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) Author Knapp, Sandra Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, United Kingdom s.knapp@nhm.ac.uk Author Vorontsova, Maria S. Comparative Plant and Fungal Biology, Royal Botanic Gardens, Kew, Richmond, Surrey TW 7 3 AB, United Kingdom text PhytoKeys 2016 2016-07-13 66 1 142 http://dx.doi.org/10.3897/phytokeys.66.8457 journal article http://dx.doi.org/10.3897/phytokeys.66.8457 1314-2003-66-1 704E9F435B62FFD93345FFD55178FFE6 128816 Solanum madagascariense Dunal, Prodr. [A.P. de Candolle] 13(1): 99. 1852. Figures 1C , 17 Solanum nitens Baker, J. Bot. 20: 220. 1882. Type. Madagascar. Fianarantsoa: "chiefly in Betsileo-land", received in K July 1880, R. Baron 145 (holotype: K [K000414183]; isotypes: E [E00193275], P [P00348975]). Solanum apocynifolium Baker, J. Linn. Soc. 20: 213. 1883. Type. Madagascar. "Central Madagascar", Oct 1882, R. Baron 1767 (holotype: K [K000414194]; isotype E [E00193276]). Solanum madagascariense Dammer, Bot. Jahrb. Syst. 38: 184. 1906. nom. illeg., non Solanum madagascariense Dunal, 1852. Type. Madagascar. Fianarantsoa: Ivohimanitra forest, Nov 1894, C.I. Forsyth-Major 15 (lectotype, designated here: K [K000414184]; probable isolectotype: BM [BM000887181]). Solanum clerodendroides Hutch. & Dalziel, Fl. W. Trop. Afr. 2: 206. 1931. Type. "Southern Nigeria, Eket District" [clearly incorrectly labelled], 1912-1913, Mr. & Mrs. P.A. Talbot 3211 (holotype: K [K000414057]). Solanum antalaha D'Arcy & Rakot., Fl. Madag., Fam. 176: 68. 1994. Type. Madagascar. Antsiranana: R.N-II, Ambohitsalonana, district Antalaha, 23 August 1950, Reserves Naturelles Madagascar [Zaty] 2738 (lectotype, designated here: P [P00349362]; isolectotypes: P [P00346372], MO [MO-277611], MO [MO-277612]). Solanum marojejy D'Arcy & Rakot., Fl. Madag., Fam. 176: 107. 1994. Type. Madagascar. Antsiranana: Reserve Naturelle de Marojejy; E-facing slope the Manantenina River, N of Mandena, 14°27'S 49°47'E , 230-550 m, 21 October 1989, J.S. Miller & A. Randrianasolo 4329 (holotype: MO [MO-150888]; isotypes: AD [AD-99109143], K [K000212283], P [P00352676], PRE [n.v., fide D'Arcy & Rakotozafy 1994], TAN [TAN000700], WAG [n.v., fide D'Arcy & Rakotozafy 1994]). Solanum madagascariense Dunal var. nitens (Baker) D'Arcy & Rakot., Fl. Madag., Fam. 176: 105. 1994. Type. Based on Solanum nitens Baker. Type . Madgascar. "In Madagascar ", Collector Unknown ( holotype : G-DC [G00144951]) . Description . Liana or small tree ( Gentry 11852 ) to 8 m. Stems terete, flattened or faintly ridged, glabrous to sparsely puberulent with simple unicellular papillae ca. 0.05 mm long, or variously pubescent with a mixture of uniseriate dendritic or arachnoid trichomes to 0.5 mm long, glabrescent or the pubescence less dense with age; new growth glabrous or pubescent with dendritic or arachnoid trichomes like those of the stems; bark of older stems smooth or longitudinally ridged, reddish brown, brown, or yellowish brown, somewhat corky on stems> 1 cm in diameter. Sympodial units plurifoliate, the leaves not geminate, evenly distributed along young branches. Leaves simple, (1.5) 3-9 (12) cm long, (1.2) 2-3 (4.5) cm wide, obovate to oblong, thick-chartaceous to thin-coriaceous, often shiny, concolorous to somewhat discolorous, glabrous, sometimes with dendritic