Brazil. “Ex Brasilia” [probably collected in Mucuri River drainage in the State of Bahia, see discussion], no date, G.W. Freyreiss s.n. (lectotype, designated here: S [acc. # S-R-5812]).

Shrubs up to 3.5 m, erect, moderately branched. Stems terete, densely glandular-pubescent and prickly, the trichomes porrect-stellate or somewhat multangulate, sessile to long-stalked, the stalks to 1 mm long, the rays 5–20, 2–3-celled, unequal in length, distally glandular, the mid-point 2–3-celled, equal to or twice the length of the longest ray, distally glandular, the prickles to 1.7 cm long, 2–3 mm in diameter at the base, straight, acicular or only slightly flattened, yellowish-red, basally somewhat pubescent with stellate trichomes like those of the stems and some small, stalked, uniseriate glandular trichomes; new growth densely glandular pubescent with long-stalked stellate or multangulate trichomes and acicular prickles like those of the stems; bark of older stems greyish-dark brown. Sympodial units difoliate to plurifoliate, the leaves not geminate. Leaves simple, lobed; blades (10)20–42 cm long, (8)20–38 cm wide, ca. 1–1.2 times as long as wide, broadly elliptic to broadly ovate, membranous, discolorous, prickly on both surfaces along the veins, the prickles to 1 cm long, straight; adaxial surface densely glandular pubescent, the lamina always visible, the larger trichomes porrect-stellate or multangulate, short- to long-stalked, the stalks to 1 mm long, the rays 4–11, unequal in length, eglandular and 1-celled or 2–3-celled and gland-tipped, the mid-points 2–3-celled, usually longer than the rays, these mixed with smaller sessile to short-stalked porrect-stellate eglandular trichomes, the stalks if present to 0.1 mm long, the rays 2–5, 1-celled, ca. 0.5 mm long; abaxial surface densely glandular-pubescent, the lamina barely visible, the trichomes porrect-stellate to multangulate like those of the adaxial surface, but denser and much more delicate; principal veins 5–7 pairs; sparsely to moderately armed on both surfaces, the prickles 1–1.7 cm long, 1.3–1.8 mm in diameter at the base, straight, somewhat laterally compressed, usually larger abaxially; base cordate, not decurrent, the two major basal lobes 2.5–7 cm long at the longest point, obtuse to rounded, often overlapping each other across the petiole; margins lobed, the lateral lobes 1.5–4.8 cm long, 4–9 cm wide at base, obtuse or rounded or less often acute at the apex, both basal and lateral lobes sometimes with small secondary lobes; apex obtuse or rounded or less often acute; petioles 4.5–19.5 cm long, densely stellate-pubescent with trichomes like those of the stems, usually densely prickly. Inflorescences subopposite the leaves or internodal, 4.5–12 cm long, usually unbranched, rarely forked or trifurcate, with 11–35 flowers, up to 3 open at a time; axes densely glandular-pubescent and prickly, the trichomes porrect-stellate to multangulate, hyaline to yellowish-brown like those of the stems, the prickles ca. 1 mm long, straight, like those of the stems and leaves ; peduncle 2.6–6 cm long; pedicel scars generally unequally spaced, closely packed to spaced 2.3 cm apart; pedicels 4.8–18 mm long, densely pubescent and prickly with trichomes and prickles like those of the stem, but these often purple-tinged, articulated at base. Buds ellipsoid to globose-ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis, but enclosed in the calyx lobes. Flowers 5-merous, heterostylous, with basal long-styled co-sexual flowers and functionally staminate short-styled flowers that vary in proportion between inflorescences, the plants andromonoecious. Calyx with the tube 4.5–8.2 mm long, 9.4–15.2 mm in diameter, broadly cup-shaped to somewhat urceolate, inflated, purple-tinged (mainly along the margins and apex of the calyx lobes) to green, armed, densely pubescent with trichomes like those of the stem, but these sometimes purple and with some eglandular rays, the lobes 7.5–15.6 mm long, 6–9 mm wide, triangular, the margins plane to strongly undulate and revolute, the apices acute to caudate. Corolla 2.3–3.9 cm in diameter, purple to lilac or bluish-lilac, white in some stages of development, shallowly stellate to stellate, lobed 2/5 to 1/2 the way to the base, interpetalar tissue absent, the lobes 10.9–15 mm long, 8.8–13.4 mm wide, deltate to triangular, spreading at anthesis, abaxially glandular stellate-pubescent with trichomes like those of the leaves, adaxially almost glabrous with only a few stellate trichomes sparsely distributed along the veins and near the tips, the apex acute, slightly apiculate. Stamens equal; filament tube 1–2.1 mm long; free portion of the filaments 1.3–2.9 mm long, glabrous; anthers 7.5–10 mm long, 2.8–4.3 mm wide, broadly lanceolate and tapering, connivent, glabrous, yellow, abaxially swollen in the lower half (gibbous) and somewhat papillate, poricidal at the tips, the pores directed distally, slightly extrorse, not elongating to slits with age. Ovary conical to somewhat cupuliform, densely stellate-pubescent and glandular at the apex, becoming glabrous with age, the trichomes porrect-stellate, sessile, 2–7-rayed, with a 2–4-celled, eglandular or glandular mid-point longer than the 1-celled rays; style 13.7–15.9 mm long in long-styled flowers, 1.2–3.7 mm long in short-styled flowers, straight, glabrous; stigma large-capitate to clavate, up to 1.4 mm long in long-styled flowers, the surface papillose, green when fresh. Fruit a globose to somewhat compressed globose berry, 1–1.1 cm long, 1.2–2.3 cm wide, pale green to white, glabrous, but with scattered stellate trichomes at the apex, the pericarp somewhat shiny when dry, the berry almost completely enclosed in the saccate fruiting calyx; fruiting pedicels 1.4–2.2 cm long, 1.5–2 mm in diameter at the base, woody and somewhat deflexed from the weight of the fruit, armed with sparse straight prickles like those of the flowering pedicels; fruiting calyx strongly accrescent and inflated, completely enclosing the berry, the tube 1.6–2.1 cm long, 1.9–3.4 cm in diameter at the widest point, the base somewhat plicate and invaginate, the lobes 1.1–2.2 cm long, 1.3–1.9 cm wide, usually somewhat overlapping, densely glandular-pubescent with porrect-stellate to multangulate trichomes. Seeds ca. 230 per berry, ca. 2 mm long, 2.4 mm wide, flattened-reniform, dark brown, the testal cells sinuate in outline; stone cells absent. Chromosome number: not known.

(Fig. 3). Solanum aciculare is endemic to eastern Brazil. Records are mostly concentrated along the Mucuri River watershed, ranging from the Municipality of Teófilo Otoni, in north-eastern Minas Gerais State, to Mucuri on the southern coast of Bahia. A single collection (J.G. Jardim et al. 3151; CEPEC, NY) is known from further north, in Mun. Caatiba of the south-central region of Bahia State.

Solanum aciculare inhabits edges of small forest fragments, especially those at the base or on granitic outcrops (inselbergs) or in disturbed sites near these rock outcrops, such as borders of unpaved roads and pastures. It is also found in herbaceous to arboreal vegetation growing along the Brazilian sandy coastal lowlands (restinga sensu Araújo (1992)), where plants grow in open disturbed areas dominated by grasses and at the edge of forest fragments (Fig. 2A). Habitats vary from environments subject to periods of drought (e.g. the edge of small seasonal semi-deciduous forest fragments or vegetation islands on inselbergs) to constantly wetter environments, at the edge of coastal evergreen forests, where the climate is under a strong oceanic influence. Plants have been collected from sea level to about 900 m elevation.

None recorded.

(IUCN 2020). EOO (38,277 km², NT); AOO (56 km², EN). Despite the relatively large range of S. aciculare, all collections are from only three broad localities and all are outside protected areas; vulnerability of the habitats in which the species occurs and the small number of localities suggest S. aciculare should be considered Endangered using the criteria B2 a, b (ii, iii, iv).

