Like Solanum restingae S.Knapp but differing in smaller flowers with narrowly deltate to long-triangular calyx lobes, unwinged stems and usually somewhat auriculate leaves.

Brazil. Bahia: Mun. Tancredo Neves, Estrada para os distritos de Água Branca e Julião, ca. 14. 1 km de Tancredo Neves, 554 m, 13°26'36"S, 39°30'40"W, 12 Sep 2005 (fl), A.M. Amorim, J. Jardim, J. Paixåo, S. Sant’Ana & E. dos Santos 5210 (holotype: CEPEC [CEPEC-110253]; isotypes: BHCB [BHCB002643, BHCB019062]).

Shrub to small treelet 0.5–3 m tall; young stems terete, glabrous or minutely puberulent with simple uniseriate trichomes to 0.5 mm long; new growth glabrous; bark of older stems smooth, greenish brown. Sympodial units difoliate, geminate; leaves of a pair not differing in shape. Leaves simple, the major leaves 8–10(-15) cm long, 2–3(-5) cm wide, elliptic to obovate, usually widest near the middle or in the distal half, glabrous on both surfaces, fleshy in texture; primary veins 8 pairs, usually paler than the lamina; base sessile and more or less auriculate; margins entire; apex attenuate; petiole absent or < 0.1 cm long; minor leaves 3–5 cm long, 1–2 cm wide, differing from the majors only in size. Inflorescence 0.1–0.3 cm long, opposite the leaves, unbranched, with 4–7 flowers, glabrous; peduncle < 0.1 cm long; pedicels ca. 0.8 cm long, 0.5 mm in diameter at the base and apex, filiform, nodding at anthesis, glabrous, articulated at the base; pedicel scars tightly packed and almost overlapping. Buds ellipsoid to rounded, the corolla exserted ca. halfway from the calyx tube just before anthesis. Flowers 5-merous, perfect. Calyx tube 1.5–2 mm long, conical, the lobes 2–3 mm long, ca. 1 mm wide, narrowly deltate to long-triangular with a 1–1.5 mm long projection that in live plants is a fleshy knob, glabrous. Corolla 0.8–1 cm in diameter, white, stellate, lobed ½ to 2/3 of the way to the base, the lobes ca. 0.4 cm long, 0.2 cm wide, planar at anthesis, minutely puberlent at the tips and along margins. Stamens 3–4 mm long; filament tube ca. 0.5 mm long, the free portion of the filaments <0.5 mm long, glabrous; anthers 2.5–3.5 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores elongating to longitudinal slits with age. Ovary glabrous; style 4–5 mm long, glabrous; stigma minutely capitate, the surface minutely papillose. Fruit a globose or depressed globose berry, ca. 1 cm in diameter, green or pale whitish green, glabrous, the pericarp thick, not markedly shiny; fruiting pedicels ca. 1.5 cm long, ca. 3 mm in diameter at the apex, woody, deflexed; calyx lobes in fruit persistent and slightly elongating, occasionally breaking off but always with > 1 mm remnants. Seeds ca. 30 per berry, not known from mature fruit.

Figure 1. 

Photograph of living plants of S. amorimii, S. apiahyense and S. filirhachis. A Immature fruit of S. amorimii (Giacomin et al.1962) B Flowers of S. amorimii (Amorim et al. 5210) C Inflorescence with flower and fruit of S. apiahyense (Giacomin et al. 1086) D Habit of S. apiahyense (Giacomin et al. 1086) E Inflorescence, flower and leaves of S. filirhachis (Giacomin et al. 1854) F Fruit (immature) of S. filirhachis (Giacomin et al. 1854). Photographs: A (S. Knapp), B (A.M. Amorim), C–F (L.L. Giacomin).

Figure 2. 

Isotype specimen of S. amorimii (Amorim et al. 5210, BHCB).

Endemic to eastern Brazil in the states of Minas Gerais and Bahia, known from northernmost Minas Gerais and southern Bahia (Figure 3).

Figure 3. 

Distribution of S. amorimii.

Solanum amorimii is found in the understory of wet Atlantic forests (Floresta Ombrófila Densa, Mata Atlântica; IBGE 2012) from 50–1000 m, most commonly found at around 500–900 m elevation in very preserved sites.

Flowering specimens have been collected from July to October but appears to peak in August; fruiting specimens have been collected from September to April.

The species epithet honours André M. Amorim, curator of the herbarium at CEPEC in Ilhéus, Bahia, and collector of the type specimen, whose knowledge of the flora of Bahia has helped many botanists in the region and beyond.

Near-threatened (NT) B1, 2a, b (ii, iii); EOO 20,663 km² (NT); AOO 40 km² (EN). Although the large extent of occurrence (> 20,000 km²) places S. amorimii out of the vulnerable category, the small number of locations (5–10) and the fragmentation of its forest habitat mean it is of some conservation concern. Populations occur within several private protected areas (in Minas Gerais the only population is within a private reserve) so the species is afforded some protection. On the other hand, the known collections suggest the species is restricted to pristine sites, which are becoming increasingly rare. As with all Geminata species, it is possible that more populations remain to be collected; these plants are inconspicuous in the deep forest understory and usually occur in small, sparsely distributed populations.

Solanum amorimii is morphologically very similar to the sympatric S. restingae, but can be distinguished by its much smaller flowers with long-triangular calyx lobes and by its unwinged stem. Both species grow in the understory of mostly undisturbed forests and can be small shrubs or treelets. Solanum restingae has markedly cucullate corolla lobes, and the calyx lobes are so small as to be almost non-existent, especially in fruit. Bud shape also differs between the two species, with those of S. amorimii being globose to somewhat ellipsoid and those of S. restingae more elongate with a distinct “nipple” from the cucullate corolla tips. In fruit the two species can be difficult to distinguish, but the winged stems of S. restingae and the presence of calyx lobes in S. amorimii should enable identification.

Leaves of S. amorimii are usually somewhat auriculate at the base, with the base not surrounding the stem but enlarged to a very short petiole. Plants grow in forest understory, sometimes in open places such as treefall gaps. From overall morphology this species would belong to the S. arboreum species group of Knapp (2002a), but its relationships have not been tested using molecular sequences.

