Solanum memphiticum J.F.Gmel., Syst. Nat. ed. 13[bis] 2(1): 385. 1791

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

6. Solanum memphiticum J.F.Gmel., Syst. Nat. ed. 13[bis] 2(1): 385. 1791 Figures 19 , 20

Solanum nigrum L. var. hirsutum Vahl, Symb. 2: 40. 1791.

Solanum nigrum hirsutum Type. Based on Forsskål’s "S. aegyptiacum b) Fructu nigro; foliis integris villosissimus" (= Solanum memphiticum J.F.Gmel.)

Solanum hirsutum (Vahl) Dunal, Hist. Nat. Solanum 158. 1813.

Solanum nigrum hirsutum Type. Based on Solanum nigrum L. var. hirsutum Vahl

Solanum grossidentatum A.Rich., Tent. Fl. Abyss. 2: 101. 1850, as “grossedentatum”.

Solanum nigrum hirsutum Type. Ethiopia. “Tchélikote” [Chelicote], R. Quartin-Dillon & A. Petit s.n. (lectotype, designated by Lester 1997, pg. 285: P [P00344046]; isolectotypes: GOET, P [P00344047], Z; possible isolectotype: K [K001156454]).

Solanum nigrum L. var. rigidum Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852.

Solanum nigrum hirsutum Type. Yemen. Sin. loc., 1837, P.E. Botta s.n. (lectotype, designated by D’Arcy 1974a, pg. 738: P [P00055305]).

Solanum hirsutum (Vahl) Dunal var. abyssinicum Dunal, Prodr. [A. P. de Candolle] 13(1): 58. 1852.

Solanum nigrum hirsutum Type. Ethiopia. Tigray: Adigrat, prope Adoam [Adoa] "nomen Abyssinicum: Alam tch’aguar”, G.H.W. Schimper 46 (lectotype, designated by Edmonds 2012, pg. 143: G [G00015053]; isolectotypes: BM [BM000942983], BR [BR0000008422200], GH [GH00139634], HBG [HBG511468], K [K000922192], L [L0700161], LE [LE00016944, LE00016943], LG [LG0000090028793], M [M-0105602], P [P00344000, P00343998, P00343999, P00344001, P00344002], TUB [TUB003999], W [1889-0283797]).

Solanum pruinosum Dunal var. pilosulum Dunal, Prodr. [A. P. de Candolle] 13(1): 59. 1852.

Solanum nigrum hirsutum Type. Sudan. Blue Nile: Sennaar, 1831, G. Acerbi s.n. (holotype: G-DC [G00144592]).

Solanum subuniflorum Bitter, Repert. Spec. Nov. Regni Veg. 10: 546. 1912.

Solanum nigrum hirsutum Type. Tanzania. Marangu near Mount Kilimanjaro, G. Volkens 2108 (neotype, designated by Edmonds 2007, pg. 665: BR [BR0000008800091]).

Solanum plebeium A.Rich. var. grossidentatum (A.Rich.) Chiov., N. Giourn. Bot. Ital. 26: 159. 1919, as “grossedentatum”.

Solanum memphiticum Type. Based on Solanum grossidentatum A.Rich.

Solanum nigrum L. var. grossidentatum (A.Rich.) De Wild., Pl. Bequaert. 1: 431. 1922, as “grossedentatum”.

Solanum nigrum hirsutum Type. Based on Solanum grossidentatum A.Rich.

Solanum memphiticum J.F.Gmel. var. abyssinicum (Dunal) Cufod., Bull. Jard. Bot. État Bruxelles 33(3): 872. 1963.

Solanum memphiticum Type. Based on Solanum hirsutum (Vahl) Dunal var. abyssinicum Dunal

Type.

Locality unknown [ Yemen or Saudi Arabia, but most likely Yemen], Herb. Forsskål 421 (lectotype, designated by Edmonds 2007, pg. 665: C [C10013456]).

Description.

