Solanum piluliferum, Dunal, Prodr.

Gouvêa, Yuri & Stehmann, João, 2019, A Revision of the Solanum asterophorum Species Group (subg. Leptostemonum), Systematic Botany 44 (1), pp. 210-232 : 228-229

3. SOLANUM PILULIFERUM Dunal, Prodr. [A. P. de Candolle] 13(1): 265. 1852.

Type: Brazil. “In Brasilia ˆcirca Novo Friburgo provinciae Rio de Janeiroʺ , P. Claussen 58 (holotype: P [ P00368472 ]! ; isotype G [ 343348 ]!).

Solanum densiflorum Sendtn., Fl. Bras. [Martius] 10: 93, Table 6, Figs. 63–66. 1846. nom. illegit., non M. Martens & Galeotti (1845). Solanum piluliferum Dunal var. densiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 265. 1852.

TYPE: BRAZIL. “In aquaticis ad Sta. Cruz et Rozário, prov. Sebastianopolitana, e, frequens, Decembri florensʺ, C.F.P. von Martius s.n. (lectotype, here designated: M [M0165943]!)

Shrubs up to 3 m, the branches spreading to erect; young stems terete, densely stellate-tomentose; the trichomes stramineous to ochraceo-ferruginous, sometimes reddish in young plants, porrect, sessile to long-stalked, the stalks to 1.6(–1.9) mm long, multiseriate, 2 cells wide, the rays (4–)6–8(–10), 1-celled, the midpoints 1-celled, obsolete to 2/3 the length of the rays; sparsely armed (less often with unarmed young stems), the prickles 1–3 mm long, 1.1–3 mm wide at base, deltoid, straight, flattened, stramineous to ochraceous or brown, with stellate trichomes like those of the stem on the base; bark of older stems glabrescent to moderately stellate-tomentose, dark brown. Sympodial units difoliate, geminate, the leaves of a pair anisophyllous. Leaves entire (sometimes lobed in juvenile plants); the major leaves 7.5–19.5 cm long, 5–8.5 cm wide, elliptic; base cuneate, asymmetric; the margins entire, less often sinuate; apex acuminate; primary veins 6–9 pairs; minor leaves 2.3–7.5 cm long, 2.4–6.4 cm wide, elliptic to nearly circular; base attenuate to rounded, often symmetric; margins entire; apex obtuse to rounded, less often acute to acuminate; major and minor leaves all chartaceous, markedly discolorous, drying green to dark brown above, with the indumentum giving an ochraceous to ferruginous appearance, and green, greenish-gray or dark brown beneath, with the indumentum giving a stramineous to ochraceous appearance; the adaxial surface moderately stellate-puberulent to densely stellatetomentose, the epidermis always visible, the trichomes stramineous to ferruginous, porrect, sessile to long-stalked, the stalks to 0.7 mm long, multiseriate, 2–3 cells wide, multiradiate, the rays (3–)4–8(–10), 1-celled, the midpoints 1-celled, 1/6 the length to longer than rays, usually oblique, the multiradiate trichomes more rayed, these usually ferruginous; the abaxial surface stellate-tomentose (always denser than the adaxial one), the trichomes hyaline, stramineous or ochraceous, porrect, sessile to long-stalked, the stalks to 0.8 mm long, multiseriate, 2–3 cells wide, the rays 4–8(–11), 1-celled, the midpoint 1-celled, 1/6 the length to almost same length the rays; unarmed above (sometimes armed in juvenile forms), rarely armed along the midrib beneath, the prickles 0–4, deltoid, straight and flattened; petiole of major leaves 0.5–2.5(–3) cm, densely stellate-tomentose, with trichomes like those of the stem, usually unarmed, less often armed with