The name Solanum aciculare has not been used previously in treatments of Brazilian Solanum (Sendtner 1846) and, until recently, had been considered an ‘unassigned’ name (Flora e Funga do Brazil 2024). Examination of the type specimen and analysis of the collecting trajectory of Georg Freyreiss (see below or in Taxonomy) make clear that this plant is identical to that described as S. kollastrum (Gouvêa et al. 2018). Sendtner (1846) had not seen the specimen on which S. aciculare was based and his description was of another plant (Sellow s.n.) that he admitted did not completely correspond to Swartz’s description (“Solano subscandenti non dissimile in unico specimine suppetente a diagnosi Swartziana paullisper recedens. Proferamus descriptionem nostro specimini accomadatam, momenti, quo discrepat Swarztius haud immemores” - Not dissimilar to Solano subscandens in the only specimen available, departing for a moment from the Swartzian diagnosis. Let us present the description attached to our specimen, the importance of which Swartzius disagrees, not unmindful of it: transl. SK). The specimen described by Sendtner as S. aciculare is S. cordifolium Dunal, a member of the Erythrotrichum clade.

Amongst the species in this group, only S. aciculare and S. sublentum have cordate leaf bases coupled with glandular trichomes throughout the stems and leaves. Decurrent leaf bases of S. aciculare are only seen in the first leaves of the seedlings, with the subsequent leaves gradually changing shape to become cordate and non-decurrent. In contrast, the leaf bases in S. hexandrum and S. stagnale remain decurrent throughout plant development, varying in shape from attenuate to truncate.

Figure 3. 

Distribution of S. aciculare.

Solanum aciculare closely resembles S. sublentum, from which it can be readily distinguished by the robust long-stalked (up to 1 mm) stellate-glandular trichomes with all rays having a glandular distal cell (some rays may lose the glandular cell through breakage or by the disruption of the gland wall) on young stems, petioles and inflorescence axis; trichomes in S. sublentum are usually simple. Solanum aciculare and S. sublentum have very similar floral morphologies, sharing well-developed calyces that are strongly accrescent in fruits, showy purple to lilac corollas and robust anthers. Fruits of S. aciculare are completely enclosed in the accrescent calyx, whereas those of S. sublentum are exposed. Their leaves also resemble each other: both are lobed (with secondary lobes or not), elliptic to ovate (or broadly ovate in S. aciculare) and have cordate bases (varying from truncate to cordate or sagittate in S. sublentum). In the field, S. aciculare has notably larger leaves than those of S. sublentum; however, usually only the apices of the branches are collected, with the fully developed leaves not represented in herbarium material, so this character is often not apparent from herbarium specimens. Although S. aciculare and S. sublentum occur in similar environmental conditions (associated with outcrops or at edges of lowland forests), they have not been observed in sympatry.

Both S. aciculare and S. phrixothrix have densely bristly stems and elongate slender inflorescences. They can be distinguished by the glandular long-stalked stellate pubescence of S. aciculare (versus eglandular bristles and lack of long-stalked stellate trichomes in S. phrixothrix) and flower shape and colour (purple and stellate in S. aciculare versus white and rotate in S. phrixothrix).

The German collector Georg Freyreiss (also sometimes spelled Freyreis), who was principally an ornithologist and Friedrich Sellow, a botanist, travelled and collected in Brazil in the first decade of the 19^th century. Sellow’s botanical collections comprise many hundreds of specimens and were the basis for many new species (Urban 1893; Moraes 2023). In 1815, Freyreiss and Sellow were planning a trip to the north of Rio de Janeiro. They joined forces with Prince Maximilian of Wied, a member of the German aristocracy who was interested in natural history and had been inspired by Baron Alexander von Humboldt to travel to South America, particularly to Brazil (Moraes 2009, 2011). Wied was keen to travel via the, at the time, relatively unexplored coast and, with Freyreiss (Sellow having stayed in what is now the State of Espirito Santo), reached the Rio Mucuri near the border of Espirito Santo and Bahia States, then continued to Salvador in today’s State of Bahia. The type specimen of S. aciculare was probably collected by Freyreiss in the Mucuri River drainage during this voyage.

Brazil. [Rio de Janeiro]: “habitat silvis nondum cultis”; (lectotype, designated by Knapp et al. (2015), pg. 831: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_125] and later published in Vellozo, Fl. Flumin. 2: tab. 122. 1831).

Shrubs (0.5-)1–2.5 m tall, erect or occasionally somewhat spreading, strongly armed. Stems terete, glabrous to densely pubescent and/or bristly, sparsely to densely prickly, the stellate trichomes long-stalked, eglandular, porrect-stellate or less often appearing simple due to the complete lack of rays, the stalks 1–5 mm long, multiseriate, the rays 4–7, ca. 1 mm long, the mid-point ca. 0.5 mm long, always shorter than the lateral rays, with age the trichomes becoming thicker and the stems then densely bristly, the bristles often tipped with stalks and rays, underlying pubescence of minute papillate trichomes dense, more apparent on more glabrous individuals, the prickles 0.3–2 cm long, yellowish-golden, broad-based, the base 1.5–2 mm in diameter; new growth glabrous to densely stellate-pubescent and bristly, the multiseriate stalks of stellate trichomes usually shorter than the rays, but lengthening with leaf expansion; bark of older stems dark brownish-black in herbarium specimens, dark brown in live plants. Sympodial units unifoliate or difoliate, the leaves not geminate. Leaves simple or shallowly lobed (in some specimens, for example, de Paula 641, Giacomin 1827 deeply lobed) and repand, the blades 12–35(-40) cm long, 8–26(-30) cm wide, ca. 1.3–1.5 times as long as wide, broadly elliptic to narrowly obovate, usually widest in the basal half, membranous, concolorous, usually prickly on both surfaces with scattered straight prickles to 0.4–1.5 cm long, the prickles occasionally absent; adaxial surface glabrous to sparsely to moderately and evenly pubescent with long-stalked porrect-stellate trichomes, the stalks 1–1.5 mm long, multiseriate and arising from an expanded base, the rays 4–7, 1–2 mm long, the mid-point 0–1 mm long, always shorter than the rays, in some individuals, the rays often lost and the trichomes then appearing to be composed of a multiseriate base 1–1.5 mm long with a single celled tip often bent at 90° to the leaf surface; abaxial surfaces glabrous to evenly and densely pubescent with similar porrect-stellate long-stalked trichomes, but the stalks thinner and shorter than those on the adaxial surfaces and the rays occasionally more numerous, the trichomes denser along the veins, the surface densely dotted with crystal sand (inclusions of calcium oxalate, this not visible on the upper surfaces); principal veins 4–6 pairs, prickly or not, the prickles, if present, 0.4–1.5 cm long, on both surfaces; base attenuate on the petiole with a wing of ca. 2 mm wide along half the petiole length, sometimes to base of petiole; margins entire to 6-lobed, the lobes usually shallow, the sinuses less than 1/4 of the distance to the mid-rib; apex acute to attenuate; petiole (0.1-)2–10 cm long, glabrous to densely stellate pubescent and bristly, usually sparsely prickly. Inflorescences opposite the leaves or internodal, (1.5-)2.5–6(-8) cm long, unbranched (rarely furcate), with 3–10 flowers; axes glabrous to densely stellate-pubescent and prickly like the rest of the plant, the bristles and trichomes grading into each other and not distinct in morphology; peduncle (0.5-)2–7 cm long; pedicels 1–2 cm long, 1–1.5 mm in diameter at the base, 1.5–2 mm at the apex (excluding trichomes), erect to spreading, glabrous to densely stellate-pubescent and bristly, if prickly, the prickles ca. 1 mm long and thinner than those of stems and leaves, articulated at the base; pedicel scars more or less evenly spaced 5–7 mm apart on mature inflorescences, more tightly packed distally. Buds globose to broadly elliptic, the corolla completely included in the saccate calyx tube until just before anthesis, the younger buds less bristly and prickly than older ones. Flowers 5–6-merous, mostly co-sexual, but a few distal flowers are sometimes short-styled and probably functionally staminate. Calyx with the tube 4–7 mm long, 7–10 mm in diameter, cup-shaped and often completely closed in bud sometimes until just before anthesis, green or purple-tinged, the lobes 5–10 mm long, irregularly tearing at anthesis, but generally broadly triangular to deltate, acute to acuminate apically, glabrous to bristly and prickly with long-stalked bristles/trichomes, these with or lacking rays, the multiseriate stalk often purple-tinged. Corolla 3–6 cm in diameter, purple (rarely white), stellate, lobed halfway to the base, interpetalar tissue thin, glabrous, the lobes 15–21 mm long, 8–15 mm wide, deltate, spreading at anthesis, abaxially sparsely to densely pubescent with long-stalked porrect stellate trichomes, the stalks to 1 mm long, these denser at the tips and along the petal mid-vein, pubescence of corollas often purple-tinged, adaxially glabrous or with a few weak stellate trichomes on the mid-vein, the mid-veins often white adaxially. Stamens equal; filament tube minute to 0.5 mm long, glabrous; free portion of the filaments 1–1.5 mm long, glabrous; anthers (7-)9–11 cm long, 2–3 mm wide, broadly lanceolate and tapering, connivent, glabrous, yellow, abaxially swollen in the lower half (gibbous) and somewhat papillate, poricidal at the tips, the pores directed distally, not elongating to slits with age. Ovary conical, sparsely to densely pubescent with sessile stellate trichomes with rays 2–3 mm long, these soon deciduous; style 10–15 mm long in long-styled flowers (in rare short-styled flowers, the style 3.5–7 mm long), straight, glabrous; stigma clavate or broadly capitate, the surface minutely papillose. Fruit a globose to flattened globose berry, 2–2.5(-3.5) cm in diameter, green or pale whitish-green, glabrous, the pericarp somewhat shiny when dry, the berry almost completely enclosed in the accrescent saccate calyx; fruiting pedicels 1.8–2.3 cm long, 1.7–2.5 mm in diameter at the base, woody and spreading to somewhat deflexed from the weight of the fruit; fruiting calyx strongly accrescent, inflated or not, almost completely enclosing the berry, the tube 1.5–2 cm long, the lobes 1.5–2 cm long, irregular, usually overlapping, glabrous to sparsely to densely bristly and prickly with multiseriate bristles occasionally topped with porrect rays. Seeds ca. 100 per berry, 2–3.5 mm long, 1.5–2 mm wide, flattened reniform to somewhat ovoid, unwinged, reddish-brown or dark brown when dry, the surface minutely pitted, the testal cells pentagonal in outline, equal in size over the entire seed surface; stone cells absent. Chromosome number not known.