BRAZIL. Bahia: Mun. Arataca, RPPN Caminho das Pedras, Serra do Peito-de-Moça, entrada a 9.5 km no Assent. Santo Antonio, mais 8.9 km ate a sede da RPPN, trilha de acesso ao topo da serra, após a Mormaço, 15°10'27"S, 38°20'22"W, 900–936 m, 26 Nov 2006 (fr), A.M. Amorim et al. 6608 (CEPEC); Mun. Arataca, Serra do Peito-de Moça, estrada que liga Arataca a Una, ramal ca 22.4 km de Arataca com entrada do Assentiamento Santo Antonio, RPPN Caminho das Pedras, 15°10'25"S, 39°20'30"W, 1000 m, 20 Jan 2007 (fr), A.M. Amorim et al. 6730 (CEPEC); Mun. Arataca, Serra do Peito-de Moça, RPPN do IESB, rodovia Arataca/Una, entrada a 9.5 km de cidade, mais 8.9 km de entrada, trilha do mormaço, 15°10'27"S, 39°20'22"W, 700–900 m, 12 Aug 2009 (fl), L. Daneu et al. 81 (CEPEC); Mun. Arataca, Serra Novo Javi, RPPN do IESB, rodovia Arataca/Una, entrada a 9.5 km N, mais 8.9 km até a sede da RPPN, trilha da Serra, acesso ca. 1.5 km NE da sede, Topo da Serra, 15°10'42"S, 39°20'09"W, 12 Sep 2009 (fl), L. Daneu et al. 96 (CEPEC); Mun. Arataca, Serra Novo Javi, RPPN do IESB, rod. Una/Arataca, entrada 9.5 km N, mais 8.9 km até a sede da RPPN, trilha da serra, acesso ca. 1.5 km NE da sede, topo da serra, 15°10'42"S, 39°20'09"W, 759 m, 12 Sep 2009 (fl), L. Daneu et al. 121 (CEPEC); Mun. Camacan, RPPN Serra Bonita, trilha da pousada, 15°23'26"S, 39°33'55"W, 835–1000 m, 25 Aug 2007 (fl), F.M. Ferreira et al. 1326 (CEPEC); Mun. Arataca, Serra do Peito-de Moça, Serra do Peito-de Moça-Serra das Lontras, estrada Arataca-Una, ramal 22.4 km de Arataca, assentamento Sto. Antonio, RPPN Caminho das Pedras, 15°10'25"S, 39°20'30"W, 1000 m, 23 Sep 2007 (fl), F.M. Ferreira et al. 1452 (CEPEC); Reserva Pratigi, 28 km de Itamarati, 6 km no ramal a direita, sentido Gandu, 13°53'52"S, 39°27'26"W, 670 m, 22 Oct 2007 (fl), F.M. Ferreira et al. 1563 (CEPEC); Mun. Uruçuca, estrada de Itacaré para Serra grande, pouco após km 43, ramal à direta após acesso para a cachoeira do Tijuipe, área explorada do plano de manejo, 14°23'12"S, 39°04'45"W, 4 Apr 2004 (fr), P. Fiaschi et al. 2249 (CEPEC); Mun. Arataca, Serra Novo Javi, RPPN do IESB, Rod. Una/Arataca, entrada 9.5 km N, mais 8.9 km até a sede da RPPN, trilha da Serra acesso ca. 1.5 km NE do sede, Topo da Serra, 15°10'42"S, 39°20'09"W, 759 m, 12 Oct 2008 (fr), J.G. Jardim et al. 5408 (CEPEC); Mun. Una, Rodovia BA-265, a 23 km de Una, 50–75 m, 26 Feb 1978 (fr), S.A. Mori et al. 9299 (CEPEC, MO, NY); Mun. Almadina, Serra do Concavado, Rod. Almadina/Coaraci, ca. 5 km, 14°42'13"S, 39°36'09"W, 300 m, 19 Mar 2006 (fr), J.L. Paixão et al. 838 (CEPEC);. Mun. Wenceslau Guimarães, ca. 3 km W of Nova Esperança, W edge of Reserva Wenceslau Guimarães, 13°36’ S, 39°43’ W, 500–600 m, 14 May 1992 (fr), W.W. Thomas et al. 9244 (CEPEC, MO, NY, RB); Mun. Camacan, RPPN Serra Bonita, 9.6 km NNW of Camacan on road to Jacaraci and Jussari, then 6 km up road to Serra Bonita, 820 m, 21 Sep 2004 (fr), W.W. Thomas et al. 14224 (NY). Minas Gerais: Mun. Santa Maria do Salto, Distrito de Talismã, RPPN Loredano Aleixo (Fazenda Duas Barras), 16°24'01"S, 40°03'24"W, 873 m, 31 Oct 2013 (fl, fr), L.L. Giacomin et al. 1962 (BHCB, BM, UT); Mun. Santa Maria do Salto, RPPN Duas Barras, ca. 27 km do distrito de Talismã, trilha em direçåo a divisa com a Bahia, 16°14'56"S, 40°08'58"W, 8 Sep 2008 (fl), R.P. Oliveira et al. 1636 (HUEFS).

Brazil. São Paulo. Apiahy, Feb 1891(fl), J.I. Puiggari 3711 (lectotype, designated here: WU [WU0037965]).

Small erect shrubs, to 50 cm tall, often rhizomatous with a horizontal woody branch bearing several adventitious roots; young stems moderate to densely pubescent, with 4–8-celled hyaline trichomes to 2 mm long; new growth drying dark, densely pubescent; bark of older stems pale gray, glabrescent, not exfoliating. Sympodial units 3-plurifoliate, normally not geminate, if geminate, with leaves differing only in size. Leaves simple, 3.4–11 × 0.8–4 cm, elliptic to narrowly elliptic, membranous, slightly discolorous, shiny green adaxially when fresh, drying pale green beneath, dark above, not shiny, both surfaces moderate to densely pubescent with hyaline simple uniseriate trichomes 1–2 mm long with up to 5 cells, sometimes with a multicellular base (but see comments); primary veins 5–7 pairs, the midrib and primary veins darker abaxially, raised; base attenuate to acute, slightly decurrent onto the petiole, mostly symmetric; margins entire, not revolute, ciliate with antrorse hyaline trichomes; apex attenuate to acuminate; petioles 2.5–15 mm long, densely pubescent, with trichomes like those of the stems and leaves. Inflorescences 1.7 to 3.3 cm long, mostly lateral or less often strictly opposite the leaves, unbranched, with 3–5 flowers, moderate to densely pubescent, with hyaline trichomes like those of the stems and leaves; peduncle 4–15 mm long; pedicels 5 to 11 mm long, articulated at base; pedicel scars closely spaced ca. 1 mm apart. Buds globose to slightly elongate, the corolla mostly included in the calyx tube, exserted only just before anthesis. Flowers all perfect, 5-merous. Calyx tube up to 1 mm long, conical, getting reflexed, the lobes up to 0.9 mm long in flower, to 1.7 mm long in fruit, approximately 1.6 mm wide, acuminate and discretely keeled, adaxially, glabrous or papillose, covered with tiny 1–2-celled glandular trichomes, abaxially densely pubescent, with trichomes as those of the stem, or sometimes even longer, with 2.5 mm, and normally 5–6 cells. Corolla 1.5–1.7 cm in diameter, white, stellate, membranous, lobed from 2/3 to 3/4 of the way to the base, the lobes 7.5–9 mm long, 3–3.5 mm wide, reflexed at anthesis, deltate to lanceolate, glabrescent adaxially, abaxially sparsely pubescent, with 3–4-celled delicate simple trichomes of ca. 0.5 mm along the midvein, with tufts of few celled tiny trichomes less than 0.1 mm long on the tips and margins. Stamens 3.2–3.6 mm long; filament tube ca. 0.5 mm long, the free portion of the filaments up to 0.6 mm long equal in length or slightly unequal, and when so, one filament slightly longer (barely visible in dried material), glabrous; anthers 2.6–2.8 mm long, 1.6–1.8 mm wide, ellipsoid, slightly connivent, yellow, slightly sagittate at the base, the pores directed introrsely, opening into longitudinal slits at maturity. Ovary glabrous; style 4.2–5 mm long, white, straight, glabrous; stigma capitate, light green. Fruit a globose berry 0.7–1.4 cm in diameter (immature?), dull green, drying dark, the pericarp glabrous and not markedly shiny; fruiting pedicels 1.2–2 cm long, ca. 0.7 mm in diam. at the base, to 1.1 mm at the apex, with a slight constriction at the receptacle; calyx lobes in fruit somewhat enlarged. Seeds approximately 70 per fruit, known only from very young fruits, possibly flattened and with a marginal wing when fully developed.

