Solanum scabrum Mill., Gard. Dict. ed. 8, no. 6. 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

15. Solanum scabrum Mill., Gard. Dict. ed. 8, no. 6. 1768 Figures 46 , 47

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

Solanum scabrum Type. "Solanum guineense fructu magno instar cerasi nigerrimo umbellato" cultivated in England, at James Sherard’s garden in Eltham (Hortus Elthamensis) (lectotype, designated by Edmonds 1979b, pg. 224: Dillenius, Hort. Eltham. 2: 366, t. 274, f. 354. 1732).

Solanum guineense (L.) Mill., Gard. Dict., ed. 8, no. 7. 1768, nom. illeg., not Solanum guineense L. (1753)

Solanum scabrum Type. Based on Solanum nigrum L. var. guineense L.

Solanum melanocerasum All., Auct. Syn. Meth. Stirp. Hort. Regii Taur. 664. 1774.

Solanum scabrum Type. "Solanum guineense, fructo magno, instar cerasi nigerrimo, umbellato" cultivated in England at John Sherard’s garden in Eltham (Hortus Elthamensis), Herb. Dillenius 336 (neotype, designated by Edmonds 2012, pg. 129 [as lectotype]: OXF [Dill. HE-274-234]).

Solanum guineense (L.) Lam. Tabl. Encycl. 2: 18. 1794, nom. illeg., not Solanum guineense L. (1753), Solanum guineense (L.) Mill. (1768)

Solanum scabrum Type. Based on Solanum nigrum L. var. guineense L.

Solanum triangulare Lam., Tabl. Encycl. 2: 18. 1794.

Solanum scabrum Type. "Ex Ind. Orient", Herb. Lamarck s.n. (lectotype, designated by D’Arcy 1974a, pg 735 [as type]: P-LAM [P00357626]).

Solanum quadrangulare Lam. var. triangulare (Lam) Pers., Sys. 225. 1805.

Solanum scabrum Type. Based on Solanum triangulare Lam.

Solanum melanocerasum Willd., Enum. Pl. (Willdenow) 1: 237. 1809, nom. illeg., not Solanum melanocerasum All. (1773)

Solanum scabrum Type. Cultivated in Berlin Botanical Garden, from "Europa australi" [southern Europe] (no original material labelled as “melanocerasum” found in B-W; neotype, designated here: B-W [BW04368010]).

Solanum pterocaulum Dunal, Hist. Nat. Solanum 153. 1813, nom. illeg. superfl.

Solanum scabrum Type. Based on Solanum scabrum Mill. (cited in synonymy).

Solanum fistulosum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 749. 1814.

Solanum scabrum Type. "Originaire de l’Isle de France [Mauritius], est cultivée en Amerique [Brazil]", Herb. Richard s.n. (lectotype, designated by D’Arcy 1974a, pg. 735: P [P00335259]).

Solanum memphiticum Mart., Pl. Hort. Erlang. 63. 1814, nom. illeg., not Solanum memphiticum J.F.Gmel. (1791)

Solanum scabrum Type. Cultivated in Erlangen, Bavaria, Germany, origins not known (no specimens cited; no original material located, possibly at ER?).

Solanum nigrum L. var. melanocerasum (Willd.) Dunal, Solan. Syn. 12. 1816.

Solanum scabrum Type. Based on Solanum melanocerasum Willd.

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

Solanum scabrum Type. Czech Republic. Prague, "in Semubackern por dem Reuthore Prag", P.M. Opiz 10/840 (no herbaria cited; no original material found, perhaps at PR?).

Solanum oleraceum Dunal var. macrocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852.

Solanum scabrum Type. Brazil. Bahia: Ilheus, 1841, C.F.P. Martius 1255 (lectotype, designated by Edmonds 1972, pg. 108 [as holotype]: G-DC [G00144295]; isolectotype: P [P00366815]).

Solanum tinctorium Welw., Apont. 590. 1859 [1858].

Solanum scabrum Type. Angola. Golungo Alto, 1856, F.M.J. Welwitsch 6103 (lectotype, designated here: BM [BM000942995]; isolectotypes: BM [BM000942996], K [K001029777]).

Solanum nigrum L. var. pterocaulum (Dunal) Schur, Enum. Pl. Transsilv. 478. 1866, as ' Solanum pterocaulon '.