trichomes on the midvein on both sides of the lamina; major veins 5-12 pairs, spreading at 60-90° to the midvein and forming loops, the finer venation usually faint or not visible; base cuneate to truncate, sometimes attenuate; margins entire, rarely lobed with a single lobe basally on each side, the lobes up to 8 mm long, rounded, with shallow sinuses; apex acute to acuminate, rarely obtuse and apiculate; petiole 0.4-2 cm long, canaliculate, usually glabrous, less often finely dendritic-pubescent, flexuous and sometimes twining around supports (e.g., Humbert 31597 , Miller & Randrianasolo 4339 ). Inflorescences terminal at the apex of branches, (2.5)5-20(25) cm long, furcate or several times branched, with (8)15-45(100) flowers, glabrous or with variable dendritic pubescence paralleling that of the stems; peduncle 1.5-5 cm long; pedicels 0.4-1.1 cm long, apically dilated, always glabrous, when the rachis is pubescent a clear boundary line visible between the glabrous petiole and the pubescent rachis (a few trichomes occasionally found at the pedicel base), articulated 0-2 mm from base; pedicel scars irregularly spaced 1-4 mm apart, often appearing as apically dilated pegs. Buds globose or broadly ellipsoid, the corolla soon exserted from the calyx tube. Flowers 5-merous, apparently all perfect. Calyx tube 2-3 mm long, broadly cup-shaped or conical, the lobes up to 1 mm long, 1.5-2 mm wide at base, often irregular or almost absent, broadly deltate, rounded to cuspidate at the tips, glabrous or with sparse short simple trichomes; margin often thickened, with dense tufts of simple hairs at the tips. Corolla 1-2 cm in diameter, white to violet, stellate, lobed almost to base, the lobes 4-10 mm long 1.5-2.5 mm wide, narrowly deltate to linear, sometimes aristate with a puberulous appendage ca. 0.5 mm long arising from the adaxial surface of the lobe just below the apex, glabrous adaxially, glabrous to puberulous abaxially with dendritic trichomes that are longer at the lobe tips. Stamens equal; filament tube 0.5-1 mm; free portion of the filaments 1-2 mm; anthers 2.5-4 mm long, ca. 1.5 mm wide, broadly ellipsoid, usually somewhat connivent, smooth or papillose abaxially, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated and not lengthening with age. Ovary conical, glabrous; style 6-12 mm long, protruding 1.5-4 mm beyond the anthers, straight or curved, glabrous; stigma clavate to capitate, the surface smooth to minutely papillose. Fruit a globose berry, sometimes ellipsoid, 0.5-1.2 cm diameter, black or purplish black at maturity, the pericarp thin, collapsing on drying to reveal the outline of the seeds, glabrous; fruiting pedicels 1-1.2 cm long, 0.5-0.7 mm diameter at base, pendent to spreading; fruiting calyx slightly accrescent, the lobes becoming woody with a light-coloured margin. Seeds 20-40(+) per berry, 1.5-4 mm long, 1-2.5 mm wide, ovoid reniform or somewhat flattened, golden-orange or reddish brown, the surface deeply pitted, the testal cells pentagonal or slightly sinuate in outline. Figure 17. Solanum madagascariense Dunal. A Flowering branch B Berry showing thin pericarp through which seeds are visible (based on: A, Bosser 16852 B Homelle s.n. ). Adapted from D'Arcy and Rakotozafy (1994) with permission of Museum National d'Histoire Naturelle. Distribution (Figure 