Solanum terminale Forssk., Fl. Aegypt.-Arab. 45. 1775.

Knapp, Sandra & Vorontsova, Maria S., 2016, A revision of the " African Non-Spiny " Clade of Solanum L. (Solanum sections Afrosolanum Bitter, Benderianum Bitter, Lemurisolanum Bitter, Lyciosolanum Bitter, Macronesiotes Bitter, and Quadrangulare Bitter: Solanaceae), PhytoKeys 66, pp. 1-142 : 70-98

Solanum terminale Forssk., Fl. Aegypt.-Arab. 45. 1775. Figures 1E , 2C & D , 25

Solanum bifurcatum Hochst. ex A.Rich., Tent. Fl. Abyss. 2: 98. 1851. Type. Ethiopia. Tigray: Adoa [Adwa], 14 Jun 1837, G.H.W. Schimper 201 (lectotype, designated by Lester 1997, pg. 281; P [P00344566]; isolectotypes: BM [BM000847558], BR [BR0000006289041, BR0000008422248], E [E00193277], G [G00301682, G00343569], G-DC [G00144683], GH [GH00139638], GOET [GOET003592], HBG [HBG511409], HOH [HOH009830], LE [3 sheets], K [K000232603], M [M0105581, M0105582], MPU [MPU011243, MPU011244], NY [NY00169749], OXF, P [P00344587, P00341599, P00344570, P00344567], PAL, REG [REG000401], S [S-09-33851, S-09-33853, S-115601], STU [STU000029, STU000028], U [U0113932], US [US-945142, barcode US00730876], W [W0000632, W1889-0283815]).

Solanum bifurcum Hochst. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 77. 1852, nom. illeg. superfl., orthographic variant. Type. Based on Solanum bifurcatum A.Rich.

Solanum inconstans C.H.Wright, Bull. Misc. Inform. Kew 1894: 127. 1894. Type. Equatorial Guinea. Bioco: "Fernando Po", Dec 1859, G. Mann 62 (lectotype, designated here: K [K000414056]; isolectotype: P [P00341583]).

Solanum phytolaccoides C.H.Wright, Bull. Misc. Inform. Kew 1894: 126. 1894. Type. Ethiopia. [Amhara/Tigray]: "ex Tigrè v. Begemder", 24 Sep 1862/13 Sep 1863, G.H.W. Schimper 310 (lectotype, designated here: K [K000232602]; isolectotypes: BM [BM000838174, BM000838175], E [E00526857], US [US-806525]).

Solanum welwitschii C.H.Wright, Bull. Misc. Inform. Kew 1894: 126. 1894. Type. Angola. Cuanza Norte: Golungo Alto, F.M.J. Welwitsch 6098 (lectotype, designated here: BM [BM000838177]; isolectotype: K [K000414110], LISU [LISU220797]).

Solanum welwitschii C.H.Wright var. oblongum C.H.Wright, Bull. Misc. Inform. Kew 1894: 127. 1894. Type. Cameroon. Sud-Ouest: Ambas Bay, G. Mann 10 [s.n., No. X] (lectotype, designated by Edmonds 2012, pg. 113: K [K000414053]; isolectotype: P [P00331492]).

Solanum welwitschii C.H.Wright var. strictum C.H.Wright, Bull. Misc. Inform. Kew 1894: 127. 1894. Type. Equatorial Guinea. Bioco: "Fernando Po", 1860, G. Mann 274 (lecctotype, designated here: K [K000414054]; isolectotype: P [P00331491].

Solanum campanuliflorum C.H.Wright, Bull. Misc. Inform. Kew 1894: 127. 1894. Type. Angola. Cunene: Cunene [River], Sep 1883, H.H. Johnston s.n. (lectotype, designated here: K [K00414073]).

Solanum nakurense C.H.Wright, Bull. Misc. Inform. Kew 1897: 275. 1897. Type. Kenya. Rift Valley: Masai [Nakuru ex protologue], 1893, G.F. Scott-Elliot 6800 (holotype: K [K000096900]; isotype: BM [BM000847730]).

Solanum symphyostemon De Wild. & T.Durand, Ann. Mus. Congo, Sér. 2, Bot., ser. 2, 1: 44. 1899. Type. Democratic Republic of the Congo. Équateur: Bolombo, près de Ngala (Ngali), 21 Sep 1986, F. Thonner 96 (lectotype, designated here: BR [BR0000008993137]; isolectotypes: BR [BR0000008994462], K [K000414043]).

Solanum lujaei De Wild. & T.Durand, Bull. Soc. Roy. Bot. Belgique 38: 209. 1899. Type. Democratic Republic of the Congo. Bas-Congo: "Sona Gunga, cataractes", 21 Nov 1898, É. Luja 112 (lectotype, designated here: BR [BR0000014800313]).

Solanum togoense Dammer, Bot. Jahrb. Syst. 38: 59. 1906 [ “1907”]. Type. Togo. Maritime: Badja, Mar 1900, F.R.R. Schlechter 12974 (lectotype, designated by Edmonds 2012, pg. 110: K [K000414055]; isolectotypes: BM [BM000838103], BR [BR0000006288921], P [P00341582]).

Solanum buchwaldii Dammer, Bot. Jahrb. Syst. 38: 180. 1906 [ “1907”]. Type. Tanzania. Tanga: Usambara [Mtns., Lushoto District], Muafa, Apr [1900], J. Buchwald 542 (lectotype, designated by Bitter 1917, pg. 450: B, destroyed, no duplicates found).

Solanum bilabiatum Dammer, Bot. Jahrb. Syst. 38: 181. 1906 [ “1907”]. Type. São Tomé and Principe. São Tomé: Sin. loc., 2-7 m, [1885], A.F. Moller 146 (type in B [?], destroyed; no duplicates found).

Solanum comorense Dammer, Bot. Jahrb. Syst. 38: 181. 1906 [ “1907”]. Type. Mayotte (French Overseas Department). Maore: forêt du Combani, 6 Oct 1884, L. Humblot [1]284 (neotype, designated by D’Arcy & Rakatozafy 1994, pg. 126: P [P00184319]; isoneotypes: BM [BM000887171], K [K000788514]).

Solanum laurentii Dammer, Bot. Jahrb. Syst. 38: 182. 1906 [ “1907”], nom. illeg., non Solanum laurentii De Wild., 1905. Type. Democratic Republic of the Congo. Sin. loc., " Laurent s.n. " (type at B [?], destroyed; no duplicates found). Democratic Republic of the Congo. Équateur: environs d’Eala, 11 May 1905, M. Laurent 701 (neotype, designated here: BR [BR0000014801273]).

Solanum plousianthemum Dammer, Bot. Jahrb. Syst. 38: 180. 1906 [ “1907”] Type. Tanzania. "Usambara, im Gebüsch niederer Hügel”, Jul 1892, C. Holst 3731 (lectotype, designated by Bitter 1917, pg. 457): B, destroyed; no duplicates found).

Solanum suberosum Dammer, Bot. Jahrb. Syst. 38: 182. 1906 [ “1907”]. Type. Cameroon. Sud-Ouest: Barombi Station between Station and Ninga town, Mar 1889, G.R Preuss 18 (type, B [?], destroyed; no duplicates found); Barombi-Bache, SW of Station, 1400 m, Apr 1889, G.R. Preuss 171 (type, B [?], destroyed; no duplicates found); Buea, 1400 m, May 1891, G.R Preuss 885 (type, B [?], destroyed; no duplicates found); Buea Station, 1000 m, Apr 1898, Lehmbach 211 (type, B [?], destroyed; no duplicates found); Yaounde, May 1894, G.A. Zenker & A. Staudt 328 (type, B [?], destroyed; no duplicates found); Yaounde, Apr 1890, G.A. Zenker 268 (type, B [?], destroyed; no du plicates found);northern Cameroon, Bangwe, Jun, G. Conrau 200 (type, B [?], destroyed; no duplicates found).

Solanum preussii Dammer, Bot. Jahrb. Syst. 38: 183. 1906 [ “1907”]. Type. Cameroon. Sud-Ouest: "Barombi Station", Aug 1890, P.R. Preuss 397 (lectotype, designated here: P [P00331480]; isolectotype: E [E00193273]).

Solanum lykipiense C.H.Wright, Fl. Trop. Afr. [Oliver et al.] 4, 2: 220. 1906. Type. Kenya. Rift Valley: Lykypia, 6000-8000 ft., J. Thomson s.n. (lectotype, designated here: K [K000096902]).

Solanum mangaschae Pax, Bot. Jahrb. Syst. 39: 648. 1907. Type. Ethiopia. Southern Nations, Nationalities and Peoples: "Amaniel am Gazerit", 2300, 2 Apr 1905, F. Rosen s.n. (holotype: WRSL, not seen).

Solanum subcoriaceum T.Durand & H.Durand, Bull. Jard. Bot. État Bruxelles 2: 394. 1909. Type. Based on and nom. nov. for Solanum laurentii Dammer, not Solanum laurentii De Wild.

Solanum leucanthum Dammer, Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-1908), Bot., 2: 284. 1911 [ “1914”]. Type. Rwanda. Western: Rukarara, Rugege Forest, 2900 m, Aug 1907, G. Mildbraed 894 (B [?], destroyed; no duplicates found); "Kissenye, Bugoyer, Hügelland, Hecken", 2100 m, 27 Oct 1907, G. Mildbraed 1431a (B [?], destroyed; no duplicates found).

Solanum aculeolatum Dammer, Bot. Jahrb. Syst. 48: 237. 1912, nom. illeg., non Solanum aculeolatum M.Martens & Galeotti, 1845. Type. Kenya. "Escarpment, in Lichtungen", 2500 m, Feb 1903, F. Thomas s.n. (type in B [?], destroyed; lectotype, designated here: E [E00193280]).

Solanum penduliflorum Dammer, Bot. Jahrb. Syst. 48: 255. 1912. Type. Kenya. Rift Valley: "Mau Plateau, common railway cuttings and open places, Molo, Ravine, Londiani and Njoro", 2300-3000 m, G.S. Baker 133 (type in B [?], destroyed; no duplicates found).

Solanum bansoense Dammer, Bot. Jahrb. Syst. 48: 237. 1912. Type. Cameroon. Nord-Ouest: “Bansso-Gebirge”, 1700 m, Oct 1909, C.L. Ledermann 5778 (type in B [?], destroyed; no duplicates found).

Solanum massaiense Bitter, Repert. Spec. Nov. Regni Veg. 11: 18. 1912. Type. Based on and nom. nov. for Solanum aculeolatum Dammer, not Solanum aculeolatum M.Martens & Galeotti.

Solanum holtzii Dammer, Bot. Jahrb. Syst. 53: 329. 1915. Type. Tanzania. Morogoro: "Uluguru, Bez. Morogoro, Waldreservat Banduki II", Mar 1913, W. Holtz 3148 (type in B [?], destroyed; no duplicates found).

Solanum rhodesianum Dammer, Bot. Jahrb. Syst. 53: 326. 1915. Type. Zimbabwe. Manicaland: Melener, Chirinda Forest, 9 Oct 1905, C.F.M. Swynnerton 86 (lectotype, designated here: K [K000414111]; isolectotypes: BM [BM000838178], K [K000414112]).

Solanum stolzii Dammer, Bot. Jahrb. Syst. 53: 327. 1915. Type. Tanzania. Mbeya: Rungwe Crater, Kyimbila Station, 2000 m, 19 Dec 1911, A. Stolz 1035 (lectotype, designated here: M [M0105595]; isolectotypes: B [B 10 0165205, received 1990], C [C10003110], G [G00343580], K [K000096901], S [S09-36468], W [W19150004892].

Solanum meyeri-johannis Dammer, Bot. Jahrb. Syst. 53: 328. 1915. Type. Tanzania. "Ussagara, Bez. Kilossa, Bugaberge" Nov-Dec 1911, Dr. Houy 1242 (type in B [?], destroyed; no duplicates found).

Solanum lateritium Dammer, Bot. Jahrb. Syst. 53: 325. 1915. Type. Tanzania. Mbeya: Kyimbila Station, 600-800 m, Aug 1912, A. Stolz 1514 (lectotype, designated here: JE [JE00004286]; isolectotypes: G [G00442975], K [K000413992], S [S-G-5699], U [U0113933], WAG [WAG0003361], W [W19150008146]).

Solanum leucanthum Bitter & Dammer, Bot. Jahrb. Syst. 54: 456. 1917, nom. illeg., non Solanum leucanthum Dammer, 1911. Type. Rwanda. Western: Rukarara, Rugege Forest, 2900 m, Aug 1907, G. Mildbraed 894 (no herbarium cited, B [?], destroyed; no duplicates found).

Solanum hemisymphyes Bitter, Bot. Jahrb. Syst. 54: 477. 1917. Type. Democratic Republic of the Congo. Nord-Kivu: Kwa Muera, Fort Beni, Jun 1908, J. Mildbraed 2238 (type in B [?], destroyed; no duplicates found).

Solanum nakurense C.H.Wright var. lykipiense (C.H.Wright) Bitter, Bot. Jahrb. Syst. 54: 449. 1917. Type. Based on Solanum lykipiense C.H.Wright

Solanum plousianthemum Dammer subsp. holtzii (Dammer) Bitter, Bot. Jahrb. Syst. 54: 467. 1917. Type. Based on Solanum holtzii Dammer

Solanum plousianthemum Dammer subsp. kasima Bitter, Bot. Jahrb. Syst. 54: 468. 1917. Type. Tanzania. Mbeya: Neu-Langenburg, Kyimbila, 1350 m, Oct 1912, A. Stolz 355 (holotype: B, destroyed; lectotype, designated here: G [G00070223]; isolectotype: K [K000413993]).

Solanum plousianthemum Dammer var. angustifrons Bitter, Bot. Jahrb. Syst. 54: 462. 1917. Type. Tanzania. Kilimanjaro: Mount Kilimanjaro, Marangu, 1500 m, Apr 1894, G. Volkens 2109 (type in B [?], destroyed; no duplicates found).

Solanum plousianthemum Dammer var. buchwaldii (Dammer) Bitter, Bot. Jahrb. Syst. 54: 459. 1917. Type. Based on Solanum buchwaldii Dammer

Solanum plousianthemum Dammer var. commixtum Bitter, Bot. Jahrb. Syst. 54: 464. 1917. Type. Tanzania. Kilimanjaro: Mount Kilimanjaro, ca. 1600 m, May 1894, H.H.Johnston s.n. (lectotype, designated here: K [K000413995]; isolectotype: BM [BM000838176]).