Figure 4. 

Lectotype specimen of S. apiahyense (Puiggari s. n., WU). Reproduced with permission of the University of Vienna.

In the Serra do Mar mountain range in the Brazilian states of Paraná, Santa Catarina and São Paulo (Figure 5).

Figure 5. 

Distribution of S. apiahyense.

Solanum apiahyense is a rare and inconspicuous shrub of the understory and edges of well preserved and secondary fragments of the montane Brazilian Atlantic rainforest (Floresta Ombrófila Densa of IBGE 2012; Mata Atlântica), from 600 to 900 m. Although most collections are from well preserved sites, S. apiahyense is not exclusively associated with shaded environments. The species is also found along unpaved roadsides close to the type locality.

Fertile specimens are known from September to February. Mature fruits were observed only in October.

The epithet refers to the type locality, the city of Apiaí in southern São Paulo state.

Endangered (EN) B1; B2 ab (ii, iii, iv). EOO 3,208 km² (EN); AOO 16 km² (EN). Although the species occurs in a wide latitudinal range, it is locally rare, and is known from only six localities. None of the known populations are from within protected areas.

Solanum apiahyense, described more than a century ago (Witasek 1910), has not been assigned to any infraspecific group of Solanum so far. Recent phylogenetic analysis using molecular data (Giacomin 2015) has shown it to be closely related to S. trachytrichium, which was previously assigned to the Geminata clade (Knapp 2002a, 2008) and to its own subsection when originally described (subsect. Silicosolanum Bitter; Bitter 1919). Bitter (1919) based this on the unusual trichome morphology of hooked cells arising from a flattened multicellular base that give the leaves a feeling of sandpaper in herbarium specimens. Although molecular data support a close relationship between S. apiahyense and S. trachytrichium, the affinities of this clade are not clear-cut. Data from combined markers place it as sister to all other Geminata clade species, but with low support. In analyses of individual markers, it emerges as sister to either the Brevantherum or Geminata clades depending upon the marker used (Giacomin 2015).

Morphologically both taxa are easy to distinguish from most other Geminata species, and have the following assemblage of characters: both are small shrubs with leaves mostly not geminate, they have leaf trichomes with an expanded multicellular base and relatively large flowers (>1.5 cm in diameter). Among them, Solanum apiahyense and S. trachytrichium are easy to distinguish: S. trachytrichium has a unique scabrous indumentum on the leaf surfaces and stems, composed of short unicellular hooked trichomes on a mound-like multicellular base, while in S. apiahyense the surface is not rough to the touch, and although some trichomes with multicellular bases can be seen on leaves, these are translucent, very long (ca. 2 mm) and mostly 5-7-celled. These long trichomes of S. apiahyense are easily seen on the new growth, while S. trachytrichium trichomes are not visible to the naked eye. In addition, the flowers of S. apiahyense are slightly smaller, 1.5–1.7 cm in diameter versus 1.6–2.2 cm in S. trachytrichium.

In the past, the epithet S. apiahyense has been applied to more than one species of the S. inornatum group (part of the Brevantherum clade; Giacomin and Stehmann 2014) by various Solanum taxonomists, although they are now known to not be closely related. Although members of the S. inornatum group (e.g., S. inornatum Witasek, S. bradei Giacomin & Stehmann and relatives) and S. apiahyense are similar in habit and in having pubescence of long, translucent trichomes, they can be readily distinguished by close examination of the trichomes; those of S. apiahyense are multicellular with 5-7(8) cells while those of members of the S. inornatum group are mostly 3-celled (probably representing modified stellate hairs, Giacomin and Stehmann 2014). Fruiting specimens of S. apiahyense have peduncles longer than 1 cm and the pedicels are strongly apically expanded and constricted just beneath the calyx lobes (see Figure 1C), while in the species of the S. inornatum group species, the peduncles do not exceed 1 cm and the pedicels are never apically expanded with a distal constriction. Examination of trichomes with a 10× hand lens will allow easy identification of both flowering and fruiting material.

The type material found at WU (Puiggari 3711) consists of a single sheet, and does not match the photograph of a dried specimen in the original publication (Witasek 1910: tab. 30, fig. 2). It should therefore be treated as an isotype (Mentz and Oliveira 2004). As no further material could be found in other possible herbaria where J.I. Puiggari deposited his collections, the specimen at WU is here designated as a lectotype.

BRAZIL. Paraná: Mun. Cerro Azul, Serra Paranapiacaba, 20 Nov 1970 (fl), G. Hatschbach & O. Guimarães 25528 (MBM, RB); Mun. Doutor Ulysses, Barra do Teixeira, 16 Sep 2006 (fl, fr), J.M. Silva (HUFU, MBM, RB). Santa Catarina: Mun. Vidal Ramos, Mina Bugre, 27°21'35"S, 49°19'12"W, 598 m, 22 Sep 2009 (fl, fr), A. Korte & A. Kniess 243 (BHCB, FURB). São Paulo: Mun. Bom Sucesso de Itararé, Estrada de terra para Bom Suceso de Itararé, Próximo a Mineração de ouro São Judas (3 km após), 24°19'13.19"S, 49°12'49.49"W, 891 m, 11 Oct 2009 (fl, fr), L.L. Giacomin et al. 1097 (BHCB, BM, NY, RB); Mun. Bom Sucesso de Itararé, Estrada Bom Sucesso de Itararé, 2 km antes da Mineração São Judas, 24°19'13"S, 49°13'04"W, 15 Dec 1997 (fr), J.M. Torenzan et al. 647 (IAC, ESA, FUEL, SPSF, UEC).

Differs from the sympatric S. campaniforme Roem. & Schultes in its deep forest habitat, leaves with ruffled margins, flowers less than 1 cm in diameter, pedicels with a constriction at the distal end that are swollen in fruit, and few seeds.

Brazil. Espírito Santo: Mun. Santa Teresa, Comunidade de Santo Antônio, Propriedade do Sr. Boza, fragmento de floresta ombrófila densa após plantação de eucalipto, à direita da entrada, descendo o vale, 19°54'32"S, 40°35'26"W, 740 m, 8 Jun 2012 (fl, fr), L.L. Giacomin, L. Bohs, Y.F. Gouvêa & F.Z. Saiter 1854 (holotype: BHCB [2 sheet holotype: sheet 1 (fl) BHCB019056; sheet 2 (fr) BHCB019057]; isotypes: BM, MBML, NY, RB).