Annual or short-lived sprawling perennial herbs to 1.5 m tall, woody and branching at base. Stems spreading to decumbent, terete or occasionally very slightly angled, green, older stems green or straw colour, not markedly hollow; new growth densely viscid-pubescent with simple, spreading, uniseriate, mixed glandular and eglandular trichomes, these 3-10-celled, 0.5-2 mm long, with a terminal single-celled gland if glandular; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, (1.5-)2-9 cm long, (0.8-)1.2-5.5 cm wide, elliptic to ovate and widest in the basal third, membranous, concolorous, foul-smelling when crushed; adaxial surfaces moderately viscid-pubescent with a mixture of glandular and eglandular simple uniseriate trichomes 0.5-2 mm long like those of the stems, these denser along the veins; abaxial surfaces densely viscid-pubescent with similar glandular and eglandular simple uniseriate trichomes, these evenly distributed on veins and lamina; base acute, then attenuate and decurrent on to the petiole; margins entire or more often irregularly toothed, the teeth 2-4 mm long, acute; apex acute to acuminate, the tip often blunt and usually somewhat rounded; petioles 0.5-1.5 cm long, winged from the decurrent leaf base. Inflorescences 1-2.5(-3) cm long, internodal, simple, umbelliform to sub-umbelliform, with (2-)3-5(-8) flowers clustered at the tip, densely viscid-pubescent with mixed glandular and eglandular simple uniseriate trichomes like those of the stems; peduncle 0.9-2(-2.3) cm long, straight; pedicels 7-9 mm long, ca. 0.5 mm in diameter at the base, 0.5-0.8 mm in diameter at the apex, spreading, densely to moderately viscid-pubescent like the inflorescence axis, articulated at the base; pedicel scars clustered at the tip of the inflorescence, the scar from the basal flower spaced 1-2 mm from the rest. Buds globose to ovoid, the corolla exserted more than halfway from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 1-2.5 mm long, deeply conical, the lobes 1-1.5(-2) mm long, 0.5-0.8 mm wide, long-triangular, tips rounded, densely viscid-pubescent with mixed glandular and eglandular simple uniseriate trichomes to ca. 0.5 mm long. Corolla (8-)10-12 mm in diameter, white, rotate-stellate to stellate, lobed ca. 2/3 of the way to the base, the lobes 4-5 mm long, 3-4 mm wide, spreading or reflexed at anthesis, minutely pubescent-papillate abaxially with simple eglandular trichomes ca. 0.2 mm long, these white when dry. Stamens equal; filament tube minute; free portion of the filaments 0.5-1 mm long, glabrous or occasionally with a few tangled simple uniseriate trichomes adaxially; anthers (2-)2.5-3 mm long, 0.75-1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 4-6 mm long, minutely puberulent in the lower 1/4, merely papillate in plants with glabrous filaments, exserted ca. half the length of the anthers; stigma capitate-globose, bright green in live plants, the surface minutely papillate. Fruit a globose berry, 7-10 mm in diameter, black when ripe, the pericarp thin, matte and somewhat translucent; fruiting pedicels 0.8-0.9 cm long, ca. 0.75 mm in diameter at the base and to ca. 2 mm at the apex, somewhat woody, deflexed to spreading (often appearing like spokes of a wheel), dropping off with mature fruits, not persistent; fruiting calyx somewhat accrescent, the tube 2.5-3 mm long, the lobes 2.5-4 mm long, ca. 1.5 mm wide, somewhat spathulate, appressed to spreading, covering ca. 1/2 of the berry. Seeds 10-30 per berry, 1.5-2.5 mm long, 1.5-2 mm wide, flattened and tear-drop shaped with a subapical hilum, pale brown, the surfaces minutely pitted, the testal cells more or less pentagonal with some sinuate margins. Stone cells 2 per berry, 0.3-0.7 mm in diameter, usually borne near the base of the berry. Chromosome number: 2n =6x=72 ( Bhiravamurty and Rethy 1983; Olet et al. 2015).

Distribution

(Figure 21 ). Distributed from the Arabian Peninsula (Saudia Arabia and Yemen) to Egypt and south in eastern Africa to Kenya and Tanzania, we have only seen a single collection from eastern Democratic Republic of the Congo.

Ecology.

Grows in open areas and along streams in forests and dry areas, also cultivated in eastern Africa; from 800 to 3,000 (-3,500) m elevation.

Common names.