Solanum plousianthemum Dammer var. conglutinans Bitter, Bot. Jahrb. Syst. 54: 461. 1917. Type. Tanzania. Tanga: Usambaras, Kwa Mshuza, “Handei”, 1570 m, Aug 1893, C. Holst 8927 (lectotype, designated here: P [P00341605]; isolectotypes: COI [COI00077085], K [K000413989], LE, W [W18940006582]).

Solanum plousianthemum Dammer var. devians Bitter, Bot. Jahrb. Syst. 54: 466. 1917. Type. Rwanda. "Mpororo (Rufua), Issenge-Posten, an nordlichen Zuflussen Kagera", Jul 1907, J. Mildbraed 336 (type in B [?], destroyed; no duplicates found).

Solanum plousianthemum Dammer var. endosiphonotrichum Bitter, Bot. Jahrb. Syst. 54: 465. 1917. Type. Rwanda. Southern: Mount Niansa [Nyanza], 1700 m, Dr. Kandt 147 (holotype: B, destroyed; no duplicates found).

Solanum plousianthemum Dammer var. microstelidium Bitter, Bot. Jahrb. Syst. 54: 461. 1917. Type. Rwanda. Western: Lake Kivu, Lubengera, Mugarura Island, H. Meyer 909 (type in B [?], destroyed; no duplicates found).

Solanum plousianthemum Dammer var. epapillosum Bitter, Bot. Jahrb. Syst. 54: 464. 1917. Type. Tanzania. Kilimanjaro: Mount Kilimanjaro, Useri, 2200 m, Mar 1894, G. Volkens 1991 (type in B [?], destroyed; no duplicates found).

Solanum plousianthemum Dammer var. gracilifilum Bitter, Bot. Jahrb. Syst. 54: 463. 1917. Type. Tanzania. Kilimanjaro: Mount Kilimanjaro, Marangu, 2000 m, May 1894, G. Volkens 2265 (lectotype, designated here: BR [BR0000006495794]).

Solanum plousianthemum Dammer var. rhodesianum (Dammer) Bitter, Bot. Jahrb. Syst. 54: 461. 1917. Type. Based on Solanum rhodesianum Dammer

Solanum plousianthemum Dammer var. subtusbarbellatum Bitter, Bot. Jahrb. Syst. 54: 467. 1917. Type. Kenya. Eastern: [Kitui County], Galunka [Galinka], 25 May 1902, L.C.T. Kässner 804 (holotype: B, destroyed; lectotype, designated here: BM [BM000847733]; isolectotype: K [K000788864]).

Solanum plousianthemum Dammer var. ugandae Bitter, Bot. Jahrb. Syst. 54: 460. 1917. Type. Uganda. Sin. loc., 1 Jan 1891, F. Stuhlmann 1329 (lectotype, designated here: K [K000413994])..

Solanum plousianthemum Dammer var. kundelunguense Bitter, Bot. Jahrb. Syst. 54: 467. 1917. Type. Democratic Republic of the Congo. Katanga: Kundelungu, 15 May 1908, L.C.T. Kässner 2760 (holotype: B, destroyed; lectotype, designated here: BM [BM000847803]; isolectotypes: K [K000414041], P [P00341576]).

Solanum ruandae Bitter, Bot. Jahrb. Syst. 54: 471. 1917. Type. Rwanda/Democratic Republic of the Congo: Kissenye, "Bugoyer Hügelland, Hecken", 2100 m, 27 Oct 1907, J. Mildbraed 1431a (holotype: B, destroyed; no duplicates found).

Solanum suberosum Dammer var. calvum Bitter, Bot. Jahrb. Syst. 54: 486. 1917. Type. Cameroon. Sud-Ouest [?]: "Nordwestkamerum, Spitze des Kamerumberges" [Mount Cameroon], A. Weberbauer 38, 60 (syntypes: B, destroyed; no duplicates found).

Solanum suberosum Dammer var. ramosivelutinum Bitter, Bot. Jahrb. Syst. 54: 486. 1917. Type. Democratic Republic of the Congo. Orientale: Ituri, Irumu-Mawambi, Apr, J. Mildbraed 3038 (type in B [?], destroyed; no duplicates found).

Solanum sychnoteranthum Bitter, Bot. Jahrb. Syst. 54: 472. 1917. Type. Democratic Republic of the Congo[?]. "lava plain", 18 Aug 1908, L.C.T. Kässner 3250 (holotype: B, destroyed; lectotype, designated here: E [E00193274]; isolectotypes: BM [BM000847732], P [P00331494]).

Solanum plousianthemum Dammer var. angustatum Bitter, Repert. Spec. Nov. Regni Veg. 18: 302. 1922. Type. Tanzania. Morogoro: “Ulugurus”, 19 Oct 1013, W. von Brehmer 891 (no herbarium cited; no duplicates found).

Solanum plousianthemum Dammer var. kyimbilense Bitter, Repert. Spec. Nov. Regni Veg. 18: 302. 1922. Type. Tanzania. Mbeya: Bex, Neu-Langenburg, Kyimbila, near Madehani, 2000 m, 3 Dec 1913, A. Stolz 2315 (lectotype, designated here: K [K000413990]; isolectotypes: A [A00139637, A00219304], BM [BM000838180], BR [BR0000006495107], CAS [CAS- 130438], K [K000414991], P [P00341613]).

Solanum bansoense Dammer var. episporadotrichum Bitter, Repert. Spec. Nov. Regni Veg. 18: 303. 1922. Type. Cameroon. Est: at junction of Sanaga [River?] with the Djerem [River], ("hylaea um Deng-Deng"), 250 km NE of Yaounde, between Mbo’s and Sardi, southwest of Deng-Deng, 8 Mar 1914, G. Mildbraed 8555 (holotype: B, destroyed; no duplicates found).

Solanum bansoense Dammer subsp. sanaganum Bitter, Repert. Spec. Nov. Regni Veg. 18: 304. 1922. Type. Cameroon. Est: at junction of Sanaga [River?] with the Djerem [River], ("hylaea um Deng-Deng"), 250 km NE of Yaounde, Mar 1914, G. Mildbraed 8619 (holotype: B, destroyed; lectotype, designated here: K [K000414042]).

Solanum welwitschii C.H.Wright var. laxepaniculatum Bitter, Repert. Spec. Nov. Regni Veg. 18: 306. 1922. Type. Cameroon. Est: [Upper Nyong District], Lomie District, Dscha region, in Tugumedjo’s village, 13°30-12°25 W latitude, 18 May 1911, G. Mildbraed 5265 (holotype: B, destroyed; lectotype, designated here: HBG [HBG511510]).

Solanum lianiforme De Wild., Pl. Bequaert. 1: 427. 1922. Type. Democratic Republic of the Congo. Nord-Kivu: Walikale, 7 Jan 1915, J. Bequaert 6531 (lectotype, designated here: BR [BR0000008993489]; isolectotypes: BR [BR0000008994448, 2 sheets]).

Solanum butaguense De Wild., Pl. Bequaert. 1: 424. 1922. Type. Democratic Republic of the Congo. Nord-Kivu: Butagu, Ruwenzori, ca. 2200 m, 13 Apr 1914, J. Bequaert 3616 (lectotype, designated here: BR [BR0000008994431]; isolectotypes: BR [BR0000008993502, BR0000008993472]).

Solanum balboanum Chiov., Racc. Bot. Miss. Consol. Kenya 87. 1935. Type. Kenya. Central: Tusu, 1 Feb 1911, P.G. Balbo 579 (lectotype, designated here: FT [FT003031]).

Solanum terminale Forssk. subsp. inconstans (C.H.Wright) Heine, Kew Bull. 14: 247. 1960. Type. Based on Solanum inconstans C.H.Wright Solanum terminale Forssk. subsp. sanaganum (Bitter) Heine, Kew Bull. 14: 248. 1960. Type. Based on Solanum bansoense Dammer subsp. sanaganum Bitter

Solanum terminale Forssk. subsp. welwitschii (C.H.Wright) Heine, Kew Bull. 14: 248. 1960. Type. Based on Solanum welwitschii C.H.Wright

Type.

yemen. “Mokhaja” [Mukhajah =Jebel Barad], P. Forsskål s.n. (lectotype, designated here: C [C10003105] Herb. Forsskal 419; isolectotype: C [C10003106] Herb. Forsskal 406).

Description.

Shrub to high-climbing woody liana, from ca. 1 m tall to canopy height. Stems terete, to 10 cm in diameter in liana forms, glabrous with a few simple uniseriate or dendritic trichomes to densely pubescent with simple and/or dendritic uniseriate trichomes, these soon deciduous; new growth sparsely to densely pubescent with simple and/or dendritic 4-6-celled uniseriate trichomes 0.5-1 mm long. Bark of older stems pale yellowish white, corky on larger woody stems> 0.5 cm in diameter. Sympodial units plurifoliate, the leaves not geminate, evenly distributed along branches. Leaves simple, (1.5)4-16 cm long, (1)2-8 cm wide, very variably sized on individual stems, elliptic, the adaxial surfaces glabrous to sparsely pubescent with simple or less commonly dendritic uniseriate trichomes 0.5-1 mm long, these denser on the veins, the abaxial surfaces glabrous to densely and evenly pubescent with dendritic and/or simple uniseriate 4-8-celled trichomes 1-1.5 mm long; major veins 5-7 pairs, usually somewhat yellowish in dry material; base acute to truncate, very occasionally somewhat cordate, not decurrent on the petiole; margins entire; apex acute to acuminate; petiole (0.5)2-7(11) cm long, highly variable from leaf to leaf even in leaves of the same size, sparsely to densely pubescent with simple and/or dendritic trichomes like those of the leaves. Inflorescences terminal, 1-20(+) cm long, many times branched, in some forms the branches very short and the inflorescence appearing spicate, with (5)10-100 flowers in clumps at the ends of branches, glabrous or sparsely pubescent with simple and/or dendritic uniseriate trichomes ca. 0.5 mm long, these denser at the base of flower clumps; peduncle to 10 cm long; pedicels 0.5-1 cm long, ca. 0.5 mm in diameter at the base and apex, glabrous to sparsely pubscent in parallel with the rest of the inflorescence, spreading at anthesis, articulated at the base leaving a tiny raised lump on the rachis when flower falls; pedicel scars in tightly packed clusters at the ends of long branches or along the main axis. Buds ovoid to narrowly elliptical (see commentary), the corolla strongly exserted from the calyx tube long before anthesis. Flowers 5-merous, apparently all perfect. Calyx tube 1-1.5 mm long, open-conical, the lobes 1-2 mm long, deltate with acuminate tips (exceptionally to 4 mm long in Strid 3113), glabrous or pubescent with simple or dendritic trichomes like those of the inflorescence axis. Corolla 1-2.5 cm in diameter, lilac to white with varying degrees of lilac towards the centre, deeply stellate, divided 3/4 to nearly to the base, the lobes 5-9 mm long, 2-2.5 mm wide, narrowly ligulate, spreading to reflexed at anthesis, the adaxial surface glabrous or densely pubescent/papillate with 2-3-celled simple trichomes, the abaxial surface densely pubescent with 2-3-celled weak-walled simple trichomes, these denser along the margins and at the tips, sometimes purple-tinged, the corolla on herbarium specimens with a mealy appearance. Stamens equal; filament tube <0.1 mm long to absent; free portion of the filaments 0.5-1 mm long, glabrous; anthers (3-)3.5-5 mm long, 1-2 mm wide, ellipsoid to narrowly ellipsoid, loosely connivent to tightly fused due to internal papillae, smooth or papillate abaxially, somewhat sagittate at the base, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous or occasionally minutely hispidulous with short erect uniseriate trichomes (Gossweiler s.n., coll. 1910); style 8-10 mm long, exserted beyond the anthers approximately equal to the anther length, glabrous; stigma capitate, the surfaces minutely papillate. Fruit a globose to elongate and fusiform berry, 5-7 mm in diameter, bright red when ripe, the pericarp very thin and shiny; fruiting pedicels 0.5-1.5 cm long, ca. 1 mm in diameter, spreading at anthesis; fruiting calyx lobes not markedly enlarging. Seeds 12-20 per berry, 1.5-2.5 mm long, 1-2 mm wide, ovoid reniform, reddish brown, the surfaces deeply pitted, the testal cells pentagonal to rectangular, the lateral cell walls composed of elongate filamentous projections to 0.5 mm long, giving the seeds a hairy appearance.

Distribution

(Figure 26 ). Widely distributed in continental sub-Saharan Africa (see also Table 2 ); also found in the Comoro Islands and on the Arabian peninsula in Yemen. We have seen specimens from Angola, Burundi, Cameroon, Central African Republic, Comoros, Côte d’Ivoire, Democratic Republic of the Congo, Equatorial Guinea, Eritrea, Ethiopia, Gabon, Ghana, Guinea, Guinea Bissau, Kenya, Liberia, Malawi, Mozambique, Nigeria, Republic of Congo, Rwanda, São Tome e Principe, Sierra Leone, South Africa, South Sudan, Tanzania, Togo, Uganda, Yemen, Zambia, and Zimbabwe.

Ecology and habitat.

A wide variety of forest habitats; also from savannah; near sea level to 3300 m elevation. Swamp forests, humid forest, sclerophyllous woodland, grassland savannah. Plants growing in the open in savanna habitats area shorter and more shrub-like than those of forest edges or interiors.

Common names and uses.

Many common names are recorded on herbarium specimens of Solanum terminale , some with the language, and some without. We have listed here those that occur multiple times, with a single voucher for each. All common names associated with individual specimens can be found in the data on the Natural History Museum Data Portal (https://doi.org/10.5519/0039445) and on the Solanaceae Source website (http://www.solanaceaesource.org). Burundi. agatoretore (Kirundi language, Michel 987), umuhanuraguba (Becquet 861); Democratic Republic of the Congo. belele (Lombo language, Louis 981), belemba (Lombo language, Louis 6362), itotofo (Lombo language, Jeanty 48), itotofo i boliki (Lombo language, Louis 14146), kampako (Bofunda language, Corbisier-Baland 1034), kedekede (Giorgi 745), kongalo n pako (Ngala language, Claessens 453), mampoko (Mbole language, Leonard 802), muhwehwe (Tembo language, Troupin 10162), qwadja (Reygaert 226), umuranuhankuba (Kinyarwanda language, Spitaels 119); Kenya. mutandambuga (Meru language, Graham 1752); Rwanda. umuranuhankuba (Kinyarwanda language, Troupin 6631); Tanzania. kadaru (Bantu language, Koritschoner 866), luvuvu (Kirangi language, Burtt 1141); Uganda. omuhanurankuba (Nkore-Kiga language, Purseglove P-1654). The fruit said to be poisonous and the leaves eaten in the Central African Republic ( Burkhill 2000).