Figure 4. 

Solanum hexandrum (pubescent individuals) A habit B inflorescence C–E buds showing differences in colour and shape F open flower G long-styled flower with 6 lobes H short-styled flower with 5 lobes I developing fruits enclosed in purple-tinged calyx J mature berries with tightly appressed accrescent calyx (A, B Stehmann et al. 4513; C Giacomin et al. 875; D, G, I Agra et al. 7359; E, F, H Giacomin et al. 1827; J Gouvêa & Salino 514). Photos: A, B João R. Stehmann D, G, I Leandro L. Giacomin E, F, H Lynn Bohs J Yuri F. Gouvêa.

Figure 5. 

Solanum hexandrum (glabrous individuals) A habit at forest edge B inflorescence C–F buds showing variation in fusion of calyx during development G open long-styled flowers H open short-styled flower I open short-styled flowers showing corolla expansion (enlargement) during flowering J mature berry with calyx removed K upper leaf surface with prickles and no stellate trichomes L upper leaf surface with prickles and rayless stellate trichomes with bent mid-points M lower leaf surface with broad-based prickles along the veins (A, B, H Giacomin et al. 1833; C Giacomin et al. 1844; D, I, J Gouvêa & Falcão 137; E, L Gouvêa & Falcão 135; F Gouvêa & Stehmann 158; G, K, M Gouvêa & Stehmann 159). Photos: A, C Lynn Bohs B, H Leandro L. Giacomin D, E, I, J, L Yuri F. Gouvêa F, G, K, M João R. Stehmann.

(Fig. 6). Solanum hexandrum is endemic to the south-eastern region of Brazil and is known from the States of São Paulo, Rio de Janeiro, Minas Gerais, Espirito Santo and Bahia.

Solanum hexandrum grows in the wet forests of the Mata Atlântica, often in openings and along roads and streams; it occurs from almost sea level to 1,600 m elevation.

Brazil. Minas Gerais: juá-bravo (a widely used vernacular name for any spiny solanum in Brazil). No uses have been recorded.

(IUCN 2020). EOO (327,280 km², LC); AOO (460 km², EN). Solanum hexandrum is the most collected of any of these species and all forms of the variation are known from many localities, several of which are within protected areas (e.g. Parque Nacional da Serra dos Orgãos, Parque Nacional do Caparaó, Parque Estadual da Pedra Selada in Rio de Janeiro State; Parque Estadual da Serra do Brigadeiro in Minas Gerais State; RPPN Cafundó, Parque Estadual Mata das Flores in Espírito Santo State; RPPN Serra do Teimoso and RPPN Serra Bonita in Bahia State; Estação Ecológica de Bananal in São Paulo State). Nevertheless, given its high degree of variability that needs further study, we suggest it be assigned a preliminary status of Near Threatened based on criteria (B 2 a, b (ii,iii,iv).

Figure 6. 

Distribution of S. hexandrum.

Solanum hexandrum is the most variable species in the clade in terms of calyx and corolla shape and degree of pubescence, but is otherwise remarkably uniform. More glabrous individuals from the more northerly part of the species range have been called S. echidniforme Dunal, although the type of that species (G00131228) is a sparsely bristly plant and fits within the overall circumscription as we treat this species here. The more glabrous individuals of S. hexandrum (mostly from the State of Espirito Santo) are strikingly different looking morphologically from more pubescent individuals (see Fig. 5), but there is a continuous gradation from glabrous to densely pubescent when specimens from across the range are examined. In several localities, individuals of both types are found and it is not clear if the differences are due to environmental or genetic factors.

The pubescence variability in S. hexandrum is extreme, with glabrous and pubescent individuals at first glance appearing to be completely different morphologically. Some other variation seems to be correlated with lack of rayed stellate trichomes; glabrous plants often have the calyx lobes fused until just before anthesis (see Fig. 5B, D), but this is not completely consistent (see Fig. 5C). The calyx in these glabrous plants is also often more saccate than in populations from elsewhere in the range (Fig. 5J versus Fig. 4J), but this also varies and is difficult to assess on herbarium sheets when all reproductive stages are not present. The single specimen of S. hexandrum which we have seen with a branched inflorescence (Brotto et al. 3265, MBM) comes from amongst these glabrous plants.

The unusual stellate trichomes with the single-celled mid-point bent at an approximately 90° angle to the multiseriate base (see Fig. 5L) are found on the leaves in many populations of S. hexandrum, even if very sparsely. Leaf shape also varies across the range, with some populations from Rio de Janeiro having very narrow, more deeply lobed leaves (e.g. Giacomin 1827). The range of variation in trichome morphology in S. hexandrum warrants further study at a population and genetic level to determine if these variants represent distinct entities. Loss of prickles (derived from stellate trichomes) has been shown to be common across spiny solanums (Satterlee et al. 2024) and it may be that the extremes seen in S. hexandrum could be genetically quite simple.

Solanum hexandrum was described and illustrated in the late 18^th century by Brother José Mariano Conceição da Vellozo (1742–1811) for his "Flora Fluminensis". Vellozo was a parish priest in Rio De Janeiro and completed his work in 1790, but, unfortunately, this work was not published until long after he had died in 1811 (Carauta 1969, 1973). The somewhat telegraphic descriptions of Solanum (Vellozo 1829) referred to original illustrations now held in the library of the Biblioteca Nacional, Rio de Janeiro; printed illustrations, based on these originals, were published several years later (Vellozo 1831). Typification of names of Solanaceae in Flora Fluminensis was treated by Knapp et al. (2015); the original illustration that is the lectotype for S. hexandrum is unambiguous and clearly shows the 6-parted corolla that inspired the specific epithet.

Many herbarium specimens of S. hexandrum are annotated as S. maroniense Poit., a species described only a year after (Poiteau 1830). The description of S. maroniense is of a cultivated plant and is quite telegraphic, but is clearly stated to come from “fleuve Maroni” (the River Maroni) in French Guiana. Although, from the description alone, S. maroniense could correspond to S. hexandrum, the locality suggests it represents a plant of S. crinitum Lam. Solanum crinitum is common in the Guianas and a neotype will be selected as part of upcoming monographic work on the Crinitum clade.