Erect shrubs to small trees, up to 3 m tall, normally branching close to the apex, the upper stems ascendant; young stems terete, glabrous; new growth brownish, glabrous. Bark of older stems turning pale greyish brown, glabrous, not exfoliating. Sympodial units difoliate, mostly geminate, with leaves not differing in shape or size. Leaves simple, 4.6–15.9 cm long, 1.3–4.9 cm wide, narrowly elliptic, membranous to chartaceous, slightly discolorous when dry, the adaxial surface glabrous, dark green and somewhat shiny in live plants, the abaxial surface sparsely pubescent with simple uniseriate 7–12-celled trichomes to 1 mm long in tufts in the primary vein axils, occasionally extending to the midrib; primary veins 5–9 pairs, yellowish green, discretely raised above, raised beneath; base attenuate to acute, slightly decurrent onto the petiole, sometimes asymmetric; margins entire, slightly undulate (ruffled) and revolute, apex long-attenuate to acuminate; petioles 1–9 mm long, glabrous. Inflorescences 3.5 to 26 cm long, opposite the leaves or internodal, unbranched, slender and very delicate, with 18–60 flowers, but bearing normally with 4–10 flowers at a time, glabrous; peduncle 1.8–3.8 cm long; pedicels 7–18 mm long, ca. 0.4 mm in diam. at the base, ca. 0.9 mm in diameter at the apex, with a constriction at the receptacle, articulated at base, unevenly spaced 1.7 to 10 mm apart. Buds globose, the corolla completely exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube to 1 mm long, conical, the lobes ca. 0.2 mm long, ca. 1.5 mm wide, acuminate and somwewhat keeled, papillose adaxially, glabrous abaxially. Corolla 6–8 mm in diameter, normally whitish purple adaxially, light purple abaxially, stellate, membranous, lobed more than ¾ the way to the base, the lobes 4–5 mm long, 1–1.7 mm wide, spreading at anthesis and becoming reflexed in older flowers, deltate to lanceolate, glabrous on both surfaces, minutely papillose at tips and margins. Stamens 2.5–3 mm long; filament tube ca. 0.3 mm long, the free portion of the filaments up to 0.2 mm long, equal in length or slightly unequal, and when so, two filaments slightly longer (barely visible in dried material), glabrous; anthers 2–2.5 mm long, 1.2–1.5 mm wide, ellipsoid, slightly connivent, yellow, poricidal at the tips the pores directed introrsely, elongating to longitudinal slits with age. Ovary glabrous; style 4–6 mm long, white, straight, glabrous, the stigma light grayish green, capitate. Fruit a globose berry 1–1.5 cm in diameter, dull green at maturity, with irregular black spots (Figure 1F) drying grayish brown, the pericarp glabrous, not shiny; fruiting pedicels 2.0–2.4 cm long, clearly obconical, ca. 0.5 mm in diam. at the base, widening markedly towards the apex to ca. 2.5 mm in diam.; calyx lobes in fruit ca. 1.5 mm long, commonly broken off in dried fruiting material. Seeds 20–25 per berry, 2.5–4.5 mm long, 2–3.3 mm wide, ovoid-reniform to somewhat flattened towards the margins, light to dark brown, the surface irregularly pitted, the testal cells undulate.

Figure 6. 

Holotype specimen (sheet two) of S. filirhachis (Giacomin et al. 1854, BHCB019057). Reproduced with permission of the Universidade Federal de Minas de Gerais.

Restricted to the state of Espírito Santo (Figure 7), in south-eastern Brazil. Collections are known from the central and northern parts of the state, from both sides of the Rio Doce.

Figure 7. 

Distribution of S. filirhachis.

Rare in the understory of well-preserved fragments of the sub-montane and montane Brazilian Atlantic coastal rainforest (Floresta Ombrófila Densa; IBGE 2012), normally in formations where granitic outcrops are present or close by, in elevations ranging from 200 to 750 m.

Fertile specimens of Solanum filirhachis are known mostly from the rainy season (from November to March), but the type collection from June indicates that the species might be fertile for a longer period. Mature fruits were observed in specimens from November and June.

The epithet refers to the long and slender inflorescence rachis, which is not observed in any of the Brazilian sympatric species, although a common feature in some species of the S. confine group from Colombia, Ecuador and Venezuela (Knapp 2002a).

Endangered (EN) B1, B2 ab (ii, iii, iv); EOO 1,136 km² (EN); AOO 20 km² (EN). Solanum filirhachis is currently known from only five localities, and all collections are from within private properties, where agriculture (both large and small scale) is known to occur. Despite the fact that it inhabits higher elevations that are usually harder to access and not always suitable for agriculture, we strongly recommend that further efforts to map new populations of the species should be undertaken, mainly within protected areas with similar forest types. Although the type locality of Santa Teresa in central Espirito Santo has several well preserved fragments of forest, the landscape has been rapidly transformed in the last few decades to Eucalyptus and coffee plantations, and summer vacation homes (cottages).

Solanum filirhachis is remarkably similar to a suite of species of the Geminata clade with ruffled leaf margins (see Figure 1E) and long filiform inflorescences (S. leptorhachis Bitter and S. nematorhachis S.Knapp from the W Andean slopes in Colombia and Ecuador and S. tenuiflagellatum S.Knapp of Venezuela). Knapp (2002a, 2008) treated these as members of her S. confine species group, all of whose members have a thin inflorescence rhachis, small flowers and leaves with ruffled (undulate) margins, although this latter character is impossible to see in herbarium specimens. Solanum filirhachis differs from those species in its distribution and in the tufts of trichomes in the abaxial leaf vein axils (domatia); other members of this morphologically similar set of species are glabrous or have fine, golden pubescence. The only Brazilian species Knapp (2002) placed in this group was S. stipulatum which can be easily distinguished from S. filirhachis by its shorter inflorescences, flowers with reflexed corolla lobes and winged stems with anisophyllous difoliate, geminate sympodial units. Solanum stipulatum is usually a shrub of watercourses, and often grows amongst rocks and is submerged in floods, while S. filirhachis is a slender treelet of forest understory. The relationships of the S. confine group have not yet been tested using molecular markers.

Another Brazilian species with which S. filirhachis could be confused is S. campaniforme that has similar (but somewhat stouter) elongate inflorescences and tufts of uniseriate trichomes in the abaxial leaf vein axils. Solanum filirhachis has leaves with ruffled margins tht normally dry pale green and smaller flowers (0.6–0.8 cm in diameter) that (at least in the type specimen) are tinged purple; S. campaniforme has leaves with entire, non-ruffled margins that normally dry black or brownish black and larger flowers (1.2–1.8 cm in diameter) with strongly cucullate corolla lobes.

We have designated a two sheet holotype for S. filirhachis in order to represent both flower and fruit in the type sheets.

BRAZIL. Espírito Santo: Mun. Águia Branca, Assentamento 16 de Abril, 18°54'25"S, 40°44'05"W, 150-200 m, 15 Mar 2006 (fl), V. Demuner et al. 1919 (MBML, BHCB); Mun. Santa Leopoldina, Colina Verde (Morro do Agudo), prop. Israel Elias Ramos (trilha da casa), 20°06'12"S, 40°26'30"W, 250-370 m, 29 Nov 2007 (fl, fr), V. Demuner et al. 4628 (MBML, BHCB); Mun. Santa Leopoldina, Pedra Branca, mata na Serra Santa Lucia, prop. Cristiano Bremencampi, 20°01'36"S, 40°29'32"W, 300-600 m, 30 Nov 2007 (fr), V. Demuner et al. 4655 (MBML, BHCB). Mun. Águia Branca, Rochedo, Trilha do Córrego, prop. Ailton Corteleti, 18°57'21"S, 40°48'05"W, 300-400 m, 19 Dec 2007 (fl, fr), V. Demuner et al. 4817 (MBML, BHCB).

