Solanum asterophorum, Mart.

Gouvêa, Yuri & Stehmann, João, 2019, A Revision of the Solanum asterophorum Species Group (subg. Leptostemonum), Systematic Botany 44 (1), pp. 210-232 : 217-226

1. SOLANUM ASTEROPHORUM Mart. ,

in Flora 21, Beibl. 2: 79. 1838.

Type: Brazil. “ prope Sebastianopolin, ibi Decembri florens . Ad Lagoa fea Sept . Dryasʺ, 1834, P. Luschnath s.n. (lectotype, here designated: BR [ BR0000008366306 ]!) .

Solanum asterophorum Mart. var. tomentosum Sendtn., Fl. Bras. [Martius] 10: 98. 1846.

TYPE: BRAZIL. F. Sellow s.n. (lectotype, here designated: P [P00324598]!; isolectotype: LE [LE00016840]).

Solanum melancholicum Salzm. ex Dunal, Prodr. [A. P. de Candolle] 13 (1): 200. 1852.

TYPE: BRAZIL: Bahia. “In collibus circa Bahiamʺ, 1830, P. Salzmann 390 (holotype: G-DC [G00145876]!; isotype: P [P00337131]!).

Solanum tetricum Dunal, Prodr. [A. P. de Candolle] 13 (1): 205. 1852.

TYPE: BRAZIL. Sin. loc., C. Gaudichaud 515 (lectotype, here designated:P [P00384256]!; isolectotypes:P [P00445098]!, P [P00384254]!, P [P00384255]!, G-DC [G00145836]!, F [F0073432F]!, F [F0073434F], F [F0073435F], W-0022476!, BR [BR0000008290564]!).

Solanum gomphoidellum Moric. ex Dunal, Prodr. [A. P. de Candolle] 13 (1): 215. 1852.

TYPE: BRAZIL. Bahia. “Circa Moritibaʺ [Muritiba], J. S. Blanchet 3473 (lectotype, here designated: G two sheets [G00405700, G00405700_a]; isolectotypes: C [C10019216]!, G [G00405785], P [P00324600]!, W-0001939!).

