Herbs, vines, scandent or prostrate shrubs, shrubs or small trees; armed or apparently (to almost completely) unarmed; young stems stellate-pubescent, the trichomes multangulate or more often porrect-stellate with the rays all in a single place, occasionally lepidote, sometimes glabrescent; prickles absent, sparse or dense, straight or curved; bark of older stems grey or brown, sometimes glabrescent. Sympodial units plurifoliate or difoliate, the leaves geminate or not. Leaves entire to deeply pinnately lobed, usually stellate-pubescent on both surfaces, sometimes glabrescent or the trichomes very sparse; pubescence of porrect, multangulate or lepidote stellate trichomes, these sessile or variously stalked, with or without midpoints, the midpoints if present very short and bump-like to elongate. Inflorescences lateral and internodal or opposite the leaves, unbranched to many times branched, usually pedunculate, not bracteate, variously stellate-pubescent; petioles articulated near the base. Flowers actinomorphic or zygomorphic, bisexual (hermaphroditic) or unisexual and the plants andromonoecious; calyx 5-parted (rarely 4-parted or more than 5-parted in cultivated forms of S. melongena), usually stellate-pubescent, sometimes glabrescent; corolla 5-parted (rarely 4-parted or more than 5-parted in cultivated forms of S. melongena), rotate-stellate to deeply stellate, white, lilac or deep purple, some species polymorphic for flower colour, interpetalar tissue absent to copious; stamens 5 (rarely 4 or more than 5 in cultivated forms of S. melongena), the filaments equal or unequal, usually glabrous; anthers equal or unequal, yellow or cream, blunt to strongly tapered, tightly connivent or spreading, dehiscing by terminal pores, the pores usually not lengthening to slits with age; ovary bicarpellate (fasciated in cultivars of S. melongena), globose to conical, glabrous or stellate-pubescent; style straight or curved, in andromonoecious species shorter than the anthers and held within the anther cone; stigma minutely to large-capitate or clavate, sometimes bilobed. Fruit a berry, usually globose or nearly globose (but variously shaped in S. mammosum and S. melongena), the pericarp dry, fleshy or leathery, glabrous or stellate-pubescent, the surfaces shiny or matte; fruiting calyx lobes not enlarging or often accrescent, sometimes completely enclosing the berry. Seeds flattened-reniform, the testal cells sinuate or polygonal in outline. Chromosome number: n = 12, 24, 36 (see Scaldaferro et al. 2012).

Members of the Leptostemonum Clade are found worldwide, in all habitats but the group is most diverse in the tropics (see Whalen 1984; Bohs 2005; Gagnon et al. 2022). They are often weedy plants of disturbed areas.

The Leptostemonum Clade is the largest monophyletic group in the genus Solanum. The clade is characterised by the possession of stellate trichomes (sometimes modified), prickles (sometimes absent) and long-attenuate, tapering anthers with distally directed pores that do not elongate to laterally dehiscing slits (as do those of other groups of Solanum, see Knapp et al. 2019a). Here we have chosen to treat the species occurring in tropical Asia as a group to fill a significant gap in the understanding of the species level taxonomy of spiny solanums. As for other geographically based groupings in spiny solanums, Asian members of the Leptostemonum Clade do not form a monophyletic assemblage (e.g., Aubriot et al. 2016a; Vorontsova and Knapp 2016); when the species treated have been included in phylogenetic studies, their hypothetical evolutionary relationships are provided in species discussion sections. For further discussion and previous assignments of the taxa treated here to species groups see sections on Taxonomy and Phylogeny above and Table 1.

We have cited types but not reproduced synonymy and complete descriptions of the introduced species here; these can be found in the cited monographic works and on Solanaceae Source. Several of these names are lectotypified here because they were incorrectly cited as having holotypes in previous works. We have cited all synonyms based on specimens from tropical Asia. Many of these introduced taxa are expanding in range with increased disturbance and environmental damage, so the distribution given in Table 2 is what we know now; we expect some of these to be found elsewhere in the future.

Cultivated in Vienna, Austria, of unknown origin, Anonymous s.n. (lectotype, designated by Vorontsova and Knapp 2016, pg. 41: W [acc. # 0022470]).

Vorontsova and Knapp (2016: 41–45); http://www.solanaceaesource.org/solanaceae/solanum-aculeatissimum.

Solanum aculeatissimum occurs only sporadically in tropical Asia (see Table 2); it is native to Brazil (Nee 1979) and adventive and widely distributed in Africa, it is less common in tropical Asia than the similar S. viarum.

China. ka xi qie (Zhang et al. 1994). Malaysia/Singapore. tĕrong perat, tĕrong asam hutan, tĕrong puyoh, tĕrong bĕlanda, tĕrong pusat, tĕrong piat (Burkill 1935), tèrong kori, tèrong tènang (Sundanese, Burkill 1935). Vietnam. cà ung (Hul and Dy Phon 2014). Common names from Burkill (1935) may also refer to S. viarum.

Solanum aculeatissimum is a member of the Acanthophora clade (sensu Stern et al. 2011; Gagnon et al. 2022) as are several other introduced species in tropical Asia (S. capsicoides, S. mammosum, S. viarum) and is most similar to S. viarum. It can be distinguished from that species in its long-acuminate (versus deltate) calyx lobes and its minutely stipitate-glandular (versus puberulent) ovary. Solanum viarum is more finely and evenly glandular pubescent than is S. aculeatissimum.

The herbarium of the French botanist and clergyman Augustin A.H. Léveillé was acquired by the Scottish botanist George Forrest from whence it passed to the Royal Botanic Garden Edinburgh. Vorontsova and Knapp (2016) incorrectly cited a collection in E as the holotype, no herbarium was cited in the protologue. We have selected the specimen at E (E00284478) that corresponds to the description, collector, and locality (Léveillé 1908), as the lectotype for S. cavaleriei.

See Suppl. materials 1–3.

Ethiopia. Sin. loc., J. Burser vol. 9 no. 17 (lectotype, designated by Hepper and Jaeger 1985, pg. 391: UPS).

Vorontsova and Knapp (2016: 49–56); http://www.solanaceaesource.org/solanaceae/solanum-aethiopicum-0.

We have recorded S. aethiopicum from China and India, it is recorded from Sri Lanka (as S. integrifolium Poir.) by Hepper (1987); it is a plant from tropical sub-Saharan Africa (native) but is not known in the wild; widely cultivated in South America, Europe, and Asia.

China. hong qie (Zhang et al. 1994).

Solanum aethiopicum is the scarlet eggplant or gilo. It is sporadically cultivated for its fruits (and possibly leaves) in southwestern China and adjacent India for its edible fruits. Cultivars from tropical Asia are usually glabrescent, with large, ribbed and sometimes flattened berries. It can be easily distinguished from the cultivated brinjal eggplant S. melongena by its smaller white flowers and red or reddish orange mature berries. It could possibly be confused with the widespread S. violaceum (that is somewhat similar to S. anguivi Lam., the wild progenitor of S. aethiopicum, see Vorontsova and Knapp 2016), but is more glabrescent than that species, the berries are much larger, and the fruiting pedicels are not as strongly spreading.

Figure 5. 

Introduced species of Solanum. Solanum aculeatissimum Jacq. A habit (Bidault et al. 3627, Gabon) B detail view of a flower (Bidault et al. 3627, Gabon) C detail view of a fruit (Bidault et al. 3627, Gabon). Solanum aethiopicum L. D detail view of a flower (field photograph, unvouchered, in cultivation at Radboud University, Nijmegen) E detail view of fruits (field photograph, unvouchered, in cultivation at the Max Planck Institute for Plant Breeding Research, Cologne). Solanum capsicoides All. F detail view of a flower (Coronado González 5457, Nicaragua) G detail view of a fruit (Coronado González 5457, Nicaragua). Solanum chrysotrichum Schltdl. H detail of a fertile branch (field photograph, unvouchered, India). Photograph credits: A–C E. Bidault D, E, H S. Knapp F, G I. Coronado González.

See Suppl. materials 1–3.

Somalia. Banaadir: Mogadischo, 30 May 1913, G. Paoli 137 (neotype, designated by Vorontsova and Knapp 2016, pg. 74: FT [FT003945]; isoneotype: FT [FT003946]).