up to 5 prickles, these deltoid, straight, and flattened; petiole of minor leaves 0.36–1.3 cm, densely stellate-tomentose, with trichomes like those of the stem, unarmed or with up to 3 prickles. Inflorescences a reduced monochasial cyme, 0.8–4 cm long, unbranched, leaf-opposed or nearly so, with (3–)5–20 flowers, 1–2 flowers open at a time; inflorescence axis (peduncle plus rachis) densely stellate-tomentose, the trichomes porrect, sessile to long-stalked, the stalks to 2.5(–3) mm long, multiseriate, 2–5 cells wide, sometimes multiangular, the rays 4–10, the midpoints 1-celled, ½ the length to longer than rays, unarmed; peduncle (0–) 1.5–7 mm long, straight; the rachis curved, 2–33 mm long; pedicel insertion points closely spaced, to 1 mm apart, often obscured by the dense and relatively long indumentum of the inflorescence axis; pedicels usually straight, 4–8 mm long at anthesis, articulated at base, unarmed, densely stellate-tomentose, with trichomes like those of the inflorescence axis. Flowers 5-merous, heterostylous, basal flowers long-styled and hermaphroditic, short-styled and functionally male flowers produced distally in the inflorescence (see Sexual Expression and Inflorescence). Calyx tube cupuliform, (2.2–) 2.5–4.3 mm long, densely stellatetomentose, often denser than the pedicels and inflorescence axis, with trichomes like those of the inflorescence axis, unarmed; calyx lobes deltate to shallowly triangular, 1–1.8 mm long, 1.8–3 mm wide at base. Corolla 1.8–2.9 cm in diameter, white, stellate, with interpetalar tissue, lobed for 1/5–1/2 of its length, the lobes 7–8.2(–9.8) mm long, 5–9.8 mm wide, deltate, the apex cucullate, densely stellate-tomentose along whole length abaxially, the trichomes hyaline to ferruginous, porrect, sessile to short-stalked, multiseriate, the rays 8–10, tortuous, the midpoints 1-celled, 3/5 the length to longer than rays, the adaxial surface sparsely to moderately stellate-tomentose at the apex, usually with trichomes like those of the abaxial surface becoming gradually sparser towards the base, the basal half glabrous or nearly so. Stamens equal; filament tube 1.4–2 mm long; free portion of the filaments 0.6–1.8 mm long; anthers 5.7–10 mm long, 1.5–2.1 mm wide, elliptic to lanceolate, sagittate at base, narrowed towards the apex, dehiscing by apical pores, connivent or not. Ovary short-cylindrical, convex at apex, with small glandular trichomes at apex, sometimes with sessile stellate trichomes; style of long-styled flowers 10.3–12.8 mm long, white, cylindrical, straight to gently curved distally, usually with some sessile, stellate trichomes at base, in short-styled flowers the style 2.5–6 mm long, straight; stigma 0.8–0.9 mm long, rounded to bilobed at apex, green, the surface papillose. Fruit a spherical to obloid berry, 9.8–14 mm long, 12.5–15 mm wide, the pericarp smooth, glabrous, the exposed portion whitish to pale green at maturity; fruiting pedicels 8.5–13.5 mm long, 1.2–2.1 mm in diameter at base, unarmed; fruiting calyx accrescent, covering 1/ 2 to 1/4 of the mature fruit, truncate to rounded at base, the lobes (3.9–) 4.2–7.5 mm long, 5.7–8.8 mm wide at base. Seeds ca. 20–50 per berry, 3–4 mm long, 2.7–3.3 mm wide, flattened, reniform, stramineous to brown. Chromosome number: not known. Figures 2 , 13 .