Preliminary conservation status

( IUCN 2014). Least Concern (LC). EOO 21,154,570 km2 (LC), AOO 2,316 km2 (NT). Solanum terminale is widespread across continental Africa in a wide variety of habitats, and occurs in several protected areas. The near threatened status suggested by the AOO calculation is likely due to collection bias; Solanum terminale is a very common plant. Genetic differentiation across this wide range, however, may be important for population level conservation or national conservation priorities.

Discussion.

Solanum terminale is the most widespread and common species of the ANS clade on the African mainland, and is often recognised as three distinct entities; Solanum nakurense , Solanum terminale s.s. and Solanum welwitschii (e.g., Edmonds 2012). Bitter (1917, 1922) recognised 20 species and 23 infraspecific taxa, while Heine (1960) amalgamated all these into a single species with three infraspecific taxa in western Africa, and suggested that a similar course of action needed to be taken throughout the continent. We are recognising these populations as a single species because the characters used to distinguish them are continuously variable and field observations indicate that habitat plays a significant role in overall morphology (see below).

Solanum terminale is not broadly sympatric with any other species of the ANS clade, although in eastern South Africa Solanum africanum also occurs, but not strictly sym patrically; Solanum runsoriense also occurs in the same regions as Solanum terminale but usually at higher elevations. Solanum terminale is easily distinguished from others in the clade by its flowers with densely papillate abaxial corolla lobes, flowers clustered at the ends of inflorescence branches (of varying length) and bright red berries. Populations from western Africa tend to have shorter inflorescence branches (see Fig. 25D ), tightly connate anthers and elongate berries and have been recognised as Solanum welwitschii , while those from eastern Africa have more open inflorescences (see Fig. 25B, C ), more loosely connate anthers and globose berries and have been called Solanum terminale . Small plants with few-flowered inflorescences primarily from eastern Africa have been called Solanum nakurense (Fig. 25A ). The leaves of Solanum terminale are usually pubescent to some degree, but are occasionally completely glabrous (e.g., the type of Solanum welwitschii ). Dense tufts of trichomes in the vein axils (domatia) do not occur in Solanum terminale , but the pubescence is often denser near the midvein.

In order to determine if there was any geographical component to the morphogical variation seen in Solanum terminale , we measured the key characters used to distinguish Solanum terminale from Solanum welwitschii in a range of specimens from across continental Africa. Long thin buds that open to flowers with fused anthers, and spicate inflorescences with short to non-existent branches are thought to be typical of Solanum welwitschii (e.g., Edmonds 2012). Neither character segregated clearly into two classes in our simple analysis (Figure 27 ). Bud shape is similarly distributed across longitude, but the longest, thinnest buds occur in western Africa (see point at top left-hand corner of Fig. 27A ). Spicate inflorescences are somewhat more common in the wet forests of western Africa (Fig. 27B ). The type of Solanum welwitschii var. laxepaniculatum (Mildbraed 5265 - http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.hbg511510), however, is a good example of a " terminale " type inflorescence in western Africa. Anthers on specimens from wet forests of western Africa (red dots in Fig. 27 ) were more often fused than those from drier forests of eastern Africa. Anther connation may be a way of protecting pollen in damp, humid environments, but this remains to be tested. The characters used to distinguish the taxa we here recognise as the single species Solanum terminale appear to be the extremes of continuous variation, and perhaps represent fixation of particular morphologies in certain populations. Field study of these plants coupled with molecular analysis at the population level will be essential to untangle the complex morphological variation with Solanum terminale .

In their catalogue of Forsskål’s plant collections Hepper and Friis (1994) cited both specimens in C as “type” of Solanum terminale ; of these two sheets Herb. Forsskål 419 (C10003105) has the larger inflorescence with more flowers and we select it here as the lectotype. They considered Herb. Forsskål 406 (C10003106) a duplicate ( Hepper and Friis 1994); we concur and consider it an isolectotype.

Both Udo Dammer and Georg Bitter described many infraspecific taxa in what we now recognise as the widely distributed and polymorphic Solanum terminale . Both botanists worked in Berlin before the Second World War and it is probable that much of the material upon which they based their descisions is no longer extanct ( Vorontsova and Knapp 2010). Bitter (1917) acknowledged his lack of ability to travel to other herbaria due to military service and war as a short-coming of his treatment of this polymorphic complex. We have searched extensively for duplicates of this material throughout the course of the PBI Solanum project, and in some cases (e.g., see Solanum plousianthemum var. subtusbarbellatum above) have encountered duplicates, while in many, if not most, others we have not been successful. Rather than neotypifying these many infraspecific names we cite type collections in the hope that duplicates will eventually be encountered in the collections we have not yet been able to visit. Dammer (1906, 1912, 1915) never cited particular herbaria in association with specimens he used in describing new taxa (see Vorontsova and Knapp 2016), Bitter (1917, 1922) occasionally did so. Where he cited herbaria in association with type specimens we have indicated this, otherwise no herbarium was cited and we chose lectotypes from duplicates we have seen. Our choices of lectotypes are based on the quality of the particular sheet selected. When syntypes were cited, our choice has been made based on the number of herbaria in which particular syntype collections now are found; where more duplicates of a collection are extant, we selected that as the lectotype.

In his treatment of the species of the ANS clade in Solana Africana Bitter (1917) re-circumscribed many of Dammer’s (1906, 1915) species and excluded some of the specimens cited by Dammer as types of new infraspecific taxa. He explicitly cited types for Dammer’s names, something that later authors did not realise, resulting in some superfluous later lectotypifications (e.g., Edmonds 2012). Solanum leucanthum was first described by Dammer (1906) then explicitly redescribed as " Solanum leucanthum Bitter & Dammer n.sp." by Bitter (1917). Dammer (1906) originally cited two syntypes from Cameroon, Mildbraed 894 and Mildbraed 1431a, while Bitter restricted his concept to only Mildbraed 894. Bitter (1917) recognised Mildbraed 1431a as a distinct species Solanum ruandae . We have found no duplicates of either of these collections that were probably in Berlin and destroyed.

We have not selected a neotype for Dammer’s (1906) Solanum suberosum ; of the seven syntype collections cited we are certain that duplicates will be found eventually. If not, the description and comments on the corky stem morphology will be easy to match, and a recent and widely distributed collection from Cameroon should be selected as a neotype.

Dammer’s (1906) illegitimate name Solanum laurentii (not Solanum laurentii De Wild., 1905) was described using a single specimen collected by “Laurent” from an unspecified locality in “Congo”. This sheet was in all likelihood at B and is no longer extant; the lack of specifics makes duplicates very difficult to trace. Durand and Durand (1909: 394) coined the replacement name Solanum subcoriaceum , but again did not specify a herbarium, sheet or locality; nor did they specify which Laurent, Émile or Marcel, was the collector of the type. We did not find a sheet annotated with this name in BR. We have selected a neotype in the Brussels herbarium collected by Marcel Laurent in 1905 (BR0000014801273) for Solanum laurentii Dammer (= Solanum subcoriaceum De Wild. & Durand) that matches the description in Dammer’s (1906) protologue.

Dammer (1906) described Solanum plousianthemum citing three collections made by C. Holst in Tanzania (Holst 232, 3731, 8927). Bitter (1917) later cited Holst 232 and Holst 3731 as elements in his redefinition of Solanum plousianthemum , and effectively lectotypified the name by stating "letztere Nr. der Typus zu Dammer’s originaldiagnose" - the latter number [Holst 3731] the type of Dammer’s original diagnosis). He cited Holst 8927 as the type for his variety conglutinans without citing a herbarium. Edmonds (2012) superfluously lectotypified Solanum plousianthemum with the Kew duplicate of Holst 8927, none of the other collections cited in Dammer’s protologue have been found. We select the Paris duplicate (P00341605, with flower and fruit) of Holst 8927 as the lectotype of Bitter’s var. conglutinans .

Similarly C.H. Wright (1894, 1906) rarely cited herbaria in new taxon descriptions, but because he worked at the herbarium of Kew, we have designated lectotypes at that institution unless a much higher quality specimen was available elsewhere (e.g., Solanum welwitschii where of the two syntypes Welwitsch 6081, 6098 the BM sheet of the better documented syntype Welwitsch 6098 has both flowers and fruits). In some cases (e.g., Wright 1897) the herbarium at Kew was mentioned in the introductory material in the publication, thus Scott-Elliot 6800 at Kew (K000096900) is the holotype of Solanum nakurense . However, in the case of Solanum lykipiense where Wright (1894) states " the specimen from which this description is drawn is about 6 in. long, but sufficient to show that the species is a very distinct one", we have lectotyified the name with the K sheet which answers to this description rather than taking it as the holotype because there is no evidence that there were not several sheets with small fragments. Two collections, Schweinfurth 3498 and Mann 274 were cited in the protologue of Solanum welwitschii var. strictum ( Wright 1894: 127); we select the Kew duplicate (K000414054) of Mann 274 as the lectotype because another sheet is in Paris and we have only seen one duplicate of Schweinfurth 3498. For Solanum phytolaccoides , we have selected Schimper 310 as the lectotype, rather than the other collection (Johnston s.n.) cited, so that can be used as the lectotype of Solanum plousianthemum var. connmixtum . In herbaria where we have examined duplicates of Schimper 310 two different collection dates have been written on sheets, but the sheet in US (labelled as "ex Berol.") has both dates; all duplicates of Schimper 310 are here cited as isolectotypes, regardless of collection date.

De Wildeman and Durand (1899), Durand and Durand (1909) and De Wildeman (1922) worked at Brussels, but again did not cite herbaria in their species descriptions. For these names we have selected lectotypes at BR with the best quality duplicate available; for example, the lectotype for Solanum lianiforme is not the one without the original label, the sheet we selected (BR0000008993489) has label in De Wildeman’s hand with the species name and is a flowering specimen. The late Richard Lester affixed holotype labels to specimens at BR (e.g., Thonner 96 the type collection of Solanum symphyostemon ), and his “designations” were followed by Edmonds (2012). We have, in general, used these sheets as the lectotypes, unless duplicates in BR are of better quality.

Two collections were cited in the description of Solanum balboanum ( Chiovenda 1935); Balbo 425 and Balbo 579. We have selected Balbo 579 (FT003031) because it has both flowers and fruits.

Selected specimens examined.

ANGOLA. Cuanza Norte: Munza , 8 Apr 1870, Schweinfurth 3498 (K) . Cuanza Sul: Kumbira Forest , 12 Jun 2014, Goyder & Maiato 7749 (K) . Huíla: Bumbo, Serra da Chela , Oct 1859, Welwitsch 6032 (BM) . Luanda: Cazengo , 1903, Gossweiler s.n. (BM) . Malanje: Pungo Andongo, Welwitsch 6107 [a] (BM, K, P); Mata de Cabondo , Feb 1857, Welwitsch 6107 [b] (BM, G) . Zaire: Ganji , 1910, Gossweiler 5092 (BM) .

BURUNDI. Bujumbura: Karonge , 17 Jun 1966, Lewalle 981 (BR, EA) . Bururi: Kwitaba , 13 Dec 1977, Reekmans 6747 (BR, EA, K, P) . Cibitoke: Mayabi, territoire Bubanza; Mabaye , riv. Lua , frontière du Rwanda, 2 Mar 1969, Lewalle 3249 (BR) ; Gitega: pâturages naturelles de Kitego [= Gitega ], 8 Dec 1922, Elskens 239 (BR); territoire Bubanza; riv. Nyamagana, 21 Mar 1971, Lewalle 5343 (BR); Nyabiraba, 4 Jun 1971, Reekmans 698 (BR) . Karuzi: Karuzi , 19 Dec 1957, Lebrun 11269 (BR) . Makamba: Kininya, Kininga [=Kininya], Mosso , 1 Jul 1952, Michel 3159 (BR) . Muramwya: Teza , 23 Jan 1980, Bouharmont 14177 (BR); territoire Muramvya; Bugarama, 2 Dec 1967, Lewalle 2446 (BR); Matara, 9 Sep 1973, Reekmans 2939 (EA); Kiganda, 4 Jan 1978, Reekmans 6848 (BR, EA, K) . Ngozi: Gundava , Mar 1935, Becquet 861 (BR) . Rutana: Kiofi, 15 Mar 1952, Michel & Reed 1572 (BR); Kininya, Mosso , 18 Jun 1952, Michel 2915 (BR) . Ruyigi: Nyabitare, 1 Jun 1952, Michel 2604 (BR); Kigamba , 12 Oct 1978, Reekmans 7184 (BR, EA, K) .