Solanum scabrum Type. Based on S. pterocaulum Dunal ["excl. German floras"]

Solanum boerhaavii Thell., Rep. Bot. Soc.& Exchange Club Brit. Isles 8: 187. 1927, as " Boerhaavii ".

Solanum scabrum Type. Based on Solanum nigrum L. var. guineense L. [as replacement name for Solanum guineense (L.) Mill.]

Solanum nigrum L. var. pterocaulum (Dunal) Domin, Biblioth. Bot. 89: 1127. 1928.

Solanum scabrum Type. Based on S. pterocaulum Dunal

Solanum intrusum Soria, Baileya 7: 33. 1959.

Solanum scabrum Type. Based on (replacement name for) Solanum nigrum L. var. guineense L., and Solanum guineense (L.) Lam.

Solanum scabrum Mill. subsp. laevis Olet, Novon 16(4): 510. 2006.

Solanum scabrum Type. Uganda. Buganda: Kampala district, Kawempe div., Kawempe North, Kalerwe, Tula road, 1220 m, 14 Feb 2001, E.A. Olet 88 (holotype: MHU; isotypes: H, K, MO [ex descr.]).

Type.

Cultivated in Chelsea Physic Garden , said in protologue to "grow naturally in North America", Herb. Miller s.n. (lectotype, designated by Henderson 1974, pg. 61 [as type]: BM [BM000847083]) .

Description.

Annual or short-lived erect or ascending perennial herbs to 1.5 m tall, often woody at the base. Stems spreading, terete, ridged or winged, green to purple, if ridged or winged the stems later spinescent, older stems with or without prominent pseudospines, usually somewhat hollow; new growth puberulent with simple, spreading, uniseriate, translucent, eglandular trichomes, these 2-8-celled, 0.3-0.8 mm long; older stems glabrescent. Sympodial units difoliate, the leaves usually not geminate, but if leaves paired, then one is usually smaller. Leaves simple, 4-15(-20) cm long, 3-10(-16) cm wide, broadly ovate to elliptic, very variable in size depending on cultivars and growth conditions, membranous to somewhat fleshy, green to dark green to somewhat purple coloured above, slightly paler below, without smell; adaxial and abaxial surfaces glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem mainly along veins and scattered along lamina; major veins 3-6(-8) pairs, paler green or often purple tinged; base abruptly acute or truncate, narrowly winged on to the petiole; margins entire or rarely shallowly sinuate; apex rounded to acute; petioles 1-5(-8) cm long, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the stem. Inflorescences 1-2(-4) cm long, internodal, simple, furcate or many times branched (in cultivars), sub-umbelliform, with 4-10(-30+) flowers clustered towards the tip(s) of the rhachis, glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem; peduncle 1-5(-8) cm long, erect and thick, much thickened at apex, subwoody; pedicels 0.4-1 cm long, 0.3-0.5 mm in diameter at the base, 0.75-0.9 mm in diameter at the apex, abruptly expanding to the calyx tube, stout, erect and/or spreading, articulated at the base; pedicel scars tightly clustered near the tip of the rhachis, spaced 0-2 mm apart, sometimes with short stumps ca. 0.5-1.0 mm long. Buds globose to subglobose, the corolla exserted 1/2-1/3 from the calyx tube before anthesis. Flowers 5-merous or occasionally fasciated and 6-7-merous in cultivars, all perfect. Calyx tube 0.9-1.1 mm long, abruptly cup-shaped with a broad base, the lobes slightly unequal, 0.9-1.5 mm long, 0.8-1.4 mm wide, broadly deltate with a rounded tip, green or purple-tinged, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the pedicels, the margins often drying scarious and white. Corolla 7-12 mm in diameter, white, purple-tinged or occasionally lilac to dark purple, with a yellow basal star, stellate, lobed ca. 1/2 way to the base, the lobes 2.5-4 mm long, 1.5-3 mm wide, spreading or reflexed, densely papillate on tips and margins. Stamens equal; filament tube very short, to 0.1 mm long; free portion of the filaments 0.5-0.8 mm long, glabrous or pubescent with tangled uniseriate simple trichomes; anthers 2-3 mm long, ellipsoid or slightly tapering towards the tips, yellow, orange or brown, poricidal at the tips, the pores lengthening to slits with age and drying, the connective often becoming brownish-black in dry specimens. Ovary rounded, glabrous; style 2.5-5 mm long, densely pubescent with simple uniseriate trichomes 0.2-0.5 mm long in the basal 1/2 where included in the anther cone, exserted beyond anthers 0-1.5 mm; stigma capitate, the surface minutely papillate. Fruit a globose to depressed-globose berry, 10-20 mm in diameter, purplish-black at maturity, the pericarp thick, shiny or somewhat matte, not transparent; fruiting pedicels 0.7-1.5(-2.0) cm long, 0.5-1 mm in diameter at the base, 1.1-1.5 mm in diameter at the apex, stout, erect and spreading, purple or brown, usually not falling with the fruit, remaining on the plant and often persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.5-2 mm long, usually tearing unevenly, the lobes 2-3 mm long, usually with thicker white margins in dry material, appressed or spreading to slightly reflexed. Seeds (20-)100-150 per berry, 2-2.8 mm long, 1.5-1.8 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow-brown or purple, the surfaces minutely pitted, thin and the embryo clearly visible, the testal cells rectangular to pentagonal in outline. Stone cells absent. Chromosome number: 2n =6x=72 ( Soria and Heiser 1961 [as S. melanocerasum ]; Heiser et al. 1965 [as S. melanocerasum ]; Henderson 1974; Edmonds 1977, 1983; Symon 1981; Jacoby and Labuschagne 2006; Bukenya 1996; Olet et al. 2015).