Erect shrub to small tree, 2–6 m, densely prickly. Young stems erect, robust, densely stellate-pubescent and densely prickly, with porrect to multangulate, variously stalked trichomes, the stalks to 0.15 mm long, the rays 7–8(–15), 0.05–0.2 mm long, the midpoints reduced or absent, the prickles 5–10 mm long, 2.5–8 mm wide at the base, strongly curved, flattened, pale yellow to orange, densely stellate-pubescent in the lower 1/3; bark of older stems glabrescent to moderately stellate-pubescent, reddish brown. Sympodial units plurifoliate. Leaves simple, lobed, the blades 2–4(–7) cm long, 1.5–2.5(–3.5) cm wide, 1.5–2 times longer than wide, elliptic, concolorous, armed with 2–5 acicular, straight prickles to 12 mm long on both surfaces; adaxial surface glabrescent, with trichomes only at the base or along the midvein; abaxial surface glabrescent to sparsely stellate-pubescent with porrect, sessile or stalked trichomes, the stalks to 0.2 mm long, the rays 7–9, 0.1–0.2 mm long, the midpoints to 0.3 mm long; principal veins 3–5 pairs, further venation not visible or faint; base cuneate to rounded; margins lobed, the lobes 2–4 on each side, 0.2–0.7(–1.5) cm long, deltate to ovate, sometimes with secondary lobing, apically rounded, sometimes obtuse, the sinuses extending 1/4–2/3 of the distance to the midvein; apex broadly acute to rounded; petiole 0.05–0.5 cm long, less than 1/5 of the leaf blade length, densely stellate-pubescent, unarmed or with a few prickles. Inflorescences apparently lateral, 2–3(–4) cm long, unbranched or forked, with 2–8 flowers; 1–3 flowers open at any one time, densely stellate-pubescent, with 0–2 prickles; peduncle 0–4 mm long; pedicels 0.45–1 cm long, erect, articulated at the base, densely stellate-pubescent, unarmed; pedicel scars spaced 1–4 mm apart. Flowers 5-merous, heterostylous and the plants andromonoecious, with the lowermost 1–2 flowers long-styled and hermaphrodite, the distal flowers short-styled and staminate. Calyx 5.5–9 mm long, the lobes 3–4 mm long, deltate, apically acute to acuminate, sparsely stellate-pubescent, unarmed or with a few prickles. Corolla 2–3.2 cm in diameter, mauve to purple, stellate, lobed for ca. 4/5 of its length, the lobes 7–14 mm long, 3.5–4.5 mm wide, narrowly deltate, spreading, densely stellate-pubescent abaxially, the trichomes porrect, sessile or stalked, the stalks to 0.2 mm, the rays 6–9, 0.2–0.3 mm long, the midpoints reduced or to 0.5 mm long. Stamens equal; filament tube ca. 1.5 mm long; free portion of the filaments 0.5–1.5 mm long; anthers ca. 5.5 mm long, yellow, connivent, tapering, poricidal at the tips, the pores not lengthening to slits with age. Ovary densely stellate-pubescent in the upper 1/3; style 13–17 mm long in long-styled flowers, stout, curved, weakly stellate-pubescent in the lower 1/3, ca. 2 mm long in short-styled flowers; stigma clavate, minutely papillate. Fruit a globose berry, 1–2 per infructescence, 1.8–3 cm in diameter, the pericarp smooth, dark green with pale green and cream markings when young, yellow at maturity; fruiting pedicels 0.8–1.5 cm long, 1.5–2 mm in diameter at the base, woody, pendulous, usually unarmed but occasionally with a few prickles; fruiting calyx not markedly accrescent, but elongating to 7–12 mm long, somewhat fleshy and covering ca. 1/4 of the mature fruit (reflexed in herbarium sheets and on old fruits), unarmed or with up to 10 prickles. Seeds ca. 20–40 per berry, 2.2–3.2 mm long, 1.8–2.2 mm wide, flattened-reniform, dull yellow to orangish brown, the surfaces minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.

Figure 6. 

Solanum arundo Mattei A herbarium specimen collected in India in 1963 (Safni CNH-2488) B detail view of flower (Vorontsova et al. 80, Kenya) C detail view of fruit (Vorontsova et al. 80, Kenya). Photograph credits: A © The Director, Botanical Survey of India, Kolkata B, C M.S. Vorontsova.

(Fig. 7). Solanum arundo is an African species, found primarily in eastern Africa from Somalia to Tanzania, in tropical Asia it occurs only in far western India on the Kathiawar peninsula in the state of Gujarat.

Solanum arundo is a plant of grasslands and savannahs in Africa (Vorontsova and Knapp 2016); in India it is found in seaside vegetation (Sen Gupta 1963).

There have been no common names or uses recorded for S. arundo, but in Africa it is often used as a hedge plant, due to its ability to form thick impenetrable thickets (Vorontsova and Knapp 2016).

Figure 7. 

Distribution of S. arundo.

(IUCN 2019). Least Concern (LC); EOO (1,277,240 km²), AOO (216 km²). Solanum arundo has a broad distribution in Africa, and although it is relatively poorly represented in India, it is not of particular conservation concern based on its worldwide distribution. The distribution and occurrence of S. arundo on the Kathiawar peninsula need to be confirmed with modern collections.

Solanum arundo is a distinctive small extremely prickly tree, in Africa it is often associated with human habitation (Vorontsova and Knapp 2016). It is easy to distinguish from other spiny solanums in western India by the combination of thick, curved, and flattened prickles on the densely stellate- or multangulate-pubescent stems, and long, straight prickles on the leaves. Solanum forskalii is sympatric with S. arundo and has similar white-pubescent stems, but it is a smaller, scandent plant, with truncate to cordate rather than cuneate leaf bases, smaller corollas (1.3–2.4 versus 2–3.2 cm in diameter) and much smaller fruits (0.6–0.9 versus 1.8–3 cm in diameter). In 2018, one of us (SK) erroneously identified the specimens of S. arundo in CAL as S. forskalii (Fig. 6).

Vorontsova et al. (2013) resolved S. arundo as sister to the more narrowly distributed African endemic S. dennekense Dammer, a similar treelet with curved stem spines, white stems, and large, leathery berries.

Sen Gupta (1963) suggested that S. arundo had arrived in India with Arab traders in the nineteenth century, as part of the extensive Gujarati trade routes established across the Arabian Sea. Its restricted distribution on the Kathiawar peninsula certainly supports this idea. Solanum arundo has not been collected, to our knowledge, since the 1960s, re-evaluation of its presence and distribution in western India is necessary.

See Suppl. materials 1–3.

Myanmar. Tanintharyi Region: “Tavoy” [Dawei], Aug 1827, N. Gómez 9071 (lectotype, designated by Hul and Dy Phon 2014, pg. 34: K [K000759381]; isolectotypes: GZU [GZU000255503], LE).

