<figure><img src="image_74.png" /><figcaption>**Figure 59.** *Solanum pubescens* Willd. **A** herbarium specimen collected in India (Wight 2629b, BM000900098) **B** habit (*Sampath Kumar et al. 126956*, India) **C** inflorescence (*Sampath Kumar et al. 126956*, India) **D** immature fruit (*Sampath Kumar et al. 126956*, India). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B–D** X. Aubriot.</figcaption></figure>

unlobed, the blades 2.5–12 cm long, 1.5–7 cm wide, ca. 1–1.5 times longer than wide, ovate to broadly triangular, widest in the lower third, chartaceous, more or less concolorous, unarmed, the leaves of lower stems much larger than those of distal branches; adaxial surface evenly and densely pubescent with sessile and very short-stalked porrect-stellate trichomes, the rays 4–8, to 0.5 mm long, glandular at the tips, the midpoints 2–4-celled, to 2 mm long, glandular at the tips; abaxial surface with similar porrect-stellate trichomes, but these denser especially along the veins; major veins 3–4 pairs, densely pubescent especially abaxially; base abruptly truncate to cordate, somewhat oblique; margins entire or slightly sinuate, not lobed; apex acute; petioles 1–4 cm long, 1/2 of the leaf blade length, unarmed, more densely glandular stellate-pubescent than the stems, but the trichomes of the same morphology. Inflo-
rescences to 3 cm long, internodal and lateral, unbranched, with 5-12 flowers, only 1 or 2 flowers open at any one time, densely glandular pubescent with mixed multangulate and stellate-porrect trichomes like those of the stems; peduncle 0.3-0.5 cm long; pedicels 1.2-1.6 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading at anthesis, glandular stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars irregularly spaced 1-3 mm apart. Buds elongate ellipsoid and tapering, strongly exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 2-2.5 mm long, conical, the lobes 3.5-5 mm long, ca. 1.5 mm wide, long-triangular to lanceolate, apically acute, densely stellate-pubescent abaxially with mixed glandular multangulate and porrect-stellate trichomes like those of the pedicels. Corolla 2-2.2 cm in diameter, violet or deep purple, stellate, lobed 3/4 of the way to the base, minimal interpretalar tissue present, the lobes 6-7 mm long, 4-4.5 mm wide, spreading at anthesis, mostly glabrous adaxially or with a few stellate trichomes along the petal midvein, densely stellate-pubescent abaxially with densely tangled sessile trichomes where exposed in bud, these densest at the tips. Stamens markedly unequal, with 4 short and one long and curved; long anther 7.5-9 mm long, 1-1.5 mm wide, strongly curved and tapering, short anthers 5-7 mm long, 1-1.5 mm wide, straight, all anthers 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 9-10 mm long, strongly curved inwards and held adjacent to the long anther, glabrous; stigma capitate or slightly clavate, the surfaces minutely papillose. Fruit a globose berry, several per infructescence, 1-1.6 cm in diameter, orange-red when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 2.5-3.5 cm long, ca. 1.5 mm in diameter at the base, 2-2.5 mm in diameter at the apex, somewhat woody, spreading to pendent from weight of berries; fruiting calyx not accrescent, the lobes often breaking off. Seeds 20-30 per berry, 4-5 mm long, 3.5-4 mm wide, flattened reniform, yellowish or reddish brown, the surfaces minutely pitted, the testal cells with sinuate margins. Chromosome number: 2n = 24 (Chennaveeraiah and Krishnappa 1971; Rao and Rao 1987).

**Distribution (Fig. 60).** *Solanum pubescens* occurs from India to Saudi Arabia and Yemen on the Arabian Peninsula.

**Ecology and habitat.** *Solanum pubescens* is found in a variety of dry forest types, such as thorn forest, often occurring in open areas and along roadsides, from 300 to 1,000 m elevation.