Shrubs up to 3 m, the branches erect; young stems terete, sparsely stellate-puberulent to densely stellate-tomentose; the trichomes hyaline to yellow or brown-tinged, rarely ochraceous near the meristems, porrect, sessile to long-stalked, the stalks to 0.8 mm long, multiseriate, 2–4 cells wide, the rays 4–8(–11), 1-celled, the midpoints 1-celled, obsolete to the same length as the rays, straight to oblique; moderately armed, the prickles 2.4–5.5 mm long, 1.2–7.2 mm wide at base, recurved, flattened, stramineous to brown, sometimes becoming ferruginous toward the apex, with stellate trichomes like those of the stems and some small, subsessile, glandular trichomes at base, these often drying dark; bark of older stems glabrescent to moderately stellate-tomentose, green to dark brown. Sympodial units difoliate, geminate, the leaves of a pair anisophyllous. Leaves lobed, rarely entire, repand in large-leaved plants; the major leaves 10.5–29.5 cm long, 5–20.5 cm wide, elliptic to obovate, rarely ovate; base narrowly cuneate to cuneate, less often rounded, generally asymmetric; margins shallowly to deeply lobed, rarely entire, the lobes (0–)1–6 per side, deltate, to 2.7(–4.9) cm long, 1–5.4 cm at base, acuminate to roundish apically; apex acute to obtuse; primary veins 6–10 pairs; minor leaves (4.5–) 6.3–17.5 cm long, 3.3–10 cm wide, elliptic to nearly circular; base acuminate to rounded, generally symmetric; margins entire to lobed, the lobes 0–4 on each side, to 2.3 cm long, to 5 cm wide at base; apex acuminate to rounded-mucronulate; primary veins 5–7 pairs; major and minor leaves all membranaceous to chartaceous, matte or shiny above when fresh, weakly to markedly discolorous, drying green, brown or black above, and pale green to pale brown or grey beneath; the adaxial surface sparsely stellatepuberulent to densely stellate-tomentose, the epidermis always visible, the trichomes hyaline, yellowish or browntinged, rarely ochraceous, porrect, sessile to short-stalked, the stalks to 0.6 mm long, multiseriate, 3–4 cells wide, the rays (3–) 4–8(–11), 1-celled, the midpoints 1-celled, obsolete to longer than the rays, straight to oblique; the abaxial surface sparsely stellate-puberulent to densely stellate-tomentose, epidermis visible or not, the trichomes like those of the adaxial leaf surface, but rarely also with long-stalked ones; unarmed to moderately armed along the midrib and the primary veins on both surfaces, the prickles straight to slightly oblique, flattened, 2.5–10 mm long, 0.7–3.9 mm wide at base, 0–10 above and 0–10 beneath; petiole of major leaves 1.5–5.8 cm, sparsely stellate-puberulent to densely stellate-tomentose with stellate trichomes like those of the stems, the prickles 0–7, straight; petiole of minor leaves (0.5–) 0.7–2.7 cm, sparsely stellatepuberulent to densely stellate-tomentose, with stellate trichomes like those of the stems, the prickles 0–6; leaves of juvenile plants usually larger, more densely armed and lobed, becoming smaller, less prickly and with fewer lobes in older plants. Inflorescence a reduced monochasial cyme, 1.2–25 mm long, unbranched, leaf-opposed or nearly so, with (2–)5– 12(–21) flowers, 1–2 flowers open at a time; inflorescence axis (peduncle plus rachis) sparsely stellate-puberulent to densely stellate-tomentose with stellate trichomes like those of the stems, unarmed; peduncle nearly obsolete to 4.5 mm long; rachis (1.2–) 4–20.5 mm, usually gently curved at the tip; pedicel insertion points very closely spaced, usually overlapping, 0–1.4 mm apart; pedicels curved downward with the flower buds pointing down before anthesis, 8.2–17 mm long and straight at anthesis, usually distally geniculate, articulated at base, armed or not, sparsely stellate-puberulent to densely stellate-tomentose, the trichomes hyaline to yellow or browntinged, rarely somewhat ochraceous, porrect, sessile to longstalked, the stalks to 0.8(–1.2) mm long, multiseriate, 2–3 cells wide, the rays (4–)6–8(–10), 1-celled, the midpoints 1-celled, obsolete to 2/3 the length of the rays. Flowers 5-merous, rarely 4-merous (anomalous flowers of plants from southern coast of Bahia), heterostylous, generally long-styled (functionally hermaphrodite), basal flowers long-styled and hermaphroditic, short-styled and functionally male flowers produced distally in the inflorescence (see Sexual Expression and Inflorescence). Calyx tube obconic, 4–7.6 mm long, sparsely stellate-puberulent to densely stellate-tomentose, with trichomes like those of the pedicels, prickles 0–30; calyx lobes narrowly oblong, elliptic or lanceolate, 1.6–5.9(–9.8) mm long, often unequal in length, 1.5–2.8(–4) mm wide, the apices acuminate to rounded, often reflexed at anthesis. Corolla 2.8–4 cm in diameter, white, stellate, with well-developed and wavy interpetalar tissue, lobed for 2/5 to 3/5 of its length, the lobes 8–10.9 mm long, 8.9–14.3 mm wide, deltate, often apiculate at the apex, moderately to densely stellatetomentose along the whole length abaxially, the trichomes hyaline to brown-tinged, porrect to multiangulate, sessile to short-stalked, the stalks to 0.4 mm long, multiseriate, 2–4 cells wide, the rays 4–8(–11), tortuous, usually forming a crab-like shape rather than a circle, the midpoints 1-celled, variable in length, the adaxial surface moderately to densely stellatetomentose at the apex with trichomes like those of the abaxial surface becoming gradually less dense towards the base, the basal half glabrous or nearly so, the lobe tips usually cucullate and reflexed at anthesis. Stamens equal; filament tube 1.3–3.2 mm long; free portion of the filaments 0.9–1.6 mm long; anthers 6.2–12.1 mm long, 1.2–2.7 mm wide, lanceolate, sagittate at base, narrowed towards the apex, dehiscing by apical pores, connivent or not. Ovary short-cylindrical, convex at the apex, with some glandular trichomes; style of long-styled flowers 10.7–14.6 mm long, white, cylindrical, straight to gently curved distally, glabrous to moderately stellate-tomentose at base, in short-styled flowers the style 6–8.2 mm long; stigma 0.8–1.4 mm long, sometimes bilobed at the apex, green, the surface papillose. Fruit a spherical to broadly depressed ovate berry, 9.5–14.1 mm long, 11.6– 15.8 mm wide, the pericarp smooth, glabrous, the exposed portion pale green, pale yellow, or white, the portion covered by the calyx pale green to green at maturity; fruiting pedicels 1.3–2 cm long, armed or unarmed; fruiting calyx accrescent, covering 1/3–3/4 of the mature fruit, the base rounded, the

lobes 3.9–8.4 mm long, 5.4–8.3 mm wide at the base. Seeds ca. 30–55 per berry, 3.5–4.2 mm long, 2.8–3.4 mm wide, flattened, reniform, stramineous to brown. Chromosome number: not known. Figures 1 , 4 , 11 .