Herbs to small shrubs to 1 m tall, heavily armed. Stems erect, terete, densely prickly and pubescent, the pubescence deciduous with age leaving only prickles and bristles on older stems; prickles of varying sizes to 0.5 cm long, broad-based, straight, yellowish tan in dry material, purplish black or yellowish tan in live plants, grading into long-stalked bristles and stellate trichomes; pubescence of porrect-stellate trichomes, mixture of sessile, short- and long-stalked, often recorded as purple-tinged or blackish red, the stalks multiseriate, to 3 mm long, the rays 4–6, 0.5–1 mm long, thin and brittle, the midpoints absent on stalked trichomes, the sessile trichomes with elongate midpoints to 2 mm long, always longer than the rays; new growth densely pubescent with mixture of short- and long-stalked stellate trichomes; bark of older stems greyish brown. Sympodial units difoliate, the leaves not geminate, or occasionally unifoliate in very small plants. Leaves simple, shallowly lobed, the blades (7–)12–30 cm long, (5–)10–17 cm wide, 1.5–1.7 times longer than wide, broadly elliptic to somewhat ovate, chartaceous, discolorous (paler green beneath fide Maxwell 91-651), sparsely armed along the midrib and major veins with prickles to 1 cm long; adaxial surface dark green, sparsely pubescent with sessile porrect-stellate trichomes, the rays 0–4, 0.1–0.5 mm long, the midpoints 2–3 mm long, delicate and thin; abaxial surface with similar sessile porrect-stellate trichomes, but these denser, especially along the veins; major veins 4–7 pairs, densely pubescent especially abaxially, in live plants often purple-tinged; base abruptly truncate, somewhat oblique; margins shallowly lobed, the lobes 4–7 on each side, occasionally somewhat secondarily lobed, to 1 cm long, more or less broadly deltate, apically acute, the sinuses less than halfway to the midrib; apex acute; petioles 3–9 cm long, half as long to almost as long as the leaf blades, prickly and sparsely pubescent, the prickles to 1 cm long, straight, grading into bristles that themselves grade into to a few long-stalked stellate trichomes, the pubescence largely of sessile stellate trichomes with 5–6 rays 0.5–1 mm long and midpoints to 2 mm, always longer than the rays. Inflorescences (1–)3–10 cm long, internodal and lateral, unbranched, with 10–40 flowers, few flowers open at any one time, densely bristly and pubescent with trichomes like those of the stems, the bristles sparser distally with early deciduous rays at the apex grading into prickles; peduncle 0.5–2 cm long, unarmed or armed with a few prickles; pedicels 0.5–0.8 cm long, 0.5–0.7 mm in diameter at the base and apex, slender and nodding at anthesis, densely bristly and stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars irregularly spaced 3–4 mm apart. Buds long-tapering and somewhat fusiform, almost completely enclosed in the foliaceous calyx. Flowers 5-merous, heterostylous and the plants weakly andromonoecious, with most flowers long-styled and only the most distal short-styled, these drop after flowering and the whip-like inflorescence axis tip persists in fruit. Calyx with the tube 2.5–5.5 mm long, deeply cupulate to somewhat urceolate, densely bristly with bristles to 5 mm long, these elongating in fruit, the lobes 5–8 mm long, 3–4 mm wide, lanceolate to broadly triangular, densely stellate-pubescent abaxially with sessile porrect-stellate trichomes with 4–6 weak rays and an elongate midpoint. Corolla 1.8–2.2 cm in diameter, white or violet (usually adaxially white and abaxially violet, but not always), deeply stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 8–10 mm long, 1.5–3 mm wide, narrowly deltate, spreading to recurved at anthesis, glabrous adaxially, densely stellate-pubescent abaxially with sessile trichomes with elongate midpoints, these densest at the tips. Stamens slightly unequal, with 2 slightly longer than the rest; anthers 8.5–10 mm long, 2 slightly longer, ca. 1 mm wide, strongly tapering, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments minute, glabrous, the anthers almost sessile. Ovary conical, glabrous; style (long-styled flowers) 8–10 mm long, glabrous, ca. 3 mm long in short-styled flowers; stigma capitate, the surfaces minutely papillose, bright green in live plants. Fruit a globose berry, 6–12 per infructescence, 1–1.4 cm in diameter, pale greenish white, completely enclosed in the bristly calyx tube, the pericarp thin and more or less shiny, glabrous; fruiting pedicels 0.7–0.9 cm long, ca. 1 mm in diameter at the base, 1.7–2 mm in diameter at the apex, somewhat woody, spreading to pendent; fruiting calyx greatly expanded to enclose the berry, the tube densely bristly and surrounding the berry, the lobes to 1 cm long, 0.3 cm wide, stellate-pubescent. Seeds 20–30 per berry, 2.5–5 mm long, (1.5)2–3 mm wide, flattened reniform, pale tan or yellowish brown (white in live plants), the surfaces minutely pitted, the testal cells pentagonal in shape. Chromosome number: not known.

Figure 8. 

Solanum barbisetum Nees A herbarium specimen collected in Laos in 1932 (Poilane 20487, P00055977) B habit and leaves (Suksathan et al. PS 3832, Thailand) C detail of the calyx with stellate trichomes and bristles (Suksathan et al. PS 3832, Thailand) D many flowered inflorescence (Suksathan et al. PS 3832, Thailand) E detail view of a flower (Suksathan et al. PS 3832, Thailand). Photograph credits: A CC-BY, Muséum national d’Histoire naturelle, Paris B–E D. Pedersen.

(Fig. 9). Solanum barbisetum occurs from southern China (Yunnan) and northeastern India to Laos and Thailand.

Solanum barbisetum grows in open areas, in fields and along forest edges in mixed broadleaf forests; usually from 300 to 2,000 m elevation, more rarely at sea level.

China. ci bao qie (Zhang et al. 1994). Thailand. Nakhon Si Thammarat: ma uk pa (Rabil Bunnag 204); North: ma khûa chê pâ [Lao] (Winit Wanadorn 1424).

Fruits (ripe berries) said to be eaten in India (Jain and Borthakur 1986).

(IUCN 2019). Least Concern (LC); EOO (1,244,077 km²), AOO (248 km²). Solanum barbisetum is widely distributed and is a plant of disturbed areas such as field and forest edges; the small AOO certainly reflects collection bias.

Figure 9. 

Distribution of S. barbisetum.

Solanum barbisetum is a distinctive species with its elongate inflorescences, berries enclosed in an accrescent calyx and copious long-stalked stellate trichomes on all vegetative parts. It is morphologically most similar to S. praetermissum and to a lesser extent S. involucratum; it shares with both those species berries enclosed in accrescent calyces and large leaves. Solanum involucratum is not sympatric with S. barbisetum and is a more robust plant with pubescent berries and shorter inflorescences. Solanum praetermissum is more or less sympatric with S. barbisetum and has been treated as a variety of S. barbisetum in the past (Clarke 1883, see description of S. praetermissum). Solanum barbisetum differs from S. praetermissum in its bristlier pubescence, with the calyx trichomes becoming stiff and prickly as the fruit matures and its more attenuate leaf bases. The calyx pubescence of S. barbisetum is often purple-tinged; this is even visible in quite old herbarium sheets.

Solanum barbisetum was treated under this species name for Bhutan in Mill (2001), but with the note that “Most specimens from our area labelled ‘S. barbisetum’ belong, or appear to belong to S. griffithii” (Mill 2001: 1059). The correct name for S. griffithii (Prain) Wu & Huang is S. praetermissum (see description); care should be taken with the identification of material from Bhutan for these taxa.

The phylogenetic affinities of S. barbisetum are unclear. In the analyses of Aubriot et al. (2016a) it is resolved as sister to the ‘S. expedunculatum and relatives’ clade (S. expedunculatum Symon, S. heteracanthum Merr. & L.M.Perry, S. involucratum and S. procumbens) but this is only supported with Bayesian inference. It might also be closely related to the morphologically similar S. praetermissum but that relationship is not clearly supported in molecular analyses (Vorontsova et al. 2013; Aubriot et al. 2016a).

See Suppl. materials 1–3.

Vietnam. Khánh Hòa: “Province Nha Trang, dunes littorales de Cam Ranh, My La face à la lagune de Bau Ro”, 7 Mar 1961, Lê Công Kiêt 94 (holotype: P [P00055921]; isotype: P [P00055922]).

Scandent shrubs, to 1.5 m tall, unarmed or sparsely prickly. Stems more or less erect, terete, unarmed or occasionally with a few scattered tiny prickles, sparsely to densely stellate-pubescent; prickles to 1.25 mm long, to 1 mm in diameter at the base, straight or curved, acicular to deltate, orange, glabrous; pubescence of mixed sessile and very short-stalked porrect-stellate trichomes, the stalks to 0.1 mm long, the rays 6–9, 0.1–0.4 mm long, the midpoints absent or up to 0.1 mm long; new growth densely stellate-pubescent, light brownish in dry material; bark of older stems brownish grey, glabrescent. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, not lobed, the blades 2–3.5 cm long, 1.5–2.5 cm wide, ca. 1–1.5 times longer than wide, broadly ovate to suborbicular, chartaceous, slightly discolorous, unarmed; adaxial surface densely stellate-pubescent, the stellate trichomes porrect, sessile to stalked, the stalks to 0.1 mm, the rays 6–8, 0.1–0.4 mm long, the midpoints to 0.1 mm; abaxial surface densely stellate-pubescent with trichomes like those of the adaxial surface; major veins 3–5 pairs, drying yellowish light-green; base truncate to subcordate; margins entire or shallowly sinuate; apex obtuse; petioles 0.4–1 cm long, 1/5–1/3 of the leaf blade length, unarmed and densely stellate-pubescent, the pubescence of sessile and short-stalked stellate-trichomes like those of the blades. Inflorescences 1.5–2 cm long, internodal and lateral, sometimes appearing terminal, unbranched, with ca. 4–7 flowers, only 1 or 2 flowers open at any one time, densely stellate-pubescent, with a mix of sessile and short-stalked porrect trichomes like those of the stems, unarmed; peduncle 0.5–2 cm long, unarmed; pedicels 5–9 mm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading to erect, unarmed, densely stellate-pubescent with porrect trichomes like those of the inflorescence axes, articulated at the base; pedicel scars irregularly spaced 0.5–4 mm apart. Buds elongate ellipsoid, more or less strongly exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube ca. 3 mm long, campanulate, the lobes 2–3 mm long, ca. 1 mm wide, deltate and constricting to a short acumen, the acumen 1/4–1/3 the total lobe length, unarmed and densely stellate-pubescent abaxially with porrect-stellate trichomes like those of the pedicels. Corolla 0.8–1 cm in diameter, blue to light purple, stellate, lobed ca. 3/4 of the way to the base, the lobes 5–7 mm long, 1.5–2 mm wide, deltate, spreading at anthesis, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal; anthers 4–6 mm long, ca. 0.75 mm wide, tapering, orange, glabrous, poricidal at the tips, the pores not lengthening to slits with age; filament tube <0.5 mm long, glabrous; free portion of the filaments 0.5–1 mm long, glabrous. Ovary globose, with minute glandular hairs; style 7–8.5 mm long, slender, curved at the apex, glabrous; stigma capitate, minutely papillate. Fruit a globose berry, several per infructescence, 0.5–0.8 cm in diameter, the pericarp thin and smooth, red when mature, glabrous; fruiting pedicels 0.7–1 cm long, 0.5–1 mm in diameter at the base, 1.6–2 mm in diameter at the apex, woody, erect to spreading, straight to slightly deflexed, unarmed; fruiting calyx lobes not expanding, 1/2–2/3 the length of the mature fruit, broadly deltate, reflexed, unarmed. Seeds 8–10 per berry, 2.5–3 mm long, 2–2.5 mm wide, flattened-reniform, dull yellow, the surface minutely pitted, the testal cells pentagonal in outline. Chromosome number: not known.