Brazil. Sin loc. [probably Rio de Janeiro] “Silvis nondum cultis ad rivulae, vel stagna crescit” (no specimens located; lectotype, designated here: Vellozo, Flora fluminensis icones 2: tab. 93. 1831; epitype, designated here: Brazil. Rio de Janeiro: Mun. Nova Friburgo, RPPN Bacchus, Macaé da Cima, near Nova Friburgo, owned by David and Isabel Miller, Trilha da Aguada, 22°23'34.4"S, 42°30'03.4"W, 1470 m, 29 Oct 2012 (fl, fr), M.F. Agra, L. Bohs & L.L. Giacomin 7298 (RB [RB00718282, accession number 551172]; duplicates in BHCB, JPB, UT).

Shrub or small treelet 1–3 m (occasionally as small as 25–30 cm or as tall as 5 m); young stems terete, glabrous; new growth glabrous or minutely papillate; bark of older stems pale brown, with prominent paler lenticels. Sympodial units difoliate, geminate or more usually not geminate; leaves of a pair usually differing in size but not in shape. Leaves simple, 9.5–25 cm long, 3.5–9 cm wide, narrowly obovate, widest in the distal half, membranous, glabrous on both surfaces, the abaxial surface paler in dry specimens; primary veins 6–10 pairs, drying dark abaxially; base attenuate; margins entire; apex bluntly acute to attenuate; petiole 1–3 cm long, glabrous; minor leaves, if present, differing only in size from the majors. Inflorescences 0.1–0.5 cm long, terminal, more or less leaf-opposed or internodal and appearing pseudoaxillary, unbranched or occasionally furcate, with 5–10 flowers, glabrous; peduncle 0.1–0.5 cm long, the flowers in an apical clump; pedicels 0.9–1.1 cm long, < 0.5 mm in diameter at the base and apex, filiform, spreading at anthesis, glabrous, articulated at the base, with a constriction at the apex just below the calyx lobes, this becoming more pronounced in fruit; pedicel scars congested and overlapping at the tip of the very short inflorescence. Buds ovoid, the corolla strongly exserted form the calyx tube before anthesis. Flowers 5-merous, perfect. Calyx tube ca. 0.5 mm long, conical, the lobes 0.5–0.75 mm long, ca. 0.5 mm wide, deltate, with scarious margins and rounded tips, glabrous. Corolla 0.9–1 cm in diameter, white, stellate, lobed ca. 2/3 of the way to the base, the lobes 3–4.5 mm long, 1.5–3 mm wide, spreading or somewhat reflexed at anthesis, the tips and margins minutely papillose. Stamens 2.5–3 mm long; filament tube ca. 0.5 mm long, the free portion of the filaments < 0.5 mm long, glabrous; anthers 1.5–2 mm long, ca. 1 mm wide, ellipsoid to almost globose, yellow, poricidal at the tips, the pores lengthening to longitudinal slits with age. Ovary glabrous; style ca. 4 mm long, glabrous; stigma minutely capitate, the surface papillose. Fruit a globose to somewhat ellipsoidal berry, 0.5–1 cm in diameter, greenish white, occasionally pointed at the apex, the pericarp thin, shiny, brittle when dry; calyx lobes in fruit not markedly enlarging; fruiting pedicels 1–1.3 cm long, 0.5–1 mm in diameter at the base, enlarging gradually to 1.5–2 mm in diameter at the apex, with a slight constriction just below the calyx lobes, not markedly woody, pendant; calyx lobes in fruit not markendly enlarged. Seeds 10–20 per berry, 3–4 mm long, 2–3 mm wide, somewhat flattened-reniform (perhaps immature?), dark to blackish brown, the surfaces minutely pitted, the margins paler and thickened; testal cells pentagonal in outline.

Figure 8. 

Photograph of living plants of S. lacteum, S. psilophyllum and S. verticillatum. A Inflorescence and flower of S. lacteum (Agra et al. 7284) B Habit of S. lacteum (from Linhares, ES; no voucher) C Habit of S. psilophyllum showing rhizomatous growth (Giacomin et al. 186) D Flowers and young stems of S. psilophyllum (Giacomin et al. 186) E Fruit of S. psilophyllum (Giacomin et al. 186) F Immature fruit of S. verticillatum, inset shows pseudo-verticillate branching pattern (Giacomin et al. 2016). Photographs: A–E (J.R. Stehmann), F (S. Knapp).

Figure 9. 

Lectotype of S. lacteum. Vellozo (1831) Volume 2, plate 93. Reproduced with permission of the Natural History Museum Library.

Figure 10. 

Epitype specimen of S. lacteum (Agra et al. 7284, RB). Reproduced with permission of the Jardim Botânico do Rio de Janeiro.

south-eastern Brazil in the states of Espirito Santo, Minas Gerais and Rio de Janeiro (Figure 11).

Figure 11. 

Distribution of S. lacteum.

Solanum lacteum grows in wet Atlantic forests (Mata Atlântica, Floresta Ombrófila Densa) in forest understory of well preserved sites, from 600 to 1500 m elevation.

No apparent pattern in flowering or fruiting; specimens are often collected with only inflorescences, each plant is very few-flowered.

The species epithet was coined by Vellozo (1829) to refer to the whitish colour of the plant – “Color albescens totius plantae nomen triviale dedit” (the white color of the entire plant gives it its trivial name [epithet]). We have not observed entire plants that are white in colour, but suspect Vellozo (1829) was referring to the congested inflorescence that is completely white.

Near Threatened (NT) B1, 2 a, b(ii, iii); EOO 32,466 km² (NT); AOO 28 km² (EN). In spite of its large extent of occurrence, S. lacteum is only known from six locations and we consider it to be at risk due to the fragmentation and loss of its primary forest habitat. Populations in all three states of occurrence, however, are from within protected areas. It is possible that it is more common than it appears, considering that the flowers are so small and inconspicuous that it is easily overlooked.

Solanum lacteum is characterized by its tiny inflorescences with tightly packed flowers and the difoliate sympodia that are usually not conspicuously geminate. The leaves are narrowly obovate and widest in the distal third. They dry a characteristic blackish brown above and paler brown beneath. The inflorescences often occur internodally and are completely white, including the peduncle and pedicels. The colour of the leaves on herbarium specimens is similar to that of S. caavurana and S. campaniforme, but those species always have leaf pubescence on the lower leaf surfaces and more elongate inflorescences. The highly congested inflorescences of S. lacteum are distinctive and the species is not easily confused with any other growing sympatrically. It is somewhat similar to S. psilophyllum, which is similarly glabrous; differences between these two species are noted in the discussion of S. psilophyllum.