**Common names and uses.** India. sonde, hucchu sonde, savadangi, cherichunda (Malayam), kaattu sundai kaai (Tamil), usthi kaai (Telugu) (https://www.flowersofindia.net). It is recorded as being used for bowel and joint pains (see https://www.flowersofindia.net).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (996,977 km², LC); AOO (272 km², EN). Like many other spiny solanums in tropical Asia, *S. pubescens* is widely distributed in a variety of habitats, although the AOO suggests some concern, this is a common species.
<figure><img src="image_75.png" /><figcaption>**Figure 60.** Distribution of *S. pubescens*.</figcaption></figure>

**Discussion.** *Solanum pubescens* is morphologically similar and probably closely related to *S. vagum*, sharing with that species zygomorphic flowers at anthesis, heteromorphic anthers and shiny berries on erect or slightly pendulous pedicels. Hepper (1987) treated material of *S. vagum* from Sri Lanka as *S. pubescens* in the “Revised Handbook to the Flora of Ceylon”. *Solanum pubescens* differs from *S. vagum* in its glandular pubescence, its narrowly elliptic leaves with attenuate bases and slightly smaller, violet (rather than white) flowers. In *S. pubescens* the lamina of young leaves is obscured by the dense covering of glandular stellate to multangulate stalked trichomes with elongate midpoints and the leaves are described as “oily to touch” (Singh 1988), while in *S. vagum* the pubescence of adaxial leaf surfaces is of very sparse sessile stellate trichomes with midpoints usually equal to the rays, and the lamina is clearly visible. The two species are sympatric in southern India.

*Solanum pubescens* and *S. vagum* share a zygomorphic androecium, with one anther distinctly longer than the rest. From herbarium sheets it appears that this difference becomes more pronounced with flower age. Post-anthesis anther expansion occurs in the unrelated *S. turneroides* Chodat (Brevantherum clade, see Stern et al. 2013) of southern South America. Buds of the African species *S. somalense* Franch. also have the anthers of more or less equal length that become different with age (Vorontsova and Knapp 2016). This phenomenon needs study with populations in the field and lab.
The Solanaceae from Nees van Esenbeck's personal herbarium are held in Graz at GZU (Stafleu and Cowan 1981). The sheet in GZU is the only one we have found of the Wallich Herb. "Madras" gatherings cited in the protologue of *S. calycinum*. We therefore designate it (GZU000255511) as the lectotype of *S. calycinum*.

Dunal (1852) coined the replacement name *S. conanthum* for *S. calycinum* Nees citing "non Dunal" in reference to his own *S. calycinum* (= *S. macrocarpon* L. of Africa, the Gboma eggplant, see Vorontsova and Knapp 2016) that is a later homonym of Nees van Esenbeck's name.

**Specimens examined.** See Suppl. materials 1–3.

**36. Solanum putii** Kerr ex Barnett, Kew Bull. 16: 485. 1963.
Fig. 61

**Type.** THAILAND. Prachuap Khiri Khan: Thap Sakae District, Hui Yang [Huai Yang], 6 Oct 1920, *Put Phraisurind 3227* (holotype: K [K000922027]; isotypes: BK [257531], BM [BM000886104]).