Habitat and Distribution — Solanum asterophorum is widely distributed along the Atlantic Forest domain in the northeastern and southeastern regions of Brazil, except in Ceará, northeastern Rio Grande do Norte and southeastern S~ ao Paulo states, where there is no record of this species ( Fig. 7 ). It grows in damp sites with indirect light, such as wet forest edges, roadsides and partially deforested places near forests, less commonly being found in forest understory; sea level to ca. 800 m.

Phenology —Flowering and fruiting specimens have been collected in all months, with a flowering peak from November to April and the fruiting peak from May to October.

Preliminary Conservation Status —Despite the relatively small area of occupancy (AOO: 476 km 2), the large extent of occurrence (EOO: 540,573 km 2) and the numerous collection localities and known populations of S. asterophorum suggest that it is likely to meet an IUCN red list status of least concern (LC) ( IUCN 2016).

Etymology —The protologue lacks specific details of the name’ s etymology, but the epithet comes from the Greek “aster(-o),ʺ star, and “phoro,ʺ bear, carry, a possible reference to the stellate trichomes covering the plant.

Additional Specimens Examined — Brazil. — ALAGOAS: Mun. Maceió, Parque Municipal de Maceió, 22 Nov 2011 (fl), Sampaio et al. 37 ( UFP); Mun. Murici, Bananeiras, 9°14, 5ʺS, 35°52, 61 ̎ ʺW, 513 m, 16 Mar 2000 (fl), Carvalho et al. 7141 ( CEPEC, MAC, NY); Mun. Pilar, Fazenda Lamarao~, 09°36, 12 ̎ ʺS, 35°55, 09 ̎ ʺW, 79 m, 19 Nov 2011 (fr), Sampaio & Araujo 29 ( UFP); Mun. Quebrangulo, REBIO Pedra Talhada, mata próxima a sede da REBIO, 9°15, 16ʺS, 36°25, 50 ̎ ʺW, 664 m, 25 Jan 2012 (fl), Sampaio et al. 55 ( UFP).— BAHIA: Mun. Almadina, 5.3 km from Almadina on road to Ibatup~ a, then left 7.9 km on road to Serra dos Sete Paus, 14°44, 11 ̎ ʺS, 39°41, 57 ̎ ʺW, 500–650 m, 04 Apr 1997 (fr), Thomas et al. s.n. ( HUEFS, MBM, NY); Mun. Amargosa, próximo à “Jacubinhaʺ, na casa do Sr. Arlindo, 13°7, 0ʺS, 39°39, 5ʺW, 630–900 m, 16 Nov 2007 (fl), Perdiz et al. 237 ( BHCB, CEPEC, HUEFS); Serra do Timbó, trilha para o jequitibá, área de estudos do Projeto Timbó/Centro Sapucaia, 13°7, 3ʺS, 39°39, 50ʺW, 809 m, 28 Jan 2007 (fl), Cardoso et al. 1688 ( CEPEC, HUEFS); Mun. Amélia Rodrigues, 4 km SE de Amélia Rodrigues, 12°26, 29ʺS, 38°44, 02ʺW, 20 Mar 1987 (fl), Queiroz & Crepaldi 1464 ( HUEFS, PEUFR); Mun. Aureliano Leal, fragmento de mata próxima à torre da Embratel, 14°25, 13ʺS, 39°16, 16ʺW, 561 m, 23 Sep 2008 (fl), Giacomin et al. 205 ( BHCB); Mun. Belmonte, distrito de Barrolandia ˆ, Estaç~ ao Expe. Gregório Bondar ( EGREB), Rodovia Itapebi/Belmonte km 51, a 5 km a oeste de Barrolandia ˆ, 30 Mar 1988 (fl, fr), Santos 853 ( CEPEC); Mun. Cachoeira, depois de Balém, na estrada para Sto. Antonio ˆ, Fazenda JOMEI, a 4 km da rodovia, 12°32, S, 39°5, W, 27 May 2003 (fl), Oliveira 889 ( HUEFS, JPB);Mun. Camaçari, margem da BA-093, próximo ao Pólo Petroqu´ımico, 28 May 2011 (fl), Barletta-Mattos & Abreu 4 ( ALCB); Mun. Dias D’ avila, acesso da BR-093, 5 Aug 1994 (fl, fr), Guedes et al. s.n. ( ALCB); Mun. Eunápolis, plantio de Eucalyptus, 22 Apr 1994 (fl,fr), Guedes et al. s.n. ( ALCB); Projeto Sapucaieira, 16°22, 00ʺS, 39°34, 00ʺW, 200 m, 19 Jan 1997 (fl), Guedes et al. 4254 ( ALCB, UFP); Mun. Itacaré, estrada