Solanum lacteum grows in the understory of undisturbed forest and can vary from being a tiny subshrub (see Figure 8B) to a small treelet ca. 5 m tall. This variation in height is common in members of the Geminata clade and may have to do with plant age and maturity.

Vellozo’s (1831) illustration (Figure 9) is not particularly clear, but the congested inflorescences and swollen fruiting pedicels with a slight distal constriction are clearly depicted. In addition, S. lacteum usually has prominent lenticels on the stems; these are also depicted in Vellozo’s plate. We have selected an epitype from Rio de Janeiro State to support this suboptimal plate with material that is fertile and shows the key characters (Agra et al. 7298).

We have recognised S. cormanthum here as a synonym of S. lacteum; after detailed study we consider the plate of S. cormanthum (t. 113) to represent flowering material of the same taxon as that shown in fruit in Vellozo’s plate of S. lacteum (t. 93). Solanum cormanthum was used by both Sendtner (1846) and more recently in the Lista de Especies de Flora do Brasil (Stehmann et al. 2014) to refer to a different taxon we here recognise as a narrow endemic from Minas Gerais (see S. psilophyllum below). Both these authors expressed reservations about the correct application of this name. As is the case with the plate of S. lacteum, the depiction of the plant is not particularly clear, but the small flowers, small anthers and inflorescences that appear axillary (although they are not) are characteristic of S. lacteum. The locality cited for S. cormanthum (“silvis maritimis Regii Praedii S. Crucis”; Vellozo 1829: 86) is well within the geographic range and habitat of S. lacteum, although today it is part of the city of greater Rio de Janeiro.

Sendtner’s (1846) plate of S. glomuliflorum (f. 11–15) clearly shows the scarious-margined calyx with rounded lobes and very plump anthers characteristic of S. lacteum. In his protologue Sendtner (1846) cited two collections of S. glomuliflorum; a flowering specimen of Schott from “Serra d’Estrella” (Serra de Estrela, in Rio de Janeiro State) and a fruiting specimen of Sellow’s from an unspecified locality in Brazil (F neg. 2823; presumably from Berlin]. We select here the Schott specimen at F (accession number 874710; barcode F0073278F) as the lectotype of S. glomuliflorum, as it bears a label with the locality and collector in J.F. MacBride’s handwriting and presumably comes from Berlin where the original is now destroyed. The collection number 5412 noted on this sheet was not mentioned by Sendtner (1846), but he rarely mentioned collection numbers in his citations.

BRAZIL. Sin. loc., Herb. Miers 2724 (BM). Espírito Santo: Mun. Cariacica, Reserva Biologica Duas Bocas, Alegre, trilha do Pau Oco, 20°17'29"S, 40°31'10"W, 600 m, 4 May 2008 (fr), A.M. Amorim et al. 7324 (BHCB); Mun. Cariacica, Reserva Biológica de Duas Bocas, localidade de Alegre, trilha do Pau-Oco, 20°17'29"S, 40°31'10"W, 600 m, 20 Jul 2008 (fr), A.M. Amorim et al. 7563 (BHCB, CEPEC, MBML, RB, UPCB); Mun. Santa Teresa, São Lourenco, Mata do Martinelli, trilha subindo o rio lado direito, 11 Apr 2000 (infl), V. Demuner et al. 885 (BHCB); Mun. Linhares, Reserva Florestal Linhares, km 0, 23 Jun 1999, D.A. Folli 3441 (BHCB); Mun. Santa Teresa, Nova Lombardia, terreno de Sr. Furlani, 19°48'14"S, 40°32'17"W, 813 m, 3 Feb 2011 (infl), L.L. Giacomin et al. 1200 (BHCB); Mun. Santa Teresa, Santo Henrique, terreno Waldecir Frey, 15 Apr 2005 (fr), L. Kollmann & A.P. Fontana 7642 (BHCB); Mun. Santa Teresa, Nova Lombardia, Reserva Biologica Augusto Ruschi, corrego entre os marcos 130 e 131, 2 Apr 2003 (fl), R.R. Vervloet & E. Bausen 2110 (BHCB). Minas Gerais: Mun. Matão, Estação Biológica de Caratinga, 23 Sep 1984 (fl, fr), P.M. Andrade & M.A. Lopes 346 (BHCB); Mun. Coronel Pacheco, Estação Experimental de Café Coronel Pacheco, 12 Aug 1941 (fl), E.P. Heringer et al. 702 (VIC); Mun. Caratinga, Fazenda Montes Claros, Estação Biológica de Caratinga, mata do Rafael, 19°43'53"S, 41°49'02"W, 5 Sep 1998 (fr), J.A. Lombardi et al. 2334 (BHCB); Mun. Caratinga, Fazenda Montes Claros, 10 Jan 1991 (st), J.R. Stehmann & C.V. Mendonça s.n. (BHCB); Mun. Tombos, Fazenda de Cachoeira, 12 Jul 1935 (fl), Mello Barreto 1577 (BHCB); Mun. Tombos, Mata do Banco, 13 Jul 2007 (fl), L. Leoni 6947 (BHCB). Rio de Janeiro: Mun. Nova Friburgo, RPPN Bacchus, Macaé da Cima, near Nova Friburgo, owned by David and Isabel Miller. Trilha da Antena, 22°22'31"S, 42°29'47"W, 1420 m, 29 Apr 2010 (fl, fr), M.F. Agra et al. 7296 (JPB, UT); Mun. Rio de Janeiro, Caminho do Macaco, 8 Aug 1878, A.F.M. Glaziou 9549 (B); Mun. Nova Friburgo, 1883, A.F.M. Glaziou 14177 (G).

Like Solanum evonymoides Sendtn. but differing in smaller flowers, inflorescences that are unbranched or branch only once near the base, pedicels with a constriction at the apex just below the calyx lobes and ovoid-reniform seeds.

Brazil. Minas Gerais: Mun. Mariana, Mina de Fazendão, em mata, próximo à ferrovia, 20°08'43.7"S, 43°24'48.4"W, 875 m, 29 Jul 2008 (fl, fr), L.L. Giacomin, J.R. Stehmann, S.G. Resende & F. Pena 186 (holotype: BHCB [BHCB019054]; isotypes: BHCB [BHCB019055], BM, NY, RB).