Sendtner (1846) described a variety (“β minax”) citing a manuscript name of Martius, citing several specimens from various collectors, as well as plants from the garden in Munich. A sheet at Munich (M-0171650) exactly matches the very precise locality information and date given in the protologue for a specimen collected by C.F.P. von Martius and we select this as the lectotype; it is the most unambiguous of the syntypes and appears to have been annotated by Sendtner. Another sheet at Munich (M-0171847) has a description and the annotation “Solanum minax” in Martius’ hand is probably also a syntype; this sheet was collected in “Oct” while the lectotype was collected in “Dec” – the protologue states “Octobri ad Decembrem florens”. Other collections cited have no or less unambiguous localities (see Suppl. materials 1, 2). Sendtner (1846) suggested that S. latifolium Poir. (= S. rigidum Lam.) might be a synonym of S. hexandrum; this species is endemic to the Cape Verde Islands and related to the brinjal eggplant (Knapp and Vorontsova 2013).

Dunal (1852: 324) cited Gaudichaud 501 from Rio de Janeiro in “h. Mus. Paris” in his circumscription of S. polytrichum Moric. var. longifolium along with Blanchet 602 from Bahia in “h. DC”. Solanum polytrichum var. longifolium is illegitimate and superfluous as he (Dunal 1852: 324) cited an earlier name at the varietal level, S. polytrichum var. grandifolium Sendtn. (Sendtner 1846), in synonymy. The three specimens of Gaudichaud 501 in P (P00368657, P00368658, P00368659) are of a plant of S. hexandrum and, although P00368657 has been annotated as an “Isosyntype of Solanum polytrichum Moric. var. longifolium Dunal”, this collection has no status as a type. Both collections (Gaudichaud 500, 501) cited by Dunal (1852) may actually have been collected by Frederich Sellow (see Moraes (2023)).

Gaudichaud 500 in “h. Mus. Paris” was the only element cited in the protologue of S. polytrichum var. enoplocalyx (Dunal 1852). Nee (1996) cited as “holotype” Gaudichaud 500 in P and cited two photographs taken by C.V. Morton (Morton neg. 8300, 8301, both held in US). Two sheets of this collection are in P, one bears the original locality label (P00368655), while the other (P00368656) has an undated label in what appears to be Dunal’s hand. Both are scrappy specimens with tiny apical leaves, but P00368655 has two inflorescences, one in flower and one in early fruit. As the more complete specimen, we select the sheet P00368655 of Gaudichaud 500 as the second step lectotype for S. polytrichum var. enoplocalyx. This sheet is labelled as an isotype. Both of these sheets are very similar morphologically to those labelled Gaudichaud 501 and may have been collected together.

Brazil. Minas Gerais: Mun. Teófilo Otoni, afloramento rochoso lado esquerdo da MG 418, cerca de 30 km norte de Teófilo Otoni, 560 m alt., 17°51'22"S, 41°15'39"W, 27 Jan 2014, L.F.A. de Paula, L. Azevedo, R. Fernandes & J.R. Stehmann 669 (holotype: BHCB [BHCB053358] ; isotype: RB [RB01472905]).

Shrubs 1–1.5 m tall, erect, armed. Stems terete, directed upwards and spreading, moderately to densely pubescent and sparsely to moderately prickly, the trichomes eglandular, porrect-stellate, variably short- to long-stalked, the stalks 0.5–1.5 mm long, the rays 4–8, 0.5–1 mm long, the mid-points 1– to 2–celled, always shorter than the rays, the prickles 4–6 mm long, 2–6 mm wide at the base, broad-based and recurved; new growth densely stellate pubescent and sparsely prickly, the trichomes pale yellow to dark brownish-red; bark of older stems glabrescent, drying greenish-brown to dark brown. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, nearly entire to shallowly lobed; blades 2.8–12.1(21.8) cm long, 2.2–7.5(10.1) cm wide, ca. 1.2 to 2 times as long as wide, elliptic to ovate, membranous, slightly discolorous, both surfaces prickly along the mid-rib and veins; adaxial surface densely to moderately stellate-pubescent and prickly, brown to dark green when dry, the trichomes like those of the stem, but with (1–)4–6 rays, the prickles along the mid-rib and major veins to 5.5 mm long and 1 mm wide at the base, straight and laterally compressed; abaxial surface more densely stellate-pubescent than the adaxial surface, whitish-green when dry, the trichomes like those of the adaxial surface, the prickles like those of the adaxial surface, but to 6.5 mm long and 2 mm wide at the base; base attenuate to truncate or rounded, less often with 1 or 2 basiscopic lobes, decurrent on to the petiole, sometimes asymmetrical; margins shallowly lobed, the lobes (0)3–5 on each side, 1–12(14.8) mm long, 3.2–11(23) mm wide at base with usually acute, sometimes rounded or obtuse apices, the sinuses 3.2–8.5 mm deep; apex acute to acuminate; principal veins 4–6 pairs, more prominent beneath, prickly on both surfaces, the prickles 5–6 mm long, straight; petioles 0.6–3.3 cm long, densely to moderately pubescent with porrect-stellate trichomes like those of the leaves, usually armed with 1–5 prickles. Inflorescence internodal, to 6 cm long, unbranched, with 4–10 flowers, up to 2 flowers open at a time; axes glabrescent to densely pubescent, usually unarmed, the stellate trichomes like those of the stem, but these sometimes with the mid-point as long as the rays; peduncles 0.4–2.3 (-3.8) cm long; pedicels 3–17 mm long, 0.5–0.8 mm in diameter at the base, to 1.5 mm in diameter at the apex, spreading to slightly deflexed, pubescent with trichomes like those of the inflorescence axes, unarmed, articulated at the base; pedicel scars evenly spaced 1–7 mm apart. Buds ovoid to ellipsoid, with the corolla enclosed in the calyx until just before anthesis. Flowers 5-merous, heterostylous with long-styled flowers (co-sexual) at the base of inflorescence, short-styled (functionally staminate) flowers more distally, the plants andromonoecious. Calyx with the tube 2.6–4.3(6) mm long, 6.5–8 mm in diameter, broadly obconical to cupuliform, the lobes 3–7 mm long, 3–5.5 mm wide, triangular to deltate, with acute to acuminate apices, glabrous adaxially, densely pubescent abaxially with bristly purple-tinged, hyaline or reddish-brown porrect to multangulate long-stalked stellate trichomes, the stalks 1.1–3.8 mm long, rays 4–8, to 1.5 mm long, the mid-points 1–2 celled, shorter than or the same length as the rays, armed or unarmed, if present, the prickles 2.8–4 mm long, 0.5–1 mm in diameter at the base, straight, acicular. Corolla 2.4–3 cm in diameter, white, often with a greenish-yellow star at the base, shallowly stellate, lobed ca. halfway to the base, interpetalar tissue nearly absent, the lobes 5.9–8.8 mm long, 9.9–12.2 mm wide, pubescent abaxially on the petal mid-vein and/or apices with sparse delicate short-stalked porrect-stellate trichomes with stalks to 0.9 mm long, the apices acute to apiculate. Stamens equal; filament tube to 1 mm long; free portion of the filaments 0.7–1 mm long, glabrous; anthers 6.5–8 mm long, 2.5–3 mm wide, broadly lanceolate and tapering, connivent or slightly divergent at the tips, glabrous, yellow, abaxially swollen in the lower half (gibbous) and somewhat papillate, poricidal at the tips, the pores directed distally, slightly extrorse, not lengthening to slits with age. Ovary somewhat conical, glabrous; style 8–10 mm long in long-styled flowers, ca. 3 mm long in short-styled flowers, straight, glabrous; stigma clavate to bilobed, the surface papillose and irregular, the style and stigma poorly developed in short-styled flowers. Fruit a globose berry, 0.9–1.8 cm in diameter, green to whitish-green at maturity, drying dark brown, glabrous, the pericarp matte; fruiting pedicels 1–1.5 cm long, 1–2 mm in diameter at the base, usually unarmed, deflexed from the weight of the fruit; fruiting calyx partially accrescent, the tube tightly investing 1/2–3/4 of the fruit at maturity, the lobes 5.8–8 mm long, 7–9.6 mm wide, not overlapping, pubescent with long-stalked porrect-stellate trichomes often with the base of the stalks markedly expanded and bristly, the stalks to 4.8 mm long. Seeds ca. 250 per berry, 2.2–2.6 mm long, 1.6–2 mm wide, pyriform to reniform, not markedly flattened, the surface irregularly pitted, the testal cells pentagonal in outline; stone cells absent. Chromosome number not known.

Figure 7. 