que liga a torre da Embratel com a BR-101/ Itacaré, a 5.8 km da entrada, cerca de 25 km a SE de Ubaitaba, 15 Jun 1979 (fl), Mori & Carvalho 12023 ( CEPEC, NY); Mun. Ituberá, Assentamento Limoeiro, proximidade da sede, área de reserva, 13°59, 33ʺS, 39°17, 11ʺW, 16 Dec 2001 (fl), Alves et al. 341 ( ALCB, CEPEC); Mata do Ponto Alto, 13°43, S, 39°08, W, 28 Nov 2005 (fl), Guedes et al. 11907 ( ALCB, CEPEC); Mun. Jaguaquara, estrada para Apuarema 5.7 km de Jaguaquara, 13°34, 48ʺS, 39°55, 51ʺW, 808 m, 24 Apr 2002 (fl), Oliveira et al.790 ( HUEFS); Mun. Porto Seguro, rodovia para Eunápolis, km 13, 4 Feb 1972 (fl), Eupunino 194 ( CEPEC); Mun. Prado, Parque Nacional do Descobrimento, entrada a esquerda no km 12 da estrada principal que atravessa o Parque, 17°11, S, 39°20, W, 70 m, 2 Nov 2009 (fl), Matos et al. 1901 ( CEPEC); Cumuruxatiba, beira de estrada de terra que liga Cumuruxatiba a Prado, 17°07, 14ʺS, 39°11, 41ʺW, 25 m, 19 Jun 2014 (fl), Gouvea ˆet al.134 ( BHCB); Beira de estrada de terra que liga Vila Guarani à Barra do Cahy, 16°58, 57ʺS, 39°22, 13ʺW, 40 m, 19 Jun 2014 (fl), Gouvea ˆet al.128 ( BHCB); Beira de estrada de terra que liga Vila Guarani à Barra do Cahy, 16°59, 40ʺS, 39°23, 27ʺW, 108 m, 19 Jun 2014 (fr), Gouvˆea et al. 125 ( BHCB); Beira de estrada (BA-489), próximo ao Parque Nacional do Descobrimento, 17°17, 10ʺS, 39°19, 05ʺW, 55 m, 19 Jun 2014 (fl), Gouvˆea et al. 123 ( BHCB); Mun. Ribeir~ ao Largo, ca. 23 km na estrada Itambé/Encruzilhada, 15°19, 39ʺS, 40°45, 5ʺW, 870 m, 14 Aug 2001 (fr), Carvalho et al. 6909 ( CEPEC, HUEFS, JPB, NY); Mun. Salvador, Estaç~ ao Ecológica de Cotegipe, 22 Sep 1994 (fl), Guedes et al. s.n. ( ALCB); Unidade Ecológica do Cia, 17 Dec 1986 (fl), Silva 05 ( ALCB); Mun. Santa Cruz Cabrália, Estaç~ ao Ecológica do Pau-Brasil e arredores, cerca de 16 km a W de Porto Seguro, 2 Jul 1978 (fl,fr), Mori 10207 ( CEPEC, NY); Estaç~ ao Ecológica do Pau-Brasil, cerca de 16 km a W de Porto Seguro, 10 Mar 1983 (fl), Brito & da Vinha 212 ( CEPEC); Mun. S~ ao Sebasti~ ao do Passé, Lamar~ ao do Passé, 12°30, S, 38°20, W, 8 Oct 1999 (fl, fr), Guedes et al. 6500 ( ALCB, CEPEC, JPB); Mun. Uruçuca, 17-20 km de Uruçuca, na estra Uruçuca para Itacaré, 14°26, 33ʺS, 39°14, 29ʺW, 75 m, 7 Mar 200 (fr), França et al. 3359 ( FUEL, HUEFS, JPB, UFG); Mun. Wenceslau Guimar~ aes, Estaç~ ao Ecológica Estadual Nova Esperança, sede a 7 km a W do povoado de Nova Esperança, 13°35, 43ʺS, 39°43, 18ʺW, 700 m, 26 Jul 2001 (fr), Mattos-Silva et al. 4447 ( ALCB, HUEFS, UESC); 13°41, S, 39°28, W, 2 Dec 2001 (fl), Alves et al. 399 ( ALCB, CEPEC). — ESPÍRITO SANTO: Mun. Ibiraçu, Estaç~ ao Ecológica do Morro da Vargem, Trilha do Bananal, 19°53, S, 40°23, W, 300–470 m, 27 May 1990 (fl), Gomes et al. 1154 ( BHCB, MBML, VIES); Mun. Linhares, Povoaç~ ao, 17 Oct 1983 (fl), Hatschbach 46923 ( MBM); Reserva da Companhia Vale do Rio Doce, Estrada Roxinho, próximo ao aceiro catel~ a, 19°09’31”S, 40°03, 39 ̎ W, 47 m, 8 Apr 2006 (fl), Pinho-Ferreira et al. 616 ( BHCB, CVRD, ESA); Estrada Jequitibá Rosa, ca. 500 m do in´ıcio, 19°08, 20 ̎ S, 39°55, 41 ̎ W, 25 m, 11 Apr 2006 (fl), Rom~ ao et al. 1337 ( BHCB, ESA, CVRD); Degredo, floresta sobre cord~ ao arenoso, 19°20, 51 ̎ ʺS, 39°43, 46 ̎ ʺW, 14 May 2010 (fl), Ribeiro et al. 166 ( BHCB, VIES); Reserva Floresta de Linhares, 