Treelet to 4 m, rhizomatous with underground stems; young stems terete, glabrous; new growth completely glabrous, occasionally minutely papillate; bark of older stems greenish brown, slightly winged from the leaf bases. Sympodial units difoliate, geminate; leaves of a pair differing in size but not usually in shape. Leaves simple, the major leaves 10–15(-25) cm long, 4–13 cm wide, elliptic to narrowly elliptic, occasionally wider in the distal third and narrowly obovate, membranous, glabrous on both surfaces, the abaxial surface often drying paler than the adaxial surface; primary veins 8–11 pairs, drying somewhat lighter than the lamina; base attenuate, somewhat oblique; margins entire; apex acute, the tip somewhat blunt; petiole 1.5–2 cm long, glabrous; minor leaves 6–8 cm long, 2–3 cm wide, differing from the majors only in size and sometimes not present in dried specimens. Inflorescences 0.2–2 cm long, opposite the leaves or appearing to arise from the leaf axils, unbranched, but apparently sometimes with 2 inflorescences from one axil and appearing branched (Giacomin et al. 186), with 5–8 flowers, glabrous; peduncle 0.1–2 cm; pedicles 1.2–1.5 cm long, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the swollen apex with a marked constriction just below the calyx lobes, slender and expanding distally, spreading or pendant at anthesis, glabrous, articulated at the base; pedicel scars 0.5 -1 mm apart, more congested in the distal part of the inflorescence. Buds obovoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers 5-merous, perfect. Calyx with the tube 0.5–1 mm long, broadly conical, the lobes 1–1.5 mm long, deltate to triangular, reflexed at anthesis, glabrous. Corolla 1.2–1.4 cm in diameter, white, stellate, lobed 1/2 to 2/3 of the way to the base, the lobes ca. 5 mm long, 2.5 mm wide, spread at anthesis, glabrous with the tips minutely papillate. Stamens 3.5–4 mm long; filament tube ca. 0.5 mm long, the free portion of the filaments ca. 0.5 mm long, glabrous; anthers 2.5–3 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 5–6 mm long, glabrous; stigma not expanded, blunt, the surface minutely papillate. Fruit a globose berry, 1–1.3 cm in diameter, green, the pericarp not markedly shiny, thick; fruiting pedicels 1.5–1.7 cm long, ca. 1 mm in diameter at the base, 2.5–3 mm and expanded at the apex, woody and pendant; calyx lobes in fruit not markedly expanding, but distinctly differentiated from the enlarged pedicel apex. Seeds not known.

Figure 12. 

Holotype specimen of S. psilophyllum (Giacomin et al. 186, BHCB019054). Reproduced with permission of the Universidade Federal de Minas Gerais.

In the south-eastern part of the state of Minas Gerais, in islands of forest (capões) associated with iron or quartzite formations in the Iron Quadrangle and Serra do Cipó regions, in the southern limit of Espinhaço mountain range (Figure 13).

Figure 13. 

Distribution of S. psilophyllum.

Solanum psilophyllum grows in the forest understory on thin soils associated with iron-rich or quartzite formations, at elevations from 800–900 m.

Flowering specimens have been collected throughout the year; fruits have only been seen on the type specimen, collected in July. It is probable that this species flowers and fruits sporadically throughout the year.

Named for its completely glabrous leaves (from the Greek psilos smooth or bare, phyllos leaf).

Critically Endangered (CR) B1, 2 a, b(ii, iii, iv); EOO 26 km² (CR); AOO 16 km² (EN). Solanum psilophyllum is known from only two localities and its habitat is under severe pressure from mining and frequent forest fires (see Notes). The population from which the type specimen was collected, close to a private railroad, has already been destroyed. Although the area of occupancy would suggest a status of Endangered we consider the extreme threats to these populations coupled with the habitat specifity of members of the Geminata clade (see above) warrant a status of Critically Endangered.One of the known collections might be from a protected area (PARNA Serra do Cipó), although not stated on the specimen label (Campos & Belisário CFSC-13505) but appears to be from a roadside, subject to occasional fire.

Solanum psilophyllum is the species previously called Solanum cormanthum Vell. in Lista de Especies de Flora do Brasil (Stehmann et al. 2014). That name, however, has been of uncertain application since Sendtner (1846) listed a collection from Minas Gerais (“Caxoeira do Campo”) as belonging to S. cormanthum, but with reservations.

Three sheets of labelled as “Solanum cormanthum Vell.” in Martius’s hand in Brussels belong to this species as do presumed duplicates of this collection in F (F-680206) and G (G00016950) cited by Knapp (2008) as belonging to S. evonymoides Sendtn., a species now considered to only occur from coastal Bahia to northeastern Minas Gerais (see discussion of S. verticillatum below). Sendtner (1846) cites a collection in Martius’s herbarium from “Caxoeira do Campo, prov. Minarum, Martio floret: Martius”; this was probably collected by Claussen. One of the three of the sheets in BR (BR00000825373) is from Martius’s herbarium and is labelled “Mart. 1839.” Another sheet is definitely attributed to Claussen and collected in 1835, while the third is attributed (“comm. Schüch fil. 1850”) to Guilherme Schüch, the Baron of Capanema (Minas Gerais, currently the active iron mine of Capanema), who sent plants to Martius.

The Vellozo illustration of S. cormanthum (tab. 113, Vellozo 1831) has distinctly axillary inflorescences and is said to come from what is now the city of Rio de Janeiro (“Praedii S. Crucis”), an area of very different vegetation and soils than the iron or quartzite rich formations of Minas Gerais. We recognise S. cormanthum here as a synonym of S. lacteum, both on morphological and distributional grounds. Members of the Geminata clade are very similar morphologically and Vellozo’s plates are often distinctly suboptimal for secure identification. In view of the restricted distribution and habitat of these plants (see below) we prefer to describe this as new here rather than use S. cormanthum for these distinct and endangered populations.

Solanum psilophyllum has a very narrow distribution restricted to the Iron quadrangle, within areas that are today active mines, and to the Serra do Cipó region, were it was collected more than ten years ago, in forest fragments close to roadsides. The fact that no collections are known from northern areas of the Espinhaço range likely indicates that the distribution is extremely restricted to the region acround Serra do Cipó and the Iron Quadrangle. Efforts to locate new populations of this species are urgent, especially considering that most areas where it might occur are currently owned by mining companies and are subject to an intensive land use.

Solanum psilophyllum is morphologically similar to S. verticillatum (described here below), another completely glabrous species of the Geminata clade occurring in the states of São Paulo and Rio de Janeiro. It can be distinguished from that species by its longer calyx lobes and by the swollen distal portions of the pedicels that are markedly constricted just below the calyx lobes. In addition, the leaf texture of S. psilophyllum is somewhat fleshy, while leaves of S. verticillatum are brittle and chartaceous.

Solanum psilophyllum is also morphologically similar to S. lacteum from Atlantic forests in Rio de Janeiro, Espirito Santo and Minas Gerais states. It differs from that species in its larger flowers (>1 cm in diameter), longer inflorescences, elliptic rather than obelliptic leaves that do not dry a blackish brown colour and in the non-lenticellate stem. Like S. lacteum, S. psilophyllum is completely glabrous. Solanum psilophyllum has an underground stem (Figure 8C), like S. arboreum Dunal of northern South America and Central America (see Knapp 2002a); this characteristic may be more common in the Geminata clade than currently thought, as it is rare that the underground parts of these small shrubs are collected or even observed.

BRAZIL. Minas Gerais: Mun. Santana do Riacho, Serra do Cipó, Rodovia MG-010, Belo Horizonte a Conceição do Mato Dentro, ca. de 1.5 km antes da bifurcação para Morro do Pilar, pequeno capão da mata a direita, próximo a rodovia, 19 Nov 1993 (fl), M.T.V.A. Campos & A.J.M. Belisário CFSC-13505 (BHCB); sin. loc., 1835 (infl), P. Claussen s.n. (BR); sin. loc., 1839 (infl), P. Claussen s.n. (F); Caxoeira do Campo, Mar 1839 (infl), P. Claussen 200 (BR, G); Mun. Santana do Riacho, Serra do Cipó, ca. 400 m antes da bifurcação Morro do Pilar-Conceição do Mato Dentro, ca. 1.8 km da estrada, 2 Mar 2001 (fl), M. Groppo et al. 640 (BHCB); Mun. Catas Altas, Mina de Fazendão, próximo à area da cava, 20°07'38"S, 43°24'48"W, 970 m, 27 May 2008 (fl), S.G. Rezende et al. 2749 (BHCB); sin. loc., 1850 (infl), G. Schüch s.n. (BR).