Figure 10. 

Solanum camranhense Dy Phon & Hul A herbarium specimen (holotype) collected in Vietnam in 1961 (Lê Công Kiêt 94, P00055921) B inflorescence (field photograph, unvouchered, Vietnam) C detail view of the fruits (field photograph, unvouchered, Vietnam). Photograph credits: A CC-BY, Muséum national d’Histoire naturelle, Paris B, C S. Hul.

(Fig. 11). Solanum camranhense is endemic to Vietnam; the few known collections are restricted to Khánh Hòa and Bình Thuận provinces of South Vietnam.

Solanum camranhense has only been collected on coastal dunes of stabilized red sands; 15–20 m elevation.

Vietnam. Khánh Hòa: củ vè [Vietnamese] (Chevalier 38932).

(IUCN 2019). Endangered (EN [B2ab(i,ii,iii)]); EOO (555 km²), AOO 16 km²). Due to the paucity of collections, it is difficult to document the range of Solanum camranhense with confidence. However, its occurrence in the fragmented and anthropogenically disturbed habitats of coastal southern Vietnam (Wikramanayake et al. 2002) suggests it is of conservation concern.

Solanum camranhense is morphologically similar to the sympatric S. robinsonii, but differs from that species in its scandent (versus erect) habit, the denser pubescence that dries yellowish brown, shorter leaves (2–3.5 cm long versus 3–8 cm long in S. robinsonii), and smaller, more deeply lobed corollas (usually less than 1 cm in diameter and lobed 3/4 of the way to the base versus 1–2 cm in diameter and only lobed halfway to the base in S. robinsonii). Hul and Dy Phon (2014) described S. camranhense as unarmed, but we have seen specimens with a few scattered prickles on the scandent stems. The two species differ in habitat; S. camranhense is a plant of coastal dunes, while S. robinsonii occurs in coastal forests.

Figure 11. 

Distribution of S. camranhense.

Solanum camranhense is a member of the clade ‘S. camranhense and relatives’ of Aubriot et al. (2016a) with S. nienkui and S. putii; S. robinsonii has not been included in any molecular analyses to date, but we expect it to be closely related to these taxa.

See Suppl. materials 1–3.

Cultivated in Turin, Italy (Torino), Anonymous s.n. (holotype: TO [fide Nee 1979]).

Vorontsova and Knapp (2016: 108–111); http://www.solanaceaesource.org/solanaceae/solanum-capsicoides.

Solanum capsicoides is widespread but scattered throughout tropical Asia (see Table 2); it is native to Southern South America and is widely adventive and cultivated for ornament.

China. niu qie zi (Zhang et al. 1994).

Solanum capsicoides is a member of the Acanthophora clade along with several other American species introduced to tropical Asia (e.g., S. aculeatissimum, S. mammosum, S. viarum). It can be distinguished from those taxa and from any native species in the region in its absence of any stellate trichomes (all trichomes are simple, but derived from stellate ones, see Levin et al. 2005) and large dark orange berries with winged seeds.

In the description of S. bodinieri, Léveillé (1908) did not cite a herbarium or a particular specimen. The herbarium of the French botanist and clergyman Augustin A.H. Léveillé was acquired by the Scottish botanist George Forrest from whence it passed to the Royal Botanic Garden Edinburgh. Vorontsova and Knapp (2016) incorrectly assumed the sheet in E was the holotype. We here lectotypify this name using the specimen at E (E00284479) they cited as holotype.

Bonati (1915) did not cite a specimen or herbarium in his description of var. geoffrayi. Hul and Dy Phon (2014) cited “holo-, P!”, presumably referring to one of two sheets of this gathering in P. We select as lectotype the sheet at P with the original label (P00055825) that was presumably the sheet referred to by Hul and Dy Phon (2014).

See Suppl. materials 1–3.

Mexico. Michoacan or Jalisco[?]: near Las Trojes, 1825-31, C.J.W. Schiede 81 (lectotype, designated by Nee 1993, pg. 45: HAL [n.v.]).

Vorontsova and Knapp (2016: 120–122); http://www.solanaceaesource.org/solanaceae/solanum-chrysotrichum-0.

Solanum chrysotrichum has been recorded from China, India, Malaysia, and Sri Lanka; it is native Mexico and Central America. Outside of its native range it has been used as a fencing treelet and is widely adventive (see Vorontsova and Knapp 2016).

China. duo lie shui qie (Zhang et al. 1994).

Solanum chrysotrichum is a member of the Torva clade (sensu Stern et al. 2011; Aubriot et al. 2016a). It is morphologically similar to the Torva clade species native to tropical Asia (see Aubriot et al. 2016a for a discussion of this disjunction), but can be distinguished by its rusty pubescence, larger flowers and leathery green, rather than yellow, red or black berries. It is most similar to S. torvoideum, but the stellate trichomes of S. chrysotrichum are generally longer stalked than those of S. torvoideum and are more reddish brown rather than golden brown and the inflorescences of S. chrysotrichum are distinctly pedunculate, while flowers of S. torvoideum are borne right next to the stem.

In northern India S. chrysotrichum forms large stands along disturbed roadsides together with S. erianthum D.Don (Brevantherum clade), another widely distributed introduced species. Solanum chrysotrichum has often been identified as S. hispidum Pers.; that name is a synonym of the Andean species S. asperolanatum Ruiz & Pav. In the absence of voucher specimens, references to S. hispidum or S. asperolanatum in tropical Asian floristic works are difficult to assign to a species.

See Suppl. materials 1–3.

Indonesia. Java: Sin. loc., 1837, Without collector [J.C. von Hoffmannsegg] s.n. (holotype: G-DC [G00131442]; isotype: W [acc. # 1889-0135755]).