Ethiopia: alam tch’aguar; Kenya: ap-poinet, isusa, manage, nafu, ol’ momoit/olmomoi, soiyot, sonja sucha, sujet; Saudi Arabia: gebel soda; Uganda: ekyala ky’ente, kyalakyente, orushwiga, osiga.

Uses.

In eastern Africa, leaves are eaten as spinach and berries are said to be edible.

Preliminary conservation status

( IUCN 2016). Solanum memphiticum is a widespread species with AOO of 320 km2 (EN) and EOO of 4,489,278 km2 (LC); considering the weedy nature of the species and the large EOO, we suggest a preliminary status of LC (Least Concern; Table 7 ) for the species. Like other members of the group, it is a species of open, disturbed areas, but is less common than S. tarderemotum or S. villosum in eastern Africa.

Discussion.

Solanum memphiticum was long confused with S. villosum ; both are similarly villous plants with toothed leaves that are sometimes pubescent with glandular trichomes. Edmonds (2007) clarified not only the differences between the two species, but also the identity and correct name for S. memphiticum . The species can be easily distinguished in fruit; S. villosum has characteristic red-orange (or yellow) berries with strongly reflexed calyx lobes, while S. memphiticum has green or black berries with the calyx lobes somewhat accrescent and covering at least a third of the berry. Fruiting pedicels of S. memphiticum are typically spreading and, in live plants, form a small bicycle spoke-shaped structure, while those of S. villosum are deflexed. Live plants of the two species are easy to distinguish: the leaves (glandular trichomes) of S. memphiticum have a strong foetid odour of rotting meat that is markedly different from the slightly pleasant odour of S. villosum .

In flower, S. memphiticum can be distinguished by its rotate-stellate corolla with small lobes that are strongly reflexed in flower and calyx lobes that extend beyond the corolla sinuses in open flowers. The flowers of S. memphiticum are delicate and only last a short time as compared to other species (the corolla bruises easily). Leaves of S. memphiticum are generally longer and thinner than those of S. villosum and the base is more decurrent on to the petiole so the free part of the petiole in S. memphiticum is much shorter than in S. villosum .

The parental origin of the tetraploid species has not been investigated in detail. Jaeger (1985) suggested that S. memphiticum might be part of a more broadly circumscribed S. villosum , which included a wide range of fruit colours. Manoko (2007) showed that S. memphiticum (as S. grossedentatum ) does not group closely with S. villosum based on AFLP data but forms a distinct group related to accessions identified as S. tarderemotum , S. chenopodioides , S. nitidibaccatum (as S. physalifolium ) and S. tweedianum . Further molecular work, based on sequence data, will be needed to understand the relationships and parental origins of S. memphiticum , but possible diploid parents could be the morphologically closely related S. sarrachoides or S. nitidibaccatum that both occur in the Old World. Both of these species are glandular and have an enlarged calyx in the fruit, characters shared with S. memphiticum .

Solanum memphiticum was previously commonly known as S. grossidentatum or S. hirsutum (see Manoko 2007), but Edmonds (2007), in her analysis of the morelloid solanums in Pehr Forsskål’s herbarium, showed that the correct name for this black-fruited, pubescent plant is S. memphiticum Gmel. Cufodontis (1963) had used the name earlier, but some confusion occurred due to the later homonym S. memphiticum Mart., that Edmonds (2007) suggested was synonymous with S. scabrum , but we place in the synonymy of S. nigrum . Homonymy is rampant in the morelloid solanums, making name changes like this inevitable; the name S. memphiticum is now well-established for this species in Africa (e.g. Edmonds 2012).

Selected specimens examined. Burundi. Ruyigi: Musongati, FTEA region: BUR, 10 May 1974, Reekmans 3419 (EA, MO).

Democratic Republic of the Congo. Nord-Kivu: Ruindi, Nov 1937, Lebrun 8380 (K).

Egypt. South Sinai: Wadi El Sheikh, Sinai, 15 Apr 1937, Shabetai s.n. (K).