Like S. evonymoides Sendtn. but differing in being a large tree with pseudo-verticillate very shiny chartaceous leaves, smaller, sweet-smelling flowers and orange berries with large seeds.

Brazil. São Paulo: Mun. Santo André, Paranapiacaba, Estação Biológica, 23°46'-23°48'S, 46°21'-46°17'W, 800 m, 30 Jul 1980. A. Custodio Filho & A.C. Dias 305 (holotype: SP [SP002705]; isotypes: BHCB [BHCB019061], BM [BM001120381]).

Tree to 8 m, the branching appearing somewhat verticillate with branches in congested groups; young stems terete, completely glabrous, usually shiny; new growth completely glabrous and shiny, in live plants sometimes purplish green; bark of older stems pale yellow when dry, in live plants greyish brown. Sympodial units plurifoliate, the leaves clustered along the stems. Leaves simple, 4.5–16 cm long, 2–5 cm wide, elliptic to obelliptic, usually narrowly so, chartaceous and somewhat brittle, both surfaces glabrous and shiny, drying a golden brown; primary veins 6–10 pairs, drying yellowish brown, not looping in a submarginal vein; base acute to acuminate; margins entire, sometimes revolute; apex abruptly acute to attenuate; petiole (0.5-)1–2 cm long, glabrous, drying pale yellowish brown. Inflorescences 2–5 cm long, terminal, appearing axillary but this due to short internodes and congested leaves, branching 1–2 times, with 30–40 flowers, completely glabrous; peduncle 0.5–2.5 cm long; pedicels 1.5–1.7 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, filiform, spreading at anthesis, glabrous, articulated at the base; pedicel scars unevenly spaced 1–2 mm apart, usually clustered at the tips of the inflorescence branches. Buds ellipsoid, the corolla completely enclosed in the calyx when young, exserted 2/3 to 3/4 of the way just before anthesis. Flowers 5-merous, all perfect, intensely sweet-smelling (Custodio Filho 305). Calyx tube 1–1.5 mm long, conical, the lobes 0.9–1 mm long, ca. 1 mm wide, broadly deltate, with scarious margins and a central thickened keel ending in a rounded point, glabrous or the tips with a few papillae. Corolla (1.4-)1.6–1.8 cm in diameter, white, stellate, lobed nearly to the base, the lobes 6–8 mm long, 2.5–3.5(-4) mm wide, spreading at anthesis, densely papillate on the cucullate tips, otherwise completely glabrous. Stamens 4.5–6 mm long; filament tube 1 mm long or less, the free portion of the filaments minute, <0.5 mm long, glabrous; anthers (3-)4–4.5 mm long, 1–1.2 mm wide, obellipsoid with the base narrower than the distal portion, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 5–7 mm long, glabrous; stigma minutely capitate, the surface papillose. Fruit a globose berry, 1–1.2 cm in diameter, pale green and white speckled (immature) becoming yellow or orange when ripe, the pericarp shiny and leathery, shattering when pressed and dried; fruiting pedicels 2–2.5 cm long, ca. 1 mm in diameter at the base, expanding gradually to ca. 2 mm in diameter at the apex, more or less woody, hanging; calyx lobes in fruit not markedly lengthening. Seeds 10–20 per berry, 5–5.5 mm long, 3–4 mm wide, reniform and somewhat flattened, dark brown with paler margins, the surfaces minutely pitted and usually quite thin the embryo easily visible, the testal cells with sinuate margins.

Figure 14. 

Holotype specimen of S. verticillatum (Custodio Filho & Dias 305, SP002705). Reproduced with permission of Instituto de Botânica, São Paulo.

Endemic to south-eastern Brazil, in the states of Minas Gerais, Rio de Janeiro and São Paulo; in the Serra do Mar and Mantiequeira mountain chains (Figure 15).

Figure 15. 

Distribution of S. verticillatum.

Solanum verticillatum grows on the montane coastal forests (Mata Atlântica) as a small tree in forests and secondary growth from 700 to almost 2000 m elevation. Plants can be as large as 10 cm in diameter, and form part of the low canopy of these forests.

Most flowering specimens collected in the months of June and July; fruiting in November-January. Sporadic flowering and fruiting apparently occurs throughout the year, but a flowering peak occurs in the austral winter (May-August), which is also the drier season.

Named for the pseudo-verticillate nature of the stems, where many branches appear to arise from a set of closely spaced nodes (Figure 8F inset).

Least Concern (LC); EOO 75, 516 km² (LC); AOO 60 km² (EN). Although only described here, S. verticillatum is known from many localities along the Serra do Mar, many of which are from within protected areas (e.g., Reserva Biológica do Alto da Serra de Paranapiacaba in São Paulo state and Reserva Ecológica de Macaé de Cima, in Nova Friburgo, Rio de Janeiro state). Where it occurs, S. verticillatum is relatively common.

Solanum verticillatum was considered a montane form of S. evonymoides by Knapp (2008); field collections in 2013 confirmed the distinctness of this species. Solanum evonymoides is known from coastal forests in Bahia and adjacent Espirito Santo, and eastern Minas Gerais and although morphologically similar to S. verticillatum is distinct in both habitat and in several morphological features. Solanum verticillatum differs from S. evonymoides in its tree habit, branches that appear verticillate due to short internodes (Figure 8F inset), smaller sweet-smelling flowers (< 2 cm in diameter), shiny chartaceous leaves, and orange berries.

Solanum verticillatum also resembles S. psilophyllum (another set of specimens previously recognised as S. evonymoides by Knapp 2008) in its glabrous shiny leaves. It differs from that species in its more broadly deltate calyx lobes, its distinctly pedunculate inflorescences (versus inflorescences that branch only very near the base in S. psilophyllum), its berry that is orange or yellow-orange when ripe, and in its flattened rather than ovoid seeds. These two species can be very difficult to distinguish, but the marked constriction just below the calyx lobes at the distal end of the swollen pedicel occurs only in S. psilophyllum.

This species was commonly collected until approximately the 1980s and populations from the Paranapiacaba reserve are well represented in SP. It is strange that more recent collections do not seem to have been made; this may be due to the tree habit of S. verticillatum and to its similarity to the more common species S. campaniforme and S. pseudoquina A.St.Hil. It can be distinguished from S. campaniforme by its shiny, completely glabrous leaves (the leaves of S. campaniforme have tufts of trichomes in the vein axils abaxially) and from S. pseudoquina by its equal anthers (those of S. pseudoquina are markedly unequal). It differs from both species in its yellow or orange berries and pseudoverticillate branching. Most specimens of S. verticillatum at SP were previously identified as S. pseudoquina.

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