CAMEROON. Centre: 2 km NE of Nguélémendouka, near Bamo River, 9 Apr 1962, Breteler 2744 (K, P); Mefou Proposed National Park, Ndanan 1, 100m after bridge, 10 Mar 2004, Cheek & Nana 11597 (BR, K); Ndokononoro, a 15 km au SW de Ndikiméki, 20 Dec 1971, Letouzey 10894 (P); Nkol Emana, pres Mebo-Mezoa, 12 km SE Ngomedzad, soit a 45 km au SSW de Mbalmayo, 27 Jun 1972, Letouzey 11385 (P); Colline Nkolandjom, près Ngoakélé, à 25 km à l’W de Ngoulémakong (route Mbalmayo-Tbolowa), 12 Jul 1972, Letouzey 11471 (P); Mefou Proposed National Park, along the dry Mbeme river bed to Ndaligan 1, 25 Mar 2004, Tadjouteu et al. 575 (BR, K). Est: Somalomo, 69 km SE Akonolinga, 17 Jun 1981, Asonganyi 306 (P); 15 km N of Bertoua along road to Deng Deng, 15 Mar 1961, Breteler 1205 (K, P); Gounté, ca. 27 km from Bertoua, along road to Betaré-Oya, 11 Dec 1961, Breteler 2200 (BR, K, P); Djaposten, 38 km N of Lomié, right bank of Dja River, near bridge in road to Abong Mbang, 17 Sep 1965, Leeuwenberg 6779 (BR, P); Yokadouma, 6 km along road to Batouri, 21 Jun 1961, Breteler 1533 (BR, K, P); a 4-8 km de 'Ene de Djolempoum, 26 Apr 1961, Letouzey 3933 (P); Laye, 30 km W de Batouri, 5 Apr 1962, Letouzey 4661 (P); Mopwo, 16 km a l’WNW de Mopwo, village située au km 22 de la route Yokadouma-Batouri, 4 Jun 1963, Letouzey 5221 (P); entre Ngatto & Zoubalot, a 95 km au SW de Yokadouma, 31 Mar 1973, Letouzey 12219 (BR, K x2, P); Boubara, 40 km NE de Batouri, 11 Apr 1962, Letouzey 4712 (P); Deng Deng, Apr 1914, Mildbraed 8838 (K); entre riv. Bango et Djombi, 27 km E de Batcha Ilo, village située a 25 km N de Moloundou, 9 Apr 1971, Villiers 636 (NY, P); 5 km W Bateka Mahambe village situé à 26 km NE de Moloundou, 16 Apr 1971, Villiers 654 (BR, K, P). Littoral: Mt Nlonako, flanc NW, 25 Dec 1967, Bamps 1564 (BR, K, P). Ouest: Ndé, Bamena, about 15 km W of Banganté, 29 Apr 1964, de Wilde & de Wilde-Duyfjes 2327 (BR). Nord-Ouest: Bambili, 3 Mar 1970, Bauer 44 (K); Boyo, Tum, from Aboh, path towards Tum, 22 Nov 1996, Etuge & Pollard 3576 (K); Bamenda, Fongom, May 1931, Maitland 1690 (K, P); Mezam, Bali Ngemba F.R [Forest Reserve], 10 Apr 2002, Onana 1982 (K); Mezam, Mantum, Bali Ngemba Forest Reserve, 12 Apr 2002, Pollard et al. 953 (K); Bafut-Ngemba F.R. on the secondary bush by the side of Forest Nursery no. 1, near the boundary, 7 Apr 1951, Ujor FHI - 30026 (K). Sud: Lolodorf, route a Ebolowa, 17 Jun 1918, Annet 401 (P); Colline Ongongondje, près Akonekye, 15 km NW de Amban, 23 Mar 1970, Letouzey 10216 (P); Okpweng, 40 km route Djoum-Oveng, 17 Nov 1966, Mesili 56 (BR); N’Koemvone, 14 km on the road from Ebolowa to Ambam, along the Seng river, 11 Jan 1975, de Wilde 7893 (BR, P); Ebolowa, ca. 15 km S of Ebolowa, 1 Mar 1964, de Wilde & de Wilde-Duyfjes 2023 (P); Bipinde [Bipindi], 1913, Zenker, G. 4675 (BM, G, GOET, LE, P, S). Sud-Ouest: Mount Cameroon, Bimbia, 4 km SE Limbe (TB), 3 Aug 1993, Baker 231 (K); Mount Kupe, Kupe Village, path between Kupe Rock and main trail to summit, 16 Jul 1996, Cable et al. 3860 (K); Mount Cameroon, Bakingili, above Bakingili on hunters path leaving village behind school, through extensive old German bamboo plantation to 'red hill’ a steeply ascending ridge of red soil, 21 Dec 1993, Cheek et al. 5843 (K); Ngombombeng, near Ngombombeng village, north of Nyassosso, Apr 1986, Etuge & Thomas 31 (BR, K, P); Mount Cameroon, Mount Etinde, Fako Division, Buea District, near Ekonjo, 1 Sep 1993, Hunt 64 (K); Mount Cameroon, Bimbia, Mabeta-Moliwe, 2 Apr 1992, Jaff 39 (K); Mount Cameroon, Likombe, 15 May 1992, Mbani 119 (K); Fako, Buea, Upper Boando, between U. Boando and Etome, 25 May 1992, Mbani 142 (K); Mount Cameroon, Boa, Operation Raleigh plots, 5 May 1994, Ndam 1347 (K, SCA); Konye Esamanga Road, western Bakossi, 17 Feb 1988, Nemba & Mambo 786 ( US); Mount Cameroon, Etome, along an old timber exploitation road SW`of the Camp, 31 Jan 1997, Nning et al. 200 (K, SCA, YA); Mount Cameroon, Mabeta, Mabeta-Moliwe Reserve area 1, 12 Jan 1992, Sunderland 1055 (K, SCA, YA); W Bamboutos Mts, Fosimondi, Fossimondi village, alongthe road leading to government school, 28 Apr 2005, Tchiengue et al. 2232 (K); Mount Cameroon, Mount Etinde, Batoke Path, 28 Oct 1992, Tchouto 478 (K, SCA, YA); Barombi Camp, 5 km S of Kumba on Buea/Douala Road, 21 Jun 1983, Thomas 2176 (NY); Fako, Bakingili, trail up Mt Cameroon, 9 Jul 1984, Thompson et al. 1555 (BH); Nyasoso, Mount Kupe, 4 Jun 1996, Zapfack et al. 682 (K).

CENTRAL AFRICAN REPUBLIC. Haute-Kotto: 60 km W de Yalinga, Le Testu 3910 (BM, BR, P); 39 km S de Yalinga route de Bangassay , 30 Jun 1922, Le Testu 3971 (BM, BR, P) ; Lobaye: Boukoko, Riviere Dungwa, 20 Mar 1923, Tisserant 1251-1040 (P); Mbaiki, Boukoko, Nzua-Motoli (Lissango), 4 Aug 1950, Tisserant 1861 (BM, G, P x2); Mbaiki, Boukoko, Station Centrale de Bounoko , Nzua-Motoli (Lissango), 16 Apr 1951, Tisserant 2078 (BM, G, P x2) . Mambéré-Kadéï: Berbérati, forêt de la mission Catholique , 1 May 1937, Tisserant 3490 (BM, P) . Nana-Mambéré: region de Bouar, Sep 1960, Koechlin 6257 (P) . Ouaka: Riv. Ngumanga, Bambari, region L’Oubangui, Jun 1921, Tisserant 286 (P); Moronbas , 20 Mar 1923, Tisserant 1040 (BM, P) .

CÔTE D’IVOIRE. Lagunes: Yapo-Nord , ad orientem-oppidi, 15 Mar 1962, Bernardi 8681 (G, P, US) ; Orumboboka, 40 km S of Toumodi, 13 Jun 1968, Bokdam 2789 (BR); forêt du Yapo, 11 May 1967, Cremers 643 (BR, P); km 9 of Yakassé Mé-Kodiosou road, 31 Jul 1970, Leeuwenberg 8068 (BR, K, P). Bas-Sassandra: Soubré, 70 km S of Soubré, 3 Apr 1968, Geerling & Bokdam 2489 (BR); 61 km N. of Sassandra, W of Niapidou, 520 Jan 1959, Leeuwenberg 2518 (K). Comoé: Mbasso, Indénié, Region de Moyen-Comoe , 25 Dec 1909, Chevalier 22652 (P). Montagnes: Mont Tonkoui, 6 Mar 1968, Aké Assi 9938 (G); Mont Nimba, sur la crête, entre le point E et le point 1380, 10 Feb 1989, Gautier-Beguin & Gautier 1133 (G); Mont Nimba, près de la base IFAN du Mt. Nimba (Guinee francaise), 23 Dec 1946, Roberty 6680 (G); Mont Tonkoui, Jan 1950, Schnell 4079 (P); Monts Nimba [border with Guinea], Apr 1950, Schnell 5341 (P);); piste Guessabo, Mont Do, 31 Jan 1969, Bamps 1995 (BR) .