200 m após o Córrego Rancho Alto, 11 Dec 1991 (fl, fr), Zortea 6 ( BHCB, CVRD, MBML); Aceiro com Nivaldo, canto G, ao lado do aceiro, 23 Jan 1995 (fl), Folli 2494 ( BHCB, CVRD); Estrada Guapuruvú, lado esquerdo indo para o norte, 21 Apr 1993 (fl), Folli 1865 ( BHCB, CVRD); A 500 m do Córrego Rancho Alto, ao lado da estrada, 10 Apr 1993 (fl, fr), Folli 1862 ( BHCB, CVRD); Estrada Gávea, entrada do Bloco C do Ensaio de Prod. Sustentada, 12 Oct 1990 (fl), Menandro 272 ( BHCB, CVRD, UEC); Aceiro com Adair Campo, próximo a divisa com José Catel~ a, 10 Nov 1998 (fl, fr), Folli 3277 ( BHCB, CVRD); Mun. Santa Teresa, trilha subindo o morro ao lado do Country Club, 25 Feb 1996 (fl), Lombardi & Temponi 1129 ( BHCB); Vale do Cana~ a, 16 Apr 1984 (fr), Boone 48 ( BHCB, MBML); 4 Dec 1985 (fl), Vimercat 321 ( BHCB, MBML); 8 Nov 1985 (fl), Boone 864 ( BHCB, MBML); Rio Saltinho, 26 Apr 1984 (fl, fr), Pizziolo 2 ( BHCB, MBML); Em plantaç~ ao de banana próximo à rodovia ES- 261, 19°56, 22 ̎ ʺS, 40°30, 21 ̎ ʺW, 527 m, 15 May 2015 (fl), Gouvea ˆ& Falc~ ao 189 ( BHCB); 16 Sep 2014 (fl), Gouvea ˆ& Falc~ ao 140 ( BHCB); Estaç~ ao Biológica de Santa Lúcia, 7 Nov 1985 (fl), Fernandes 1601 ( BHCB, MBML); Trilha do Indaiá-açú, 19°58, 04 ̎ ʺS, 40°32, 15 ̎ ʺW, 622 m, 6 Feb 2011 (fl), Giacomin et al. 1214 ( BHCB); Nova Lombardia, Estrada de terra que leva de Santa Teresa para Nova Lombardia, 19°50, 59 ̎ ʺS, 40°30, 56 ̎ ʺW, 606 m, 16 Sep 2014 (fl), Gouvea ˆ& Falc~ ao 141 ( BHCB); Reserva Biológica de Nova Lombardia, 2 Aug 1984 (fl), Pizziolo 145 ( MBML, RB); Reserva Biológica Augusto Ruschi, 800 m, 16 Oct 2001 (fl), Kollmann & Bausen 4867 ( BHCB, MBML); Estrada para Goipabo-açu, parte final, 800 m, 24 Oct 2002 (fl), Vervloet et al. 1293 ( BHCB, MBML); Divisa com propriedade Vanildo Bragacha, 23 Jan 2003 (fr), Vervloet & Bausen 1734 ( BHCB, MBML); Dra. Marlene, antiga estrada, 23 Jul 2002 (fr), Vervloet et al.502 ( BHCB, MBML); Estrada partindo da Casa da Pedra, 11 Dec 2002 (fl), Vervloet et al. 1491 ( BHCB, MBML); Propriedade do Sr. Furlani, próximo a plantaç~ ao de banana, 19°48, 22 ̎ ʺS, 40°32, 19 ̎ ʺW, 720 m, 9 Jun 2012 (fl, fr), Giacomin et al. 1868 ( BHCB, NY); Fragmento de Floresta Ombrófila preservada anexo à plantaç~ ao de bananeira, 19°48, 16 ̎ ʺS, 40°32, 18 ̎ ʺW, 803 m, 7 Feb 2011 (fl), Giacomin et al.1223 ( BHCB); Mun. S~ ao Mateus, Reserva Biológica do Sooretama, Lago do Macuco, ca. 30 m, 15 May 1977 (fl), Martinelli 2157 ( BHCB, RB). — MINAS GERAIS: Mun. Ataleia, estrada que leva da Comunidade Cana~ a a Ataleia, pasto, 18°00, 51 ̎ ʺS, 41°09, 21 ̎ ʺW, 286 m, 15 Jun 2014 (fl), Gouvea ˆet al. 100 ( BHCB); Mun. Caratinga, Estaç~ ao Biológica de Caratinga, 11 Oct 1987 (fl, fr), Costa et al. 296 ( BHCB); 06 Apr 1990 (fl), Costa et al. s.n. ( BHCB); Fazenda Montes Claros, beira de estrada, 11 Jan 1991 (fl), Stehmann s.n. ( BHCB); APA Lagoa Silvana, 28 Jun 2002 (fl), Pivari & Cortes ˆ116 ( BHCB, CESJ, MBM, SPF); F.M. C. Mata do Rafael, 26 Apr 1984 (fl, fr), Andrade & Lopes 371 ( BHCB, UEC); Mun. Descoberto, Reserva Biológica da Represa do Grama, 09 Jun 2001 (fl, fr), Castro et al. 466 ( BHCB, CESJ, MBM); 21°25, 53 ̎ ʺS, 42°56, 48 ̎ ʺW, 563 m, 15 May 2015 (fl, fr), Gouvˆea & Falc~ ao 192 ( BHCB); Mun. Dion´ısio, Lagoa do Jacaré, 19°49, 15 ̎ ʺS, 42°39, 48 ̎ ʺW, 276 m, 06 Jun 2003 (fr), França 