A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae) Author Knapp, Sandra Department of Life Sciences, The Natural History Museum, Cromwell Road, London SW 7 5 BD, United Kingdom s.knapp@nhm.ac.uk text PhytoKeys 2013 2013-05-10 22 1 432 http://dx.doi.org/10.3897/phytokeys.22.4041 journal article http://dx.doi.org/10.3897/phytokeys.22.4041 1314-2003-22-1 FFC8FFF7FF8EFFFE9F6EFFEEFFFF3860 576167 12. Solanum cutervanum Zahlbr. , Ann. K. K. Naturhist. Hofmus. 7: 7. 1892 Figure 31 Solanum angustifolium Ruiz & Pav., Fl. Peruv. 2: 33, t. 163b. 1799, non Solanum angustifolium Miller, 1768, nec Solanum angustifolium Lam., 1793. Type: Peru. Huanuco : Acomayo, H. Ruiz & J. Pavon s.n. (lectotype, designated by Knapp 2008c , pg. 310: MA [MA-747093]; isolectotypes: F, MA [MA-747094, MA-747095]). Solanum pulverulentum Pers., Syn. 1: 223. 1805. Type: Based on Solanum angustifolium Ruiz & Pav., Fl. Peruv. 2: 33, t. 163b. 1799, non Solanum angustifolium Miller, 1768, nec Solanum angustifolium Lam., 1793. Solanum aureum Dunal var. angustelanceolatum Bitter, Bot. Jahrb. Syst. 54, Beibl. 119: 13. 1916. Type: Peru. Huanuco : Chaglla, 3100-3200 m, c. 9°46'S, 1909-1914, A. Weberbauer 6700 (holotype: B, destroyed; lectotype, designated by Knapp 1989 : 90: F [F-628476, F neg. 69667]; isolectotypes: GH [GH00077582], MOL, US [US-1444961]). Solanum aureum Dunal var. latelanceolatum Bitter, Bot. Jahrb. Syst. 54, Beibl. 119: 13. 1916. Type: Peru. Huanuco : Chaglla, 3100-3200 m, c. 9°46'S, 1909-1914, A. Weberbauer 6700 (holotype: B, destroyed; lectotype, designated here: US [US-1444961]; isolectotypes: F, MOL, US). Type. Peru. Cajamarca: Cutervo, C. von Jelski 30 (holotype: W [W 1891-0004325, F neg. 33065]). Description. Shrubs to small trees, 1-7 m tall. Stems and leaves densely covered with loosely branching golden tree-like trichomes; leaf scars somewhat raised, the stem not winged; new growth densely pubescent with golden tree-like trichomes above and below. Bark of older stems dark reddish-brown, sparsely pubescent with the tree-like trichomes of the young stems. Sympodial units plurifoliate. Leaves simple, 6.5-13 cm long, 1.7-5 cm wide, elliptic or occasionally narrowly elliptic (type), the upper surfaces of the blades drying dark, sparsely pubescent with golden tree-like trichomes, these mostly along the veins, the lower surfaces pubescent with golden trichomes like those of the upper surfaces, the pubescence denser than that above; primary veins 8-12 pairs, sparsely pubescent; base acute, not winged on to the petiole; margins entire, not markedly revolute; apex acute; petiole 0.5-2 cm long, densely golden pubescent. Inflorescences terminal, later appearing lateral or in the fork of the branches, 4-10 cm long, pyramidal, branching ca. 10 times, with 10-20 flowers densely pubescent with loose golden tree-like trichomes like those of the young stems; peduncle 1-4 cm long; pedicels 0.7-1.3 cm long, ca. 0.5 mm in diameter at the base tapering to an apical diameter of 1 mm, slightly nodding at anthesis, densely pubescent with golden tree-like trichomes, articulated at the base and inserted in a sleeve ca. 0.5 mm long; pedicel scars closely spaced and congested at the inflorescence branch tips. Buds ellipsoid, the corolla strongly exserted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube 1.5 -2 mm long, conical, the lobes deltate, 1.5-2 mm long, densely pubescent abaxially with golden tree-like trichomes, densely pubescent adaxially with dendritic and simple trichomes. Corolla 1.5-1.8 cm in diameter, violet or occasionally white, lobed 3/4 of the way to the base, the lobes 7-8 mm long, 4-5 mm wide, planar at anthesis, densely pubescent abaxially with tiny dendritic trichomes, these denser at the tips of the lobes, adaxial surfaces glabrous. Filament tube absent; free portion of the filaments 1-1.5 mm long, occasionally slightly pubescent near the base; anthers 