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 17. Solanum flaccidum Vell. , Fl. Flumin. 87. 1829 Figure 43 Solanum triphyllum Vell., Fl. Flumin. 88. 1829. Type: Brazil. Rio de Janeiro: Boavista, Anon . (no specimens cited; lectotype, designated here: Vellozo, Fl. Flumin. Icones 2, tab. 120. 1831). Solanum convolvulus Sendtn. in Mart., Fl. Bras. 10: 48. 1846 . Type: Brazil. Southern Brazil, F. Sellow [46] (lectotype, designated here: P [P00335275]; isolectotypes: C [F neg. 22880], F fragment [F-617553], LE, M [Morton neg. 8689], P [Morton neg. 8165], W [W-1889/0294607, W-0003282]). Solanum fultum Schrank ex Sendtn. in Mart., Fl. Bras. 10: 49. 1846 . Type: Brazil. Minas Gerais: Sao Joao Batista, Sept, C. Martius [1071] (lectotype, designated here: M [M0171876, F neg. 6531]). Solanum delilei Dunal, Prodr. [A.P. de Candolle] 13(1): 81. 1852. Type: France: Cultivated in Paris from material said to have come from Mexico, Anon. s.n. (lectotype, designated here: G [G00357858, F neg 23109]). Solanum convolvulus Sendtn. var. pilosum Dunal, Prodr. [A.P. de Candolle] 13(1): 83. 1852. Type: Brazil. No locality or specimens cited, only reference is to Sendtner's original description. Solanum uncinellum Lindl. var. atrosanguineum Dunal, Prodr. [A.P. de Candolle] 13(1): 76. 1852. Type: Brazil. Sao Paulo: Sin. loc., C. Gaudichaud 312 (holotype: P [P00335286, Morton neg. 8358]). Solanum convolvulus Sendtn. var. heterophyllum Witasek, Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 79: 328. 1910. Type: Brazil. Sao Paulo: Sorocaba, 1902, M. Wacket s.n. (lectotype, designated here: W [W-1922-0001503]). Solanum flaccidum Vell. var. heterophyllum Witasek, Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 79: 329. 1910. Type: Brazil. Parana : Rio Cachoeira, Antonina, 1904, M. Wacket s.n. (lectotype, designated here: WU [WU0037973]; isolectotype: WU [WU0037974]). Type: Brazil. Rio de Janeiro: Sin. loc., Anon . (no specimens cited; lectotype, designated here: Vellozo, Fl. Flumin. Icones 2, tab. 115. 1831). Description. Woody vine, twining by petioles. Stems glabrous to densely white pubescent with simple uniseriate trichomes 0.25-0.5 mm long; new growth densely white pubescent with simple uniseriate trichomes ca. 0.5 mm long. Bark of older stems reddish brown. Sympodial units plurifoliate, not geminate. Leaves simple or more rarely pinnatifid with 1-3 lobes, 2.5-9 cm long, 1-4 cm wide, ovate or obovate to triangular or elliptic, membranous, the upper surfaces glabrous to sparsely pubescent with simple uniseriate trichomes, these always present on the midrib, the lower surfaces pubescent with simple uniseriate trichomes ca. 0.5 mm long, these in tufts in the vein axils to densely distributed over the entire lamina; primary veins 7-9 pairs, the midrib pubescent adaxially; base truncate to slightly cordate, often asymmetric; margins entire to lobed, the lobes to 1 cm long, pinched in at the base; apex acute; petioles 1-5 cm long, glabrous to densely white-pubescent, curling and twining. Inflorescences terminal or very occasionally lateral, 1-9 cm long, many times branched, with 10-50 flowers, glabrous to densely pubescent with simple uniseriate trichomes ca. 0.5 mm long, the transition from stem to inflorescence not always clear; peduncle ca. 1 cm long; pedicels 1-1.5 cm long, slender, ca. 0.5 mm in diameter at the base, 0.5-1 mm in diameter at the apex, spreading at anthesis, glabrous to sparsely pubescent with simple trichomes, articulated at the base from a small sleeve; pedicel scars irregularly spaced 0.5-1 cm apart. Buds ellipsoid to fusiform, the corolla strongly exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 2-2.5 mm long, conical, the lobes 1-1.5 mm long, broadly deltate, glabrous or sparsely pubescent. Corolla 2.3-2.5 cm in diameter, violet or purple, fading to white, rotate-stellate, lobed ca. 1/2 way to the base, the lobes 6-9 mm long, 5-8 mm wide, spreading or planar at anthesis, densely pubescent abaxially with simple uniseriate trichomes <0.25 mm long, glabrous adaxially. Filament tube minute, free portion of filaments unequal, one filament 3-3.5 mm long, the others 2-2.5 mm long, the long filament equalling the others early in anthesis, lengthening with flower age, all filaments glabrous; anthers 4.5-5 mm long, 1-1.5 mm wide, ellipsoid, loosely connivent, yellow, poricidal at the tips, the pores lengthening to slits with age or not. Ovary glabrous; style 8-10 mm long, glabrous; stigma clavate, the surface minutely papillose. Fruit a globose berry, 0.9-1 cm in diameter, green (immature?), probably shiny at maturity, glabrous the pericarp thin; fruiting pedicels 2-2.5 cm long, ca. 1 mm in diameter at the base (immature?), woody, pendent. Seeds not observed from mature berries. Chromosome number: not known. Figure 43. Solanum flaccidum Vell. ( A-D drawn from Nee & Bohs 50809 E drawn from Nee & Bohs 50768 ). Illustration by Bobbi Angell. Distribution ( Figure 44 ). Coastal eastern Brazil inland to Paraguay; 40-1800 m. Figure 44. Distribution of Solanum flaccidum Vell. Ecology. In forests and along forest margins and riversides. Common names. Brazil. Santa Catarina: joa-cipo ( Smith and Downs 1966 ). 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 flaccidum is a relatively common species where it occurs, and in Brazil it is found sympatrically with Solanum odoriferum , also a vining member of the Dulcamaroid clade. The two species can be distinguished by their leaf morphology (glabrous and shining in Solanum odoriferum , variously pubescent in Solanum flaccidum ), calyx shape (truncate in Solanum odoriferum , lobed in Solanum flaccidum ) and anther morphology. Solanum flaccidum has one anther with the filament longer than the other four, while in Solanum odoriferum , the filaments are all equal throughout anthesis. In early anthesis in Solanum flaccidum , the filaments appear to be more or less equal, so care must be taken to examine fully mature flowers. In Paraguay Solanum flaccidum is sympatric with Solanum uncinellum , which also has one anther borne on a longer filament. Solanum flaccidum has more rotate flowers than the latter species, and in general is less pubescent with simple, rather than dendritic, trichomes. The long filament in flowers of Solanum uncinellum is twice as long as the other filaments, while in Solanum flaccidum the long filament is less than twice as long as the others; in addition, the anthers of Solanum uncinellum are tapered and pointed, while those of Solanum flaccidum are ellipsoid. As with many species in this group, Solanum flaccidum has heteromorphic leaves, with simple and pinnatifid leaves borne on the same stems. Once a stem is flowering, it appears that the leaves are all simple, so very few collections have been made of plants with pinnatifid leaves (but see Mueller 311 at K). It may be, as with other species of this clade, that juvenile foliage is more likely to be pinnatifid than is foliage on reproductive shoots (see Solanum dulcamaroides ). In the absence of specimens ( Carauta 1973 ), lectypification of Vellozo names is best done using the plates from that work ( Vellozo 1831 ); they are distinctive and in this case have all the distinguishing features of the species (i.e., twining petioles