Solanum hydroides A habit B flowering branch C inflorescence D open long-styled flower E mature berry with only partially accrescent calyx F cross-section of 4-locular mature berry showing fleshy mesocarp and brown seeds (A, C–F Gouvêa et al. 492; B Gouvêa & Santos 325). Photos: Yuri F. Gouvêa.

(Fig. 8). Solanum hydroides is endemic to the south-eastern region of Brazil, with records in four localities in north-eastern Minas Gerais (Mun. Teófilo Otoni and Conselheiro Pena) and northern (Mun. Nova Venécia) and central (Mun. Santa Teresa) Espírito Santo States.

Solanum hydroides grows at the edge of seasonal semi-deciduous tropical rainforests associated with granitic or gneissic rock outcrops (inselbergs) and somewhat disturbed sites at their bases, such as roadsides and clearings; from 300 to 600 m elevation. It also occasionally grows in epilithic vegetation patches lying on the flatter parts of inselbergs.

None recorded.

Figure 8. 

Distribution of S. hydroides.

(IUCN 2020). EOO (12,549 km², VU); AOO (24 km², EN). Solanum hydroides is known from only four disjunct localities in vegetation remnants associated with inselbergs: these rock outcrops harbour the last remnants of forest fragments (Martinelli 2007) in areas where they occur. Only one of these localities is within a protected area (APA Pedra do Elefante, Espiríto Santo State). Although we have seen a few more collections than were used in the original assessment (Gouvêa et al. 2020), we concur with their evaluation of S. hydroides as Endangered (B 2 a,b ii,iii,iv).

Solanum hydroides is a comparatively smaller plant than other species of the group, except S. sublentum; its smaller leaves and thinner stems, petioles and inflorescence axes give it a more delicate overall aspect. Solanum hydroides can, however, be readily distinguished from S. sublentum by its pubescence of stellate eglandular trichomes (Fig. 7A, B) and by the widely obconical to cupuliform shape of the calyx at anthesis. In S. sublentum, the indumentum is of both conspicuous simple glandular trichomes and stellate eglandular trichomes (Fig. 13E–G), with the stellate trichomes usually much less numerous than the simple ones and often early deciduous (i.e. present only in new growth). Calyces of S. sublentum are somewhat urceolate, inflated and prominently plicate at the top of the calyx tube (Fig. 13J), whereas, in S. hydroides, calyces are tightly adherent to the berry at maturity and not notably plicate, especially in live plants.

Although being a markedly less robust plant, S. hydroides can be very similar to some specimens of S. hexandrum, the most variable species in the clade, with which it shares the indumentum of few-rayed stellate eglandular trichomes on the stems, leaves, inflorescence axis and calyces. Solanum hydroides differs from S. hexandrum in its white and smaller corollas (13–21.5 mm total length), shorter corolla lobes (5.9–8.8 mm long; Fig. 7D) and accrescent, but not inflated, fruiting calyces that only partially cover the mature fruit (Fig. 7E). Solanum hexandrum has corollas in various shades of lilac to purple and are larger (24.3–40 mm long), with longer corolla lobes (12.6–25 mm long; Figs 4G, H, 5G, H, I) and the fruiting calyces are accrescent and inflated, completely enclosing the mature fruit (Figs 4J, 5J). The corollas of S. hydroides are thin and membranous and easily tear apart between the lobes during the drying process, which can make the lobes on herbarium specimens seem longer than they really are. Care is needed to ensure correct measurements from herbarium specimens.

Leaf measurements are also useful for distinguishing S. hydroides from S. hexandrum. The leaves of S. hydroides are usually smaller (7.5–13.6 cm long and 5–8.7 cm wide) than those of S. hexandrum (17–45 cm long and 10.5–32 cm wide). Nevertheless, leaves of S. hydroides are larger in plants growing in shade and in young individuals (see Roe (1966) for other examples in Solanum) and we have seen plants with leaves to 22 cm long and 11 cm wide. Specimens of S. hydroides growing in shade are less densely pubescent, with less robust (i.e. stalks with fewer series of cells) and slightly shorter trichomes on stems, leaves and calyx. Corollas of these shade plants are usually larger in relation to the other flower parts (e.g. stamens and calyx).

Trichome morphology in S. hydroides is not particularly variable within individual plants and amongst plants of the same population; however, there is a significant variation in the number of trichome rays between some populations, as is seen also in S. hexandrum. Trichomes of specimens from the southernmost-known population (in Santa Teresa Municipality, Espirito Santo State) are mostly 6- to 8-rayed and usually denser, whereas those of plants from the other populations (Teófilo Otoni and Nova Venécia Municipalities of Minas Gerais State) are mostly 4-rayed. Within individual plants, the variation in trichome morphology is limited to a reduction in the number of rays and is especially evident in plants with four-rayed trichomes. In these plants, the trichomes may lack one to almost all rays, sometimes with only the mid-point or a lateral ray remaining and the trichome appearing to be unbranched, but with a basal multiseriate stalk, as is also seen more dramatically in S. hexandrum. This kind of variation has been reported in other Solanum groups, such as the Brevantherum clade or members of the Acanthophora clade (Nee 1991; Levin et al. 2005; Stern et al. 2013).

Solanum phrixothrix differs from all other members of the S. hexandrum group in its rotate, white corollas. It is similar to S. aciculare and S. sublentum in its cordate non-decurrent leaf bases, but differs from both in its eglandular pubescence. The eglandular bristle-like trichomes completely lacking lateral rays distinguish it from S. aciculare and its densely bristly stems distinguish it from S. sublentum. It differs from S. hexandrum, S. hydroides and S. stagnale in its cordate non-decurrent leaf bases and from S. stagnale, it is acicular, rather than broad-based and usually curved prickles.

Figure 9. 

Solanum phrixothrix A flowering branch B section of stem with copious bristles C inflorescence with two open flowers D adaxial surface of calyx E open short-styled flower F short-styled flower with corolla bent back to expose the tapering anthers (A–F Folli 7560, BHCB). Photos: Yuri F. Gouvêa.

Brazil. Espirito Santo: Mun. Linhares, bairro Bebedouro, mata de tabuleiro, estrada ES-245, sentido a Regência, mata de cabruca (Cacau), UTM: 394627, 7851005 [19°25'57.1"S, 40°00'13.5"W], 12 Jun 2017, D.A. Folli 7560 (holotype: BHCB [BHCB221244]; isotype: CVRD [acc. # 15743]).