Distribution

(Figure 48 ). Native to tropical Africa, but introduced worldwide as a cultivated plant.

Ecology.

Grows in open areas, in a wide variety of habitats, in wet forests or drier areas, along roads and at field edges; often cultivated; between sea level and 2,300 m elevation.

Common names.

Benin: odu, ogomo, feibii ( Essou and Hermans 2006); Cameroon: houlohada, legume vert, njebanyoon, ossan, tindar noon, wikitiniho, zom/zoum; China: mu long kui ( Zhang et al. 1994); Côte d’Ivoire: foue; Equatorial Guinea: hierba mora, nahú, seba [Bubi people], sisa; Ghana: nsusua; Madagascar: anamamy, brede; Nigeria: awa ibo, gautan kaji, kumbi, odu, ogunmo, ugumakbe; São Tome e Principe: losua; Sierra Leone: a-kempa,borisuguli, filami, kemba, kembei, kholekoleden-na, magboli, reinpa, timbainyi; Slovenia: čučoried kovitý ( Bertová and Goliašová 1993); South Africa: nastagal; South Sudan. nzuobire; Tanzania: isonga (Kinyakyusa language); Uganda: eshwiga, eshwiga enzungu, kujikuji, nswiga ya kizungu; United Kingdom: garden huckleberry ( Stace 2010).

Uses.

Leaves used as greens (spinach) and cooked, sold in markets; berries used as ink; in Benin, the powdered leaves are used to cure dysentery ( Essou and Hermans 2006).

Preliminary conservation status

( IUCN 2016). Solanum scabrum is widespread across Africa and an important leafy vegetable in cultivation; it can be assigned a preliminary status of LC (Least Concern; Table 7 ), but it may become important to conserve local populations in order to preserve genetic variation for plant breeding.

Discussion.

Solanum scabrum is the most commonly cultivated and widespread of the African black nightshades. It is the most important indigenous leafy vegetable in the black nightshade group and is commonly known as the African nightshade ( Edmonds and Chweya 1997; Dinssa et al. 2016). Solanum scabrum shows great variation in growth form, leaf shape and size and berry number, in part due to the significant local variation introduced by human selection in cultivated populations.

The species has also been introduced to Europe, North America, Australia and New Zealand. In the United States of America, S. scabrum is known as "Garden Huckleberry" and its identity, origin and suitability to human consumption were the subject of great interest in the 1960s (as S. guineense , e.g. Soria and Heiser 1959, 1961; Heiser 1969).

Different cultivars are recognised, the late flowering plants cultivated for their larger abundance of leaves with smaller number of fruits and earlier flowering plants that have larger inflorescences which are cultivated for their fruits ( Manoko 2007). Cultivated forms of S. scabrum stand out as quite distinctive; all forms have larger flowers and anthers that are often brownish in colour, while the forms cultivated for their leaves have larger, longer petiolate leaves and those cultivated for their fruits have larger, more numerous and shinier berries (the size of cherries) on erect or spreading pedicels. The pericarp in S. scabrum berries in all varieties is quite thick compared to other black nightshades, but mature berries of cultivars show variation in anthocyanin content where some individuals have dark purple mesocarp and others pale green. One of the major limitations to cultivation of S. scabrum as a leaf vegetable is the relatively low leaf yield due to early flowering and excessive fruiting ( Ojiewo et al. 2013; Ojiewo et al. 2006). In response to this, enhanced varieties are being developed and new registered varieties are being released ( Ojiewo et al. 2006; Ojiewo et al. 2013) such as "Medium leaf long-lasting" released in Kenya in 2006. Breeding is also focussing on local variation and drought resistance ( Dinssa et al. 2016).