DEMOCRATIC REPUBLIC OF THE CONGO. Bandundu: Kaniama, Jan 1939, Brédo 2721 (BR); Kibulu, [=Kibubu], 7 Feb 1950, Callens 2432 (BR); Bolobo, Demeuse 451 (BR); Lupatapata [Luhatahata], NE Dibaya, Bakura Sumpi, crête partage Lubi-Lubilash (plateaux), 16 Feb 1937, Gillardin 202 (BR, K); Bokoro, 12 Nov 1947, Jans 621 (BR); Loozi, [Lozi], 19 Feb 1904, Luja 151 (BR); Kikwit, 28 Oct 1993, Masens 1341 (BR); Bas-Katanga, Kaniama-Haut Lomami, 24 Feb 1947, Mullenders 1520 (BR); Popokabaka, 29 Nov 1958, Pauwels 778 (BR); Lukombe, Oct 1910, Sapin s.n. (BR); Kikwit, Jan 1914, Vanderyst 2881 bis (BR); region Babunda, Mar 1921, Vanderyst 8980 (BR); Ipami [Ipamu] a Kikval, Jul 1921, Vanderyst 9814 (BR); Lemfu, Kistanu-Kwango [=Inkisi], Vanderyst 14229 (BR); Kwango, Goa, 1926, Vanderyst 19168 bis (BR). Bas-Congo: Thysville, [Mbanza-Ngungu], 3 Jun 1915, Bequaert 7747 (BR); Luki [Luki Biosphere reserve, Mayombe Forest], Brixhe 19 (BR); Kingedi, Terr. Lukula, 19 Aug 1959, Compère 128 (BR); Mission de Lemfu, 30 Mar 1944, Germain 2007 (BR); Kinanga, 1874, Gillet s.n. (BR); Twataba, 1909, Gillet s.n. (BR); Kisantu, [=Inkisi], Gillet 717 (BR); Sanda, Apr 1903, Gillet 3016 (BR); Gimbi, 2 May 1958, Laurent 562 (BR); Sona Gangu, cataractes [Sona Gungu], 21 Feb 1898, Luja 112 (BR); Ndembo, 14 Apr 1911, Vanderyst s.n. (BR); Kinkosi, 26 Feb 1907, Vanderyst s.n. (BR); Lazaret du Sacré Coeur [Kisantu, =Inkisi], Mar 1911, Vanderyst s.n. (BR); Monte Ngala, Kisantu [=Inkisi] - Kwango, Jun 1925, Vanderyst 14444 (BR); Temvo, 17 Feb 1919, Vermoesen 964 (BR); Boma, INEAC- LUKI, 16 Feb 1959, Wagemans 2172 (BR); Takalu, 29 Aug 1921, Wellens 286 (BR). Équateur: Bongabo, terr. Gemena, 8 Jun 1971, Breyne 1703 (BR); Lokolenge, Bruneel 46 (BR); Eala-Bamania, 18 Aug 1912, Chevalier 28197 (P); Mbandaka, Eala, 19 Apr 1931, Corbisier-Baland 1034 (BR, K, P); Station Inéac Boketa, 14 Sep 1955, Evrard 1788 (BR); Monkoto, route Wafania-Biyanga, 13 Nov 1957, Evrard 2975 (BR, K); Likimi, Mobeka, Likimi, Jan 1913, de Giorgi 69 (BR); Yambata, de Giorgi 1344 (BR); Yambata, de Giorgi 1824 (BR); Bikoro, environs, Lac Tumba, Dec 1920, Goossens 1522 (BR); Pasha, environs, Lac Tumba, Dec 1920, Goossens 2384 (BR); Monsole, Jul 1923, Goossens 4209 (BR); Sud Ubangi, Kalo, Dist. de Mbangi, May 1924, Goossens 4318 (BR); Bokuma, 028 Feb 1941, Hulstraert 567 (BR); Wafanya, 20 Dec 1941, Hulstraert 592 (BR); Porte de Moma, à un kilometre au porte sur la route vers la rive de la Tshuapa, Nov 1910, Jespersen s.n. (BR); Tshuapa, Belo, (Ikela) porte de Belo, 1Oct 1910, Jespersen s.n. (BR, K); Bakasu, 17 Sep 1905, Laurent s.n. (BR); Wendji, environs de Coquilhatville [Mbandaka], May 1930, Lebrun 307 (BR); Bolombo, Eala, May 1930, Lebrun 438 (BR); entre Bokuma et Bokatola, Jul 1930, Lebrun 1354 (BR, EA, P); Businga (Ubangi), Jan 1931, Lebrun 1972 (BR); Buta (Uele-Gamberi), Mar 1931, Lebrun 2483 (BR); Bili (Uele-Gambiri), May 1931, Lebrun 2828 (BR); Mobwasa, 15 May 1913, Lemaire 202 (BR); Mbandaka, Eala, route de Coq, 7 Oct 1946, Leonard, A. 802 (BR, EA, P); Mongo, 25 Sep 1935, Louis 146 (BR, EA, FT); Likimi, environs a Likimi, 9 Sep 1910, Malchair 300 (BR); Mbandaka, Coquilhatville [Mbandaka], 019 Jul 1906, Pynaert 274 (BR); Dundusana, Reygaert 226 (BR); Parc National Monkoto, Parc National Monkoto, au sud de Booke (terr. Kole) [=PN de la Salonga], May 1958, Robin 67 (BR); Eala, route de Bolombo, Territ. Coquilhatv[ille], Sep 1925, Robyns 684 (K); Equateur, Mabali, Lac Tumba, I.R.S.A.C., Terr. Bikoro, 14 Feb 1957, Thonet 154 (BR); Bokondji, 7 Sep 1959, de Wankel 148 (BR). Kasaï-Occidental: Kasai, Luiza, Nov 1921, Achien 639 (BR); Hempt St Ben, 1 Jan 1917, Callewaert s.n. (BR); Kakenge, terre du Bakuba, Dec 1937, Gillardin 317 (BR, K); Concession Elkasai a Tumba (Terr. Luisa), 26 Feb 1957, Liben 2571 (BR); Kasongo, Bateteur [Kasongo Batetela], 1906, Sapin s.n. (BR); Tshibangu, Feb 1910, Sapin s.n. (BR); Tshibangu, Jan 1910, Sapin s.n. (BR). Kasaï-Oriental: Sankuru, Kole, Oct 1904, Claessens 299 (BR); Katako-Kombe, (Kindu), Jun 1860, Claessens 453 (BR); Sankuru, Bena Dibele, Oct 1904, Claessens 949 (BR); Mukumari, Lomela, May 1939, Gillardin 586 (BR); Sankuru, entre Lusambo et la Lomami, Dec 1898, Laurent s.n. (BR); Bombaye [Bombaie], 27 Nov 1903, Laurent & Laurent s.n. (BR); Gandajika, [Gandajika], 10 Dec 1956, Liben 2065 (BR); Mwenetontole [Mwene Tonto], 1 Feb 1936, Luxen 384 (BR). Katanga: a Kaziba, col. Kachika (district Mitwaba), 2 Sep 1958, Laurent 542 (BR); Rivière Mwera, 10 km S de Kiembe (Haut Shaba), 5 Feb 1982, Malaisse & Robbrecht 2378 (BR); Kapanga, Nov 1933, Overlaet 1086 (BR); Kankima River, galerie de Kankima, Jul 1939, Quarré 39 (BR); Gorge de Kalaba, Jul 1939, Quarré 698 (BR); Muhila, [Bugana-Muhila] Ferme Bertrand, Oct 1945, Quarré 7051 (BR); Elisabethville, [Lubumbashi], Jun 1920, Rogers 26252 (G); Lualaba, Dilolo, 1935, Vin 45 (BR). Kinshasa: Tshangu, Maluku, route Mabana km 3, 27 Nov 1970, Breyne 981 (BR); Leopoldville, Territ. Lukula, route Kimbuya-Kingedi, 24 Aug 1959, Compère 160 (BR, K); Leopoldville, [Kinshasa], 30 Jan 1907, Vanderyst s.n. (BR). Maniema: Tschopo, Namoya, Mwendamboko Hill, 12 Apr 2008, Bytebier & Luke 2784 (BR); Kindu, chemin forestier de Bweni, 15 Apr 1959, Gaillez 281 (BR); Usega, Waniema [Maniema], Jul 1932, Lebrun 5705 (BR); Kamusuku, territoire de Shabunda, 18 Aug 1959, Leonard 5974 (BR, K); Muhulu, de Kyampsumba, a 42 km au NNW de Kolweli, 6 Dec 1983, Schaijes 2088 (BR); Pope, Kasonga Lunda, 1925, Vanderyst 17358 (BR). Nord-Kivu: route Goma-Costermanville, environs de Goma, 24 Jun 1953, van der Ben 548 (BR, K); Beni, 2 Apr 1914, Bequaert3344 (BR, K); Irangi, 70 km W of Lake Kivu, 26 Aug 1959, Cambridge Congo Expedition 1959 428 (BR, US); Kahele, Irangi, pres Camp ICRSAC a Irangi, km 110 route Kavamu-Walikale, 13 Aug 1956, Christiaensen 1835 (BR, CAS); Bwito, Station Kikondo, Colline Shinga, 23 Mar 1954, Deru 102 (BR); Colline Kashaka, route Goma-Sake, 11 Jan 1945, Germain 3323 (BR); Kokola, km 100 Irumu-Beni, Aug 1939, Gille 267 (BR); Kililema, Terr. Masisi [=Kilima], 29 Aug 1957, Gutzwiller 1541 (BR); Lugandamo, Biega [Mont Biega], May 1947, Hendrichx 4602 (BR); plaines de lavas entre les lacs Kivu et Edouard, Apr 1929, Humbert 7915 (BR,P); chaine des Virunga (Kivu), Volcan Karisambi, Lebrun 4933 (BR, G, K); Kasindi-Lubango (Kibali-Ituri), chaine W de Lac Edouard, 1932, Lebrun 4788 (BR, K); Nyiragongo, entre Kibali et Nyiragongo, Nov 1937, Lebrun 8692 (BR); Lac Muguga [Mugunga], Nov 1937, Lebrun 8856 (BR); Rutshuru, Kivu, Dec 1937, Lebrun 8979 (BR, K); Kirambo, Terr. Masisi [=Kilumbo], 15 Jan 1959, Leonard 2562 (BR); Ruzizi, Kahuzi, [Kahuzi], 28 May 1960, Meurillon 931 (BR); Musisi, route Bukavu-Bitale, 13 Mar 1959, Pierlot 2771 (BR, K); km 193 route Butavu-Goma, abords de cette ville, 3 Jun 1959, Pierlot 2947 (BR); Kisharo, km 30 route Rutshuru-Katwe ( près de la frontière de l’Uganda), territoire de Rutshuru, 8 Jun 1959, Pierlot 3090 (BR); Irangi, Territ. Kalehe, route Kavumu-Walikale, vers km 110, Irangi, Reserve IRSAC, 10 Jul 1957, Troupin 3795 (BR, K); Mutsora, Parque Nacional Albert, Talya, 17 Apr 1953, de Witte 8843 (BR, K); Mont Muhete, près Mutsora, 24 Feb 1954, de Witte 9866 (BR); Lusilube, affl. droit Semliki (P.N.A.=Virungas National Park), 13 Sep 1954, de Witte 11084 (BR); N’Jomaluini, marais près N’Zinga, P.N.A. [Virungas National Park), 12 Mar 1955, de Witte 11977 (BR); Tsliaberimu Mt. [Tschiaberimu], 15 Apr 1955, de Witte 12153 (BR); Saya-Sago, rig. Mt. Huyo [Mount Hoyo], 114 Jul 1955, de Witte 12531 (BR); Kideduya, près riv. Lusilube, 13 Sep 1955, de Witte 12689 (BR); Riv. Talya affl. Lumi, près Mutsora, 29 Mar 1956, de Witte 13040 (BR); Riv. Kenyamoshii, affl. droit Murve (Kashubu), 7 Aug 1956, de Witte 13224 (BR). Orientale: Ituri, Foret de Djugu, Riv. Ruida, 17 Apr 1958, Bamps 160 (BR); Yangambi, île Esali, 4 Mar 1959, Bamps 392 (BR); Barumbu, 30 Oct 1913, Bequaert 1036 (BR); Yambuya, 21 Nov 1913, Bequaert 1242 (BR); Avakubi, 13 Jan 1914, Bequaert 1915 (BR); Ituri, Irumu, 9 Mar 1914, Bequaert 2859 (BR); Yafake Commuinity, Lilanda, 7 May 2010, Boyekoli Ebale Congo Expedition 2010 271 (BR); Bomane, Aruwimi River, 21 May 2010, Boyekoli Ebale Congo Expedition 2010 658 (BR); Bambesa, et environs, 1933, Brédo s.n. (BR); Haut-Uele, Dingila, 4 Jul 1933, Brédo 135 (BR); Isangi, Yangambi ( réserve forestière), 12 Feb 1979, Breyne 3604 (BR); Barumbu, Dec 1920, Claessens 27 (BR); Bafwaboli, Claessens 308 (BR); Nebasa, Apr 1921, Claessens 531 (BR); Dinda, May 1921, Claessens 634 (BR); Ituri, Lekwa, Djugu, 6 May 1959, Deville 341 (BR, CAS); Basankusu, terr. de Mbale, District Tshuapa, Jun 1934, Dubois 432 (BR); Stanleyville [=Kisangani], 16 Dec 1898, Duchesne 2 (BR); Rucu (Mahagi), 27 May 1959, Froment 451 (BR); Ituri, Berunda, (Mahagi), 12 Jun 1959, Froment 539 (BR); km 24 route Kole-Kanwa, Terr. de Banalia, 13 Jun 1932, Galdermans 11 (BR); Tukpwo, 1 Apr 1954, Gerard 1286 (BR); Bambesa, 12 Nov 1956, Gerard 2519 (BR); Nadabu, (Zobia), 15 Apr 1957, Gerard 2812 (BR); Kurukwata, (Aba), 10 Aug 1957, Gerard 3562 (BR); Kurukwata, (Aba), 2 Sep 1957, Gerard 3681 (BR); Rubi, (Buta), 28 May 1958, Gerard 3918 (BR); Ituri, zone de Mambasa, along trail to Aketu Hill from abandoned coffee plantation in Nzaro (about 90 km N of Mambasa on road to Isiro, 8 Mar 1994, Gereau et al. 5457 (EA, MO). Bambesa, 3 Jul 1942, Gilbert 465 (BR); Haut-Uele, Riv. Guruba (Doruma), May 1936, de Graer 550 (BR); Mulimba, Mutongo, territoire Walikale, 1 May 1958, Gutzwiller 2641 (BR); Kaseke, Mutongo, territoire Walikale, 2 May 1958, Gutzwiller 2703 (BR); Ibucha, Mutongo, Terr. Walikale, 12 May 1958, Gutzwiller 2883 (BR); Mumu, Mutongo, Terr. Walikale 19 Jun 1958, Gutzwiller 3174 (BR); Epulu, Zone de Mambasa, Ituri, 114 Jul 1988, Hart 863 (BR); Bamanya [Bamanga], Bosako, 31 Mar 1945, Hulstraert 1392 (BR); Ituri, Djugu (Tibali - Ituri), Jul 1931, Lebrun 3941 (BR, P); 500 m au SE de Kabondo, Kisangani, 30 Apr 1977, Lejoly 1444 (BR); Ile Kongolo, a la confluence avec le fleuve Zaire [Congo], 10 Feb 1979, Lejoly 4774 (BR); Kitshanga, Terr. Walikale, 17 Feb 1959, Leonard 2826 (BR); Haut-Zaïre, route Kisangani-Opala, 2 Jun 1973, Lisowski 18587 (BR); Tschopo, Yangambi, réserve-flore-Isalowe, 13 Jan 1938, Louis 7498 (BR, NY); Lilanda, route d’Isangi, près de la basse Lilanda, 9 May 1938, Louis 9280 (BR, FT); Ile Idjwi, Lake Kivu, 1 Mar 1939, Loveridge 639 (K); Mobwasa, Reygaert 1360 (BR); Haut-Uele, Garamba National Park, piste Wilibadé, source Nambia-Boisquet, 18 Jul 1948, Robyns 3125 (BR); bords Kidi - Gombari, 26 Jan 1906, Seret 576 (BR); Ituri, Reserve de Djugu, 29 Oct 1951, Smeyers 72 (BR); Tshopo, Panga, Popoy, km 139 de la route Banalia-Panga, près dernier pont avant Panga, 20 May 1987, Szafranski 1277 (BR); Mbolohu, Chefferie Wanundi, piste Eringete, 17 Apr 1944, de Wilde 367 (BR, CAS); Mont Mangalaba, près Talia Nord, rég. Tshiaberimu, 17 Mar 1954, de Witte 9919 (BR); Bikito, près Malibotu affl. Ilimbo, 2 Jul 1954, de Witte 10827 (BR). Sud-Kivu: Lukando, 15 Feb 1936, Babault 845 (P); Idjwi, Lac Kivu, Réserve C N Ki, Kalambo, 229 Sep 1953, van der Ben 909 (BR); Tahibati, Kabare Territ., 14 Aug 1953, Christiaensen 5 (BR, NY); Kabare, Lwiro, Centre ICRSAC, 4 Jul 1958, Christiaensen 2444 (BR, CAS); Lubarika, Plaine de la Ruzizi, 30 Jan 1950, Germain 5920 (BR,K); Kalongé, Ruwenzori, Aug 1932, Hauman 199 (BR); Tshizimwho [Mont Kahuzi], 1 Oct 1940, Hendrickx 1288 (BR, K); Ndjondohe, Tshirumbi, Jan 1946, Hendrickx 3844 (BR); Kavumu, Luhasagula, May 1946, Hendrickx 4060 (BR); entre Kahuzi et Biega, Nov 1946, Hendrickx 4246 (BR); Ile Idjwi, Lac Kivu, May 1929, Humbert 8333 (P); Parc National Virunga, massif de Ru wenzori, entre Mutwanga et la gite Kalonge, 28 Dec 1977, Lejoly 2339 (BR); Mikonzi, vallée de la Tshinginda, km 42 route Kavamu-Walikale, territoire de Kahele, 12 Feb 1957, Pierlot 1461 (BR); forêt de la Musisi, (Kahusi), territoire de Kabare, 7 May 1958, Pierlot 2006 (BR); Kahuzi, an der Strasse Bukavu-Walikale (Kahuzi Sattel), 14 Jan 1955, Stauffer 1043 (BR, K, P); Centre Iwiro, IRSAC, riv. Iwiro [Lwiro], Kabare Territ., 21 Jun 1958, Troupin 7517 (BR, NY); le long de la Butahu, pres de Kalongé, Parc National Albert, 19 Aug 1952, de Witte 8006 (BR, CAS); Kalongé, versant droit de la vallée de la Butahu, en fase de Kalongé, 13 May 1953, de Witte 9015 (BR).

EQUATORIAL GUINEA. Bioko Norte: Fernando Po , El Pico, 10 Dec 1951, Boughey 190 (K); carretera del pico Basilé, km 6-7, 12 Dec 1989, Carvalho 4232 (K, MA) . Bioko Sur: Gran Caldera de Luba, descenso en al dirrecion a Moraca 28 Feb 1990, Carvalho 4274 (K, MA); Musola, Nov 1942, Guinea 1094 (MA); Fernando Po, Missola [Musola], trocha de servicio Agronómico, 10 Jan 1947, Guinea 1144 (K); Finca Puente, km 17, carretera de Musola a Moka, 5 Mar 1950, Guinea 1666 (MA); Homiga Camp, 23 Jan 2009, Luke 13206 (EA, K, MA) . Centro Sur: Parque Nacional de Monte Alén, Ocuam , 26 Nov 1998, Pérez Viso 623 (BM, K, MA) . Litoral: Bata-Niefang, estrada Km 35, en zonas de Adjape e de Comayá, entre os Rios Sama e Comayá, 18 Feb 1995, Carvalho 5814 (BM, K, MA) .

ERITREA. Maekel: Hamasen, Asmara , 15 Oct 1909, Baldrati 407 (FT); Hamasen, Asmara, 15 Oct 1909, Baldrati 3532 (FT) .

ETHIOPIA. Addis Ababa: Addis Ababa, 23 Jan 1963, Tekle 220 (K). Amhara [?]: Lötho, 11 Nov 1857, Schimper 480 (E, FT, G x2, P). Amhara: Agew Awi, Injiabare [Injiabara] to Chilga, 31 Oct 1995, Friis et al. 7011 (K); Dessie, Wollo province, 23 Aug 1946, Hall 98 (BM); 34 km N of Bahar Dar, along the road to Gondar, counted form the bridge crossing the Blue Nile River, 17 Sep 1970, de Wilde 7173 (BR). Gambella: [Gojam floristic region] 30 km W of Tepi, 21 Jul 1993, Renya 77 (ETH); Illubabor region , Bishan Waqa Lake [Bishan Wak’a Hayk’], 24 Sep 1996, Tesfaye 408 (ETH). Oromia: Bale, Robie [Bale Robe], 11 May 1975, Ash 2932 (BR, K, US) ; West Shewa, Ambo [Hagre Hiwof], Sep 1938, Benedetto 26 (FT); Illubabor, Gore, towards Tepi , 4 Nov 1986, Gilbert & Friis 8417 (ETH, K); Harerge, 22 Feb 1932, Gillett 5149 (K); Belleta Forest , 718 Apr 1997, Pavlov 97113 (ETH); East Welega , 5 km E of Nekemt [e], 16 Apr 1965, de Wilde & de Wilde-Duyfjes 6315 (ETH). Southern Nations , Nationalities and Peoples Region : Keffa, 20 km SW of Mezan Tefari (Aman), 29 Nov 1984, Friis et al. 3842 (ETH, K); Kibre Mengist to Wondo, 12 Apr 1975, Gilbert & Jones 187 (K); Gidole, [Dirashe], 22 Jan 1985, Haugen 440 (ETH); Leliso River , 18 Nov 1979, Mesfin Tadese 759 (ETH); Jimma, 5 Dec 1955, Stewart B - 19 (BM, EA, K); Keffa prov. , about 8 km along the new road for Bonga to Mizan Tefari (counted from turnoff to Bonga), 2 Aug 1969, de Wilde 5310 (BR); [Nolaita] Soddo, 5 Mar 1938, Vatova 1896 (FT). Tigray: Temben [Tembien], Solazzo 284 (FT) .