Shrubs to ca. 2 m tall, erect to somewhat spreading, strongly armed and bristly. Stems terete, conspicuously fistulose (at least the younger ones), densely bristly and prickly; the bristle-like trichomes simple, 3.4–19.2 mm long, with a long multiseriate stalk and a shorter uniseriate mid-point at the tip, the stalks 2.5–16 mm long, the mid-points 0.9–3.2 mm long, 1–2-celled, underlying pubescence of sparse, very tiny papillate trichomes, these drying golden yellow; the prickles 0.5–1.2 cm long, the base ca. 1 mm in diameter, straight to retrorse, somewhat laterally compressed, yellowish-golden at base, becoming yellowish-brown towards the tip; the epidermis densely dotted with crystal sand (inclusions of calcium oxalate); new growth densely bristly and prickly, with simple bristle-like trichomes and prickles like those of the stems; bark of older stems not known. Sympodial units plurifoliate, the leaves not geminate. Leaves lobed, the blades 21–30 cm long, 20–24 cm wide, ca. 1–1.25 times as long as wide, broadly ovate to broadly elliptic, usually widest in the basal half, membranous, concolorous, sparsely to moderately prickly on the mid-rib and major veins of both surfaces with prickles like those of the stems, but usually smaller; adaxial surface densely to moderately pubescent to hirsute with simple bristle-like trichomes similar to those of the stems, but smaller, 0.8–6.5 mm long, the stalks 0.2–4.5 mm long, the mid-points 0.7–2 mm long; abaxial surfaces more sparsely pubescent with trichomes like those of the adaxial surface, these restricted to the mid-rib, major and minor veins and usually smaller and thinner-walled; principal veins 5–7 pairs; base cordate to angular-cordate often with a prominent pair of basiscopic lobes, not decurrent on to the petiole, symmetrical; margins 4–6-lobed, leaves of young plants or new growth also with secondary lobing, the sinuses 1/3–1/5 of the distance to the mid-rib; apex acute to obtuse; petiole 4.8–9.5 cm long, densely bristly, moderately to densely prickly with trichomes and prickles like those of the stems. Inflorescences opposite the leaves or internodal, 12–15 cm long, unbranched, with 19–25 flowers; axes densely to moderately bristly and prickly with trichomes and prickles like those of the stems, but the prickles sometimes thinner; peduncle 2.7–3.6 cm long; pedicels 1–2 cm long, 0.6–0.9 mm in diameter at the base, same diameter at the apex (excluding trichomes), erect to directed downwards, moderately to densely bristly with bristle-like trichomes like those of the stems, prickly or not, articulated at the base; pedicel scars more or less evenly spaced 1–5 mm apart on mature inflorescences, more closely packed distally. Buds ellipsoid to narrowly ellipsoid, the calyx lobes soon splitting, exceeding the length of the corolla until just before anthesis. Flowers 5-merous, heterostylous, the proximal flowers long-styled (co-sexual) and distal ones short-styled (functionally staminate), the plants andromonoecious. Calyx with the tube 4.3–7.5 mm long, 4–8 mm in diameter, shallowly cup-shaped to obconical, the lobes 4.5–11.3 mm long, triangular to somewhat lanceolate, sometimes varying in size in a single flower, densely bristly with trichomes like those of the stems, prickly or not. Corolla 3–3.6 cm in diameter, white, rotate, shallowly campanulate or tubular, lobed less than 1/8 of the way to the base, interpetalar tissue indistinguishable from the rest of the lobe or nearly so, copious and reaching nearly to the tips, the lobes 3.8–5 mm long, 16.5–25 mm wide, rounded to retuse, glabrous with pubescence restricted to the lobe apices on both surfaces, the trichomes minute, simple. Stamens equal; filament tube 1.5–1.7 mm long, glabrous; free portion of the filaments 1.4–1.8 mm long, glabrous; anthers 6.5–10 mm long, 2.2–3 mm wide, broadly lanceolate and tapering, connivent, glabrous, yellow, abaxially swollen in the lower half (gibbous) and somewhat papillate, poricidal at the tips, the pores directed distally, not elongating to slits with age. Ovary cup-shaped, moderately to densely pubescent, the trichomes simple, glandular, very tiny papillate and sessile to 0.6 mm long, 1–5-celled, thin-walled, the longer ones less abundant, pubescence sometimes restricted to the ovary apex; style 8.8–12 mm long in long-styled flowers, 1.2–3.6 mm long in short-styled flowers, straight, sparsely to moderately puberulent with very tiny papillate trichomes; stigma clavate, the surface minutely papillose. Fruit and seeds not known. Chromosome number not known.

(Fig. 10). Solanum phrixothrix is endemic to the south-eastern region of Brazil and known only from two collections; one with a specific locality from Espírito Santo State and another made by A. St.-Hilaire that is likely to be from Minas Gerais State (see Dwyer (1955)).

The only known collection with locality and vegetation type information of S. phrixothrix is from wet evergreen forests of the lower Rio Doce drainage, at approximately 13 m elevation. This collection is from a “cabruca” (cacao plantation). These cacao plantations retain the upper strata of the forest for shade, but the understorey is significantly damaged by shrub and herb removal, along with significant cacao leaf litter.

None recorded.

The species epithet is derived from the Greek, meaning with bristling (or horrid) hairs.

(IUCN 2020). Solanum phrixothrix is known from only two collections, gathered 200 years apart and so must have a preliminary assessment of Data Deficient. That said, however, it is imperative that more populations be sought to better assess its range and population density. It is likely to be of conservation concern, as the single collection with an accurate locality (Folli 7506) is from a highly disturbed anthropogenic site (cacao plantation), not within a protected area. It is near the Floresta Nacional de Goytacazes in Espiríto Santo State and should be sought there. Other protected areas close to this collection, such as Reserva Biológica de Sooretama and Reserva Natural Vale are well-inventoried and we have seen no specimens of S. phrixothrix from them in any of the many herbaria we consulted.

Solanum phrixothrix is a distinctive, densely bristly plant that has only been collected twice, once by Auguste St. Hilaire in the early 19^th century and more recently in 2017 (Folli 7506) along the Rio Doce in Espiríto Santo State. It differs from other taxa in the group in its rotate to rotate-pentagonal corollas (Fig. 9E, F) and densely bristly stems with no stellate trichomes present (Fig. 9B). The long inflorescences and somewhat delicate pedicels are similar to those of S. aciculare, but that species has copious glandular stellate to multangulate pubescence and stellate corollas (Fig. 2F, G, I). It is surprising that this species has escaped notice for so long, but members of the S. hexandrum group often occur in very small populations at the bases of rocky outcrops and may easily overlooked despite their large size and fearsome appearance.

Figure 10. 

Distribution of S. phrixothrix.

Solanum phrixothrix shares dense bristly stems with S. aciculare but can be easily distinguished from that species by its lack of copiously glandular pubescence of long-stalked stellate trichomes. Solanum phrixothrix appears to lack long-stalked stellate trichomes on any part. White flowers are also found in S. hydroides (Fig. 7D) and S. sublentum (Fig. 13H), as well as occasionally in S. aciculare (Fig. 2F). Corollas in these three species are always stellate with variously deltate to triangular lobes, whereas those of S. phrixothrix are rotate and somewhat campanulate with the lobes reduced to tiny apiculae.

Brazil. Bahia: sin. loc., J.S. Blanchet 2085 (lectotype, designated by Whalen et al. (1981), pg. 69 [as holotype]: G [G00343733]; isolectotypes: BM [BM000617832], G-DC [G00131205], P [P00371689, P00578808, P00371690]).

Large, soft-wooded perennials 0.3 - several m tall. Stems terete, usually somewhat winged from the decurrent leaf bases, densely pubescent and sparsely prickly, the pubescence of whitish or reddish-cream porrect-stellate or occasionally multangulate (Agra 617) trichomes with multiseriate stalks 0.2–0.8 (-2.5) mm long, the lateral rays 4–9, 0.4–0.6 (-1) mm long, the mid-points much shorter than the rays, 0.05–0.2 mm long, the hairs densely interwoven and entirely concealing young stems, the prickles 1–2 cm long, 0.3–0.5 cm wide at the base, straight or recurved (e.g. Blanchet 2085) broad-based and strongly laterally compressed, often densely stellate-pubescent basally. Sympodial units difoliate, the leaves of a pair not geminate. Leaves shallowly lobed and coarsely repand; blades 9.5–30 cm long, 8–15 cm wide, ca. 1–1.5 times as long as wide, broadly elliptic to ovate, slightly discolorous, membranous or somewhat chartaceous, prickly on both surfaces along the veins with straight prickles to 15 mm long; adaxial surfaces densely pubescent with eglandular short- to long-stalked porrect stellate trichomes, the stalks 0.5–0.8 (-1) mm long, the rays 4–8, 0.5–0.7 mm long, the mid-points minute or equal in length to the rays, the lamina visible under the microscope; abaxial surfaces densely woolly-pubescent with stalked porrect stellate trichomes, the stalks 0.5–1 mm long, the rays 6–10, 0.5–1 mm long, often not in a single plane, the mid-points 0.2–0.4 mm long, much shorter than the rays; principal veins 4–5 pairs, with scattered straight prickles to 1.3 mm long, the prickles longer and larger on the mid-rib; base strongly decurrent along a winged petiole, the wing of laminar tissue to 0.5 cm wide on each side, often decurrent on to stem; margin lobed, the lobes 4–5, 1.5–3 cm long, 2–3 cm wide, deltate, acute- or round-tipped, often with irregular secondary lobing, the sinuses reaching less than halfway to the mid-rib ; apex acute or obtuse; petioles 0.8–5 cm, usually 1/4–1/3 the length of the blades and winged, stellate-pubescent like the stems, prickly. Inflorescences 2–7 cm long, extra-axillary or leaf-opposed, unbranched, with ca. 10 flowers, the axes densely stellate-pubescent, unarmed; peduncle 1–2 cm; pedicels 2–5 mm, 1–2.5 mm in diameter at the base, 2–2.6 mm diameter at the apex, articulated at the base; pedicel scars closely spaced 2–7 mm apart. Flowers 5-merous, heterostylous, with the lowermost long-styled (co-sexual) and the distalmost short-styled (functionally staminate), the plants probably andromonecious. Calyx with the tube 5–10 mm long, 8–10 mm in diameter, broadly obconical, the lobes 10–15 mm long, 5–8 mm wide, ovate-lanceolate or somewhat spathulate or tongue-shaped, obtuse or round apically, densely stellate-pubescent on both surfaces, often with scattered prickles on both surfaces near the mid-vein, often purple-tinged distally. Corolla 2.5–4.5 cm in diameter, white or lilac with a paler central star, stellate, lobed 2/3 to 3/4 of the way to the base, interpetalar tissue a thin edge on the lobes, the lobes 13- 20 mm long, 10–12 mm wide, ovate-lanceolate, densely stellate-pubescent abaxially, the trichomes with robust mid-points equal to or longer than the rays, glabrous adaxially, but the acute tips stellate-pubescent, the interpetalar tissue thin, glabrous. Stamens equal; filament tube minute; free portion of the filaments ca. 1 mm long, glabrous; anthers 7–10 mm long, ca. 3 mm wide, broadly lanceolate and tapering, connivent, glabrous, yellow, abaxially swollen in the lower half (gibbous) and somewhat papillate, poricidal at the tips, the pores directed distally, slightly extrorse, not elongating to slits with age. Ovary conical, densely stellate-pubescent, the trichomes with well-developed lateral rays; style 10–14 mm long, glabrous or sparsely stellate-pubescent in the lower half; stigma large and capitate. Fruit a globose to flattened-globose berry, 2–2.5 cm in diameter, whitish-green at maturity, sparsely stellate-pubescent, ultimately glabrous, the pericarp matte or slightly shiny; fruiting pedicels 0.5–1 cm long, usually less than 0.5 cm long, 3–5 mm in diameter at the base, ca. 6 mm in diameter at the apex; fruiting calyx only partially accrescent, tightly investing. but not completely covering fruit, the tube ca. 1.5 cm long, the lobes ca. 15–20 mm long, ca. 10 mm wide, not overlapping. Seeds ca. 100 per berry, ca. 2.5 mm long, ca. 1.5 mm wide, flattened reniform, dark brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Chromosome number; 2n = 24 (Bernardello et al. 1994; voucher (grown in Indiana and, therefore, should be in IND, but not seen) Carvahlo 3213, possibly a misprint for Carvalho 3219).