Eritrea. Anseba: Geleb, Gheleb-Caropebir, 16 Jan 1893, Terracciano & Pappi 2027 (FT); Debub: Adi Ugri-Mai Tacala, 4 Sep 1909, Bellini 335 (FT); Saganeiti, gorge Gona pres Addingofon, 29 Mar 1892, Schweinfurth & Riva 1319 (FT, K); Gash Barka: Badum, lungo fiume Mareb, 10 Jan 1906, Pappi 6893 (FT); Get Arba, Gret-Arba, 7 Jan 1893, Terracciano & Pappi 1706 (FT); Maekel: Asmara, 12 Sep 1912, Baldrati 3536 (FT); Asmara, 2 Aug 1902, Pappi 2099 (EA, MO, P, SI); Semienawi Keyih Bahri: Nefasit-Maha-bar, 2 Feb 1909, Fiori 1593 (FT); Illalia -Scilliki; Assaorta, 28 Mar 1893, Pappi 3601 (FT).

Ethiopia. Addis Ababa: Addis Ababa, 18 Jul 1962, Mooney 9136 (FT, K); Amhara: c. 15 km S of Debre Sina on the main rd towards Debre Berhan, 22 Nov 2000, Friis et al. 10115 (K); Dessie, Wollo Prov., 18 Aug 1946, Hall 22 (BM); Wello Prov., Azewagedel mountain, 2 km E of Desse, 11 Apr 1969, Sutherland 167 (MO); Dessie, 26 May 1938, Vatova 2417 (FT); Oromia: Borena, Mega, presso fortino nuovo, 4 May 1937, Cufodontis 627 (FT, W); Agheremariam-Dilla rd, 3 Dec 1952, Gillett 14586 (EA, FT, K); lower slopes of Mt. Cilalo, nr Asella, 10 Sep 1965, de Wilde & de Wilde-Duyfjes 8017 (K, MO, P); Southern Nations (SNNP): Gurage Mountains, above Butajira towards the village of Ageta on the track towards Endibis, 24 Feb 2000, Friis et al. 10066 (K); Tigray: Amba Alaga pass, 9 Oct 1995, Friis et al. 6640 (K); Bellaka, 8 Nov 1854, Schimper 506 (E, FT, W); Arba Tensesa, 7 Oct 1862, Schimper 523 (BM).

Jordan. East Jordan, Pelit, 30 Apr 1963, Gillett 15973 (K); Aqaba: Wadi Rum, 13 Apr 1945, Davis 8987 (E); Ma’an: Petra, 28 Dec 1935, Dinsmore 12151 (E).

Kenya. Central: Nyeri, Aberdare National Park, The Ark, 7 Apr 1975, Hepper & Field 4912 (K); Kiambu, Makuyu, Fort Hall Dist, 26 Jun 1960, van Someren 11980 (EA, K); Eastern: Makueni, Chyulu North, 21 May 1938, Bally B-7788 (K); Makueni, Chyulu North, 28 Apr 1938, Bally B-8306 (K); Makueni, Chyulu Hills North, Apr 1938, van Someren 7644 (K); Nairobi: Mathare Valley, between Mathare Police Station and Eastleigh Section One, 11 Sep 1971, Mwangangi & Kasyoki 1797 (EA, K); Nairobi, FTEA region K4, 1942, Nattrass 341 b (EA, MO); Rift Valley: Kericho, Londiani to Elburgon, Dec 1905, Baker K-346 (EA, K); Nakuru, Lake Naivasha, West shore Mennel’s Farm, 14 Feb 1971, Gillett 19300 (EA, K); Glover et al. 181 (EA, K); Kajiado, Olekairtoror Escarpment, 20 mi from Narok on Nairobi rd, 14 Jul 1962, Glover & Samuel, 3110 (EA, K); K6 Rift Valley, Suswa volcano, 1 Jun 1997, Phillipson & Bytebier 4785 (MO); Western: Nandi, Kapsoret Forest, 15 Jun 1951, Williams 240 (EA, K).

Saudi Arabia. Al Mahmoud, 35 km N of Abha, 21 May 1980, Boulos & Ads 14150 (E, K); Abha Pass (nr. W. Abha), 25 Oct 1971, Popov 71 257 (BM); Asir: Jebel Sudah, c. 18 km N of Abba, 5 Apr 1979, Collenette 1269 (K).