Habitat and Distribution —In southern Brazil, S. piluliferum is known from several localities in eastern Paraná and Santa Catarina States, and in southeastern Brazil it is known from eastern Minas Gerais and S~ ao Paulo States, and throughout the state of Rio de Janeiro ( Fig. 8 ). Solanum piluliferum grows at the edge of wet forests, roadsides, and disturbed areas near these forests; from 200 to 1500 m elevation.

Phenology —Flowering collections have been made between September and April, with a flowering peak from September through November; fruiting specimens have been collected from December to May.

Preliminary Conservation Status — Solanum piluliferum is considered of least concern (LC) due to its relatively wide distribution (EOO: 202,475 km 2), abundant populations and numerous collections across the distribution range, although the small area of occupancy (AOO: 188 km 2) suggests that it would merit being treated as endangered (EN) ( IUCN 2016).

Etymology —The specific epithet “piluliferumʺ probably derives from the Latin noun “pilaʺ meaning “ballʺ or “bullet,ʺ due to Dunal’ s mention in the original description “[…] Calyx post anthesin globosus, ovarium tegens, piluliformis.ʺ: Calyx globose after anthesis, covering the ovary, ball- or bullet-shaped.

Additional Specimens Examined — Brazil. — GOIÁS: “in prov. Goyazana ante Sobradinhoʺ, Pohl s.n. (syntype, BR [BR000005538669]); Pohl s.n. (syntype, BR [BR0000005538027]).— MINAS GERAIS: Mun. Juiz de Fora, Monte Verde/Rio Santa Bárbara, Jan 2007 (fl, fr), Viana & Maciel s.n. ( BHCB); Reserva Biológica Municipal Santa Candida ˆ, 8 Aug 1997, Lafetá 252 ( CESJ); Mun. Lima Duarte, Serra Negra, RPPN Fazenda Serra Negra , 5 Apr 2009 (fr), Oliveira et al. 53 ( CESJ); Mun. Miradouro, Córrego Alegre, 12 Jan 2001 (fl), Salino & Morais 5994 ( BHCB); Mun. Rio Preto, Serra Negra, mata atrás do Cambu´ı, 26 Jan 2007 (fl, fr), Feliciano et al. 18 ( CESJ); Mun. Santa Rita de Jacutinga, 30 Sep 1989 (fl), Grandi 2630 ( BHCB). — PARANÁ: Mun. Adrianópolis, Parque das Lauráceas, 17 Nov 1999 (fl), Barbosa & Abe 403 ( CESJ, FUEL, ICN, NY); Mun. Guaratuba, Castelhanos, 20 Jan 1994 (fl), Kummrow et al. 3334 ( BHCB, MBM, NY); Pirizal, 14 Dec 1971 (fl, fr), Dombrowski & Kuniyoshi 3886 ( BHCB, MBM). RIO DE JANEIRO: Mun. Nova Friburgo, Parada Augusto Alves, 1 Jan 1997 (fl, fr), Kinupp 121 ( BHCB, FUEL, HCF); Mun. Nova Iguaçu, Distrito de Tinguá, Rebio, Estrada do Ouro, 400–700 m, 24 Oct 2002 (fl), Bovini et al. 2213 ( BHCB, RB); Mun. Petrópolis, Estrada do Ribeir~ ao n° 68, 12 Nov 2000 (fl), Alves 1 ( CESJ); Fazenda Itaipava, beira da estrada de acesso, 17 Nov 2005 (fl), Siqueira & Gracff 38 ( BHCB, RB); Mun. Resende, Margens da rodovia BR-354 indo de Engenheiro Passos para Itamonte, ca. 2 km antes da entrada para a parte alta do Parque nacional do Itatiaia, 22°23, 12ʺS, 44°45, 12ʺW, 1439 m, 20 Nov 2013 (fl), Giacomin et al. 2023 ( BHCB); Mun. Rio Claro, Rio do Braço, 1 Dec 2004 (fl), Oliveira 1100 ( MBM, RB); Mun. Santa Maria Madalena, Alto Imbé, próximo à plantaçao~ de Eucalyptus à margem da rodovia RJ-180, 21°59, 59ʺS, 41°56, 45ʺW, 282 m, 12 Sep 2014 (fl), Gouvea ˆ& Falcao~ 138 ( BHCB); Sellow 135 (W [W1889-0291695]). SANTA CATARINA: Mun. Angelina, Linha do Chaves, 27°32, 24ʺS, 48°57, 36ʺW, 718 m, 6 Apr 2010 (fr), Stival- Santos et al. 2346 ( BHCB FURB); Mun. Apiúna, Faxinalzinho, 27°10, 49ʺS, 49°23, 37ʺW, 793 m, 17 Mar 2010 (fr), Korte & Kniess 2187 ( FURB); Mun. Jaraguá do Sul, Garibaldi, 26°33, 08ʺS, 49°10, 33ʺW, 180 m, 26 Jan 2010 (fr), Dreveck & Carneiro 1592 ( FURB); Mun. Rodeio, Sao ~ Pedro, 26°54, 37ʺS, 49°24, 41ʺW, 611 m, 30 Mar 2010 (fl, fr), Korte & Kniess 2340 ( FURB); Próximo a divisa com Benedito Novo, Morro do Ipiranga, 26°52, 19ʺS, 49°24, 13ʺW, 775 m, 15 Oct 2012 (fl), Funez 1108 ( FURB); Mun. S~ ao Bento do Sul, arredores do Cepa Rugendas-Univille, Rio Natal, 26°09’01”S, 49°13, 27 ̎ W, 7 Feb 2009 (fl, fr), Meyer 899 ( BHCB, JOI); Próximo à comunidade de Rio Natal, CEPA Rugendas, area pertencente à Univille, em margem de fragmento de Floresta Ombrófila Densa, na estrada que leva ao alojamento, 26°19, 24 ̎ ʺS, 49°18, 29 ̎ ʺW, 670 m, 10 Feb 2012 (fl,fr), Giacomin et al. 1706 ( BHCB, NY). —S AO ~ PAULO: Mun. Bom Sucesso de Itararé, estrada de Bom Sucesso a 2 km da Mineraçao~ de cal Sao ~ Judas Tadeu, 24°19, 13 ̎ ʺS, 49°13, 04 ̎ ʺW, 15 Dec 1997 (fr), Chung et al. 154 ( BHCB, ESA, IAC); Mun. Iporanga, Fazenda Intervales, Estrada para Bocaina, Trilha Gruta da M~ aozinha, 22 May 1996 (fr), Hoch et al. 29 ( BHCB, SP, UEC); Trilha do Carmo, 21 May 1996 (fr), Hoch et al. 13 ( BHCB, SP); Mun. Mamparra, Reserva Florestal Carlos Botelho, a 2km da “sedinhaʺ, beira de estrada, 15 Feb 1995 (fl), Miyagi et al. 484 ( BHCB); Mun. Ribeirao~ Grande, Parque Estadual Intervales, trilha para o mirante antigo, 9 Feb 2000 (fr), Amorim et al. 3303 ( CEPEC); Borda de estrada de S~ ao Pedro, próximo à entrada da