Figure 11. 

Solanum stagnale A habit of a young plant B habit of an older reproductive plant B, C inflorescence congested with flowers clustered due to extremely short pedicels D flower buds and short-styled flower (arrows point to spathulate calyx lobes) E long-styled flower and leaf base markedly decurrent on to the petiole F berries (almost mature) with appressed, only partially accrescent fruiting calyces (A, D Giacomin & Stehmann 1930, BHCB; B, C, E, F unvouchered field photograph, Bahia State [12°32'23"S, 38°03'08"W]). Photos: A, D Leandro L. Giacomin B, C, E, F Wagner Nogueira.

(Fig. 12). Solanum stagnale is endemic to eastern Brazil; it has been recorded from the States of Bahia and adjacent northern Minas Gerais and disjunctly in Paraíba State. This disjunct distribution is unusual, the single collection from Paraíba is from the northern side of the São Franscisco River, the site of the Pernambuco area of endemism.

Solanum stagnale occurs in sandy coastal vegetation (restinga) habitat, in sand dunes, forests, forest edges and somewhat open habitats, from sea level to 300 m elevation.

None recorded.

(IUCN 2020). EOO (157,059 km², LC); AOO (68 km², EN). Solanum stagnale is known from more than five localities, even if the widely disjunct collection from Paraíba State is not included. Only one of these is within a protected area (Estação Ecológica de Cotegipe in Mun. Salvador, Bahia). The fragmented nature of the habitat and the absence of state or national level of protection for areas where it occurs suggests it should be assigned a preliminary conservation status of Vulnerable, based on criteria B 2 a,b i,ii,iii,iv.

Like all members of this group, S. stagnale has large, repand leaves. Most collections have strongly winged petioles with a wing extending fully to the base, but occasionally the wing becomes very narrow basally (Rosas 1 from Salvador). The pedicels in both flower and fruit of S. stagnale are the shortest in the group, rarely reaching 5 mm long. Solanum stagnale is easily distinguished from S. hexandrum, with which it is most similar, by its pubescence of porrect-stellate trichome with usually more than 5 lateral rays, usually curved prickles, short, stubby pedicels usually less than 0.5 cm long, spathulate calyx lobes with rounded apices and berry that is not completely enclosed in an accrescent calyx. The trichomes of S. stagnale usually have mid-points that are shorter than or equal to the rays in length; in contrast, other species of the group have longer mid-points.

Figure 12. 

Distribution of S. stagnale.

Whalen et al. (1981) treated S. stagnale as a member, albeit anomalous, of section Lasiocarpa (Dunal) D’Arcy and Whalen (1984) later included it in his equivalent S. quitoense Lam. species group. Both groups were composed of species, with the exception of S. stagnale, which are now recognised (Gagnon et al. 2022) as the Lasiocarpa clade; they share with members of the S. hexandrum group large repand leaves. Molecular data (Gouvêa 2020; Gagnon et al. 2022), however, clearly show that S. stagnale is related to S. hexandrum and other Brazilian endemic species of this group, not to the largely Andean members of the Lasiocarpa clade.

Dunal (1852) changed the name S. stagnale to S. moricandii because he felt it was inappropriate (“Blanchet in schedis non dicit hanc specimen crescere in stagnis, ut putat Moricand, et hâc ratione, nomen stagnale mutavi” – Blanchet does not say this specimen grows in ponds, as Moricand thinks, for this reason I have changed the name stagnale”: Dunal 1852: 319) rendering the name S. moricandii illegitimate and superfluous.

Brazil. Minas Gerais: Lagoa Santa, [24 Feb 1863], E. Warming s.n. (lectotype, designated here: C [C10019316]; possible isolectotype: [note date on sheet is 1866] S [acc. # S04-2985]).