18 ). Endemic to Madagascar, throughout central and eastern parts of the island, with a few records from humid forests on the western part of the island. Figure 18. Distribution of Solanum madagascariense Dunal. Ecology and habitat. Humid and subhumid forests; sometimes found in disturbed vegetation by the roadside; 0-1500 m elevation. Common names and uses. Madagascar. Antsiranana: vahimasina ( Humbert 18109 ), vahimbingy ( RN Madagascar 2738 , Miller & Randrianasolo 4563 ), vahinazo ( RN Madagascar 8388 ), voajaboala ( Miller & Randrianasolo 4329 ); Fianarantsoa: keranzy ( Anon. 4026 ), vahimaitso ( Malcomer et al. 1581 ), vahivahy ( Randriantafika 15 ). No uses recorded. Preliminary conservation status ( IUCN 2014 ). Least Concern (LC). EOO 551,384 km2 (LC), AOO 436 km2 (EN). Solanum madagascariense is a common liana, and occurs in several different forest types, including in some protected areas. Its local rarity and patchiness of distribution, along with its range of morphological varability (see below) suggest further studies as to local abundance are necessary. Discussion. Solanum madagascariense is a liana with prominent terminal inflorescences of 15-45 white to deep purple flowers (Fig. 1C ), sturdy branches, and thick leaves. It occurs throughout Madagascar's wet forests which are situated predominantly on the eastern coast, adjacent parts of the High Plateau, and northern parts of the island. It is the most common and variable species of endemic non-spiny Solanum . Solanum madagascariense is similar and possibly closely related to the rare and local species Solanum trichopetiolatum and Solanum humblotii . It can be distinguished from Solanum trichopetiolatum by its glabrous petioles (versus petioles with long simple trichomes 0.5-0.15 mm long); Solanum trichopetiolatum also has looser inflorescences with finer branches and fewer flowers, a greater tendency towards discolorous oblong leaves and is restricted to a narrow area of Antsiranana. Solanum madagascariense differs from Solanum humblotii in its large, many-branched inflorescence with many flowers; Solanum humblotii has an unbranched inflorescence with few flowers and is restricted to the northern part of Madagascar in Toamasina. Solanum madagascariense as delimited here encompasses great range of variation. It is possible to isolate groups of specimens with small long narrow leaves, groups of specimens with wide coriaceous leaves and fewer veins, and groups of specimens with sparse inflorescences and hariy leaves, but intermedidates between all these forms are common. The name Solanum apocynifolium has been used to describe individuals with tomentose stems and smaller leaves. Variation in indumentum is continuous with that observed in other populations of Solanum madagascariense , and leaf size seems to be largely determined by ecological factors. Solanum apocynifolium was accepted by Bitter (1917) , tentatively accepted by D'Arcy and Rakotozafy (1994) with acknowledgement of continuous variation, and accepted at the level of variety by the late R.N. Lester (unpublished manuscript). Forms recognised as the two taxa are the extremes of a range of morphological variation and are here considered to be conspecific. Solanum nitens is a smaller glabrous variant of Solanum madagascariense with shiny subcoriaceous leaves