The presumed wild forms were described at the infraspecific rank by Olet et al. (2006) as subsp. Solanum scabrum laevis . This differs from the cultivated forms in having narrower leaves and smaller fruits that are globose rather than subglobose. Dehmer (2001) and Olet et al. (2011) have shown, however, that these differences are not reflected in relationships based on AFLP molecular markers. These wild forms of S. scabrum have often been called S. nigrum (e.g. Edmonds 2006a, 2006b, 2012), but they differ from true S. nigrum which, in Africa only occurs in the north along the Mediterranean, in their congested inflorescences, spreading pedicels, calyx lobes that tear unevenly and long-petiolate leaves. Olet et al. (2011) clearly demonstrated that the wild forms of S. scabrum are genetically closely related with cultivated S. scabrum accessions and form a distinct cluster away from European S. nigrum .

Solanum scabrum can be distinguished from the somewhat similar S. americanum by its larger anthers (2-3 mm long versus 0.8-1.5 mm long). In both these species, as well as S. retroflexum , the berries usually lack stone cells (some populations of S. americanum can have up to 4 stone cells) and drop without the pedicels at maturity, leaving the pedicels behind on old inflorescences. In both S. scabrum and S. americanum , berries are purple-black and shiny, while S. retroflexum has dull purple-black berries with a distinct grey bloom. The pedicels of S. scabrum are usually erect or spreading.

Plants described as S. fistulosum and S. oleraceum var. macrocarpon by Dunal (1814, 1852 respectively, see synonymy) from Brazil were almost certainly taken there from western Africa by enslaved people and were either collected from home gardens or became established in the New world.

Solanum scabrum is closely related to S. nigrum and S. villosum based on molecular data ( Manoko 2007; Poczai and Hyvönen 2011). The close relationship amongst these polyploids indicates that they might share the same diploid parent. The idea that S. villosum is the likely tetraploid parent of the hexaploids S. nigrum and S. scabrum has also been supported by evidence from crossing experiments and cytological studies ( Soria and Heiser 1961; Heiser et al. 1965; Rao et al. 1971; Jardine and Edmonds 1974; Edmonds 1977, 1978; Edmonds and Glidewell 1977; Edmonds 1978). Two possibilities still remain: one of the hexaploid species ( S. nigrum and S. scabrum ) evolved first and gave rise to the other or that the two hexaploids originated from the same parents independently ( Poczai and Hyvönen 2011). These two scenarios will be difficult to distinguish based on molecular data due to the complexity caused by high ploidy level.

The nomenclature and typification of the various synonyms of S. scabrum and the earliest name applied to the taxon, have been extensively discussed by others (e.g. Gras 1863; Thellung 1927; Polgár 1939; Stebbins and Paddock 1949; Soria and Heiser 1959; Heine 1960; Edmonds 1979b). We merely add additional notes for our lectotypifications designated here or for those that have been designated inadvertently (e.g. Prado et al. 2015).

Edmonds (2012) designated a specimen in the Dillenian herbarium at OXF as the lectotype for S. melanocerasum All.; Allioni did not cite and probably never saw these specimens; this is correctable to neotype.

D’Arcy (1974a: 737) inadvertently lectotypifed S. guineense Lam. by citing a specimen in "Herb. Lam. s.n. (P)" as “Type”; this corresponds to a specimen in the Lamarck herbarium (P00357674) that is labelled "Solanum nigrum guineense L. planta glabra.. fructu magno nigro. Sol. guineense Lam. Ill", suggesting that Lamarck was basing his name on Linnaeus’ S. nigrum var. guineense . In the protologue, Lamarck (1794) indirectly referred to Linnaeus, through citing the single element in the protologue of S. nigrum var. guineense , the illustration in Dillenius (1732). We are therefore treating these two names ( S. nigrum var. guineense L. and S. guineense Lam.) as homotypic, rendering D’Arcy’s (1974a) lectotypification superfluous.

