. nigrum L.

Solanum nigrum L. subsp. memphiticum (Mart.) Marzell, Ill. Fl. Mitt.-Eur. [Hegi] 5(4): 2593. 1927, not intended as a new name; Marzell only says "Zur gesamtart S. nigrum gehört anscheinend auch S. memphiticum Mart." [to the total S. nigrum also belongs S. memphiticum Mart.] = S. nigrum L.

Solanum nigrum L. var. merrillianum (Liou) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 383. 1958, not validly published; no direct citation of basionym (Art. 38.1) = S. americanum Mill.

Solanum nigrum L. var. villosum (Mill.) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 380. 1958, not validly published; no direct citation of basionym (Art. 38.1) = S. villosum Mill.

Solanum nigrum L. var. violaceum Chen ex Wessely, Feddes Repert. Spec. Nov. Regni Veg. 63(3): 293. 1960, nomen nudum; not intended as a new name, listed as one of the taxa accepted by Filov (1958, Kult. Fl. SSSR 20: 382). = S. americanum Mill.

Solanum nigrum L. var. vulgatum L., Sp. Pl. ed. 2: 266. 1762, evidently an orthographic variant of S olanum Solanum nigrum L. var. Solanum nigrum vulgare L. (1753). = S. nigrum L. (see Jarvis 2007).

Solanum nigrum L. var. vulgatum Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, not a new name, referring to Solanum nigrum var. vulgatum L. (and of others “auct.”) = S. nigrum L.

Solanum nodiflorum Desv. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 46. 1852, pro syn. Solanum desvauxii Ham. = S. americanum Mill.

Solanum nodiflorum Jacq. var. acuminatum Chodat, Bull. Herb. Boissier, sér. 2, 2: 811. 1902, not intended as a new name, as " Solanum acuminatum (?)", with no specimen cited. In the rest of this work the new taxa are clearly indicated with “nob.” and a specimen (or several) cited = S. americanum Mill.

Solanum nodiflorum var. fauriei (H. Lév.) O.Deg. & O.Deg., Plants Hawaii Natl. Parks 419. 1930, no evidence in any editions of Degener’s book (1930, 1945, 1974) that this name, occurring in several indices and based on Solanum fauriei H. Lév. was ever published; probably = S. opacum A.Braun & Bouché

Solanum nodiflorum Jacq. var. sativum A.Chev. Expl. Bot. Afrique Occ. Franç. 1: 464. 1920. nomen nudum; based on "Guinea. Kankan, dans les jardins cultives, 16 Mar 1899, A.Chevalier 594 " = S. scabrum Mill.

Solanum pigmaeum Larrañaga, Escritos Damaso Antonio Larrañaga 2: 88. 1923, almost certainly not intended as a new name, an orthographic variant of Solanum pygmaeum Cav. (1806). In other parts of Larrañaga’s work (e.g. S. trilobum in vol. 1: 12), he indicates new taxa with "sp. n." or gen. n."; none of the names in vol. 2 is so designated, leading us to believe he was not coining new names, but instead using those already published. This work was compiled from his notes and diaries and these were heavily annotated (see intro in vol. 1 http://bdh-rd.bne.es/viewer.vm?id=0000016651&page=1) up to his death in 1848. = S. pygmaeum Cav.

Solanum photeinocarpum Nakam. & Odash. var. violaceum C.Y.Wu & S.C.Huang, Acta Phytotax. Sin. 16(2): 72. 1978, nomen nudum; incorrectly cited in this publication as Solanum violaceum Chen ex Wessely but Wessely (1960) did not formally publish this name, she cited Filov’s (1958) list of accepted taxa. Filov did not provide Latin descriptions or diagnoses (Art. 38.1) and so all names in that work are not validly published. The varietal epithet should be attributed only to Wu and Huang (1978), but they do not provide a Latin diagnosis either (Art. 39.1) = S. americanum Mill.

Solanum pterocaulon Rchb. ex Nyman, Consp. Fl. Eur. 3: 526. 1881. (1881), pro syn. Solanum melanocerasum Willd. = Solanum scabrum Mill.

Solanum repens Noronha, Verh. Batav. Genootsch. Kunsten 5: 84. 1791, nomen nudum = S. nigrum L.

Solanum rubrum Murray, Syst. Veg., ed. 14 (J. A. Murray) 224. 1784, not intended as a new name; Murray repeats Linnaeus’ (1767) polynomial exactly; Dunal’s (1813) citation of "Murr. Syst. veg. 183" in his description of S. erythraeum Dunal (1813: 238) is clearly an error = S. rubrum L. (nom. utique rej. prop.)

Solanum villosum Opiz ex Dunal, Prodr. [A. P. de Candolle] 13(1): 51. 1852, pro syn. Solanum decipiens Opiz. = S. nigrum L.

Solanum villosum Lam. var. alatum (Moench) Marzell, in Hegi, Ill. Fl. Mitt.-Eur. 5(4): 2594. 1927, not intended as a new name, citation of Solanum alatum Moench in text = S. villosum Mill.

Solanum villosum Lam. var. glabrescens Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 592. 1819. This name was cited in Berchtold and Opiz (1843: 24), but it is not a published name; no such name was coined in Systema Vegetabilium; on page 592, however there is a "β glabrescens" in the synonymy of S. villosum Lam., but it is a reference in the synonymy "Solanum etc. B Hall. Helv. n 576 β glabrescens" - referring to Haller’s Historia Stirpium Indigenarum Helvetiae Inchoata (1768) a work in which only polynomials were used).

Solanum viridescens Hort. ex Rchb., Fl. Germ. Excurs. 391. 1831, pro syn. Solanum humile Bernh. ex Willd. = S. villosum Mill.

Solanum virgatum Endl. ex Sendtn., Fl. Bras. (Martius) 10: 13. 1846, pro syn. Solanum gracile Sendtn. = S. chenopodioides Lam.

Figures

Figure 57. Solanum villosum A Flowering habit B Detail of adaxial leaf surface C Detail of abaxial leaf surface D Bud E Dissected flower F Fruiting habit G Fruiting habit with dense indumentum H Fruit (A-E Wood 2193; F Popov GP / 72 / 31; G Wood Y / 74 / 265). Drawing by R. Wise.

Figure 58. Solanum villosum A Habit B Inflorescence in hairy individual C Flowers and buds D Immature infructescence E Fully mature fruits F Fully mature fruits showing reflexed calyx lobes (A Nijmegen acc. 884750135; B Nijmegen acc. 914750047; C Nijmegen acc. A 34750061, D Nijmegen acc. A 34750043; E Nijmegen acc. A 14750048; F Nijmegen acc. A 34750038). Photos by T. Saerkinen and G. van der Weerden.

Figure 59. Distribution of Solanum villosum in the native range (black circles) and its non-native range (white squares) in the Old World.