Habit not known, but probably shrubs or small trees, unarmed or armed with a few tiny prickles hidden by dense pubescence. Stems erect, terete, with a few tiny prickles and densely stellate-pubescent; prickles 1–3 mm long, very sparse if present, broad-based, straight, pale yellowish tan, usually absent; pubescence of mixed sessile and very short-stalked porrect-stellate trichomes, the stalks to 0.5 mm long, the rays 8–10, ca. 0.5 mm long, weak and tangled, the midpoints absent or to 0.4 mm long; new growth densely stellate-pubescent, the trichomes tangled whitish grey in dry material; bark of older stems grey (but only quite young stems seen). Sympodial units difoliate, the leaves geminate, the leaves of pair equal in size and shape or one leaf slightly smaller. Leaves simple, not lobed, the blades 5–11 cm long, 2.5–5 cm wide, ca. 2 times longer than wide, elliptic to narrowly elliptic, chartaceous, strongly discolorous, unarmed or sometimes sparsely armed along the midrib and major veins with small straight prickles; adaxial surface evenly and sparsely pubescent with erect short-stalked multangulate trichomes, the multiseriate stalks 0.1–0.5 mm long, the rays 6–8, ca. 0.4 mm long and all directly upwards (no clear midpoint), the lamina surface clearly visible; abaxial surface densely pubescent with tangled long-stalked porrect-stellate trichomes, the stalks ca. 1 mm long, the rays 8–10, to 0.6 mm long, thin and delicate; major veins ca. 5 pairs, densely pubescent especially abaxially, with a few tiny prickles abaxially; base acute-attenuate, somewhat oblique; margins entire; apex acute, the tip rounded; petioles 1–1.5 cm long, 1/8–1/4 as long as the leaf blades, sparsely prickly and densely stellate-pubescent, with 1–2 prickles to 2 mm long or more commonly prickles absent, the pubescence of sessile and short-stalked stellate-trichomes like those of the stems. Inflorescences 1–5 cm long, internodal and lateral, unbranched, with 5–15 flowers, only a few flowers open at any one time, densely pubescent with white sessile and short-stalked stellate-porrect trichomes like those of the stems, with 6–8 rays ca. 0.5 mm long and midpoints to 0.4 mm long or absent; peduncle absent and the first inflorescence branches appearing to arise directly from the stem, or to 0.3 cm long, unarmed; pedicels 1.2–1.7 cm long, ca. 1.5 mm in diameter at the base and apex, spreading to erect at anthesis, unarmed, densely stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars irregularly spaced 1.5–2 mm apart. Buds elongate ellipsoid and somewhat tapering, strongly exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect, but some distal flowers may be short-styled. Calyx with the tube ca. 3 mm long, conical, the lobes 1.5–2 mm long, 1–2 mm wide, deltate, unarmed and densely white stellate-pubescent abaxially with porrect-stellate trichomes like those of the pedicels. Corolla 2–2.6 cm in diameter, colour not recorded, shallowly stellate, lobed ca. halfway to the base, interpetalar tissue present and abundant, the lobes 6–9 mm long, 6–8 mm wide, deltate, spreading at anthesis, mostly glabrous adaxially but with a few stellate trichomes at the tips, densely stellate-pubescent abaxially with densely tangled sessile trichomes where exposed in bud, these densest at the tips, the interpetalar tissue glabrous or with a few stellate trichomes abaxially. Stamens equal or very slightly unequal, if unequal 3 longer than the other 2; anthers (longer 3) 7.5–8 mm long, ca. 1.5 mm wide, (shorter 2) 5–6 mm long, ca. 1 mm wide, all tapering, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments ca. 0.5 mm long, glabrous. Ovary conical, glabrous; style 10–13 mm long, glabrous; stigma capitate, the surfaces minutely papillose. Fruit a globose berry, several per infructescence, ca. 1.2 cm in diameter, colour not known, pericarp thin and shiny, glabrous; fruiting pedicels 1.6–1.8 cm long, 0.8–1 mm in diameter at the base, 2–2.5 mm in diameter at the apex, erect and slightly woody; fruiting calyx lobes ca. 2.5 mm long, not markedly accrescent, but covering the base of the berry and not reflexed. Seeds 20–50 per berry, ca. 3 mm long, ca. 2.3 mm wide, flattened reniform, pale yellowish tan, the surfaces minutely pitted, testal cells shape not clear. Chromosome number: not known.

Figure 12. 

Solanum comitis Dunal. Herbarium specimen (isotype) collected in Indonesia (Without collector [J.C. von Hoffmannsegg] s.n., W [acc. # 1889-0135755]). Photograph credit: CC BY, Naturhistorisches Museum, Wien.

Figure 13. 

Distribution of Solanum comitis.

(Fig. 13). Solanum comitis is endemic to the island of Java, Indonesia. No specific localities are recorded on the only two gatherings known of this species.

No habitat information has been recorded for S. comitis.

None recorded.

(IUCN 2019). Data Deficient (DD); known only from two collections of uncertain specific locality. Solanum comitis has not been re-collected since the early 19^th century, indicating it is certainly of conservation concern. Recollection of this distinctive species and discovery of any precise localities for its occurrence are priorities.

Solanum comitis is a distinctive species, with dense pubescence of multangulate trichomes that dries with a whitish grey tinge. It is superficially similar to the western Australian S. lasiophyllum Dunal with dense whitish grey pubescence but differs from that species in its smaller flowers that are probably all hermaphroditic, smaller berries and completely different habitat (tropical versus dry and seasonal). Its relationships are not known, but in the inflorescence, flower and fruit morphology S. comitis resembles members of the Torva clade that occur in the Asian tropics (e.g., S. poka, S. pseudosaponaceum) and we suspect it is a member of that group. Re-collection of this species is a priority.

Although the type specimen is attributed to J.C. Graf van Hoffmansegg, he never actually collected in Java, but rather employed a friend who made natural history collections for him (van Steenis-Kruseman 1985).

See Suppl. materials 1–3.

Yemen. Naquil Khailan between Beui Harath and Nehm, NE of Sana’a, 2400 m, 12 May 1978, J.R.I. Wood 2327 (neotype, designated by Wood 1984, pg. 136: K [K000441137]; isoneotype: BM [BM000942294]).

Scandent, sometimes erect herb to shrub, 0.3–1 m, unarmed or prickly. Young stems slender, creeping to ascendent, glabrescent, unarmed or sparsely prickly, the pubescence of multangulate, sessile trichomes visible to the naked eye as white dots, the rays 12–18, up to 0.1(–0.15) mm long, the midpoints reduced to a globular gland, the prickles, if present, 0–4 mm long, 0.1–2(–6) mm wide at the base, straight, sometimes slightly reflexed to curved, perpendicular to the stem, pale yellow to brown; bark of older stems glabrous, dark brown or dark grey to black. Sympodial units difoliate, not geminate. Leaves simple, entire, the blades 0.7–1.5(–3) cm long, 0.7–1.3(–3.5) cm wide, 1(–1.5) times longer than wide, orbicular, sometimes ovate, membranous, drying concolorous, yellowish green, glabrescent on both surfaces, with porrect, sessile, sometimes stalked trichomes, the stalks to 0.1 mm long, the rays 8–16, 0.05–0.1(–0.15) mm long, the midpoints reduced or a mere bump, unarmed on both surfaces; the primary veins not visible or 2–4 pairs, the tertiary venation not visible to the naked eye; base cordate to attenuate; apex rounded to acute; petiole 0.2–1.3 cm long, 1/3 as long to equal in length to the leaf blade, narrowly winged to almost filiform, sparsely stellate-pubescent, unarmed. Inflorescences apparently lateral, 1.5–4 cm long, unbranched, with 1(-2) flowers, 1(-2) flowers open at any one time; peduncle absent; pedicels 0.5–3 cm long, erect, filiform, protruding beyond the leaves, articulated at the base, moderately stellate-pubescent to glabrescent, unarmed; pedicel scars spaced 1–4 mm apart. Buds narrowly ellipsoid, the corolla strongly exserted from the calyx before anthesis. Flowers 4–5-merous, apparently all perfect. Calyx 2–4 mm long, the lobes 0.5–2 mm long, ca. 1 mm wide, deltate to narrowly deltate, apically obtuse to acuminate, sparsely stellate-pubescent, unarmed. Corolla 0.9–1.6 cm in diameter, mauve to purple, stellate, lobed 2/3–4/5 of the way to the base, the lobes 3–6.5 mm long, 1.2–2 mm wide, deltate, reflexed or spreading, stellate-pubescent abaxially, the trichomes porrect, sessile, the rays 8–15, up to 0.1 mm long, the midpoints shorter than the rays or reduced to bulbous base. Stamens equal; filament tube ca. 0.1 mm long; free portion of the filaments 0.5–0.7 mm long; anthers 3–5 mm long, yellow, spreading, tapering, poricidal at the tips, the pores distally directed. Ovary glabrous or with 1–2 stellate trichomes near the apex; style 0.8–0.9 cm long, filiform, curved, glabrous. Fruit a globose berry, 1(-2) per infructescence, 0.6–0.8 cm in diameter, the pericarp thin, glabrous, red at maturity; fruiting pedicels 1.5–3.5 cm long, 0.3–0.4 mm in diameter at the base, 1.5–2 mm in diameter at the apex, herbaceous, pendulous, unarmed; fruiting calyx weakly accrescent, elongating to 3–5 mm long, covering 1/4–1/3 of the mature fruit, unarmed. Seeds ca. 10–20 per berry, 1.8–3 mm long, 1.5–2.2 mm wide, flattened-reniform, dark brown to almost black, the testal cells somewhat sinuate in outline. Chromosome number: n = 12, 2n = 24 (Kumar and Subramaniam 1987, as S. gracilipes).