Trilha da Cachoeira da Pedrinhas, 24°18, S, 48°21W, 790 m, 18 Apr 2003 (fl, fr), Medeiros et al. 37 ( BHCB, HUEFS, UEC); Mun. S~ ao Jose do Barreiro, Outskirts of Parque Nacional Serra da Bocaina, road from park administrative headquarters at Sao ~ Jose do Barreiro to alojamento at entrance to park, 22°39, 50ʺS, 44°35, 33ʺW, 746 m, 2 May 2011 (fr), Agra et al. 7354 ( BHCB); Mun. S~ ao Miguel Arcanjo, Parque Estadual de Carlos Botelho, Trilha do Rio Taquaral, 24°03, 41ʺS, 47°59, 44ʺW, 853 m, 8 Dec 2011 (fl, fr), Bünger et al. 575 ( BHCB); State not indicated: according to protologue “In Brasilia australioreʺ, only “Brasiliaʺ in the herbarium labels, Sellow s. n. (syntype, K [K000590086]); Sellow s. n. (syntype, P [P00368471]); Sellow s.n. (syntype, BR [BR0000005538355]).

Notes — Solanum piluliferum is easily differentiated from other species in the S. asterophorum species group by its cupuliform calyx with short deltate lobes, which is usually densely covered with long-stalked stellate trichomes ( Fig. 2B ), entire leaves with acuminate apices ( Fig. 2C ) and sparsely armed stems with short, straight, deltate, flattened prickles. Although the adult individuals have entire leaves, juvenile plants may have lobed leaves and often possess a distinct reddish indumentum on the younger portions of their stems and leaves. In contrast to other species in the group, herbarium material made from distal reproductive shoots may lack prickles. The stramineous to ochraceo-ferruginous indumentum of the young stems also can help to differentiate S. piluliferum , especially from S. igniferum ; S. piluliferum is ocher or red in the distal and younger portions of the stems and in immature leaves, which usually become evenly stramineous when older, while S. igniferum has young stems and adaxial leaf surfaces evenly orange to ferruginous, without a marked differentiation between younger and older parts.

Solanum piluliferum is found in higher and cooler areas than other members of the group. Most specimens have been collected above 600 m elevation, and the species usually only occurs below this range in cooler regions (e.g. Gouvˆea & Falc~ ao 138, collected at 282 m elevation, near the municipality of Santa Maria Madalena in the State of Rio de Janeiro).

Solanum densiflorum Sendtn. ( Sendtner 1846) is an illegitimate later homonym of S. densiflorum M. Martens & Galeotti ( Martens and Galeotti 1845) , synonym of S. lanceolatum Cav. , a species belonging to Torva clade sensu Stern et al. (2011). Dunal’ s (1852) S. piluliferum var. densiflorum is therefore treated as a new name at infraspecific rank (Art. 6.11, McNeill et al. 2012). Sendtner (1846) cited syntypes collected by C. F. P. von Martius, J. B. E. Pohl, and F. Sellow, all of them cited without collector number in the protologue. We choose as lectotype one of the two Martius specimens from M [M0165943] because it best represents the described species.

Figures

FIG.2. Solanum piluliferum. A. Young plant; note the first trichotomous branching followed by dichotomous branching. B. Inflorescence; note the longstalked trichomes and the cupuliform calyx. C. Habit. D. Fruit. Photo credit D: L. Menini Neto.

FIG. 13. Scan of holotype of Solanum piluliferum (P. Claussen 58, P [P00368472]). Reproduced with permission of the Muséum National d’Histoire Naturelle, Paris.

FIG. 8. Distribution of Solanum piluliferum.