and unequal anthers, see Figure 45 )). The name Solanum flaccidum has been in more common use (e.g., Mentz and Oliveria 2004 ) and is therefore to be preferred over the simulataneously published Solanum triphyllum ( Figure 46 ). Solanum convolvulus is lectotypifed with one of the many sheets of the syntypes that is annotated in Sendtner's hand. Figure 45. Lectotype of Solanum flaccidum Vell. ( Vellozo 1831 : tab. 115). Reproduced with permission of the Natural History Museum Botany Library. Figure 46. Lectotype of Solanum triphyllum Vell. ( Vellozo 1831 : tab. 120). Reproduced with permission of the Natural History Museum Botany Library. Although the type of Solanum delilei was said to have been grown from Mexican seeds and the description does not mention unequal anthers, the specimen at G has the unmistakable anther morphology and is vegetatively a perfect match for Solanum flaccidum . Dunal cited material he had seen cultivated in Paris and Montpellier, but the only herbarium material unambiguously cited is a sheet from "h. Delessert", here designated as the lectotype. Mix-ups as to the provenance of plants grown in European botanic gardens were common (see Turland et al. 1996 ). Several sheets of Lhotsky 87 in G-DC have labels stating "Solanum uvareaifolium nob. 1819" in Dunal's hand; this name was never published. Several Martius collections (Martius "745" and "747" from Minas Gerais and Bahia and one Sellow collection (from southern Brazil) are syntypes of Solanum fultum ; I have chosen the sheet at M (M0171876) of Martius 745 bearing a label with a description in Sendtner's hand and with several inflorescences bearing flowers with the diagnostic unequal anthers as the lectotype. The other two syntypes are also from Minas Gerais, but are less complete ( Martius 747 , M0171877; Sellow s.n., P00335352). Witasek (1910) described variety heterophyllum under both Solanum convolvulus and Solanum flaccidum ; a sheet of the former at W (W-1922-0001503) is selected as the lectotype (no herbaria were cited in the protologue and these specimens are at either W or WU) and the sheet at WU [WU0037973] with pinnatifid lower leaves as the lectotype of the latter. The type specimen of Solanum uncinellum var. atrosanguineum at P has the ellipsoid anthers and rotate-stellate corolla of Solanum flaccidum , not the tapered anthers and deeply stellate corolla of the more widespread Solanum uncinellum . Specimens examined. Brazil . Bahia : Conquista, 1000 m, Jun 1944, Schury 616 (US); Espirito Santo : Itaguacu , Alto Limoeiro, 10 May 1946, Brade et al. 18054 (K); Muniz Freire, Rod. BR-262, 1000 m, 21 Jul 1982, Hatschbach & Guimaraes 45169 (US); Minas Gerais : Vicosa , Araponga, beyond Areponga, Fazenda do Serra, 1300 m, 3 May 1930, Mexia 4660 (BM, F, G, GH, K, MO, S, US); Ilheu , Fazenda da Tabunha, 300 m, 17 Aug 1930, Mexia 4692 (BH, BM, F, G, GH, K, MO, S, US); Caldas, Rio Capivahy, 25 Nov 1873, Mosen 983 (S); Caldas, 26 Jul 1869, Regnell II-1712 (S, US); Parana : Araucaria , 13 Nov 1972, Dombrowski 4246 (US); Roca Nova, 15 Mar 1909, Dusen s.n. (F, K, S); Serrinha, Rio Iguassu, 12 Dec 1908, Dusen 7469 (S); Porto de Cima, 28 Sep 1909, Dusen 8764 (S); Serra do Mar, desvio Ypiranga, 6 Sep 1911, Dusen 12147 (S); Piraquara, estrada entre Rio Taquari e Rio Divisa, 13 Mar 1949, Hatschbach 1217 (US); Paranagua , Sertao , 26 Aug 1951, Hatschbach 2685 (US); Guarapuava, Serra da Esperanca , 1135 m, 20 Oct 1960, Hatschbach 7329 (US); Paranagua , Encruzilhada, 18 May 1961, Hatschbach 8026 (US); Porto Amazonas, Fazenda Sao Luiz, margens do Rio Iguacu, 13 Oct 1963, Hatschbach 10892 (B, US); Antonina, Rio