Figure 14. 

Solanum cordatum Forssk. Herbarium specimen collected in India (Badami 114, E00757243). Photograph credit: Royal Botanic Garden Edinburgh.

(Fig. 15). Solanum cordatum occurs from northeastern Africa and the Arabian Peninsula to Pakistan and western India (Punjab, Gujarat, Maharashtra). The record of S. cordatum from Tamil Nadu (Ramachandran and Viswanathan 2010) is based on mis-identified specimens of S. wightii.

Solanum cordatum occurs in grassland, bushland, and open woodland on silty, sandy, or stony soil; low elevations (not recorded for India, from sea level to 1,500 m fide Vorontsova and Knapp 2016).

None recorded from the region (see Vorontsova and Knapp 2016 for Africa).

(Knapp 2021a). Solanum cordatum has been formally assessed as a species of Least Concern (LC; https://www.iucnredlist.org/species/186619213/186619254).

Solanum cordatum appears to be much less commonly collected in western India than is the very similar and sympatric S. forskalii. It differs from S. forskalii in its compact multangulate stem pubescence of trichomes with many very short (<0.1 mm long) rays, S. forskalii has porrect-stellate trichomes with fewer rays that are marginally longer (> 0.15 mm long); stems of S. cordatum appear white dotted from the stubby trichomes. Leaf petioles of S. cordatum are decurrent and narrowly winged, while leaves of S. forskalii are distinctly petiolate. The calyx lobes of S. cordatum are longer and narrower than those of S. forskalii, while anthers are shorter in S. cordatum (3–5 mm versus 4.5–7 mm).

In eastern Africa, S. cordatum is deciduous in the dry season, growing opportunistically during times of moisture (Vorontsova and Knapp 2016). This may be part of the reason for its relatively fewer collections in India than S. forskalii; this species has more robust leaves that are not deciduous. Wood (1997) suggested that S. cordatum was an “Indian species which extends west to Yemen but is only found in Africa in Somalia.” It is indeed found only in the Horn of Africa (Somalia, Ethiopia, and northern Kenya).

Figure 15. 

Distribution of S. cordatum.

Ramachandran and Viswanathan (2010) reported S. cordatum from Sethukadai in the Namakkal District of Tamil Nadu, far from its known range in India. This report is based on mis-identified specimens of S. wightii; the description in Ramachandran and Viswanathan (2010) appears to be a mixture of elements from published descriptions of S. cordatum and the actual specimens of S. wightii they used as their new record.

See Suppl. materials 1–3.

Indonesia. Java: West Java [“in oryzetis siccis montium Seribu” Curug Seribu near Bogor; from protologue], C.L. Blume s.n. (lectotype, designated here: L [L0003630]).

Herbs to small shrubs, creeping over the ground, to 1 m tall, armed. Stems decumbent, terete, black to dark brownish, prickly and sparsely stellate-pubescent; prickles to 5 mm long, to 2.5 mm in diameter at the base, straight or curved at the tip, awl-shaped to deltate, flattened, pale yellow in dry material, tan or purplish black in live plants, glabrescent; trichomes porrect-stellate, sessile to stalked, the stalks to 0.1 mm long, the rays 4–7, 0.1–0.4 mm long, the midpoints absent or to 0.4 mm long, sometimes purplish black in live plants; new growth moderately to densely stellate-pubescent, light green in dry material; bark of older stems brownish grey, sparsely stellate-pubescent. Sympodial units plurifoliate, the leaves usually not geminate. Leaves simple, more or less deeply lobed, the blades 4.5–9 cm long, 2–5 cm wide, ca. 2 times longer than wide, elliptic to ovate, chartaceous, slightly discolorous, moderately prickly with 3–7 prickles per leaf side, the prickles to 1 cm long, to 1 mm wide at the base, straight at the tip, awl-shaped, conical, pale yellow in dried material sometimes purplish black in live plants, glabrous; adaxial surface mid-green, moderately stellate-pubescent, the stellate trichomes porrect, sessile to stalked, the stalks to 0.1 mm long, the rays 3–5, 0.1–0.4 mm long, the midpoints to 1 mm long, 2–3 times longer than the rays; abaxial surface light green, moderately stellate-pubescent with trichomes like those of the adaxial surface; major veins 4–5 pairs, drying dark; base cuneate to truncate; margins shallowly to deeply lobed, the lobes 1–4 on each side, 0.5–1 cm long, deltate to oblong, apically rounded, the sinuses extending up to halfway to the midrib; apex rounded to acute; petiole 0.7–2.2 cm long, 1/8–1/5 of the leaf blade length, moderately stellate-pubescent, armed with 1–4 prickles like those of the blades. Inflorescences 1.5–3.5 cm long, apparently lateral, unbranched, with ca. 1–4 flowers, 1–2 flowers open at any one time, sparsely to moderately stellate-pubescent with trichomes like those of the stems but with longer midpoints, unarmed; peduncle 0–8 mm long, unarmed; pedicels 0.5–2 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading to erect to somewhat nodding at anthesis, unarmed or with 1–11(–16) prickles, moderately stellate-pubescent with trichomes like those of the inflorescence axes, articulated at the base; pedicel scars spaced 0.5–6(–18) mm apart. Buds globose to oval, strongly included in the calyx lobes. Flowers 5-merous, apparently all perfect. Calyx with the tube 2–4 mm long, campanulate, the lobes 2–5 mm long, 1–1.5 mm wide, narrowly deltate, apically acute, densely prickly and stellate-pubescent abaxially with numerous prickles and trichomes like those of the pedicels but with shorter midpoints. Corolla 1–1.5 cm in diameter, white or purple, stellate, lobed ca. 1/2 of the way to the base, the lobes 3–6 mm long, 2–3 mm wide, deltate, spreading or somewhat campanulate and erect (not spreading and perpendicular to the pedicel) at anthesis, glabrous adaxially, moderately to densely stellate pubescent abaxially, the trichomes often purple-tinged. Stamens equal; anthers 3–5 mm long, ca. 0.5 mm wide, connivent, tapering, orange, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments ca. 1 mm long, glabrous. Ovary conical, minutely glandular-puberulent; style ca. 5 mm long, slender, curved at the apex, glabrous; stigma capitate, the surface minutely papillate. Fruit a globose berry, 1–4 per infructescence, 1–1.6 cm in diameter, red when mature, the pericarp thin and smooth, glabrous; fruiting pedicels 2.5–4 cm long, 0.5–1 mm in diameter at the base, 1–1.5 mm in diameter at the apex, woody, deflexed and nodding, unarmed or with 1–10 prickles; fruiting calyx lobes elongating to 1.3 cm long, 1/2–3/4 the length of the mature fruit, the tips slightly reflexed, usually completely enclosing at least the lower half of the berry, with 6–17(–24) prickles, these often purplish black. Seeds 85–170 per berry, 1.75–2 mm long, 1.5–2 mm wide, flattened reniform, dull yellow, the surface minutely pitted, the testal cells sinuate in shape. Chromosome number: not known.

Figure 16. 

Solanum cyanocarphium Blume A herbarium specimen collected in Vietnam in 1929 (Lichy 18, P00055844) B detail of the leaves and prickles (field photograph, unvouchered, Vietnam) C detail view of a flower (field photograph, unvouchered, Vietnam) D detail view of a fruit (field photograph, unvouchered, Vietnam). Photograph credits: A CC-BY, Muséum national d'Histoire naturelle, Paris B–D M. Nuraliev.

(Fig. 17). Solanum cyanocarphium is widely distributed from southeastern Indochina to western Malay Archipelago (Borneo, Java, Sumatra and south Philippines).

Solanum cyanocarphium has been found growing on limestone, in riparian and secondary tropical forests as well as in open degraded vegetation; from 10 to 1,000 m elevation.

Cambodia. plone lane [Khmer] (Chassagne 138), trâp krab [Khmer] (Hul and Dy Phon 2014); Indonesia. North Sumatra: tioeng (Toroes 1639), teroeng oetan (Toroes 2926); Malaysia/Singapore. tĕrong puyoh, tĕrong tikus, tĕrong pipit (all meaning “little brinjal”, Burkill 1935); Vietnam. Dông Nai: blou xit [Mnong] (Pierre s.n.), cà co [Vietnamese] (Pierre s.n.).

Solanum cyanocarphium is said to have edible berries (Cuadra A1010) and the juice is drunk for fever (Burkill 1935). Burkill (1935) also records its use to improve the appetite of elephants.