Lamarck’s (1794) S. triangulare has traditionally been considered a synonym of S. africanum Mill. (a member of the Dulcamaroid clade, Knapp 2013); the protologue cites two elements, only one of which “?” represents S. africanum and was questioned as only possibly being the same by Lamarck. The other cited element of S. triangulare is an illustration from Herbarium Amboinense ( Rumphius 1750) that represents S. americanum . D’Arcy (1974a), however, cited the specimen in Lamarck’s herbarium ("Lamarck s.n. (P)"; P00357626) as “Type” and we must accept this as a valid lectotypification. This specimen is of a narrow-leaved plant of S. scabrum.

Willdenow (1809) did not specifically reference Allioni’s epithet in his protologue, so we are treating it as a new name. There is no material in the Willdenow herbarium labelled as S. melanocerasum , but a specimen (B-W04368010) of S. scabrum (labelled as "S. guineense" and "ex horto proprio W") was clearly grown in Berlin and seen by Willdenow. We designate this as the neotype for S. melanocerasum Willd.

Solanum pterocaulum is illegitimate because Dunal (1813) cited S. scabrum in synonymy. In the Prodromus ( Dunal 1852), his use of ' Solanum pterocaulon ' was simply an orthographic variant and is correctable (see McNeill et al.. 2012, Art. 61.5 and associated provisions in Art. 61); it does not alter the legitimacy of the name.

No original material has been traced for S. memphiticum Mart., but the emphasis placed on the dark purplish colour of the stems and leaves by Martius (1814) in the protologue, coupled with the petiolate nature of the leaves and the erect peduncles, suggests this plant was S. scabrum.

Of the several species in this group described by P.M. Opiz (see discussion under S. nigrum ), we are convinced from the description that S. nitens is a synonym of S. scabrum , rather than of S. nigrum , although Opiz (1843) places it in his “superspecies” S. nigrum . The mention of shiny berries on erect pedicels is distinctive; we have yet to confirm this with specimens from Opiz’s herbarium at PR.

The lectotype (G00144295) chosen for S. oleraceum var. macrocarpum is the better preserved of the two duplicates of Martius 1225 cited by Dunal (1852).

The lectotype chosen for S. tinctorium (Welwitsch 6103) is the more clearly duplicated of the collections cited in the protologue; we have selected the best preserved sheet (BM000942995) as the lectotype. The name apparently comes from the staining berries that were used as ink.

Selected specimens examined.

Angola. Bengo: Ambriz, Abriz , Dec 1872, Monteiro s.n. (K) ; Benguela: Hochland zwischen Ganda und Caconda, Dec 1933, Hundt 797 (BM) ; Cuanza Norte: Varzea do Isidoro ad rivum Cuango, Dto. Golungo Alto , Jul 1855, Welwitsch 6100 (BM) ; Malanje: Distr. Pungo Andongo, prope Lusillo , Jan 1857, Welwitsch 6108 (BM) ; Namibe: Mofsamedes, Herb. Moira, Habit Cavalheiros , Jul 1859, Welwitsch 6033 (BM) .

Benin. Atlantique: Wida, Dahomey , 26 Aug 1903, Estève 111 (BM) .

Botswana. Ghanzi: Ghanzi camp, 4 May 1969, Brown 6019 (K); Ghanzi camp, 4 May 1969, Cole 6019 (K) ; North West: Mutsoi NE of Nokaneng, 21 Mar 1967, Lambrecht 90 (K) .

Burkina Faso. Seno: Dori Dam , 16 Oct 2007, Sanou & Leonard BUR-596 (K) .

Cameroon. Adamaoua: Mayo-Banyo, Mambilla Plateau Cameroon, around Somie village , 21 Jun 2009, Komaromi 48 (K) ; Centre: Mimboman area, Yaounde , 16 Sep 1986, Manning 235 (MO) ; Extreme-Nord: Monts Mandara, Hossere Oupay , 15 km NNO de Mokolo, 15 Sep 1964, Letouzey 6880 (K, P) ; Littoral: Donala , 16 Aug 1986, Johns 86-481 (K) ; Nord-Ouest: Mezam, Above Bamenda , 20 Jan 1928, Migeod 358 (BM, K) ; Sud-Ouest: Mt Kupe, Kupe Village , 24 May 1996, Ryan 289 (K, MO, P) .

Central African Republic. Mboukou Griko, Territoire du Haut-Obangi, 24 Sep 1902, Chevalier 5518 (K) .