**Description.** Shrubs of unknown height, unarmed. Stems erect, terete, stellate-pubescent; pubescence of sessile to very short-stalked porrect-stellate trichomes, the rays 4–8, ca. 0.2 mm long, the midpoints absent or to 0.1 mm long, much shorter than the rays; new growth densely stellate-pubescent, the trichomes white and tangled, soon deciduous and the stems glabrate; bark of older stems pale brown. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, the blades 4–10 cm long, 1.5–5 cm wide, ca. 2 times longer than wide, elliptic, widest just below the middle, chartaceous, discolorous, unarmed; adaxial surface evenly and sparsely pubescent with mixed sessile and stalked porrect-stellate trichomes, the stalks to 0.5 mm long, the rays 4–8, to 0.5 mm long, the midpoints absent or to 0.5 mm long; abaxial surface moderately to densely pubescent with mixed stalked and sessile porrect-stellate trichomes, the stalks to 1.5 mm, the rays 8–10, to 0.5 mm long, the midpoints equalling the rays, the lamina clearly visible; major veins 4–5 pairs, densely pubescent especially abaxially; base abruptly truncate, usually strongly oblique; margins entire; apex acute to acuminate; petioles 1–2.5 cm long, ca. 1/4 as long as the leaf blades, unarmed and densely pubescent with weak-rayed porrect-stellate trichomes like those of the stems. Inflorescences 3–7 cm long, internodal and lateral, unbranched, with 10–20 flowers, apparently only a few flowers open at any one time, pubescent with porrect-stellate trichomes like those of the stems; peduncle 1.2–2.5 cm long; pedicels 0.9–1 cm long, ca. 1 mm in diameter at the base, ca. 1.3 mm in diameter at the apex, spreading and perhaps slightly nodding at anthesis, sparsely stellate-pubescent with porrect-stellate trichomes like the inflorescence axes, articulated at the base; pedicel scars evenly spaced 2.5–3 mm apart. Buds tapering, about halfway exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect (but isotype at BM has a flower dissection that is fasciated with 7 corolla lobes). Calyx with the tube 1.5–2 mm long, conical, the lobes 2–2.5 mm long, ca. 1 mm wide, deltate with a subulate tip to 1 mm long, sparsely stellate-pubescent with porrect-stellate trichomes like those of the pedicels. Corolla ca. 1.6 cm in diameter,
<figure><img src="image_76.png" /><figcaption>**Figure 61.** *Solanum putii* Kerr ex Barnett – Herbarium specimen (holotype) collected in Thailand in 1920 (*Put 3227*, K000922027). Photograph credit: © copyright of the Board of Trustees of the Royal Botanic Gardens, Kew.</figcaption></figure>
white or pale lilac, stellate, lobed ca. halfway to the base, interpretalar tissue scarce, but the petal margins thin and somewhat “ruffly”, the lobes ca. 5 mm long, ca. 5 mm wide, spreading or slightly reflexed at anthesis, mostly glabrous adaxially but with a few stellate trichomes along the petal midveins, densely stellate-pubescent abaxially with densely tangled sessile trichomes where exposed in bud, these densest at the tips. Stamens equal or slightly unequal with 2 slightly longer; filament tube ca. 0.5 mm long, glabrous and papery; free portion of the filaments ca. 1 mm long, glabrous; anthers 4–4.5 mm long, ca. 0.7 mm wide, if unequal, 2 ca. 0.2 mm longer, slightly tapering, yellow, poricidal at the tips, the pores directed distally, not elongating to slits with drying. Ovary conical, glabrous but with a few trichomes at the style base; style 6–6.5 m, glabrous; stigma tiny, a mere broadening of the style apex, the surfaces minutely papillate. Fruit a globose berry, 0.5–0.6 cm in diameter (immature?), colour not known, the pericarp thin and shiny, glabrous; fruiting pedicels 1–1.5 cm long, ca. 1 mm in diameter at the base, tapering to an apex ca. 2 mm in diameter, somewhat woody, spreading; fruiting calyx not accrescent, the lobes often breaking off. Seeds ca. 20 per berry, 2–2.5 mm long, 1.5–2 mm wide, flattened reniform, reddish tan or yellowish brown, the surfaces deeply pitted, the testal cells with sinuate margins. Chromosome number: not known.

**Distribution** (Fig. 62). *Solanum putii* is endemic to Thailand and only known from the type collection. A single collection from Vietnam (*Nuraliev NUR-1863a*)

<figure><img src="image_77.png" /><figcaption>**Figure 62.** Distribution of *Solanum putii*.</figcaption></figure>
appears to be this taxon, but we include it here with some reservation until further
studies are undertaken.