Cotia, 360 m, 17 Sep 1965, Hatschbach 12809 (US); Antonina, Cab. Rio Faisqueira, 50 m, 11 Apr 1968, Hatschbach 19019 (F, L); Campina Grande do Sul, Rio Bonito, rod. Br. 116, 31 Oct 1969, Hatschbach 22756 (F, S); Jaguariaiva , Rib. Taquaral, 12 Nov 1981, Hatschbach 44362 (MA); Serrinha, 840 m, 8 Oct 1914, Joensson 1084 a (F, K, S); Pinhaes , 30 Oct 1908, Lange 6972 (S); Pinhaes , 16 Nov 1960, Moreira 36 (US); Curitiba, Parque Iguacu , mini-pantanal, 12 Nov 1987, Silva & Cordeiro 399 (S); Curitiba, Campina do Siqueira, Rio Barigui, 20 Nov 1966, Stellfeld 1644 (US); Pernambuco : Floresta, Inaja , Reserva Biologica de Serra Negra, 21 Jul 1995, Ferraz et al. 273 (BM); Floresta, Inaja , Reserva Biologica de Serra Negra, 21 Jul 1995, Silva et al. 85 (BM); Rio Grande do Sul : Caixas do Sul, Sao Virgilio, 3a Legua, 780 m, 29 Nov 1999, Kegler 390 (US); Bom Jesus, Passo da Guarda, 14 Jan 1952, Rambo 51858 (B); Sao Francisco de Paula, Iaimbezinho, 20 Feb 1953, Rambo 53995 (B); Itaimbezinho, pr. S. Fr. de Paula, 12 Feb 1956, Rambo 58557 (B, S); San Salvador, Montenegro, 600 m, 9 Oct 1946, Sehnem 2196 (B); Rio de Janeiro : Teresopolis , 2 May 1917, Asampaio 2425 (US); Organ Mts., Feb 1837, Gardner 558 (K); province of Rio de Janeiro, 1876, Glaziou 8855 (G, K, LE); environs de Rio de Janeiro, Glaziou 12100 (K, LE); Serra das Orgaos , Sep 1832, Lhotsky 87 (G-DC); Organ Mount. Vargem, Jan 1838, Miers s.n. (BM); Organ Mts., Miers 3465 (K); prope Engo, Pasos, km 151 viae BR-2, 23 Oct 1962, Pabst 7146 (F, US); estrada para Friburgo, 4 Oct 1959, Pereira 5116 (US); Macae , 1900, Ule s.n. (US); Serra das Orgaos , Dec 1897, Ule 4302 (CORD); Santa Catarina : Itajai , 5 km W of Itajai , 100 m, 6 Jan 1974, Conrad & Dietrich 2124 (MO); Itajai , Cunhas, 10 m, 27 Oct 1955, Klein 1731 (B, G, L); Pantano do Sul, Morro do Saquinho, 27 Jul 1967, Klein & Bresolin 7525 (US); Tapera, Ribeirao , 250 m, 18 Aug 1970, Klein 8719 (US); Lajes, Encruzilhada, 900 m, 16 Dec 1967, Lourteig 2286 (US); Ibirama, Horto Florestal I.N.P, 350 m, 2 Nov 1953, Reitz & Klein 1168 (US); Campo dos Padres, 1800 m, 20 Dec 1948, Reitz 2571 (US); Brusque, 40 m, 3 Nov 1949, Reitz 3162 (S); Sao Joaquim, Bom Jardim, fazenda da Laranja, 1400 m, 13 Dec 1958, Reitz & Klein 4082 (B, G, K, L, US); Rio do Sul, Serra do Matador, 700 m, 12 Mar 1959, Reitz & Klein 8514 (K, US); Itajai , Cordeiros, 5 m, 9 Oct 1959 Reitz & Klein 9157 (US); Sao Jose , Serra da Boa Vista, 1000 m, 24 Jan 1961, Reitz & Klein 10719 (K); Ipumirim, 19 km S of the Rio Irani, 9 Dec 1964, Smith & Klein 13922 (US); Sao Paulo : Santa Ana, Nov 1912, Brade, A.C ., 6031 (US); Jundiahy, Mar 1900, Campos Novaes 167 (US); Capoeira Grande, Sep 1898, Campos Novaes 4089 (US); Atibaia, Sep 1910, Duarte s.n. (US); sin.loc, 1833, Gaudichaud 312 (P); proximo a Campinas, margens do Rio Pirapitingui , 22 Nov 1953, Grotta SPF 15112 (F); Guararema, 1 Nov 1897, Guttermans 9 (US); Butanta , 24 Nov 1919, Hoehne s.n. [3493] (F, MA, US); estrada de Pirapora a Cabreuva , 4 Dec 1924, Hoehne, 12910 (US); Guararema, km 348 da Via Dutra, 29 Oct 1956, Hoehne SPF 15754 (F); Cidade Jardim, 1 Dec 1933, Kuhlmann s.n. (US); Jundiai , ca. 10 km SW, Serra do Japi , 8 Oct 1976, Leitao Filho et al. 3228 (F); Sao Jose do Rio Pardo, 24 Oct 1889, Loefgren 1441 (US); Xiririca, 29 Oct 1894, Loefgren 2828 (US); Estacao Sagrado do Caracao , linha Sorocabana, 750 m, 3 Oct 1979, Mizoguchi 1048 (MO); Iporango, BR-2 a 40 km de Jacupiranga em direcao a Curitiba, 12 Nov 1961, Pabst 6754 (Pereira 6928) (F, US); Sao Paulo, 7 Aug 1941, Praed 5332 (US). Paraguay . Amambay : Sierra de Amambay, Jul 1912, Hassler 11288 (G, K, P).