362 ( BHCB, FUEL); CEMAS, Cia. Agr´ıcola e Florestal Santa Bárbara, 7 Feb 1986 (fl), Campos 38 ( BHCB); Mun. Gonzaga, Fazenda do Sr. Gezier Nunes, 18°47, 47 ̎, 42°27, 40 ̎, 790 m, 5 Sep 2008 (fr), Kamino & Silva 1152 ( BHCB); Mun. Mariliéria, Parque Estadual do Rio Doce, entrada do alojamento, trilha para Porto Capim, 30 Mar 1996 (fl), Lombardi et al. 1195 ( BHCB); 02 May 1997 (fl, fr), Tavares s.n. ( BHCB, FUEL, MBM); Estrada Restaurante- Laboratório, perto do laboratório, 19°46, S, 42°37, W, 3 Mar 1999 (fl), Lombardi 2548 ( BHCB, FUEL); Trilha da Campolina, 31 Oct 1992 (fl), Stehmann & Arantes s.n. ( BHCB); Mun. Novo Cruzeiro, Fazenda Araras, 17°36, 47 ̎ ʺS, 41°57, 49 ̎ ʺW, 754 m, 2 Oct 2004 (fl, fr), Stehmann et al. 3548 ( BHCB, MBM); Mun. Santa Bárbara, Estaç~ ao de Pesquisa e Desenvolvimento Ambiental de Peti, 19°53, 33 ̎ ʺS, 43°21, 55 ̎ ʺW, 21 May 2005 (fl), Ferreira & França 106 ( BHCB); 8 Mar 1988 (fl), Stehmann & Pedralli s.n. ( BHCB); Mun. Teófilo Otoni, Afloramento rochoso lado esquerdo da MG- 418, cerca de 30 km ao norte de Teófilo Otoni, 17°51, 22 ̎ ʺS, 41°15, 39 ̎ ʺW, 546 m, 8 Jan 2011 (fl), De Paula et al. 106 ( BHCB); Mun. Timóteo, Parque Estadual do Rio Doce, 19°35, 28 ̎ ʺS, 42°34, 07 ̎ ʺW, 248 m, 4 May 2004 (fl), França & Raggi 547 ( BHCB, CESJ); Mun. Viçosa, Mata do Para´ ıso near town of Viçosa, at roadside, 20°47, 59 ̎ ʺS, 42°51, 60 ̎ ʺW, 705 m, 21 Apr 2010 (fl), Agra et al. 7250 ( BHCB, JPB, UT). — PARAÍBA: Mun. Areia, Escola de Agronomia do Nordeste, em terrenos de mata, lugares altos e úmidos, 08 Jun 1953 (fl), Moraes 796 ( MBM). — PERNAMBUCO: Mun. Jaqueira, Serra do Urubu, nas proximidades da Pedra do Cruzeiro, no interior da mata, 8°43, 56ʺS, 35°50, 44 ̎ ʺW, 730 m, 1 Apr 2012 (fl, fr), Sampaio et al. 83 ( BHCB, UFP); Sampaio et al. 75 ( BHCB, UFP); Mun. Recife, Reserva Florestal do Curado, 25 Oct 1967 (fl, fr), Lira 128 ( UFP). — RIO DE JANEIRO: Mun. Casimiro de Abreu, Estrada de terra que liga uma das entradas da Reserva Biológica Poço das Antas à BR-101, beira de estrada, 22°31, 35 ̎ ʺS, 42°19, 29 ̎ ʺW, 35 m, 29 Apr 2015 (fl), Gouvea ˆ& Stehmann 179 ( BHCB); 22°31, 44 ̎ ʺS, 42°18, 49 ̎ ʺW, 44 m, 29 Apr 2015 (fl), Gouvea ˆ& Stehmann 175 ( BHCB); Mun. Rio de Janeiro, estrada da Vista Chinesa,Alto da Boa Vista, km 3, 10 Ar 1981 (fl), Carauta 91 ( RB); Vista Chinesa, 20 May 1958 (fl, fr), Liene et al. 3767 ( RB); same loc., 20 May 1958 (fl, fr), Pereira 3767 ( RB); Pedra da Gávea, Caminho das Furnas, ca. 500 m, Carauta & Kirkbride Jr 2440 ( RB); Estrada das Paineiras, ca. 500 m, 9 May 1973 (fl), Sucre 9978 ( JPB, RB); Serra da Carioca, estrada do Cristo Redentor, 25 May 1945 (fl, fr), Occhioni 354 ( RB); Botafogo, Morro Mundo Novo, vertente leste, 12 Jun 1998 (fl, fr), Amancio et al. 7 ( JPB, RB); same loc., Campus da Universidade Santa Úrsula, 27 May 1992 (fl), Amado & Konno 98 ( RB) Monte Corcovado, 400–550 m, 22 May 1969 (fl), Sucre & Plowman 5049 ( RB); same loc., 18 Dec 1945 (fl), Altamiro & Walter 186 ( RB); à 150 m. da Estrada de Ferro Corcovado, 550 m, 18 Jul 1977 (fr), Almeida 283 ( RB); Estrada Rio/Petrópolis, 13 Apr 1939 (fl), Cacerelli s.n. ( RB); Mun. Silva Jardim, Poço d’ anta, 35 m, 22 Oct 1977 (fl), Martinelli 3340 ( RB); same loc., 35 m, 14 Sep 1977 (fl), Martinelli et al.2873 ( RB);Poço d’ Antas, (fl), ( RB 382144).