and a shrubby, densely branched growth form that grows in drier, more central areas of Madagascar. It was accepted by Bitter (1917) , reduced to a variety by D'Arcy and Rakotozafy (1994) and considered a synonym of Solanum madagascariense by Lester (unpublished manuscript). We agree with Richard Lester's assessment and Solanum nitens is included in Solanum madagascariense as an arid environment variant. Solanum antalaha is a wet environment variant of Solanum madagascariense occurring further north, representing mature or shade dwelling individuals. Solanum antalaha has glabrous leaves, petioles, stems, and corolla, and leaves with fewer veins; it was described as having larger anthers but these are less than 3.8 mm long on the type specimen, within the variation range of Solanum madagascariense (2.5-4 mm long). Solanum marojejy was described as a Marojejy endemic with dorsally papillose anther surfaces and glabrous petioles; both of these features, however, are commonly observed in populations of Solanum madagascariense from other localities. Solanum marojejy is here considered to be conspecific with Solanum madagascariense . Solanum clerodendroides is a specimen of Solanum madagascariense almost certainly incorrectly labelled as collected in Nigeria (fide H. Heine 1967, in sched.) Unusually large papillae seen on the abaxial anther surface of some collections have been postulated to restrict access to pollen or provide support or orientation cues for pollinating bees ( D'Arcy 1992 ). Only three specimens with lobed leaves have been seen, but lobed leaves may be more common in juvenile plants as is true in the Dulcamaroid clade (Knapp 2103). An unusual specimen at P from a cultivated plant in the garden at Antananarivo ( s. coll. 2174 , P04063654) is from a plant with inflorescences on many short branches and only 2-3 flowers per inflorescence and somewhat resembles Solanum humblotii , but morphologically conforms to Solanum madagascariense in terms of floral form, calyx size and leaf morphology. We have chosen Forsyth-Major 15 [K000414184] as the lectotype for Solanum madagascariense Dammer (nom. illeg., a later homonym of Solanum madagascariense Dunal) as it is the only unambiguous duplicate of the type collection number we have found. It is probable that the sheet at BM labelled Forsyth-Major 55 [BM000887181] is another duplicate (and thus an isolectotype), curatorial annotation on the BM sheet suggests 55 may be an error for 15. The protologue of Solanum antalaha cites a holotype at P; of the two duplicates of the type collection P00349362 has been selected as the lectotype because it annotated as "holotype" in W.G. D'Arcy's handwriting. Selected specimens examined. Madagascar . Antananarivo : Manjakandriana, Mandraka, Aug 1906 , D'Alleizette 988 (P); Tampoketsa de Ankazobe, 5-12 km E of highway 31 km N of Ankazobe, 19 May 1974 , Gentry 11852 (MO, NY); Antananarivo-Nord, la foret a l'Est d'Ambakolaona , 11 Nov 1912 , Humbert & Viguier 1265 (P); mont Kalambatitra et ses abords, Nov 1933 , Humbert 11901 (MO, P); foret d'Ankilahila , 16.2 km SE de Tsinjoarivo, le long de la riviere d'Andrindrimbola , 21 Jan 1999 , Messmer & Andriatsiferana 744 (G, K); Andramasina, Ambohimiadana, Kelilanina, Antsararoloha, 15 Dec 2009 , Rakotonasolo et al. RNF-1535 (K); foret d'Andranomay a 2 