(IUCN 2019). Least Concern (LC); EOO 3,563,723 km² (LC), AOO 92 km² (EN). Historical collections of Solanum cyanocarphium suggest that this species is widely distributed but absence of recent collections from large areas of the historical range suggests it may merit conservation concern because of the increasing anthropogenic alteration of natural habitat throughout lowland and coastal tropical Asia (Wikramanayake et al. 2002).

Solanum cyanocarphium is a weakly climbing shrub to vine that scrambles over vegetation with hooked prickles; most specimens are quite thin-stemmed and somewhat scrappy. It is superficially similar to S. procumbens but differs from it in the strongly accrescent and spiny fruiting calyx (not accrescent in S. procumbens), larger prickles on stems and leaf blades, deeply lobed leaves (leaves usually entire in S. procumbens) and larger and many seeded berries. The accrescent calyx in fruit is similar to that of S. involucratum, but that species is more robust and has pubescent, rather than glabrous berries. Hul and Dy Phon (2014) suggested that S. cyanocarphium was introduced to Indochina but described what we recognise as a synonym (S. sakhanii) as new; we consider S. cyanocarphium to be native across tropical Asia where it occurs.

Figure 17. 

Distribution of S. cyanocarphium.

Aubriot et al. (2016a) included S. cyanocarphium in a clade called ‘S. cyanocarphium + S. sakhanii’; subsequent examination shows that the two taxa are synonyms (see above), not surprising given the very short branch lengths (Aubriot et al. 2016a).

The Blume specimen in L (L0003630) we have selected as the lectotype of S. cyanocarphium is the only one we have found that corresponds to the protologue and is likely to have been the original material that Blume used.

Nees van Esenbeck (1834) only cited a Wallich catalogue number in the protologue of S. sarmentosum, but no herbarium. The Solanaceae from his own herbarium are now held at the herbarium (GZU) of the University of Graz (Stafleu and Cowan 1981) and we have selected the specimen there (GZU000255826) as the lectotype, it has flower and fruit and an annotation “S. sarmentosum” in Nees van Esenbeck’s hand.

The protologue of S. bullatorugosum (Dunal 1852) cites two specimens at G; we have selected the more complete sheet at G-DC (G00145849) with both flowers and fruits as the lectotype.

The collection cited in the protologue of S. sparsiflorum (Elmer 1913) without specification of any herbarium (Elmer 13157) is represented in many collections; we have chosen that at GH (0007785) as the lectotype because it is the duplicate best matching the protologue and has both flower and mature fruit. Hul and Dy Phon (2014) cited a sheet in PNH as “holo-, PNH” but this does not constitute effective lectotypification.

Hul and Dy Phon (2014) cited a holotype at P for S. thorelii, but no herbarium was cited in the original protologue (Bonati 1914). We select here the best preserved of the three duplicates of Thorel 1419 held at P (P00054148) as the lectotype.

See Suppl. materials 1–3.

China. Yunnan: “Xiangping Mountain, Xishou [transl. from protologue]”, 29 Aug 1947, G. Feng 11429 (holotype: KUN [KUN484279]; isotypes: A [A00310057], PE [PE00730648], WUK [WUK0194818]).

Shrubs to 2 m tall, unarmed or rarely sparsely armed. Stems erect, terete, moderately stellate-pubescent; prickles, if present, to 5 mm long, to 3 mm wide at the base, broad-based, curved, pale yellowish tan, sparsely stellate-pubescent near the base; pubescence of sessile to very short-stalked porrect-stellate trichomes, the rays 5–7, 0.1–0.4 mm long, the midpoints absent or to 0.5 mm long; new growth densely stellate-pubescent, the trichomes white and tangled, soon deciduous and the stems glabrate; bark of older stems greyish brown to grey. Sympodial units difoliate, the leaves geminate, or not geminate, the leaves of a pair equal in size and shape. Leaves simple, shallowly lobed, the blades 5–11 cm long, 2.5–7.5 cm wide, ca. 2 times longer than wide, elliptic, chartaceous, discolorous, unarmed or sparsely armed along the midrib and major veins with small curved prickles; adaxial surface evenly and sparsely to moderately stellate-pubescent with white-cream sessile porrect trichomes, the rays 4–6, 0.1–0.2 mm long, the midpoints 0.5–1.3 mm long; abaxial surface with similar sessile porrect-stellate trichomes, but with some short-stalked stellate trichomes with stalks to 0.5 mm and longer rays to 0.5 mm long; major veins 3–4 pairs, densely pubescent especially abaxially; base abruptly truncate, somewhat oblique; margins shallowly lobed, the lobes 3–4 on each side, 0.3–1 cm long, broadly deltate, apically rounded, the sinuses less than halfway to the midrib; apex acute; petioles 1–2 cm long, 1/4–1/2 as long as the leaf blades, sparsely prickly and moderately pubescent, the prickles 0–3, like those of the stems, the pubescence of sessile stellate-trichomes like those of the stems. Inflorescences 1–3 cm long, internodal and lateral, unbranched, with 5–10 flowers, only 1 or 2 flowers open at any one time, pubescent with sessile stellate-porrect trichomes like those of the stems, with 4–6 rays ca. 0.2 mm long and midpoints to 0.5 mm long, unarmed or with a few small prickles to 3 mm long; peduncle 0.1–0.8 cm long, unarmed; pedicels 0.35–0.5 cm long, 0.5–1 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading and slightly nodding at anthesis, unarmed or with a few small prickles to 1 mm long, densely stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars irregularly spaced 3–4 mm apart. Buds ellipsoid and somewhat tapering, strongly exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect, but some distal flowers may be short-styled. Calyx with the tube 2–2.5 mm long, conical, the lobes 1–2 mm long, ca. 1 mm wide, deltate, unarmed or with a few small prickles to 3 mm long and densely stellate-pubescent abaxially with sessile porrect-stellate trichomes like those of the pedicels. Corolla 0.8–1 cm in diameter, white, deeply stellate, lobed 3/4 of the way to the base, interpetalar tissue present, the lobes 4–4.5 mm long, 2–2.5 mm wide, deltate, spreading at anthesis, mostly glabrous adaxially but with a few stellate trichomes at the tips, densely stellate-pubescent abaxially with densely tangled sessile trichomes where exposed in bud, these densest at the tips. Stamens equal; anthers 3–4 mm long, ca. 2 mm wide, slightly tapering, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments ca. 0.5 mm long, glabrous. Ovary conical, glabrous; style 5–6 mm long, glabrous; stigma capitate, the surfaces minutely papillose, bright green in live plants. Fruit a globose berry, several to many per infructescence, 1–1.2 cm in diameter, orange-red when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 0.8–1 cm long, 1–1.2 mm in diameter at the base, ca. 2 mm in diameter at the apex, somewhat woody, sharply deflexed and nodding, unarmed or with small prickles; fruiting calyx not accrescent, the lobes often breaking off. Seeds 50–60 per berry, 2–3 mm long, 2–3 mm wide, flattened reniform, pale tan or yellowish brown, the surfaces minutely pitted, the testal cells with sinuate margins. Chromosome number: not known.

Figure 18. 

Solanum deflexicarpum C.Y.Wu & S.C.Huang A herbarium specimen collected in China in 2007 (Knapp et al. 10130, BM001019378) B detail of a fertile branch (Knapp et al. 10130, China) C detail view of a flower (Knapp et al. 10130, China) D detail view of an infructescence (Knapp et al. 10130, China). Photograph credits: A CC-BY, © copyright The Trustees of the Natural History Museum, London B–D S. Knapp.

(Fig. 19). Solanum deflexicarpum is endemic to China (Yunnan province) and only known from few collections; potentially also present in northeastern Myanmar (Andrew s.n. collected in ‘Poneshee’ close to the boarder with Myanmar).

Figure 19. 

Distribution of S. deflexicarpum.

Solanum deflexicarpum grows in open vegetation and roadsides in semideciduous tropical forests, from 1,255 to 1,500 m elevation.

China. ku ci (Zhang et al. 1994); Yunnan: ku ci, wan bing ci tian qie (pedicel curved and spiny, high altitude) [Mandarin] (Feng 11429).

(IUCN 2019). Near Threatened (NT). EOO (65,755 km²); AOO (16 km²). Solanum deflexicarpum occurs over a relatively wide range, but there are very few collections; the paucity of collections suggest that the species is of some conservation concern, meriting further study.