Notes — Solanum asterophorum differs from other species of the S. asterophorum species group in the combination of obconic calyces with curved pedicels which hold the flower buds pointing down (see Figs. 1B, F ). The latter character can sometimes be difficult to observe in herbarium sheets, once the pressing process influences the position of the buds. Thus, in order to facilitate identification, this character should be noted at the time of collection and included in the label observations when possible. Solanum asterophorum is morphologically similar to S. igniferum and S. sessilantherum , with which it shares lobed leaves and the obconic calyx. Solanum igniferum differs from S. asterophorum in its notable orange rather than sparse hyaline indumentum, plagiotropic rather than erect branches, erect to spreading rather than downward pointing buds and its more lax inflorescences. From S. sessilantherum , S. asterophorum can be distinguished by its downward pointing rather than spreading buds; its congested inflorescences with unarmed axes rather than lax inflorescences with armed axes; its coriaceous rather than membranous leaves; and anthers borne on distinct filaments rather than being sessile.

Solanum asterophorum has the most variable morphology among the species belonging to the S. asterophorum species group, and also the widest latitudinal range. Occurring from the states of Rio de Janeiro State to Para´ıba, its populations vary mainly with respect to the indumentum of the stems, leaves, reproductive structures, and the coloration of the epidermis when dry. There is a great variation in trichome density, color, and length of trichome stalks and midpoints. The indumentum density ranges from nearly glabrous to dense, with interwoven trichome rays, stalks, or both. The trichomes can be sessile to long-stalked, with midpoints obsolete to longer than the rays. Trichomes also vary in coloration from hyaline to yellow or brown-tinged, and they are rarely somewhat ochraceous. When dry, the color of the leaf epidermis can range from green to black, and specimens of intermediate coloration are common.