**Ecology and habitat.** No habitat notes are recorded on the type gathering of *S. putii*, but possibly occurs in dry forest.

**Common names and uses.** None recorded.

**Preliminary conservation status (IUCN 2019).** Data Deficient (DD). *Solanum putii* is only known from the type collection and little is known about this species otherwise.

**Discussion.** *Solanum putii* is very similar and may be conspecific with *S. robinsonii*, but we hesitate to synonymise these two taxa until more material is available; the similarities were noted in the original description (notes in packet on holotype at K; Barnett 1963). The leaves of *S. putii* are more broadly elliptic, more softly pubescent, and less discolorous than those of *S. robinsonii*; they are also markedly oblique at the base, while those of *S. robinsonii* are not. The lamina beneath is clearly visible and the rays on the stellate trichomes are thinner and more delicate. A single dissected flower on the isotype sheet at BM is 7-merous and clearly fascinated, perhaps indicating this is an aberrant plant, a note by Kerr on the holotype at K states "Only flower of *Put 3227* examined was abnormal with double ovary, stamens, etc.". The fruit of *S. putii* are smaller (ca. 0.6 cm in diameter) than those of *S. robinsonii* (ca. 1 cm in diameter).

**Specimens examined.** See Suppl. materials 1–3.

37. *Solanum retrorsum* Elmer, Leaflets Philipp. Bot. 1: 342. 1908.
Fig. 63

*Solanum luzoniense* Merr., Philipp. J. Sci. Bot. 13: 58. 1918. Type. Philippines. Ilocos: Luzon, Central Luzon, Pangasinan Province, Mount Umingan, Aug 1910, *M. Ramos & G.E. Edaño* [*Bur. Sci.*] 26487 (lectotype, designated here: K [K000195878]; isolectotype: US [acc # 1376054, US00027662]).

*Solanum luzoniense* Merr. var. *glabrum* Merr., Philipp. J. Sci. Bot. 13: 59. 1918. Type. Philippines. Ilocos: Luzon, Central Luzon, Pampanga Province, Calumpit, Sep 1905, *E.D. Merrill* 4237 (lectotype, designated here: K [K000195959]; isolectotype: US [acc # 710167, US00027663]).

**Type.** PHILIPPINES. CAR: Luzon, Benguet Province, Baguio, Mar 1907, *A.D.E. Elmer* 8719 (lectotype, designated here: K [K000195910]; isolectotypes: A [0077850], E [E00243613]).