GABON. Estuaire: Mont de Cristal, km 20 road Kongouleu-Assak, 8 Oct 1986, Breteler & Lemmens 8395 (BR, P); Monts de Cristal , Mont Mela , 9 Feb 1968, Hallé & Villiers 5118 (P) . Haut-Ogooué: km 20 Moanda-Mbinda, 18 Sep 1970, Breteler 6502 (BR); SSW of Moanda, 113 Oct 1970, Breteler 6869 (BR, P); between Mouila and Yeno, about 40 km. on road from Mouila, 23 Sep 1986, Breteler et al. 8127 (BR); region de Lastourville , Miçoungangui, 6 May 1930, Le Testu 8069 (BM, BR, MA, P); ca. 17 km SE of Franceville, along the road to Ndoumou, 4 km before the village of Ndjoky, 6 Dec 1989, de Wilde et al. 9942 (BR) . Ngouie: Mimongo, route de Libamba a Etéké, 14 May 1963, Hallé & Cours 5868 (K, P) . Ogooué-Lolo: Chantier Bambidie (CEB), ca. 55 km E of Lastourville, 40 km on CEB road to Okondja , 1 Nov 2005, Sosef et al. 2176 (BR) . Ogooué-Ivindo: Station d’Ipassa, 10 km S de Makokou, 11 Mar 1977, Florence 37 (P); route de Belinga km 60, 21 Mar 1978, Florence 669 (P); Nké, campement, 4 Oct 1983, Floret & Louis 1600 (MA, P); M’Passa Field Station, near Makokou on Riviere l'Ivondo , 28 Jun 1981, Gentry 33014 (MO, P); Mekambo , 20-50 km W of Mekambo, 14 Oct 1964, Hallé 2612 (P); Belinga, 11 Nov 1964, Hallé 3125 (P); Makokou , 10 km SSW of Makokou, near Ivindo R., right bank, 1 Nov 1977, Leeuwenberg 11403 (BR, P) . Wolem-Ntem: 7 km E of Mitzic, 9 Jul 1957, Jeffrey 36 (K); Medouneu, Mitzic , 2 Aug 1957, Jeffrey 199 (K, P) .

GHANA. Ashanti: Akin, Begoro district, Jan 1930, Irvine 1172 (K); Akin Ranch, Begoro district , Mar 1932, Irvine 1812 (E, K); Asin Cocoa Station , Juaso, Jul 1957, West-Skinn 19 (K) . Eastern: Atewa Range Forest Reserve, beside path in east central part near Kibi, 5 Nov 1982, Hepper 7406 (K); Aburi, 23 Nov 1898, Johnson 158 (K); Atewa Range, starting in Awhiniasi on the Kibi-Apapam road, walking W to meet the road between Apapam and the other side of the range, 30 Jun 1994, Jongkind & Abbiw 1638 (BR); Accra-Kumasi Highway, 4 km along the forest service road that intersects the highway at the village of Sagyimase, Atewa Range Forest Reserve , 1 Jul 1995, Merello et al. 1147 (K, MO); Kade, 4 Feb 1953, Morton GC - 8387 (K); Bosuso , road between Bosuso and Begaro, 9 May 1947, Robertson K - 9 (K) . Volta: Togo Plateau Forest Reserve , 19 Jun 1972, Abbiw GC - 43712 (BH, EA) . Western: Ateiku, May 1930, Anonymous 1952 (K); Nzema East, Axim , Jul 1905, Chevalier 13825 (P) .

GUINEA. Nzérékoré: Yah River, 14 Sep 1964, Adames 536 (K); Loffa, Macenta, 14 Feb 1949, Adam 3734 (P); Zoubouromai, 20 Mar 1949, Adam 4008 (P); Sérédan, May 1936, Jacques-Felix 913 (P); Boola, Mount Vtongoy, pays des Guerzés, Montagne de Boola, 16 Mar 1909, Chevalier 20935 (P); entre Lola et Nzo, Pays des Guerzés, 24 Mar 1909, Chevalier 20992 (P); Mt. Yonon, forêt classée, 19 May 1971, Jongkind 10971 (BR); Simandou range, east slope of Pic de Fon, near the water intake of Canga East mining camp, 20 Mar 2008, Burgt et al. 1139 (K); Macenta, Kouankan, 2 km près du village, 12 Jun 1970, Camara 691 (G); forest around Whiskey 2, 25 Mar 2008, Tchiengue & Haba 3136 (K).

GUINEA BISSAU. Tombali: Bedanda-Cadique , floresta de Cantanez, 5 Jan 1962, Alves Pereira 2640 (K, MA); Cantanhez-Bedanda, 1 Sep 1962, Alves Pereira 3148 (MA); Camocote, 7 Nov 1995, Malaisse & Claes 16813 (BR) .

KENYA. Central: Nyeri, 20 Jul 1929, Balbo 425 (FT); Nyandarua, Kinangop, Belcher H - 138 - 53 (EA); Kirarkop, Aberdare Mountains, Apr 1938, Chandler 2216 (BRK); Thika, Blue Post Hotel, above Thika falls, behind the Hotel., 5 Jan 1967, Faden 67/1 (EA); Aberdare National Park, 2 Feb 1932, Fries 1565 (EA); Kiambu, Muguga, 1 Aug 1960, Greenway 9700 (EA, FT, K); Kikuyu Escarpment Forest, Lari Forest Reserve and adjacent areas, Kiambu District, 26 Nov 1972, Hansen 790 (BR, EA, FT, K); Kiambu, Banana Hills, 14 Sep 1969, Hindorf 805 (BR); Muranga, Kimakia Forest Station, Essex, Aberdares, 29 Jul 1958, Kerfoot 667 (EA); Kieni Forest, 27 Oct 1974, Mungai 77 (EA); Kiambu, Kikuyu, Mar 1944, Nattrass 345 (EA); Kikuyu Escarpment Forest, along Naivasha road, opposite Ngubi Forest Reserve Guard Post, 15 Dec 1966, Perdue & Kibuwa 8252 (EA, K); Kiambu, Limuru, 10 Jul 1918, Snowden 584 (BM, K); Muguga, 13 miles on Nairobi-Naivasha road, 6 Jun 1952, Verdcourt 653 (EA, K); Nyeri, Mt. Kenya, 800 m outside the Naro Moru Gate to Mt. Kenya National Park, 14 May 2009, Vorontsova et al. 56 (BM, EA). Coast: Taita Taveta, Ngangao Forest, Taita Hills, 15 May 1985, Faden et al. 483 (EA, K, US); Taita Taveta, Chawia Forest, Taita Hills., 22 May 1985, Faden et al. 857 (EA); Taita Taveta, Mbololo Forest, Oct 1938, Joana 8996 (EA); Taita Taveta, Taita Hills, Mgambonyi, Uminga, Mgambonyi market, 25 Jan 2006, Kirika et al. 01/2009 29 (EA, K); Ngangao Forest, northern end, 26 Aug 1999, Mbuthia & Mwangangi 185 (NY); Taita Taveta, Bura, Taita Hills, top of ridge about 5 km S of Bura Mission west side of Bura Valley, 8 Apr 1998, Mwachala et al EW-1251 (EA, K, NY); Taita Taveta, Chawia Forest, Taita Hills, 18 May 1931, Napier 1132 (EA, K); Wundanyi, Mar 1939, Opiko 8758 (EA). Eastern: Makueni, Chyulu, 14 Jun 1938, Bally 7786 (K); Meru, Mount Kenya forest, Battiscombe K - 854 (EA, K); Meru, Mount Kenya forest, about 2 km from Meru, Mt. Kenya Lodge in WNW direction, 4 Jan 1986, Beentje 2691 (EA); Meru, Chuka, 13 Jun 1932, Graham 1752 (EA, K); Chyulu Hills, main forest North of Camp 3, 18 Feb 2001, Luke & Luke 7355 (EA); Mount Kenya, from camp on Mount Kenia (2550 meters) westward to the Kasorongai River (1950 meters), 17 Oct 1909, Mearns 1821 (US); Meru, Nyambeni Hills, base of Kirima Mountain near the Nyambeni Hills Tea Estate, 11 Oct 1960, Polhill & Verdcourt 297 (BR, EA, K); Marsabit, Mount Kulal, S end, Mar 1972, Tweedie 4280 (K). Nyanza: Kisii, Sep 1933, Napier 2926 (EA); Kisii, River Riaguma, 11 May 1978, Plaizier 1321 (EA); Kisii, 30 Oct 1974, Vuyk & Breteler 51 (EA). Rift Valley: Nandi, Kapsabet 6 miles to the W, 31 Mar 1961, Archer 249 (EA); Kericho, Londiani, Mt. Blackett, 11 Mar 1952, Argyle 25 (EA); Kajiado, Ol Doinyo Orok, 6 Dec 1944, Bally 4146 (EA, K); Kericho, Sotik, Cains Farm, 1981 m, 19 Jul 1945, Bally 4577 (EA, K); Kajiado, Ngong Hills, East valley, 31 Dec 1984, Beentje 1798 (EA); Uasin Gishu, Soy, Oct 1931, Brodhurst-Hill 211 (EA, K); Trans Nzoia, Saiwa Swamp National Park, 22 Mar 1992, Cheseny 488 (EA); Narok, Aitong, Oledebesi-Lemoko hillside behind, Kapkimolwa, about 45 miles North of Aitong, 28 Apr 1961, Glover et al. 765 (BR, EA, K); Elburgon, 20 miles from Olokurto on road to Elburgon, Nakuru District, 14 May 1961, Glover et al. 1092 (BR, EA, K); Uasin Gishu, Eldoret-Elgeyo escarpment, Harger s.n. (BM); Elgeyo Marakwet, Kapsowar, Kipkunurr Forest Reserve, E end of Cherangani Hills, 14 Apr 1975, Hepper & Jaeger 4976 (EA, K, P); Trans Nzoia, Mount Elgon, Dec 1931, Jack s.n. (EA); Samburu, Leroghi Plateau, Karissia Forest, near Maralal, 30 Jul 1960, Kerfoot 2116 (EA); Kericho, Tinderet Forest Reserve, Camp 4, Nyanza Province, Londiani District, 29 Jun 1949, Maas Geesternanus 5246 (BR, K); Nakuru, Eastern Mau Forest Reserve, Camp 9, Rift Val ley Province, Nakuru District, 25 Aug 1949, Maas Geesternanus 5894 (BR, G, K); Mau Forest Reserve, Nakuru District 24 Nov 1971, Magogo 1517 (EA, K); Mau, summit (Mau Summit), 1928, Mettam 178 (DS, K); West Pokot, Kapenguria, West Suvi, 6 May 1953, Padwa 65 (EA, K); Kericho, 6.5 miles to the East in forest along the south perimeter of Sambret Tea Estate, 6 Dec 1967, Perdue & Kibuwa 9288 (BR, EA, K); Trans Nzoia, ‘Kavirondo’ [Winam Gulf of Lake Victoria, along Nzoia River and W slopes of Mt. Elgon], 1907, Skene 121 (G); Kajiado, Ngong Hills, Sep 1939, van Someren 9835 (EA); Trans Nozia, Mount Elgon, foothills, Kabyoyon Valley; Nov 1971, Tweedie 4135 (FT, K); Kajiado North, Ngong, top of Ngong Hills, 200 m from radar installations, 19 May 2009, Vorontsova et al. 93 (BM, EA, NY); Nandi, 2nd day’s march from Eldama Ravine, 1898, Whyte s.n. (K). Western: Kakamega, Yala River, Opposite Nature Reserve, near Quarry Hill, 26 Jan 1982, Gilbert 6880 (EA, K); Kakamega, Kakamega Forest, NW part of forest, 6 May 1971, Mabberley 1093 (K); Vihiga, Malaba Forest, Mar 1963, Tweedie s.n. (K).

LIBERIA. Nimba: Clairiére, 5 Jan 1965, Adam 20447 (K); Mt. Alpha Mine, upper Yiti River , 16 May 1973, Adam 27565 (P); Mt. Alpha Mine , upper Yiti River , 16 May 1973, Adam 27588 (P); Yiti Valley , crête orientale, 6 Aug 1974, Adam 28907 (P); Trig Point 36, 22 Nov 1964, Adames 779 (K); Mount Nimba , upper edges of the forest on Mount Nimba , 14 Dec 1966, Bos 2393 (BR, K, P) .

MALAWI. Central: Dedza Mountain Forest , 19 Jan 1987, Balaka 1859 - B (K, NY); Ntchisi Forest, 2 Feb 1983, Dowsett-Lemaire 618 (BR); Nchisi Mountain, 19 Feb 1959, Robson 1662 (BM, K) . Northern: Misuku Hills, Mugesse Forest, 1 Mar 1983, Dowsett-Lemaire 667 (BR); Chitipa, Muyumba, above the Waterfield’s house, Misuku District, 1926, Maitland s.n. (K); Mzuzu , Vipya Plateau, E of Champira , Lwanjati peak, 11 Jan 1975, Pawek 8927 (EA, K); Mpalampala, highest elevation on Vipya, top of Mpalampala, Nkhata Bay District, Vipya Plateau, 30 miles SW of Mzuzu, 13 Feb 1977, Pawek 12349 (K); Nkhata Bay District, 5 miles E of Mzuzu, 1 May 1977, Phillips 2164 (BH); Chikangawa , Nkhata Bay District, 3.5 miles SE of Chikangawa, 10 Sep 1978, Phillips 3899 (K); Chitipa, Misuku Hills, 10 Jan 1969, Richards 10596 (K); Mzimba, Mtangatanga Forest Reserve , Mzimba District, 5 Nov 1968, Salubeni 1195 (K) . Southern: Zomba District, Zomba Plateau, 27 Jan 1979, Blackmore 208 (BM, K); Cholo Mountain, Cholo District, 16 Feb 1970, Brummitt & Banda 8591 (K); Zomba Plateau, by Mlunguzi River near Trout Ponds, Zomba District, 18 Apr 1970, Brummitt 9975 (K); Naming’omba Tea Estate, at the source of Maponga Stream, Thyolo District, 25 Aug 1982, Chapman & Tawakali 6377 (BR, K); foot of Mt Mulanje, along Nakolu Stream about Lauderdale estate, 30 Oct 1985, Chapman & Chapman 6740 (E); Phalombi, Mount Mchese , 5 Jun 2008, Harris et al. 532 (K); on top of Majemba Hill where the flag is, under the stone, Mulanje, 9 Feb 1980, Patel & Morris, B. 479 (K) .

MAYOTTE (French Overseas Department). Mayotte [Maore], Bandrélé, 14 Apr 1999, Pignal 1143 (MO) .