Comoros. Anjouan: Anjouan (Comoro-Insel Johanna), Jun 1875, Hildebrandt 1626[b] (BM, W) ; Moheli Island: Moheli, NE center of island, 14 Aug 1987, D’Arcy 17617 (MO, P); 1 Nov 1990, D’Arcy 17765 (MO); Njazídja: Grande Comore, 9 Aug 1981, Doutrelepont 1203 (MO, P) .

Côte d’Ivoire. Abidjan: Adjame , 31 May 1973, De Koning 1747 (MO) ; Dimbokro: c. 5 km W of Dimbokro , 13 Oct 1975, van der Burg 1177 (MO); M’Bahiakro: Koffi-Akakro (s. Prikro), 25 Jul 1973, Smittenberg-Visser 61 (MO) ; Montagnes: Man, 6 km N nr Yebegouin , 27 Jan 1984, Hepper & Maley 7849 (K) .

Democratic Republic of the Congo. Katanga: Lukonzolwa, Moero, 1933, Quarré 3297 (K); Kongolo, 5 Feb 1920, Schantz 646 (K) ; Kinshasa: Maluku, Route Menkao-Kingankati, 5 Nov 1971, Breyne 2227 (MO) ; Nord-Kivu: Lubarika, 1953, Gilon 315[a] (MO); Orientale: Ituri, w. v. Albert-See, Oct 1934, Gusinde s.n. (W); Yangambi, 26 Jul 1938, Louis 10507 (K, P); Sud Kivu: Kizozi, Jul 1933, Lejeune 55 (K) .

Egypt. Giza: Faculty of Agriculture, Giza , 13 Jun 1971, Sisi s.n. (MO) .

Equatorial Guinea. Annobon: Pico de Fogo, SW side, 25 Jul 1959, Melville 188 (BM, K, MA, P) ; Bioko: entre Moca y Riaba por el camino viejo, 20 Feb 1989, Fernández-Casas 11820 (K, MA, MO, P) ; Bioko Norte: Pico Basilé, Carretera del pico Basile, 28 Mar 1990, Do Carvalho 4305 (K, MA, MO, P) ; Bioko Sur: Moca , camino de Ureca, 18 Feb 1989, Fernández-Casas 11725 (K, MA, MO) .

Eritrea. Maekel: Asmara , 1 May 1892, Terracciano & Pappi 187 [2206] (FT) ; Semienawi Keyih Bahri: Asmara, Beless, Hamasen , 4 May 1892, Terracciano & Pappi 2536 (FT) .

Gabon. Estuaire: Jardin Cenarest, Libreville, Poubelle , 16 Jan 1986, Louis 1990 (MO) .

Ghana. Ashanti: nr Mampong, Ashanti, 7 Aug 1963, Darko 5115 (K) ; Central: Aswansi, W.P, 12 Oct 1954, Darko 1032 (K) ; Eastern: Akosombo, 25 May 1970, Enti 1723 (MO); Akwapim, Mampong Scarp, 14 Jun 1953, Morton s.n. (K) ; Greater Accra: Accra, May 1961, Irvine 5095 (K) ; Volta: Adzido, Keta, 16 Sep 1960, Akpabla 2117 (K) .

Guinea. Nzérékoré: Lola, Mt. Nimba , 1 Nov 2012, Diabate & Mas 1419 (MO) .

Indonesia. Java: Central Java, Mt. Slamet, 15 Mar 2004, Hoover et al. 89 (A); Seram: Manusela National Park , 12 Sep 1987, Argent C 87-179 (A, E) ; Sumatra: E of Berastagi, Karo Highlands , 4 Jun 1928, Si Toroes 407 (A, GH) .

Lesotho. Sehlabathebe , 4 Jan 1973, Bayliss 5476 (MO) .

Liberia. Central: 3 mi NE of Suacoco, Gbarnga , 7 Feb 1951, Daniel 117 (B, BM, MO) ; Grand Gedeh: Tchien, Mim Timber Co ( Fijnhout ), 16 May 1970, de Koning 519 (MO) ; Lofa: mi along rd to Wologesi , 15 Jul 1970, Jansen 2015 (K, MO, P) ; Nimba: Nimba Mountains , 27 Jul 1962, Leeuwenberg & Voorhoeve 4668 (B, K, MO, P) .