3.4-4 mm long, 1-1.5 mm wide, loosely connivent, poricidal at the tips, the pores becoming slit-like with age. Ovary glabrous or with a few dendritic trichomes at the apex, glabrate in fruit; style 5-7 mm long, sparsely to densely pubescent at the base or along its entire length with golden dendritic trichomes; stigma bilobed, the surface minutely papillose. Fruit a globose, purplish-black berry, with thin pericarp, 1-1.2 cm in diameter; fruiting pedicels 1-1.5 cm long, woody, nodding to more or less erect, ca. 1 mm in diameter at the base. Seeds ca. 8-10 per fruit, 3-4 mm x 2.5-3.5 mm, flattened lenticular, reddish-brown, the surfaces minutely pitted. Chromosome number: not known. Figure 31. Solanum cutervanum Zahlbr. (drawn from Hutchison & von Bismarck 6571 ). Reproduced from Knapp (1989) with permission of the Natural History Museum Botany Library. Illustration by Margaret Tebbs. Distribution ( Figure 32 ). Andean Peru from Piura to Puno with a single collection known from Bolivia; 2500-3300 m. Figure 32. Distribution of Solanum cutervanum Zahlbr. Ecology. In rocky uplands, cloud forests and along trails in forest, usually growing in the open. Common names: Peru: rama de serrano ( Knapp 1989 ). Conservation status. Least Concern (LC); EOO>100,000 km2 (LC) and AOO>10,000 km2 (LC). See Moat (2007) for explanation of measurements. Discussion. Solanum cutervanum had long been confused with and placed in the synonymy of S. nitidum . Specimens of both these species were previously annotated as Solanum pulverulentum Pers., perhaps due to their superficial similarity and sympatric distribution. The two species, however, are very different. Solanum cutervanum has golden or brownish tree-like trichomes on stems, leaves and inflorescences and black berries, while Solanum nitidum has more delicate, greyish, strictly dendritic trichomes and red ripe berries. Solanum cutervanum is closely related to Solanum ruizii , also from central Peru. It differs from that species in its often dichasial branching, rounded leaf bases, smaller flowers and deltate calyx lobes. These sister taxa are sympatric and flower size may contribute to reproductive isolation. Some populations of Solanum cutervanum from N Peru have broadly elliptical leaves superficially reminiscent of Solanum aureum , a vining species with golden strictly dendritic pubescence. Unfortunately the species name Solanum pulverulentum Pers., so long in use for this species, is a later homonym of Solanum pulverulentum L., itself a superfluous name for Solanum tomentosum L., an African prickly species (see Vorontsova and Knapp, in review ). Linneaus only used the epithet pulverulentum in the 1759 edition of his Systemae Naturae , and in later publications reverted to the use of his original epithet tomentosum for the South African species. Many monographers in Solanum have stated that holotypes or lectotypes for Ruiz and Pavon names were in the Madrid herbarium (MA), but without specifying a particular sheet. This is incorrect when several sheets are present in MA. Knapp (1989) lectotypified Solanum angustifolium citing only a sheet in MA; this was rectified in 2008 by citation of the particular sheet ( Knapp 2008c ). In describing Solanum aureum vars. angustelanceolatum and latelanceolatum Bitter (1916) used sheets of Weberbauer 6700 at B for both names; he stated that within a branch leaf morphology was relatively uniform, but that between branches (probably sheets, as he never saw these or any other South American Solanum in the field) there was a great deal of variation. He compared Weberbauer 6700 to a sheet at B collected by Humboldt (an isotype of Solanum aureum ) and clearly stated ( Bitter 1916 ) that the Humboldt sheet represented the typical variety, and that various sheets of Weberbauer 6700 were his new taxa. I have selected the F duplicate as the lectotype of var. angustelanceolatum ( Knapp 1989 ) and here selected the US sheet of the same number with slightly wider leaves as the lectotype of var. latelanceolatum as no duplicate material annotated by Bitter has been found. It is likely he was using material at Berlin (now destroyed) for his description. Specimens examined. Bolivia . La Paz : Franz Tamayo, Parque Nacional Madidi, Puina Viejo, ca. 3 km rio abajo por camino al E del rio , 3345 m, 20 Jun 2005, Fuentes 8507 (NY). Ecuador . Azuay : Canton Sevilla de Oro, Sevilla de Oro, eastern Cordillera, 4-6 km N of village, 2743 m, 14 Aug 1945, Camp E-4698 (MO); Canton Cuenca, Cuenca, Cuenca, Parroquia Cumbe, 2682 m, 3 Jul 1991, Ceron 15542 (MO); Canton Sevilla de Oro, Sevilla de Oro, 2950 m, 18 Apr 1968, Harling et al. 8454 (MO); Azuay/Morona Santiago : road Gualaceo-Limon , at the pass point, La Virgen, 3500 m, 26 Feb 1993, Harling & Stahl 26713 (MO); Loja : Las Chinchas, region central, 2250 m, 12 Apr 1944, Acosta-Solis 7786 (F); Cordillera de Las Lagunitas. Amaluza-Jimbura-Zumba, Km 36, 3390 m, 22 Nov 1994, Jorgensen et al. 747 (BM, MO); Parque Nacional Podocarpus, above Cajanuma, trail from 'Centro de Informacion' toward 'Lagunas de Compadre' , 3100 m, 19 Jan 1989, Madsen 85561 (BM). Peru . Amazonas : Luya, Camporedondo, Tullanya, base Cerro Huicsocunga, 3075 m, 7 Dec 1996, Diaz & Pena 8853 (MO); Chachapoyas, Cerros de Calla-calla, uppermost slopes and summit, near kms 403-407 of Balsas-Leimebamba road, 3400 m, 18 Aug 1962, Wurdack 1704 (USM); Cajamarca : Jaen , Sallique, 3300 m, 26 Jun 1998, Campos et al. 5102 (MO); Jaen , Sallique. Quebrada Grande, camino entre La Cocha y Tablon , 2750 m, 2 Jul 1998, Campos et al. 5183 (BM, MO); Jaen , Lanchal, La Concha, Dist. Sallique, 2960 m, 16 Jun 1998, Diaz et al. 9597 (MO, USM); Jaen , Sallique, Quebrada grande, ruta entre La Cocha y Tablon , 2770 m, 30 Jul 1998, Diaz et al. 9779 (BM, MO, MOL); Jaen , Paramillo de Pomahuaca, antes del pajonal, 3200 m, 8 Nov 1999, Diaz & Campos 10908 (USM); El Pargo, 42 km E of Llama, ca. 14 km SE of Tunas Pampas, 3000 m, 8 Sep 1991, Gentry et al. 74570 (MO, USM); El Pargo, 16 km E of Tunas Pampa, ca. 42 km E of Llama on road to Huambos, 3000 m, 18 Sep 1991, Gentry et al. 74897 (MO, USM); Chota, Bosque El Pargo, entre Llama y Huambos, 3010 m, 12 Aug 1994, Leiva G. et al. 1485 (BM); Cusco : Calca, Choquecancha, Azulcocha, Dist. Lares, 3832 m, 18 Feb 2005, Valenzuela et al. 4978 (NY); Huanuco : Huamalies , Monzon , cerros al sudoeste de Monzon , 3500 m, Weberbauer 3310 (MOL); Lambayeque : Ferranafe , Canariaco , Distrito Canaris , sector Canariaco , 3083 m, 16 Aug 2008, Marcelo Pena et al. 3688 (MOL); Pasco : Oxapampa, Distrito Huancabamba, Lanturachi, sector Santa Barbara, camino a Milpo, 2824 m, 10 Oct 2003, Perea et al. 702 (BM); Piura : Huancabamba, El Tambo, 3 Jun 1961, Acleto 294 (USM); Huancabamba, Carmen de la Frontera, alturas de Nueva York, 3280 m, 27 Jul 2006, Cano et al. 16766 (USM); Huancabamba, carretera entre Canchaque y Huancabamba, 2800 m, 14 Jan 1988, Diaz et al. 2719 (MO); Huancabamba, Mitopampa (Huancabamba-Cuello del Indio), 2650 m, 22 Jul 1975, Sagastegui et al. 8251 ( MO); San Martin : Mariscal Caceres, Parque Nacional Rio Abiseo, cerro al sur de campamento Chochas, 3500 m, 30 Jun 1996, Cano et al. 7439 (USM); Distrito de Huicongo, valle de Ruibarbos, 3650 m, 12 Jun 2001, Leon & Ramirez 5200 (BM); Mariscal Caceres , Parque Nacional Rio Abiseo, Puerta del Monte, 3200 m, Young 1573 (K); Mariscal Caceres , Parque Nacional Rio Abiseo, Chochos, 3400 m, 14 Feb 1986, Young 2809 (K, MOL, USM); Mariscal Caceres , Parque Nacional Rio Abiseo, Chochos, 3450 m, 13 Jun 1986, Young 3763 (K); Mariscal Caceres , Parque Nacional Rio Abiseo, 3400 m, 3 Jul 1986, Young 3866 (K, MOL, USM); Mariscal Caceres , Parque Nacional Rio Abiseo, along trail to El Mirador, Puerta del Monte, NW corner of Park, 3100 m, 11 Jul 1987, Young & Leon 4475 (USM).