MOZAMBIQUE. Gaza: Chirinda zone, picada para o riacho Machecane no lado do picada, 18 Sep 1980, Nuvungaet al. 318 (BM, K) . Manica: Mossurize, (Espung abera), Serra de Espungabera (Manica e Sofala), 14 Feb 1966, Pereira et al. 1358 (BR); Mossurize, (Espungabera), Nova Sintra (Gogoi), Monte Maruma (proximo a fronteira entre os rios Macuha e Nhamoa) (Manica e Sofala), 1 Mar 1966, Pereira et al. 1585 (BR); Chimoio, prox. da Misao de Amatongas (Manica e Sofala), 10 Dec 1943, Torre 6283 (BR, EA); Muza, Vallée du Muza, Vasse s.n. (P) . Zambezia: Milange, Serra do Chiparana, subida pelo chefo Marraga , vertiente voltada a NE, 28 Jan 1972, Correia & Marques 2365 (BR); Gúruè, serra do Gúruè, ao km 3 a seguir a cascata picada para o pico Namúli (Gf), 23 Feb 1966, Torre & Correia 14805 (K, P) .

NIGERIA. Akwa Ibom: Eket, 1912, Talbot s.n. (BM); Bakosi Proposed Reserve. Kumba district , 19 May 1951, Olorunfemi 30589 (K) . Cross River: Sankwala area , 30 Dec 1948, Savory & Keay FHI - 25255 (K); Oban, 1911, Talbot s.n. (BM); Obudu Plateau, 1 Nov 1964, Tuley 1058 (K) . Edo: Irhue, Ehor Forest Reserve, Ehor District , 2 Jun 1974, Eimunjeze Oguntayo FHI-72752 (K) . Ondo: Akuru Forest Reserve , 15 Aug 1946, Jones s.n. (K) . Osun: Omo Forest Reserve, near PSP.208 A’s boundary, 25 May 1960, Emwiogbon FHI-44266 (K); Shasha Forest Reserve , 20 Apr 1935, Richards 3373 (BM, BR) . Taraba: Mambilla Plateau , between Kara and Titong, Sardauna District, 9 Aug 1973, Chapman 88 (K); Mambilla Plateau, 28 Apr 1972, Chapman 2777 (K) .

REPUBLIC OF CONGO. Brazzaville: Brazzaville, 22 km N de Brazzaville, Plateaux Batiké, 3 Feb 1970, Makany 1528 (P). Lekoumou: Komono, piste SW, 17 Jan 1965, Bouquet 957 (P); Meyah, 19 Dec 1969, Hallé 1715 (P); Fôret de Bangou, près de la grotte Yengo, 6 Feb 1970, Hallé 1795 (P). Niari: Niari region, Col de Bamba, 8 Dec 1990, La Croix 5027 (BR). Pool: Brazzaville, 32 km N de Brazzaville (route Nord), 20 Jan 1971, Makany 1673 (P). Sangha: Nouablé-Ndoki National Park, Goualougo Study Site, 37.92 km E de Bomassa., 17 Apr 2007, Ndolo E-52 (E); Nouablé-Ndoki National Park, Goualougo Study Site, 36.85 km E- NE de Bomassa, 18 Dec 2008, Ndolo E-410 (E).

RWANDA. Eastern: Birenga, pref. Kibungo, route de la frontière du Burundi au S de Bare (sector Igahara), 22 Jun 1978, Raynal 20648 (P); Forêt de Mukura au N et près de la roche de Ndaba, Isabinyama, Kivumu, Pref. Kibuye, 18 Mar 1980, Runyinya 958 (BR); Mutara, environ de Mimuli, reserve IRSAC, Terr. Biumba, 23 May 1957, Troupin 3196 (BR, EA, K); Mutara, colline Karukwanzi, Terr. Biumba, 9 Mar 1958, Troupin 6631 (BR,K); Mutara, colline Gikandura, Terr. Biumba, 14 Nov 1958, Troupin 8773 (BR, NY); Mutara, colline Nteko, 17 Jan 1959, Troupin 9614 (BR); Mutara, Mimuli, Reserve IRSAC, Colline Ntako, rivière Kakitumba, 10 Jun 1959, Troupin 11702 ( US). Kigali: Tarabana, Kigali, May 1933, Becquet 674 (BR). Northern: Gishwati, Forêt de Gishwati, route de Gisenyi à Kibuye, a 32 km de Gisenyi, 6 Feb 1972, Auquier 2434 (BR); Lac Bulera, riv. Kabaga, Pref. Ruhengeri, 15 Feb 1972, Bamps 3143 (BR); Rukoro, Kidalho, Ruhengeri (pref) Graniko, 23 Mar 1962, Nshorere 121 (BR). Southern: Rubona INEAC, Colline Shyunga, 21 Oct 1958, Michel 5794 (BR); Rubona INEAC, Colline Gashikeri, 2 Dec 1958, Michel 5879 (BR); Uwinka, route Bukavu-Astrida, environ d’Uwinka, colline Bunyanguru, Terr. Sangugu, 12 Mar 1959, Troupin 9826 (BR, EA). Western: forêt de Rugege (Cyangugu), au km 91 de la route Butare-Cyangugu, 24 Jul 1974, Auquier 3368 (BR); Nyungwe, Forêt de Nyungwe (Cyangugu), a km 1.3 de la route de Pindura a la frontière du Burundi, 26 Jul 1974, Auquier 3438 bis (BR); Cyangugu, route Butare-Cyangugu vers km 100, 10 Jan 1980, Bridson 187 (BR, EA, K); Gisovu, Wisumo, 17 Feb 1980, Bridson 415 (BR, EA, K); Uwinka, piste Uwinka- Rangiro, forêt de Nyungwe, pref. Cyangugu, 3 Mar 1980, Bridson 488 (K); entre Sisovu et Wisumo, pref. Kibuye, 12 Mar 1973, Troupin 14630 (BR); Karisimbi, lava field, 25 Feb 1972, van der Veken 9577 (BR).

SÃO TOME E PRINCIPE. São Tomé: Lagoa Amelia , pic de M. Café, Sep 1905, Chevalier 14284 is, (P); S. Nicolau, 1885, Moller 95 (BM); Bom Successo, Jul 1885, Moller 404 (BM); Vermelho, Roça Sandade, 1885, Moller 516 (BM); Monte Café, Lagina S. Pedro, Oct 1885, Quintas 920 (BM); Bom Sucesso, chemin Lago Amelia, station Jachere, 31 Jul 1997, Stevart & de Oliveira 8 (BR); Angra, Angolares, 1886, Juintas 1018 (BM) .

SIERRA LEONE. Troma , 16 Dec 1865, Jaeger 8531 (K). Eastern: Tingi Mountains, 12 Dec 1965, Morton & Gledhill SL - 2959 (K) .

SOUTH AFRICA. Eastern Cape: 7 miles NW of Port St. Johns, 1949, Skinner & McGough s.n. (US); Eggosa Forest , 22 Aug 1969, Strey 8863 (K) . KwaZulu-Natal: Ndandhla Forest , 10 miles SE of Nkandhla, 12 Jun 1946, Codd 1393 (K); Inanda, [Hillcrest], Apr 1879, Medley Wood 559 (K); Zwaartkop, 16 Feb 1907, Medley Wood 10241 (P); Otto’s Bluff, Jul 1931, Rump s.n. (NU); Eshowe, Saunders 9 (K) . Limpopo: Hanertsberg, 21 Oct 1938, Acocks 1374 (S); Mahonde Mission Station, 15 miles NE of Sibosa, Dist. Zoutpansberg, 19 Feb 1952, Codd 6844 (K); km 5 from Haenertsburg to Pietersburg, 5 Dec 1975, Crawford 354 (K); Zoutpansberg, Entabeni Forest , environs de Louis Trichardt (Transvaal) Zoutpansberg Mountains , 7 Sep 1933, Humbert 10617 (P); Pietersburg, Mountain Home Farm , Woodbush area , 40 miles E, 2 Jan 1945, Mogg 17435 (BM, BR, K, P); Modjadjes, Div. Pietersburg [Polokwane], Dec 1915, Rogers 18111 (K); Zoutpansberg, Louis Trichardt, Jun 1918, Rogers 21326 (BM); Rosendal, near bridge across 'Mati stream, between the waterfalls, Dist. Duiwelskloof [Modjadjiskloof], 24 Jun 1958, Scheepers 396 (K); Zoutpansberg, ca. 9 miles W of Louis Trichar [t] farm, Rustfontein, 9 Nov 1955, Schlieben 7555 (BR, K, NY, US) .

SOUTH SUDAN. Equatoria: Loubouti, Lafuka, Imatong Mtns., Mt. Baghanj , 14 Jun 1939, Andrews 1430 (K); Talanga, Imatong Mountains , 4 Dec 1980, Friis & Vollesen 683 (K); Mondalia, Mongalla, Kagelu, Hamdi 38 (K); Lotti Forest , 24 Jun 1953, Jackson 3045 (K) .

TANZANIA. Arusha: Monduli, Ketumbeine Forest Reserve, ca. 3 km S of Ketumbeine Peak on foot trail ca. 4 km N of Elang’atadapashi, 1 Apr 2000, Gobbo et al. 674 (MO, NY); Ngorogoro, E edge of Malanja, 8 Mar 1966, Henlocker 342 (K); Monduli, Ketumbeine Forest Reserve , 5 Apr 2000, Massawe et al. 708 (MO, NY); Ngorongoro, near Safari Lodge, 2 Feb 1961, Newbould 5699 (EA, K, US) ; Arumeru, Mount Meru, Arusha National Park Near Miriakamba Hut , 9 Oct 1997, Phillipson et al. 4933 (MO); Ngurdoto Crater National Park , Arusha region , 5 May 1965, Richards 20353 (EA, K); Mount Meru , Kilinga Forest , 6 Apr 1971, Richards 26922 (EA, K); Mount Meru , Jekuhama, Arusha National Park , Arusha District , 6 Feb 1970, Richards 25346 (K); Ngorongoro Crater, E side of crater, 8 Sep 1968, Robinette & Gilbert 3125 (EA); Monduli, Monduli Forest Reserve , Mwandeti Forest near Kriganyisho pond, 22 Jan 2001, Simon et al. 671 (MO, NY); Arusha National Park , Nasula camp, 18 Aug 1969, Vesey-Fitzgerald 6393 (EA). Dodoma: Kolo Hills, C. Katikati, Kolo na Beroko , Katissa mhina, 17 Jan 1938, Burtt 1141 (K). Iringa: Mufindi, forest road from Ngwazi to Lugoda, 20 Mar 1986, Bidgood & Keeley 368 (EA, K); Ilongelo, near Kiboma, Mar 1954, Carmichael 419 (EA); Luhega Forest Reserve , Iringa District , 20 Feb 1996, Frimodt-Moller et al. NG - 041 (K); Ludewa, Livingstone Mountains , on saddle N of summit of Msalaba Mountain , near foot trail from mission at Luana, 11 Jan 1991, Gereau & Kayombo 3522 (BR, G, MO); Mufindi, Lake Ngwazi , Dam end, 9 Apr 1986, Lovett et al. 632 (K, MO); Iringa Rural, New Dabaga/Ulangambi, 15 Jul 1999, Massawe DA - 75 (K, MO); Mufindi, Lulanda, village, SW of Fufu forest, 19 Oct 1998, Mwangoka et al. 66 (MA, MO, P); Uzungwe Mountains National Park , Kilolo District , W of Kidatu dam, 5 Nov 2005, Mwangoka et al. 4530 (EA); Dabaga Highlands, Ihangana Forest Reserve , near Kibengu, 18 miles South of Dabaga, 14 Feb 1962, Polhill & Paulo 1479 (BR, EA, K); Ibumu Village , about 30 miles E of Iringa and S of the Gt. N Road, 14 Dec 1961, Richards 15628 (K). Kagera: Bukoba Rural, Minziro Forest Reserve , Nyanja Etagera Forest , 8 Aug 2000, Festo 702 (MO, NY); Bukoba Dist. , Minziro Forest , Nov 1958, Procter 1055 (EA,K); Ngara Dist. , Mumwendo, Mumwendo Ferry, 11 Feb 1960, Tanner 4703 (BH, BR, EA, K); Buseke, Keza, Bushubi, Ngara District , 20 May 1960, Tanner 4930 (BH, DUKE). Kigoma: Gombe Stream Reserve , Sleeping Buffalo ridge, 14 Feb 1970, Clutton-Brock 387 (EA); Kigoma, Kakombe Valley , Gombe National Park , at base of lowest waterfall, 6 Mar 1996, Gereau et al. 5930 (MO); Kabogo Mts , 17 Feb 1963, Kyoto University Expedition 325 (EA); Gombe Stream Reserve , Middle Kakombe, 5 May 1992, Mbago & Lyanga 1109 (K); Mkenke, Gombe River , Feb 1972, Parnell 2284 (EA); Mahali Mountains , Mt. Mhensabantu , 9 Jan 1974, Uehara 1-80 (EA). Kilimanjaro: Rau, Moshi District , Feb 1928, Haarer 995 (EA, K); Masai Dist. , Ngorongoro, Conservation Area , Olmasiko on Endulen road, 15 Dec 1965, Herlocker 246 (EA); Same, Kanza Village , Gonja Ward, along Yongoma River just below bridge, 3 Dec 2000, Kindeketa 526 (MO, NY); Masai Dist. , Makuyuni, W. Usambaras, Jun 1935, Koritschoner 609 (EA, K); Marangu, Kilimanjaro, 22 May 1961, Machangu 67 (EA, K); Same, Mponde Village , South Pare Mountains. Mbaga Ward , Manka/ Mponde Mwanyesi Sacred Forest , 13 Nov 2000, Mlangwa & Kaniki 1094 (MO, NY); Same, Shengena Forest Reserve , Gonja Bombo Beat, W side of Mvaa Subvillage near forest boundary, 8 Aug 2001, Mlangwa & Msangi 1596 (MO, NY); Pare Dist. , Muheza, Amani Bustani, 28 Oct 1969, Ngoundai 404 (EA, K); Kilimanjaro, track to Shira Plateau, Moshi District , 12 Feb 1969, Richards 24031 (BR, K); Lemosho, Kilimanjaro, 1 Jan 1967, Robertson 445 (EA, K); Lyamungu, C.R.S., Moshi District , 21 Dec 1943, Wallace 1164 (EA, K); Marangu, Kibo Hotel, 7 May 1962, Wright 6 (K). Lindi: Rondo Forest Reserve , Rondo Plateau, Lindi District , 20 Feb 1991, Bidgood et al. 1674 (K); Rondo Forest Reserve , Lindi District , 22 Nov 1966, Gillett 17990 (EA, K). Manyara: Mbulu Dist. , Oldeani, 6 Sep 1968, Gilbert 3518 (EA); Ngorogoro, Oldean Mountain , 1 Jan 1989, Pócs & Chuwa 89-002/C (K); Pienaars Heights, Great North Road , 12 miles S of Arusha, Mbulu District , 5 May 1962, Polhill & Paulo 2341 (BR, EA, K); Nou Forest Reserve , Tomati, 5 Jan 1968, Trut 576 (EA). Mbeya: Mount Rungwe , W slopes, Lukuyu District , 11 Mar 1932, St. Clair-Thompson 977 (K); World’s End View, Ipinda, Mbeya Range, 6 Feb 1976, Cribb et al. 10579 (K); Poroto Mountains , Chumvi Forest , above Irambo, 8 Feb 1979, Cribb et al. 11340 (EA, K); Mbeya Peak, Myembe, Mbeya Peak Forest Reserve , 28 Oct 1958, Gaetan 93 (EA); Mbeya Range, 25 Nov 1961, Kerfoot 3251 (EA); Rungwe Dist. , Rungwe Mt. , top of mountain, 8 Mar 1950, Milton 56 (EA); Rungwe Mountain , N slope, Tukuyu District , 8 Feb 1961, Richards 14331 (EA, K); Elton Plateau, 30 Nov 1963, Richards 18480 (EA, K); Mount Mbeya , Chunya Escarpment, Mbeya District , 10 Dec 1963, Richards 18618 (K); north of Lake Nyasa , Nbaka Oberlauf, Station Kyimbila [Rungwe District], 1 Mar 1912, Stolz 1159 (BM, K). Morogoro: Nguru Mountains , 11 Dec 1966, Beesley 285 (BR, K); Chenzema, 1 km before town on roadside, Uluguru Mountains , 26 Jan 1976, Cribb & Grey-Wilson 10441 (K); Mount Mnyera , Ukaguru Mountains , Kilosa District , 31 Jan 1976, Cribb & Grey-Wilson 10489 (K); Ulanga Dist. , Mahenge , Nawenge Forest Reserve , 2.5 km SW of Mahenge, 20 Feb 2006, Gereau et al. 6891 (EA, NY); Mwere River , Uluguru Mts. , 26 Sep 1970, Harris et al. 5155 (EA); Mamiwa Forest Reserve , Ukaguru Mountains , ridge to the N of Mandege Forest Station , Kilosa District , 2 Aug 1972, Mabberley 1346 (K); Maskati Mission, Nguru Mountains , 29 Jan 1991, Manktelow et al. 91009 (K); Morogoro Rural, Uluguru Mountains , Bunduki Forest Reserve , 29 Aug 2000, Mhoro UMBCP - 474 (K, MO); Morogoro Rural, Nguru South Forest Reserve , Kwechazungu, Dikulula River , 21 Aug 2004, Mwangoka et al. 3305 (MO); Ilole Forest , Mikumi Divison, Kiluma District , 9 Feb 2007, Mwangoka & Kidanka 5121 (BR); Ukaguru Mountains , along track between Mandege and Vingwete, Kilosa District, 3 Jun 1978, Thulin & Mhoro 2916 (K). Njombe: Njombe, Matamba, Kitulo Plateau above Matamba, below waterfall on headwaters of Ndumbi River , 22 Nov 1986, Brummitt & Congdon 8112 (K); Njombe, Aug 1931, Hornby & Hornby 36 (EA, K); Kipengere Mountains , Nyarere River , Njombe District , 9 Jan 1957, Richards 7639 (K); Elton Plateau, foothills to Kepengere Mtns. [Kitulo Plateau], 26 Jan 1961, Richards 14194 (K). Ruvuma: Songea, Mitanga area , Magengu, 21 Oct 1956, Mgaza 120 (EA, K); Songea, Lupembe Hill, 29 Feb 1956, Milne-Redhead & Taylor 8909 (BR, EA, K). Shinyanga: Shinyanga, 11 Mar 1937, Burtt 5568 (BM, EA). Singida: Singida Rural, Mangonyi, Ikungi Div., about 25 km on rd E & S from Ikungi, then 1.2 km SW along old track 17 Mar 2002, Kuchar 25227 (MO, NY); Itigi, Road Itigi-Singida, Itigi thicket, 26 Mar 1965, Richards 19926 (K). Tabora: Igunga road, Tabora region , 28 Nov 1978, Lawton 2155 (K). Tanga: Vugiri, Lushoto District , 12 Jun 1963, Archbold 216 (K); Kwakomo Forest Reserve , East Usambaras , SE of Kwakoro Tea Estate , Muheza District , 28 Oct 1986, Borhidi 86260 (K); Kitivo-Lushoto, 7 Mar 1974, Damaris 2 (K); Bumbuli, West Usambaras , 1.5 miles NE of Sakarani, 6 May 1953, Drummond & Hemsley 2410 (BR, EA, FT, K); Soni, W Usambaras, Lushoto District , 24 Jun 1971, Faulkner 4603 (BR, K); Korogwe, Ambangulu Forest Reserve under TFCG control, SE of Makweli village near tea estate plot #18, 1 Feb 1999, Mwangoka 289 (MO, P); Korogwe, Tamota village in cultivated area along road to village, 13 Apr 1999, Mwangoka 418 (MO, P); Korogwe, Kieti Village , forest patch near Mbaha River and tea estate forest, 24 May 1999, Mwangoka 578 (MO); Kwetango, village, NE part of forest, Muheza district , 23 Feb 2008, Mwangoka et al. 5638 (BR); Mkusa Forest Reserve , Lushoto District , 17 Jan 1967, Richards 22005 (K); Lushoto, Camphor near Staff-quarters, 1 Mar 1969, Shabani 318 (BR, EA, K); Magamba Forest Reserve , near the Magamba Forestry Office, 23 Mar 2010, Tepe et al. 2783 (BM, DSM); Korogwe, Kunga Tea Estates, West Usambaras ; Bungu, Masange Forest , 22 Mar 2010, Vorontsova et al. 163 (BM, DSM); Bomole, 10 Aug 1916, Zimmermann 8065 (EA) .