Madagascar. Antananarivo: Grande Terre-Ouangani, Apendzo-Jivany. Barakani , 7 May 2002, Barthelat et al. 885 (K, MO, P) ; Antsiranana: Mt. d’Ambre, partie centrale, Prov. Diego-Suarez , 11 Nov 2007, Gautier 5198 (K, MO) ; Fianarantsoa: Ankafana, 1880, Cowan s.n. (BM); 10 km W of Ivato on Route 35, 25 Jan 1975, Croat 29606 (MO) ; Toamasina: au km 26 de la route de Tamatave, 18 Oct 1951, Benoist 147 (P) ; Toliara: Beroroha, Vallée du Mangoky et de l’ Isahaina aux environs de Beroroha, Oct 1933, Humbert 11309 (K, MO, P); Andohahela RNI, Mt. Trafonaomby , Taolagnaro, 7 Apr 1994, Randriamampionona 692 (MO, P) .

Malawi. Central: Salima Distr., Liganga village A. Mpemba, 14 Jun 1985, Kwatha et al. 210 (MO); Salima Distr., Luwadzi stream, 14 Jun 1985, Salubeni et al. 4242 (MO) ; Northern: Nkhata Bay, Musalowa village, Chizumulu Island , 25 Mar 1989, Balaka & Patel 2025 (K, MO) ; Southern: Malawe Hill, W of Port Herald , 23 Mar 1960, Phipps 2653 (K, MO) .

Mauritania. Trarza: Rosso Administrative area, Village Rosso , 9 Oct 1962, Adam 18728 (MO) .

Mozambique. Baroma: N’kanya, N of Zambesi River , 25 Jul 1950, Chase 2857 (BM) ; Cabo Delgado: Pemba , Jun 1909, Rogers 8256 (K) ; Manica: Lower slopes of Chimanimani Mountains , Apr 1967, Westwater 192954 (K) .

Namibia. Onjossariviers , im Ufergestrupp, 21 Jun 1957, Seydel 1162 (A, K) .

New Zealand. North Island: Bay of Plenty, Ohope, Whakatane Ecological Region, Taneatua Ecological District , 13 Mar 1979, Grant s.n. (AK) .

Nigeria. Delta: [ Ogwashi-Ukwu ], 25 Nov 1912, Thomas 2042 (K) ; Edo: Nikrowa Forest Reserve, Mid-west State, Iyekovia Distr. , 8 Oct 1973, Daramola FHI-72483 (K, MO) ; Enugu: 10 mi E of Erugu , Mar 1948, Irvine 3607 (K) ; Gongola: Gemu Distr., Mambilla plateau, 18 Aug 1977, Fagbemi 438 (MO) ; Jos: Naraguta , Lely 31 (K) ; Kaduna: Zaria , 1975, Magaji MG-725 (K) ; Kano: Kano, Wudil Distr , 50 km SE of the city of Kano, 16 Mar 1988, Etkin 63[b] (MO) ; Kogi: Odu, SW Nigeria, Van Eyenhuisen 7 (K) ; Lagos: Lagos, Dalziel 1188a (K); Lagos, Dalziel 1188b (K) ; Niger: Nupe , Barter 1054 (K, P) ; Ondo: between Ikare and Oke-Agbe, Ikare District , 2 May 1979, Daramola & Osanyinlusi 154 (K) ; Oyo: Ibadan , 7 km W of Polytechnic, 11 Jun 1977, Pilz 2108 (K, MO) ; Taraba: State of Gongola, Distr. Mambilla Plateau, Nguroje township , 23 May 1982, Odewo 130 (MO) .

São Tome e Principe. Bom Sucesso , 27 Jan 1949, Espírito Santo 218 (BM); Vanhulst ( Macambrará), 28 Oct 1932, Exell 89 (BM) .

Senegal. In paludilb[us] N’Boro nec non ins[ula] Bonavista, 1838, Brunner 108 (BM, G, W); Sin. loc., 1828, Perrottet 555 (BM, W); Perrottet 556 (BM);

Seychelles. Morne Blanc , 2 Oct 1970, Schlieben 11670 (B, K) .

Sierra Leone. Eastern: Kailahun, Musaia, 19 Dec 1946, Deighton 4571 (K); Northern: Port Loko, Rokupr, Magbema, 18 Apr 1959, Jordan, 1059 (K); Yonibana, 12 Nov 1914, Thomas 4909 (W); Western: Freetown, 28 Apr 1965, Morton s.n. (K).