Somalia. Upper Sheikh Kitchen Garden, 1 Dec 1919, Godman 67 (BM, MO); Saaxil: Ally Ully nr Sheikh, 11 May 1973, Wood S/73-65 (K).

Sudan. Darfur: Jebel Marra, Golel, c. 120 km E of Zalingei, 22 Jan 1965, de Wilde et al. 5504 (K, MO).

Tanzania. Arusha: Lake Manyara National Park, Mto ya Ukindu, 29 Nov 1963, Greenway & Kirrika 11096 (EA, K); Monduli Forest Reserve, T2. Nr Olchoropus Village, 25 Jan 2001, Simon et al. 720 (MO); Central: Kondoa, Great North rd, 11 Jan 1962, Polhill & Paulo 1130 (K); Kagera: Ngara, Murugwanza, Ibivyaza Bakobwa, Bugufi, 20 Jan 1961, Tanner 5617 (K); Kilimanjaro: Osirwa Farm, TBL Estates, 26 Jan 1994, Grimshaw 94-148 (K); Ulei, Kwa Sadala, 2 May 1994, Grimshaw 94-470 (K); Moshi, Sanya River, Mar 1928, Haarer 1209 (EA, K); Lake: Ngara, Kirushya, Bugufi, 23 Nov 1959, Tanner 4530 (K); Mbulu: Pienaars Heights, Great North rd, 120 mi S of Arusha, 5 May 1962, Polhill & Paulo 2342 (EA, K); Northern: Mbulu, Mbulumbul, Block D1, 24 Jun 1944, Greenway 6952 (EA, K).

Uganda. Central: Masaka, Kasambya, Kigezi, Feb 1948, Purseglove P-2595 (K); Eastern: Sironko, Budadiri, Bugishi, Jan 1932, Chandler 405 (K); Serere, at Tira, Jul 1926, Maitland 1290 (K); Northern: Zombo, Paidha, 28 Aug 1953, Chancellor 185 (K); Western: Kigezi D.F.I, 28 Aug 1972, Goode G-5 -72 (K); Kasese, Muhokya, Ruwenzori, 25 Dec 1925, Maitland 1790 (K); Kigezi, Kachwekano Farm, Sep 1949, Purseglove 3121 (EA, K); Kisoro, Virunga-Kette, Muhavura, Nkanda, 25 Nov 1954, Stauffer 957 (EA, K, P).

Yemen. Hadramaut: Al Mukalla, Arabia, West rd, 6 Sep 1949, Guichard KG/HAD 35 (BM); Kor Seiban, 14 Sep 2002, Killian et al. YP-3578 (B); Sa’dah: Sadah, 1 Jul 1984, Gordon 609 B (E); Sana’a: Sana’a, 12 Mar 1981, Miller 3004 (E); 14 Feb 1934, Rathjens s.n. (BM); Menacha, 7 Mar 1889, Schweinfurth 1476 (BM, GH, K, P); Ar Rowdah nr San’a, 23 Feb 1972, Wood 72-7 (BM); 10 Oct 1974, Wood Y/74-20 (BM); Ta’izz: Jabal Sabir, c. 15 km S of Taizz, 11 Jun 1982, Gordon 11 (E); Jabal Sabir, nr Taiz, 23 Sep 1977, Lavranos 15947 (E).

Figures

Figure 19. Solanum memphiticum J. F. Gmel A Habit B Detail of stem with glandular indumentum C Detail of abaxial leaf surface D Glandular trichome E Eglandular trichome F Inflorescence G Flower at anthesis H Infructescence I Seed (A-E, H de Wilde 8017; F-I Gilbert 1387; G Nijmegen acc. A 34570474). Scale bar: 3 cm (A), 3 mm (B), 4 mm (C, G), 0.8 mm (D-E), 1.5 cm (F, H) and 2 mm (I). Drawing by L. Smith.

Figure 20. Solanum memphiticum J. F. Gmel A Habit B Glandular indumentum C Flowers at full anthesis D Maturing fruits with long, appressed calyx lobes (A-B, D Nijmegen acc. A 14750032; C Nijmegen acc. A 34750474). Photos by S. Knapp and G. van der Weerden.

Figure 21. Distribution of Solanum memphiticum J. F. Gmel.