UGANDA. Central: Entebbe, Victoria Nyanza, 21 Apr 1905, Bagshawe 679 (BM); Buikwe, Mabira Forest, Kyagwe, Sep 1933, Brasnett 1388 (K); Mengo, Kajansi, Entebbe, Jun 1935, Chandler 1247 (BR, EA, K); Mabira forest, W. Umbango, Jan 1916, Dümmer 1387 (BM, P, US); Mulange, Maburu, Nov 1922, Dümmer 5608 (K); Mabira Forest, 2 km N of Namanyama-Mabira, 1 Jun 1971, Katende 964 (EA, K); Kisimbiri, on the trail from Entebbe, Victoria Nyanza, to Butiaba, Albert Nyanza, 23 Dec 1909, Mearns 2457 (BM, NY, US); Entebbe, mile 13 Kampala-Entebbe Road, Feb 1931, Snowden 1919 (K). Eastern: Teso, Serere, Sep 1932, Chandler 959 (EA, K, S); Mabura, Sep 1920, Dümmer 4492 (K); Busoga, Lolui [Dolwe Islsand], Jun 1960, Goodall 19 (EA); Mount Elgon, Dec 1931, Mrs Jack s.n. (K); Mbale, Namasindwa, Mt. Elgon, 25 May 1924, Snowden 889a (EA, K, P); Elgon, Namasindo, Snowden 889 (BM, BR); Bugishu, Mount Elgon, Namasindo, Snowden 890 (BM, K); Bunya, Busoga, Nov 1937, Webb 62 (K). Northern: Karamoja, Morongole [Mount Morungole], Jun 1946, Eggeling 5655 (EA). Western: Budongo Forest Reserve, Masindi District, Nyakafunjo Block, ca. 2.5 km N of Nyabyeya Forestry College, 18 Jun 1998, ATBP 554 (MO); Kireba Gap, Kigezi District, area between hills and camp, Dec 1938, Chandler 2644 (BR, K); Bundibungyo, Semliki Forest, 1905, Dawe 672 (K); Kinkisi, Siba Forest, Kinkizi County, Bunyoro District (NE Albert); about 4 miles S of Susingoro hill, 25 miles W of Masindi, near boundary of coupes 4 and 5 of Siba concession, 21 May 1951, Dawkins D - 751 (K); Kabula, on Masaka-Ankole border, Oct 1932, Eggeling 695 (K); Mbarara, Mar 1968, Harrington 302 (EA); Ntungamo, Ankole, Ruizi River, 21 Nov 1950, Jarrett253 (EA); Echuya Forest Reserve, 25 Apr 1970, Katende K - 221 (EA, K); Bushenyi District, Central Kashoha forest, 21 Sep 1987, Katende 3197 (MO); Kibale National Forest, edge of Cyperus swamp near Rweimba bridge, N of MUBFS, 13 Aug 2003, Knapp & Mallet 9811 (BM, NY); Bwindi Impenetrable Forest, Kayonza Sector, Ishasha Gorge, 10 Feb 1998, Eilu 247 (K); Bujuko, Mwakoto County, 2 km E of Bujuko, on Mubende road, 26 Feb 1969, Lye 1950 (K); Bunyaruguru, Aug 1938, Purseglove P - 358 (K); Kigezi District, path from Chehafi to Behungi, 2 miles from and behind lake (N.q.), 13 Jan 1933, Rogers & Gardner 358 (BR, K); Buhweju, Kamukaki-Rugongo, 31 Jul 1992, Rwaburindore 3430 (MO); Budongo Forest, Bujenje, 16 Jul 1971, Synnott 617 (EA, K); Kigezi District, Virunga Mountains, pass betwen Mgahinga and Sabinio, 21 Nov 1934, Taylor 1875 (BM); Bukenyi, Ankole, 2 Sep 1938, Thomas 2520 (BR, K).

YEMEN. Ibb: Wadi Bana, c. 15 km south of Yarim, c. 15 km east of main Sana’a to Taiz Road , near As Saddah , 13 Mar 1981, Miller 3025 (E); Jebel Badaan [Jabal Ba’dan], 26 Aug 1977, Wood 1864 (BM) . Sana’a: Bei Hajjera, Menacha , 13 Mar 1983, Müller-Hohenstein & Deil 1353 (E) .

ZAMBIA. Copperbelt: Chingola , 25 Aug 1954, Fanshawe 1480 (K); Ndola, 5 Mar 1956, Fanshawe 2817 (BR, K) . North-Western: Mwinilunga, Mwinilunga, West Lunga River at Mwinilunga , 23 Jan 1975, Brummitt et al. 14028 (K); Solwezi , 6 km W of Solwezi, 18 Mar 1961, Drummond & Rutherford-Smith 6998 (K); by R. Matonchi-Luao junction, Mwinilunga District , 18 Dec 1937, Milne-Redhead 3723 (BR,K); Mwinilunga, 56 miles south of Mwinilunga on road to Kabompo, 3 Oct 1952, White 3442 (BM, BR, K) . Northern: Mbala, Abercorn District, Tasker’s deviation Chilongowelo , 23 Jan 1955, Richards 4231 (BR, K) .

ZIMBABWE. Manicaland: Melsetter, Bridal Veil Falls, 11 Jan 1974, Bamps et al. 763 (BR); Inyanga, Nyangami, 24 Nov 1949, Chase 1839 (BM); Umtali, Norris farm, 11 Jan 1951, Chase 3660 (BM, K); Melsetter, Sona farm, 15 Mar 1953, Chase 4872 (BM, BR); Cross Hill, Commonage, Umtali District , 4 Feb 1950, Corner s.n. (E); Umtali, Bant i Forest, 4 Feb 1955, Exell et al. 191 (BM); Makone, Rusape, Dec 1919, Eyles 5202 (K); Inyanga, 30 Oct 1930, Fries et al. 2434 (BM, BR, NY); Stapleford Forest Reserve , Central Patrol , 13 Jun 1934, Gilliland B - 314 (BM); Chimanimani, Gungunyana Forest Reserve , Melsetter, top of Chiredza Gorge, Nov 1961, Goldsmith 99/61, (K); Nyanga, Rhodes Inyanga Orchards, 23 Mar 1966, Simon 769 (K) . Mashonaland East: Wedza, Wedza Mountain , 27 Feb 1964, Wild 6343 (BR, K) .

Figures

Figure 1. Flowers of species in the ANS clade. A Solanum africanum Mill. (South Africa, Rebelo s. n. - no herbarium voucher) B Solanum imamense Dunal (Madagascar, Rakotavao 5128) C Solanum madagascariense Dunal (Madagascar, Vorontsova et al. 498) D Solanum sambiranense D'Arcy & Rakot. (Madagascar, Randrianasolo 580) E Solanum terminale Forssk. (Kenya, Vorontsova et al. 93) F Solanum truncicola Bitter (Madagascar, Antilahimena et al. 7846). Photo credits: A Tony Rebelo; B Charles Rakotavao; C, E Maria Vorontsova; D Richard Randrianasolo; E Patrice Antilahimena.

Figure 2. Habit and fruits of species of the ANS clade. A Habit of Solanum sambiranense D'Arcy & Rakot. (Madagascar, Randrianasolo 580) B Fruit of Solanum imamense Dunal (Madagascar, Rakotavao 5128) C Fruit of Solanum terminale Forssk., pointed form (Angola, Goyder et al. 7749) D Fruit of Solanum terminale Forssk., globose form (Tanzania, Tepe et al. 2783). Photo credits: A Richard Randrianasolo; B Charles Rakotavao; C David Goyder; D Maria Vorontsova.

Figure 25. Solanum terminale Forssk. A Flowering branch of savanna form (nakurense - type) B Fruiting branch of climbing plant with open infructescence with globose berries (terminale - type) C Flowering branch of climbing plant with open inflorescence D Flowering branch of climbing plant with narrow, spicate inflorescence (welwitschii - type) E Flower with fused anther cone (welwitschii - type) F Flower with non-fused anther cone (terminale - type) G Corky bark of older stems H Fusiform mature berry I Globose mature berry J " Hairy " seed from mature fruit showing elongate lateral cell walls. (Based on: A Ash 2932; B Drummond 3155; C Gilbert & Friis 8417; D, E Etuge & Thomas 31; F Greenway & Kanuri 13883; G Synnot 617; H, J Mbani 142; I Mbatchou 21). Scale bar: A, C, D, G = 3 cm; B = 4 cm; E, F, H, I = 7 mm; J = 2 mm. Drawn by Lucy T. Smith.

Figure 26. Distribution of Solanum terminale Forssk.

Figure 27. Morphology of Solanum terminale Forssk. across continental Africa. All coordinates are decimal degrees. A Bud length / width ratio plotted against longitude B Length of lowermost inflorescence branch against longitude. Red points represent plants with fused (connate anthers), green points plants with free anthers; black points represent plants with no open flowers whose bud and anther characters were not measured. For details of character choice see Materials and Methods and discussion of Solanum terminale.