km a l'Est d'Andranomay et 13 km au SE d'Ankazobe , 20 Dec 1996 , Randrianaivo 47 (BR, MO); Ambohitantely RS, a 36 km au Nord Est d'Ankazobe , 1 km a l'Est des Batiments de l'Angap , 14 Jan 1997 , Randrianaivo et al. 59 (MO, NY, TAN). Antsiranana : Ampasindava, foret de Betsitsika, 12 Dec 2008 , Ammann et al.193 (G); Anamalaho, massif de Makirovana, au Nord-Ouest de Sambava, 22 Aug 2007 , Andriamihajarivo et al. 1256 (TAN); Tsaratanana Massif, along path from Mangindrano to Mahatsabory Mica, 19 Oct 2001 , Birkinshaw et al. 978 (MO, NY, P, TAN); montagnes entre le haut Sambirano et le haut Maivarano (entre Mangindrano et Ampanompia), Nov 1937 , Humbert 18109 (P); environs d'Andapa , bassin de la Lokoho, 1948, Humbert & Capuron 21939 (P); Reserve Speciale Manon garivo , E of Ankaramy, Bekolosy, 7 Dec 1992 , Malcomber et al. 1998 (BR, G, K, MO x2, NY, P); Reserve Naturelle Marojejy, along the trail to the summit of Marojejy Est, NW of Mandena between the first and second camps, 6 Oct 1988 , Miller et al. 3413 (K, MO x2, P, TAN); commune rurale de Daraina, foret de Binara, 20 Nov 2005 , Nusbaumer & Ranirison 1637 (G,K); commune rurale de Bealampona, village de Mandritsarahely, Sud-Ouest d'Andapa , 18 Oct 1994 , Ravelonarivo et al. 418 (MO, NY, P); Ambilobe, Beramanja, foret de Salabenono, sur la chaine Galoko, 7 km au sud-est d'Anketrabe , 25 Nov 2006 , Razafitsalama & Torze 1145 (MO, TAN); Marotolana, Ampanompy, 5 Apr 2001 , Razakamalaka et al. 104 (MO, TAN); canton de Manaka est, district d'Antalaha , Res . nat. 3, 9 Nov 1952 , Reserves Naturelles Madagascar 4480 (P); Marozato, district d'Ambanja , Res . 4, 18 Nov 1952 , Reserves Naturelles Madagascar 4499 (P, TAN); Sambava, Marojejy National Park, path down from camp 3 (camp Simpona) to camp 2 (camp Marojejia), 17 Oct 2011 , Vorontsova et al. 498 (K, TAN); Cap Masoala Grand Parc, haut vallee d'Anaovanandrano , 26 Sep 2003 , Wohlhauser et al. 648 (G). Fianarantsoa : Fianarantsoa rural, foret d'Ambondrombe , Anonymous 4026 (P); Betanatana Forest Reserve, Fargangana, Ankarana, Manobo, 28 Aug 2008 , Bussmann et al. 15257 (MO, TAN); foret basse au PK 298 (Sud Ambositra), 12 Dec 1974 , Cremers 3635 (MO, P); Vondrozo, province de Farangana, bord de route en foret , 9 Sep 1926 , Decary 5235 (P); Ambositra, Ambohimitombo, 21 Dec 1894 , Forsyth-Major 325 (K); Bassin de l'Itomampy , mont Papanga pres de Befotaka, Dec 1928 , Humbert 6912 (P); massif de l'Ivakoany , pentes orientales du massif, 1933, Humbert 12240 (P); foret ombrophile d'Ambatofitorahana , 2 Mar 1960 , Keraudren 229 (MO); Parc National Ranomafana, Parcelle I, south of Ambohimiera, valley of Sakavolo river, 15 Sep 1992 , Malcomber et al. 1581 (BR, G, K, MO x2, NY, P); Reserve Speciale de Manombo, parcelle 2, 37 km au SW de Farafangana, 21 Aug 1995 , Messmer & Rakotomalaza 46 (TAN); Reserve Naturelle Integrale d'Andringitra , 50 km S of Ambalavao, near abandoned meteorological station above Ambalamarina, 12 Jan 1987 , Nicoll 236 (MO, TAN); limite nord de la Reserve Speciale d'Ivohibe , 7.5 km ENE d'Ivohibe , 11 Oct 1997 , Rakotomalaza et al. 1398 (G, MO, NY); Ambositra, Ambalamanakana, 28 Jan 2005 , Ralimanana et al. 449 (K); foret dense humide de l'Est d'Andranobetokana , Ampasimadinika, Marofototra, Mananjary, 16 Jan 1999 , Randriantafika et al. 15 (MO); 2 km Ouest d'Andrambovato , bord de la riviere Tatamaly, haut versant, Fivondronana Tolongoina., 19 Oct 2000 , Randriantafika 176 (MO, NY, P); Ambalavao, 23 Jan 1958 , Reserves Naturelles Madagascar 9981 (P); 7 km W of Ranomafana, S of the Namorona River at Duke University Primate Center study site and along road N of river, eastern domain, 28 Oct 1987 , Schatz et al. 1713 (BR, K, MO, P); Ambalamanakana, 18 Dec 1959 , Schlieben 8197 (BM, BR, G, TAN). Mahajanga : ruisseau Ambatoharanana, Andranomena, Matsoandakana, 14 Feb 2008 , Bernard et al. 813 (TAN); Tsitondroina, 16 Apr 1941 , Herb Jard Bot Tananarive 4812 (P); Tsaravilona, Amparihy, Androva, suivant une ligne de crete vers le nord est, 26 Feb 2008 , Ravelonarivo et al. 3042 (K); Bealanana, Mangindrano, Ambohimirahavavy, Antsahivo, point cote , 20 Oct 2005 , Wohlhauser et al. 786 (MO, P). Toamasina : Parc National de Zahamena, Fivondronana, Antanandava, Antenina, 31 Jan 2002 , Andrianjafy et al. 273 (MO, NY, P); Aloatra-Mangoro, Ambatondrazaka, Didy, Antsevabe, Sahananto, Riviere de Sahananto, 12 Dec 2005 , Andrianjafy et al. 1535 (TAN); Ambanizana, Maroantsetra, Anjahana, Ambanizana, 16 Sep 2002 , Antilahimena 1405 (MO, NY); foret d'Analalava , sous-prefecture Tamatave II, Morarano, a 7 km du SO de Foulpointe, 10 Mar 2005 , Birkinshaw et al. 1442 (TAN); Ambatondrazaka, Cours, G. 1058 (P); Moramanga, Foret de Perinet a 7 km a l'Est de Perinet vers Tamatave, 28 Feb 1971 , Cremers 1481 - 3 (P); Ambodimanga a Tamatave, 11 Oct 1957 , Herbier de la Station Agricole de l'Alaotra 2807 (MO); Onibe, massif de l'Andragovalo au Sud-Est du lac Alaotra, Res . Nat. 3 dite de Zekamena, bassin de l'Onibe , Oct 1937 , Humbert 17746 (MO, P); Maroantsetra, massif de l'Anjanaharibe (pentes et sommet Nord) a l'Ouest d'Andapa , 1951, Humbert & Cours 24533 (P); Betampona Reserve Naturelle Integrale , 40 km NW of Toamasina, 27 Sep 1993 , Lewis & Razafimandimbison 639 (MO); Foret de Mantady, road to N (Tamatave), 29 Oct 1993 , Nek et al. 2021 (BR, TAN); Ikongo, Ranomafana, 50 km E of Fianarantsoa on Mananjary road, across R. Namorona from Ambatolahy, 4 Nov 1986 , Nicoll 139 (K, MO x3, P, TAN); bois des environs de la baie d'Antongil , 1912, Perrier de la Bathie 8686 (P); Vatomandry, Ambalabe, Ambinanindrano II, hameau villageois Tobin'i Foara, le long du sentier Nord, 4 Oct 2005 , Ranaivojaona et al. 1185 (MO,P); canton de Varaina Manatsambaliny est, district d'Ambatondrazaka , 13 Nov 1948 , Reserves Naturelles Madagascar 1598 (P); Parc National de Masoala, excursion d'Andranobe a Bedinta, entre Ambatoavo et Bedinda, Nov 2001 , Sauquet et al. 86 (P); Ankirindro Massif, slopes above the village Ambodivato, ca. 5 km NW of Ambinanitelo along the Vohimaro River, 20 Nov 2002 , Schatz & Antilahimena 4018 (MO, NY); Vohibinany, Anivoranokely, Ambidimanga, 21 Sep 1954 , Vigreux 15491 (P). Toliara : Betroka, Ivahona, Reserve Special de Kalambatritra. Foret d'Analamaro , 5 Nov 2005 , Andrianjafy et al. 497 (MO); Morondava, Bara, 1880, Cowan s.n. (BR,P); col de Tsitongabarika, 16 Nov 1932 , Decary 11017 (P); bassin de la Manampanihy, Col de Fitana, 15 Oct 1928 , Humbert 6050 (P); massif du Beampingaratra, vallee de la Maloto, 1928, Humbert 6302 (NY, P); N of Taolagnaro town; along Andranoroa River, border of Marosoy Forest and Andohahela Reserve, 2 Dec 1989 , McPherson 14591 (MO).