**Description.** Shrubs or scrambling shrubs to 1.5 m, armed or unarmed. Stems erect, terete, prickly or unarmed, moderately stellate-pubescent; prickles if present 3–5 mm long, straight (occasionally somewhat slightly curved) and downwardly pointing (retorse), yellowish tan; pubescence of mixed sessile and short-stalked porrect-stellate trichomes, the stalks to 0.2 mm, the rays 6–8, ca. 0.5 mm long, the midpoints equal to the rays, the trichomes persistent or deciduous and the stems glabrate; new growth moderately to densely stellate-pubescent, the trichomes whitish cream, mixed sessile and
<figure><img src="image_78.png" /><figcaption>**Figure 63.** *Solanum retrorsum* Elmer – Herbarium specimen collected in the Philippines in 1914 (*Mer-rill 9665*, K000195908) Photograph credit: © copyright of the Board of Trustees of the Royal Botanic Gardens, Kew.</figcaption></figure>
short-stalked porrect-stellate like those of the stems; bark of older stems whitish grey. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, not lobed or occasionally very shallowly lobed, the blades 3–15 cm long, 1–6 cm wide, 2–3 times longer than wide, narrowly elliptic to lanceolate, widest at the middle, chartaceous, somewhat discolorous, usually unarmed; adaxial surface glabrous to moderately and evenly stellate-pubescent with mixed sessile and short-stalked porrect-stellate trichomes, the stalks to 0.2 mm, the rays 6–8, ca. 0.5 mm long, the midpoints equal to the rays; abaxial surface similarly but more densely pubescent with porrect-stellate trichomes, the lamina still visible; major veins 5–7 pairs, barely visible above, stellate pubescent especially abaxially; base attenuate, usually somewhat oblique; margins entire or rarely very shallowly lobed, the lobes 1–2 on each side, ca. 0.5 cm long, broadly deltate with rounded or acute tips, the sinuses reaching less than 1/8 of the distance to the midrib; apex acute to acuminate; petioles 0.6–2.5 cm long, ca. 1/4 as long as the leaf blades or less, unarmed and densely stellate-pubescent with mixed sessile and short-stalked porrect-stellate trichomes like those of the stems, pubescence denser than that of stems. Inflorescences 1.5–4.5 cm long, internodal and lateral, unbranched or very occasionally forked, with 10–30 flowers, only a few flowers open at any one time, sparsely to densely pubescent with porrect-stellate trichomes like those of the stems; peduncle 0.3–0.8 cm long, to 1.5 cm long on each of the inflorescence branches; pedicels 0.6–0.75 cm long, ca. 0.5 mm in diameter at the base, ca. 0.5 mm in diameter at the apex, spreading and perhaps slightly nodding at anthesis, stellate-pubescent with porrect-stellate trichomes like the inflorescence, articulated at the base; pedicel scars irregularly spaced 2–4 mm apart. Buds elongate and tapering, approximately halfway exerted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 1.5–2 mm long, conical, the lobes 2–2.5 mm long, ca. 2 mm wide, deltate to narrowly deltate, densely stellate-pubescent like the pedicels. Corolla 1–1.4 cm in diameter, white, stellate, lobed nearly to the base, the lobes 4–7 mm long, 3–4 mm wide, narrowly triangular, spreading at anthesis (perhaps slightly reflexed?), glabrous adaxially, moderately stellate-pubescent abaxially where exposed in bud. Stamens equal; filament tube minute, glabrous; free portion of the filaments ca. 0.5 mm long, glabrous; anthers 3–4.5 mm long, 1–1.2 mm wide, strongly tapering, yellow, poricidal at the tips, the pores directed distally, not elongating to slits with drying. Ovary conical, glabrous; style 6–8 mm long, glabrous or with a few stalked stellate trichomes near the base; stigma capitate, the surfaces minutely papillose. Fruit a globose berry, 0.6–0.8 cm in diameter, bright red when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 0.8–1.8 cm long, 0.7–1.1 mm in diameter at the base, 1.5–2 mm in diameter at the apex, somewhat woody, erect or somewhat spreading; fruiting calyx not accrescent, the lobes often breaking off. Seeds 6–10 per berry, 3.5–5 mm long, 1.5–3.5 mm wide, flattened reniform with incrassate margins, yellowish tan, the surfaces minutely pitted, the testal cells with sinuate margins. Chromosome number: not known.

**Distribution** (Fig. 64). *Solanum retrorsum* occurs from the very northern part of Sulawesi (a single collection) in Indonesia to the Philippines, where it is much more common. The single collection from Lanyu (Orchid) Island off the southeastern edge of Taiwan is a very poor specimen and is included here with some hesitation.
<figure><img src="image_79.png" /><figcaption>**Figure 64.** Distribution of *S. retrorsum*.</figcaption></figure>

**Ecology and habitat.** *Solanum retrorsum* grows in thickets, forests and forest edges in broadleaf evergreen and semideciduous woodlands; few of the collections we have seen have specific habitat information; elevations from 500 to 1,600 m elevation.

**Common names and uses.** Taiwan. lü song qie (Zhang et al. 1994, as *S. luzoniense*, but this identification is tentative).