Shrubs 1–3 m, erect or sometimes somewhat prostrate, armed. Stems terete, densely glandular-pubescent and sparsely prickly, the trichomes weak, simple uniseriate ca. 0.5 mm long with glandular tips, mixed with sparse short-stalked porrect stellate trichomes with 6–7 rays to 1 mm long, the mid-point to 1 mm long, gland-tipped or eglandular, the prickles 0.5–1.2 cm long, slightly to strongly curved and broad-based, ca. 0.5–1 cm in diameter at the base; new growth densely glandular pubescent with mixed simple and stellate trichomes like the stems; bark of older stems pale greyish-brown, somewhat glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves shallowly lobed and repand, much smaller in younger branches; blades (5)8.5–17 cm long, (3)6–13 cm wide, ca. 1.3–1.6 times as long as wide, broadly elliptic or ovate, widest at or just below the middle, membranous, concolorous, sparsely prickly on both surfaces along the veins with straight prickles 0.3–1 cm long; adaxial surface densely pubescent with a mix of short-stalked porrect stellate trichomes with 5–7 rays ca. 1 mm long, trichomes consisting of solely unicellular or multicellular gland-tipped mid-points to 1.2 mm long (probably derived from stellate trichomes) and sessile, papillate glands composed of 4 cells; abaxial surface pubescent like the adaxial surface, but lacking the sessile papillate glands, also with delicate sessile porrect stellate trichomes with 4–5 rays ca. 0.3 mm long and mid-points shorter than the rays, these underneath the dense layer of larger short-stalked trichomes; principal veins 4–5 pairs, usually sparsely prickly on both surfaces with straight prickles to 0.9 cm long; base somewhat cordate-angular to hastate or sagittate-hastate from the basiscopically directed lowest leaf lobes, occasionally acute to abruptly attenuate, usually not decurrent on to the petiole; margins shallowly and broadly lobed, the lobes 4–5, 1–2.5 long, 2–4 cm wide, apically acute to acuminate, sometimes minutely secondarily lobed, the sinuses less than 1/4 of the way to the mid-rib; apex acute to acuminate; petiole (1-) 1.5–7 cm long, prickly with straight prickles to 1 cm long, densely glandular pubescent like the stems with a mix of simple trichomes apparently consisting of unicellular or multicellular mid-points with glandular tips and sparse porrect stellate trichomes. Inflorescences internodal, 2–9 cm long, unbranched, with 3–6 flowers. but only one open at a time; axes densely glandular pubescent like the stems with a mix of unicellular and multicellular gland-tipped simple uniseriate trichomes (derived from mid-points of stellate trichomes) and sparse porrect stellate trichomes with glandular mid-points; peduncle 1.5–5 cm long; pedicels 1.2–1.5 cm long, ca. 1 mm in diameter at the base, ca. 2.5 mm in diameter at the apex (excluding trichomes), erect to spreading, densely glandular pubescent like the inflorescence axes and stems and occasionally with a few straight prickles, articulated at the base; pedicel scars more or less evenly spaced 4–5 mm apart, further apart in fruit, distally and, in young inflorescences, more tightly packed. Buds globose to ovoid, the corolla ca. halfway exerted from the calyx tube just before anthesis. Flowers 5-merous, co-sexual or perhaps a few distal flowers short-styled and functionally staminate, the plants only weakly andromonoecious. Calyx with the tube 3.5–4 mm long, 5–6 mm in diameter, deeply to shallowly broadly cup-shaped, plicate from the fused bases of adjacent lobes, usually invaginate at the base, the lobes 7–10 mm long, 2.3–5 mm wide, long-triangular, apically acuminate, densely glandular pubescent with a mix of simple uniseriate (mid-points?) and sparse short-stalked or sessile stellate trichomes, often with a few straight prickles 0.2–1.5 mm long on the main veins abaxially. Corolla 3.5–5 cm in diameter, purple to pale violet or white, shallowly stellate, lobed ca. 1/4 of the way to the base, interpetalar tissue thin, glabrous, the lobes 9–12 mm long, 11–19 mm wide, spreading to slightly cupped, densely pubescent abaxially where exposed in bud with short-stalked and sessile stellate trichomes, these occasionally glandular, glabrous adaxially, but occasionally with a few minute prickles along the veins. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, glabrous; anthers 8.5–9 mm long, 2.6–3 mm wide, broadly lanceolate and tapering, connivent, glabrous, yellow, abaxially swollen in the lower half (gibbous) and somewhat papillate, poricidal at the tips, the pores directed distally, not elongating to slits with age. Ovary conical, glabrous; style 14–16 mm long, glabrous, widening markedly distally; stigma clavate or broadly capitate, the surface minutely papillate. Fruit a globose berry, 1.4–2 cm in diameter, green or pale whitish-green, glabrous, the pericarp matte when dry, opaque, the berry completely enclosed in the accrescent saccate calyx; fruiting pedicel 1.6–2.5 cm long, 1–1.5 mm in diameter at the base, 3–5 mm in diameter at the apex, spreading or pendent from the weight of the fruit; fruiting calyx strongly accrescent, inflated and invaginate, exceeding the length of the berry, but not completely enclosing it, the tube 1.5–2 cm long, saccate (invaginate) at the base, the lobes 0.8–0.9 cm long, often broken in dried specimens, not overlapping, densely glandular pubescent and occasionally prickly like the calyx in flower. Seeds 80–100 per berry, ca. 2.5 mm long, ca. 2 mm wide, flattened reniform, reddish-brown when dry, the surface minutely pitted, the testal cells thick-walled and sinuate in outline. Chromosome number not known.

Figure 13. 

Solanum sublentum A habitat of arboreal restinga (coastal scrub-forest transition) forest edges B habit C habitat in seasonally dry deciduous forests D habit in rocky inselberg E stem with recurved prickles and unbranched glandular pubescence F leaf with cordate base and secondary lobing G variation in leaf bases and lobing H inflorescence I long-styled flowers showing colour polymorphism J mature berry with accrescent invaginate calyx (B, F, H–J Gouvêa & Guerrero 452; D Stehmann et al. 6370; E, G Stehmann et al. 6372). Photos: A, B, H–J Yuri F. Gouvêa C, D, E, G João R. Stehmann.

(Fig. 14). Solanum sublentum is recorded to south-eastern and central Brazil, in the States of Espírito Santo, Goiás, Minas Gerais, Rio de Janeiro and São Paulo. The collection from Goiás is discontinuous from the rest of the species range (see discussion).

Solanum sublentum occupies primarily forest understory, edges and clearings of wet coastal and semi-deciduous forests in the Atlantic Forest domain (Mata Atlântica; Fig. 13A, B), as well as semi-deciduous and deciduous seasonally dry tropical forests (Fig. 13C, D) in ecotonal zones or within savannah matrices in the Cerrado domain. The forests where S. sublentum occurs can be associated with granite/gneiss (wet and semi-deciduous forests in coastal and sub-coastal regions) and limestone or basaltic (inland seasonally dry tropical forests in savannah matrices) outcrops, it is the only species in the group and one of the few American species of Solanum found in STDFs associated with limestone outcrops. Solanum sublentum grows both in deep soils and in rock cavities, in fissures and in small, shallow soil patches that accumulate on bare rocks (Fig. 13D); from sea level to 800 m elevation.

Figure 14. 

Distribution of S. sublentum.

None recorded.

(IUCN 2020). EOO (642,872 km², LC); AOO (72 km², EN). Solanum sublentum is known from eight localities (including the extremely disjunct occurrence in the State of Goiás), at least three of which are in protected areas (Parque Nacional Cavernas do Peruaçu, Estação Biológica de Caratinga in Minas Gerais State; Parque Municipal Sombra da Tarde in Espirito Santo State) with other collections from the mountains near Rio de Janeiro in what is now Parque Nacional da Serra dos Orgãos (e.g. “Organ Mount”). Even with the disjunct occurrence in Goiás excluded, S. sublentum has the largest extent of occurrence of any of the species of the S. hexandrum group. Nevertheless, due to the extreme level of habitat alteration in the region where it occurs and the paucity of recent collections (most of those used in calculating the EOO are more than a century old), we consider it of some conservation concern and suggest a preliminary status of Near Threatened, based on criteria B 2 a,b ii,iii, iv.

Solanum sublentum is easily distinguishable from other members of this group in its markedly plicate accrescent calyces that exceed, but do not completely enclose the berries and simple, glandular pubescence with a mix of much less abundant stellate trichomes.

The majority of trichomes of S. sublentum are simple, uniseriate and unicellular or multicellular. Their shape and overall morphology suggest they are structurally analogous to mid-points of stellate trichomes without rays, like those that occur in the Acanthophora clade (Nee 1979; Hilgenhof et al. 2023) and most members of the Gonatotrichum group of the Brevantherum clade (Stern et al. 2013).

The single collection of S. sublentum from Goiás (Hatschbach 34747) is from Mun. Jataí, in the extreme southwest of the State. The habitat is stated as “clareiras da mata” (forest clearings) and the area in which it was collected was a remnant of the now very restricted Atlantic Forest. The southernmost portion of Goiás State is the original limit of the Atlantic Rainforest domain in central Brazil and an ecotonal zone with the Cerrado domain, where contiguous forested formations were found in the past (IBGE 2012). These mostly semi-deciduous forests were either considered part of the Cerrado (“Cerradão”) or the Atlantic Rainforest itself. They have been largely converted first to cattle farming and later to soy and corn plantations (Oliveira 2007). No recent collections from this locality have been seen, so recollection there is a priority.

Solanum sublentum is similar to S. aciculare in possessing glandular indument and strongly accrescent, inflated and invaginate fruiting calyx lobes. The pubescence of S. sublentum is of unbranched trichomes (the bases and mid-points of modified stellate trichomes), while that of S. aciculare is of long-stalked stellate trichomes with glands on each ray tip. Prickles in S. sublentum are sparse and recurved, whereas those of S. aciculare are straight and denser especially on stems. The calyx completely covers the berry in S. aciculare (Fig. 2H) while, in S. sublentum, the berry is clearly visible (Fig. 13J).

Many different collecting dates are written on the Warming collections at C used by Hiern (1878) to describe S. sublentum. We have chosen the only sheet with both flowers and fruits (C10010316), showing the characteristic saccate calyx in fruit of S. sublentum. The many syntype collections in Copenhagen and sent elsewhere by Hiern have a confusing panoply of dates and some may be isolectotypes, but, in the absence of unambiguous dates on these sheets, we do not recognise them as such. The sheet we here cite as a possible isolectotype at S (acc. # S04-2985) has a label stating 1866, so it too is likely not a duplicate, but it is the only original material we have seen outside of Copenhagen.