Populations with certain sets of characteristics are restricted to some localities along its range, and two extremes can be noted. Plants exhibiting a sparse indumentum composed of apparently sessile, hyaline trichomes with reduced midpoints, and an epidermis shiny green when fresh and black when dry (e.g. Gouvea ˆ& Falcao~ 189; Giacomin et al. 1214; Fig. 1E–G ) are particularly found in the states of Esp´ırito Santo (Mun. Santa Teresa and near localities), Minas Gerais (Mun. Caratinga, Estaçao~ Biológica de Caratinga), and throughout the state of Rio de Janeiro. Specimens with these morphological features correspond to the type material of Solanum tetricum , a name treated as synonym of S. asterophorum by Whalen (1984) and subsequently recognized as an accepted name by Stehmann et al. (2015). Because many morphologically intermediate specimens have been collected, we agree with Whalen’ s opinion, and here treat S. tetricum as synonym of S. asterophorum .

At the other extreme, plants from the southern coast of Bahia State have dense indumentum on the young stems, leaves, inflorescence axis, and calyx consisting of short- to longstalked, interwoven, pale yellow to yellow (rarely ochraceous) trichomes. The trichomes of the adaxial leaf surface have the midpoints longer than rays (e.g. Gouvˆea et al. 130; Matos et al. 1901) and the leaf epidermis does not become black after drying.

However, most collections of S. asterophorum consist of specimens with an intermediate morphology ( Fig. 1A–D ). These plants usually exhibit an indumentum of moderate density with hyaline to brown-tinged sessile to short-stalked trichomes, these with midpoints of variable length, usually shorter than rays in specimens from southeastern Brazil (e.g. Gouvea ˆ& Falc~ ao 192), and longer than rays in northeastern specimens (e.g. França et al. 3359). The epidermis ranges from green to brown when dry.

Solanum asterophorum var. tomentosum Sendtn. ( Sendtner 1846) was described based on four syntypes, one collected by Prince Maximilian of Wied (“inter arbusta prope Campos”, M. A. P. Prinz zu Wied s.n.), in the area now known as Campos dos Goytacazes, Rio de Janeiro, and the other by Sellow, with no locality indicated. These two collections belong to different species: S. asterophorum (Sellow s.n., P [P00324598] and LE [LE00016840]) and S. igniferum (Prinz zu Wied s.n., BR [ BR836628 ]; Sellow s.n., BR [ BR836629 ]). The short description in the protologue includes morphological characters that could be associated with either of these taxa, but the yellowish-brown or canescent (rather than ferruginous) leaves better correspond to S. asterophorum . For this reason, we select the collection of Sellow s. n. (P [P00324598]) as the lectotype of S. asterophorum var. tomentosum .

Some collections of Salzmann s. n. from Bahia (“in collibus umbrosisʺ HAL [HAL0070575], “in collibusʺ MO [MO503643] and LE [LE00016982] and “in umbrosisʺ E [E00190946]) have been indicated on the sheets as type materials of S. melancholicum Dunal. In the protologue ( Dunal 1852), however, the type was clearly indicated as Salzmann 390 in G-DC, thus other un-numbered Salzmann’ s collections do not represent original materials.

Dunal (1852) described S. tetricum , a synonym of S. asterophorum , based on several syntypes: “In Brasiliae Rio de Janeiro sylvisʺ, Lhotsky s.n. (G-DC), Lund 192 (G-DC), Gaudichaud 515 (G-DC, P); and “circa Bahia,ʺ Guillot s. n. (P). From these syntypes, we have selected Gaudichaud 515 (P [P00384256]) as the lectotype because it has buds and flowers, and the sample corresponds best with the description in the protologue. Furthermore, duplicates of this collection are distributed in many herbaria.

Figures

FIG. 1. Solanum asterophorum from Minas Gerais (Gouvˆea & Falc~ ao 192, BHCB). A. Flower. B. Young inflorescence; note the pedicels curved downwards with the buds pointing down and the pedicel insertion points very closely spaced. C. Habit. D. Fruit. E. Habit of S. asterophorum from Santa Teresa, Espirito Santo (Giacomin et al. 1214, BHCB); note the shiny aspect of the stem and leaves due to the very sparse indumentum. F. Young inflorescence. G. Fruits. Photo credit E: L. L. Giacomin.

FIG. 4. Detail of root tubers in S. asterophorum (Gouvˆea & Falc~ ao 192,

FIG. 11. Scan of lectotype of Solanum asterophorum (P. Luschnath s.n., BR [BR0000008366306]). Reproduced with permission of the National Botanic Garden Meise, Brussels.

FIG. 7. Distribution of Solanum asterophorum.