Solanum deflexicarpum is morphologically similar to the sympatric and very widespread S. violaceum and to the Indian endemics S. hovei and S. multiflorum; in the analyses of Aubriot et al. (2016a) it is part of a monophyletic group with those species. It differs from S. violaceum in its condensed inflorescences with white rather than violet flowers and strongly down-curved fruiting pedicels; the fruiting pedicels of S. violaceum are more spaced, straight and are strongly spreading from the axis. Solanum deflexiflorum differs from S. hovei in its recurved fruiting pedicels (straight in S. hovei) and truncate, oblique (versus attenuate) leaf bases. The inflorescences of S. multiflorum are similarly condensed, but S. deflexicarpum differs from that species in its more compact trichomes, fewer flowers per inflorescence, and its distribution in southern China.

See Suppl. materials 1–3.

Indonesia. Malaku: Pisang Island, Moluccas Islands, [Dec 1818], C. Gaudichaud s.n. (lectotype designated by Bean 2004, pg. 672: P [P00369093]; isolectotype: P [P00369092]).

Shrub or small tree to 4 m tall, unarmed or sparsely prickly. Stems erect, unarmed or with a few scattered prickles, glabrous or very sparsely stellate-pubescent; prickles to 3.2 mm long, to 3.6 mm at the base, straight, narrowly triangular with the sides concave from a wide base, yellow-ferruginous, glabrous; trichomes porrect-stellate, sessile, the rays 6–8, ca. 0.1 mm long, the midpoints shorter to equal to the rays, white to yellow in dry material; new growth glabrous to sparsely pubescent with mixture of stellate and minute glandular trichomes, brownish to black in dry material; bark of older stems dark brownish red, glabrous. Sympodial units difoliate, the leaves geminate. Leaves simple, not lobed, the blades of major leaves 19.7–30 cm long, 9.5–16 cm wide, ca. 2 times longer than wide, ovate to elliptic, the minor leaves half as large as or the same size as the major leaves, subcoriaceous, slightly discolorous, unarmed or with a few prickles along the midrib; adaxial surface densely pubescent with a mixture of numerous minute glandular trichomes and a few porrect-stellate trichomes, the glandular hairs to ca. 0.04 mm long, the stellate trichomes with 6–8 rays, 0.1–0.2 mm long, the midpoints shorter to equal to the rays; abaxial surface moderately pubescent with similar sessile porrect-stellate and glandular trichomes; major veins 12–16 pairs, drying light brownish yellow; base short to very long attenuate, oblique; margin entire or slightly wavy; apex subacute to acute, or short acuminate; petioles 1.5–9 cm long, 1/9–1/6 of the leaf blade length, unarmed or occasionally armed with a few broad-based prickles to 3.9 mm long, to 2.5 mm in diameter at the base, straight, yellow-ferruginous, glabrous or with a few stellate trichomes like those of the blades. Inflorescence to 2 cm, apparently lateral, forked or several-branched, with ca. 50 flowers, 5–15 flowers open at any one time, glabrous to moderately stellate-pubescent on the youngest parts, with sessile porrect trichomes like those of the stems; peduncle 4–12.6 mm long, unarmed; pedicels 5.1–18.6 mm long, 0.3–0.4 mm in diameter at the base, 0.7–1.1 mm in diameter at the apex, erect, unarmed, glabrous to moderately stellate-pubescent with porrect trichomes like the inflorescence but often with longer rays, articulated at the base. Buds fusiform, exserted from the calyx before anthesis. Flowers 4–5-merous, apparently all perfect. Calyx with the tube 1.4–1.9 mm long, campanulate, the lobes 1–1.5 mm long, 1.5–1.8 mm wide, deltate, apex acuminate, the abaxial surface more or less strongly keeled along the midvein, unarmed and glabrous or sparsely stellate-pubescent abaxially with a few porrect-stellate trichomes like those of the pedicels. Corolla 1.8–2 cm in diameter, lavender to violet, stellate, lobed ca. 4/5 of the way to the base, the lobes 8.2–10.5 mm long, 2.5–2.9 mm wide, oblong, spreading at anthesis, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal; anthers 3.7–5.7 mm long, 0.8–1.3 mm wide, connivent, tapering, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying. Ovary conical, with a few stellate trichomes; style 6.5–9.5 mm long, filiform, curved towards the apex, glabrous; stigma capitate or slightly bilobed. Fruit a globose berry, several to many per infructescence, 0.8–1.2 cm in diameter, orange to red when mature, the pericarp thin and shiny, glabrous; fruiting pedicels 10.8–18.2 mm long, 0.7–0.9 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, erect; fruiting calyx lobes slightly expending to 1.7 cm long, covering 1/4 of the berry or reflexed, glabrous. Seeds ca. 40 per berry, 1.7–2.5 mm long, 2.5–2.8 mm wide, flattened-orbicular to flattened-reniform, yellow-tan to yellow-ferruginous, the surface minutely pitted, the testal cells pentagonal to sinuate in outline. Chromosome number: not known.

Figure 20. 

Solanum dunalianum Gaudich. Herbarium specimen collected in Indonesia in 1917 (Kaudern 57, L.4319974). Photograph credit: Naturalis Biodiversity Center.

(Fig. 21). Solanum dunalianum is found from Sulawesi east through New Guinea and south to the Cape York Peninsula in Queensland, Australia. It has also been collected in Vanuatu and Tonga (Symon 1985).

Solanum dunalianum typically occurs in disturbed habitats and has been found in secondary rainforest, along roads, in clearings, along streams, and gardens; from sea level to 1,200 m elevation (on New Guinea).

None recorded from the region treated here. Papua New Guinea. Eastern Highlands: gonovise (Kerenga LAE 56923); Tonga. Vava‘u: polo jongo (Soakai 1048).

(IUCN 2019). Least Concern (LC). EOO (887,043 km²); AOO (100 km²). Solanum dunalianum is at the northeastern edge of its range in tropical Asia, the species is common and widely distributed in New Guinea.

Solanum dunalianum is a species primarily of northern Australia and the island of New Guinea (McClelland 2012) that only just gets into tropical Asia. It was treated as a member of section Dunaliana Bitter by McClelland (2012) along with a number of other Pacific species such as S. viridifolium Dunal and S. labyrinthinum D.McClelland (see McClelland et al. 2020). Solanum dunalianum is one of the few species in this group that has been included in molecular analyses (e.g., Aubriot et al. 2016a) and it resolves as part of the ‘Sahul-Pacific clade’ and member of a group with S. schefferi (as S. lianoides) and S. graciliflorum. Within the area treated here, S. dunalianum is distinctive in its branched inflorescence with small red fruits and almost glabrous vegetative parts with very few, if any prickles. Other taxa in the area with several-branched inflorescences are usually densely and variously pubescent (e.g., S. giganteum, S. graciliflorum, S. pseudosaponaceum, S. torvoideum).

Figure 21. 

Distribution of Solanum dunalianum.

Figure 22. 

Introduced species of Solanum. Solanum chrysotrichum Schltdl. A detail of inflorescence (field photograph, unvouchered, India). Solanum elaeagnifolium Cav. B habit (Sampath Kumar et al. 126972, India) C detail view of a flower (Sampath Kumar et al. 126972, India) D detail view of a fruit (field photograph, unvouchered, India). Solanum jamaicense Mill. E detail view of a flower (Stern 265, Trinidad and Tobago) F detail view of fruits (Stern 265, Trinidad and Tobago). Solanum macrocarpon L. G detail view of flower (in cultivation at GAFL Avignon, unvouchered) H detail view of fruit (in cultivation at Radboud University, Nigmegen, unvouchered, material now at CGN, Wageningen). Photograph credits: A S.Knapp B–D X. Aubriot E, F S. Stern G, H S. Knapp.

The two collections from northern Papua New Guinea cited in the protologue of S. dunalianum var. puberius were destroyed in Berlin, and we have found no duplicates of these, despite extensive searches. Ledermann’s travels are well-documented (Ledermann 1919; van Steenis-Kruseman 1985; Veldkamp et al. 1988), and many of the areas he visited have not been accessed since (Takeuchi and Golman 2002), so we delay neotypifying this name until such collections are available or duplicates of Ledermann’s collections are found.

See Suppl. materials 1–3.

Cultivated in Madrid from “America calidiore” [“del viaje de los espanoles alrededor del mundo, Cult. en el R. J. Bot. 1793”], Anonymous s.n. (lectotype, designated by Knapp 2007, pg. 198: MA [MA-476348-2]; isolectotype: MA [MA-476348-1).

Knapp et al. (2017: 22–46); http://www.solanaceaesource.org/solanaceae/solanum-elaeagnifolium.