South Africa. Eastern Cape: Grahamstown , 7 Sep 1970, Bayliss 4594 (A) ; KwaZulu-Natal: 2632 CD Bella Vista grid, between Ndumu Store and the Game Reserve , 30 Oct 1969, Moll 4138 (A, K) ; Limpopo: Mopani, Shiluvane , Aug 1899, Junod 575 (K) ; North West: Okawango Delta, Okavango Delta, Delta camp, 16 Jun 1994, Cole 923 (K) ; Northern Cape: Augrabies Falls National Park, Orange river bank, 14 Mar 1978, Balsinhas & Harding 3290 (K, MO) ; Western Cape: Dowweklip, Voelklip, Hermanus , 6 Jun 1980, Williams 287 (K, MO); Skeleton Ravine , 3 Oct 1897, Wolley-Dod 3180 (K) .

South Sudan. Bahr El Ghazal: Anglo-Egyptian Sudan, Bahr El Ghazal Prov. , Ibba, 10 Mar 1934, Dandy 624 (BM, EA) ; Equatoria: R. Napere, 25 Nov 1937, Wyld 347 (BM) .

Tanzania. Dodoma: Mpwapwa, Mbuga Village, Kibakwe Division , Mbuga Ward , 31 May 2005, Kindeketa et al. 2549 (EA, MO) ; Iringa: Ruaha National Park, FTEA region T7, top of Mpululu mountain , 21 May 1968, Renvoize & Abdallah 2312 (EA, K) ; Kagera: Bukoba , 30 Oct 1992, Breteler 11599 (MO) ; Mbeya: Rungwe, Ngumbulu Village, NE part of Rungwe Forest Reserve , 14 Mar 2008, Abeid et al. 2848 (MO) ; Morogoro: Tanganyika, Ulugurus , Jan 1935, Bruce 524 (BM, K) ; Shinyanga: Shinyanga , Nov 1938, Koritschoner 2191 (EA, K) .

Togo. Lomé, 22 Sep 1976, Ern et al. 883 (B); slopes of Bauman Peak, 14 Aug 1962, Morton A 4277 (MO) .

Uganda. Central: Kyadondo Mengo (U4); Nr Kanyanya , 16 Jun 1990, Rwaburindore 2992 (MO) ; Mengo: Distr. W. Mengo , 4 mi Gayaza rd, 5 Jul 1980, Rwaburindore 701 (MO) ; Northern: Yumbe, 26 Nov 1941, Thomas 4070 (EA, K) ; Western: Kigezi DFI, 28 Aug 1972, Goode G 3-72 (K); Kigezi, Bugangari , Rhuzumbura , Kigezi , Feb 1949, Purseglove 2712 (EA, K) .

United Kingdom. England: Hertfordshire, Rye Meads Sewage Works, nr Rye House , 1 Oct 1996, Hanson s.n. (K) .

Zimbabwe. Mashonaland Central: Imayanga, Nyamaropa TTk, 16 Jan 1967, Biegel 1767 (MO) ; Matabeleland North: Binga Distr. , Sanyam R. and Zambezi R. confluence, Sep 1955, Davies 1514 (MO) ; Matabeleland North: Binga, Chizarira Game Reserve, Busi River, 10 Nov 1971, Thomson 481 (K); Wankie: Wankie Distr. , Victoria Falls , Elephant Hills Hotel, 19 Dec 1978, Mshasha 144 (MO) .

Figures

Figure 46. Solanum scabrum A Habit of wild form B Flower of wild form C Infructescence of wild form D Habit of cultivated form E Inflorescence of cultivated form F Fruit of cultivated form G Seed (A-C Pilz 2108; D-G Nee 16088). Scale bar: 4 cm (A, D), 3.3 mm (B), 1.5 cm (C, F), 7 mm (E) and 2 mm (G). Drawing by L. Smith.

Figure 47. Solanum scabrum A Common habit B Habit in taller varieties C Flowers of the larger berried variety at full anthesis D Fruits of a larger berried variety E Flowers of the smaller berried variety at full anthesis F Fruits of a smaller berried variety (A Nijmegen acc. BG 13; B Nijmegen acc. A 34750072; C Nijmegen acc. GB 22; D Nijmegen acc. H 065; E Nijmegen acc. A 34750067; F Nijmegen acc. 2010 / 3). Photos by S. Knapp.

Figure 48. Distribution of Solanum scabrum in its native range.