**Preliminary conservation status (IUCN 2019).** Near Threatened (LC). EOO (115,321 km², LC); AOO (48 km², EN). *Solanum retrorsum* is found throughout the Philippines, with populations in a variety of habitats. The range extends to Sulawesi, making it widely distributed. The status of populations outside the main range in the Philippines, however, needs further investigation and the few recent collections suggest that the species merits some conservation concern.

**Discussion.** *Solanum retrorsum* is polymorphic for prickliness; some plants are densely prickly with the downward point prickles that give the species its name, while others lack prickles entirely. These unarmed plants have been called *S. luzoniense*. Despite this polymorphism *S. retrorsum* is distinctive, with its narrowly elliptic leaves and very small ripe berries, and not easily confused with any other species in the region.

The polymorphism in prickliness seen in *S. retrorsum* is also found in *S. ratale* D.McClelland of Fiji (see McClelland et al. 2020). That species also has unarmed and densely prickly individuals with strongly retrorse prickles mainly on the stems; the plurifoliate sympodial units, trichomes with midpoints equal to the rays and bright red berries (versus difoliate sympodia, small trichomes with elongate midpoints and black to deep purple berries in *S. ratale*), as well as their distinct distributions, clearly distinguish the two species.
The description of *S. luzoniense* in Zhang et al. (1994) was taken from a mixture of material from Taiwan and the Philippines. Material from Taiwan was a mixture of *S. miyakojimense* (Yang 8701) and a very scrappy specimen from Lanyu Island (*Leu et al.* 2118) that we here tentatively assign to *S. retrorsum*, but with considerable hesitation. The occurrence of *S. retrorsum* in Taiwan remains to be confirmed.

Merrill (1918b) did not specifically designate a type for his *S. luzoniense*. He cited “Bur. Sci. 24687 Ramos & Edaño”, “Bur. Sci. 17710 Otanes” and “Merrill s.n., July 1903”; we have selected the Kew duplicate of *Ramos & Edaño* [Bur. Sci.] 26487 (K000196878) as the lectotype because it is well preserved and has both flower and fruit. For var. *glabrum*, he clearly stated that his own collection (*Merrill* 4237) was the type but did not specify the herbarium. We have selected the Kew duplicate of this collection (K000195959) as the lectotype of the varietal name because it is the best-preserved specimen of all the duplicates we have seen.

**Specimens examined.** See Suppl. materials 1–3.

38. *Solanum robinsonii* Bonati, Bull. Soc. Bot. Genève, sér. 2, 3: 311. 1914.
Figs 2C, 65

**Type.** VIETNAM. Khánh Hòa: Nha-trang and vicinity, 11–26 Mar 1911, C.B. Robinson 1082 (lectotype, designated here: P [P00054128]; isolectotype: K [K000922030]).

**Description.** Erect shrub, up to 1 m tall, unarmed. Stems erect, terete, sparse-ly to moderately stellate-pubescent, glabrescent; pubescence of mixed sessile and short-stalked porrect-stellate trichomes, the stalks to 0.08 mm long, the rays 7–8, 0.1–0.2 mm long, the midpoints to 0.03 mm long with bulbous bases; new growth densely stellate-pubescent, light brownish green; bark of older stems brownish grey, glabrous. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, entire, the blades 3–8 cm long, 1–3 cm wide, ca. 2–3 times longer than wide, elliptic to ovate, chartaceous, strongly discolorous; adaxial surface moderately stellate pubescent, the trichomes porrect-stellate, sessile and short-stalked, the stalks extremely short, the rays 6–8, 0.05–0.15 mm long, the midpoints to 0.08 mm long, with bulbous bases; abaxial surface densely stellate-pubescent with sessile and short-stalked porrect-stellate trichomes, the stalks if present to 0.5 mm, the rays 5–10, 0.2–0.5 mm long, the mid-points to 0.2 mm, stout with a bulbous base, the lamina completely obscured; major veins 4–9 pairs, markedly impressed above, drying light green; base cuneate; margins entire or somewhat undulate, never lobed; apex rounded to acute; petiole 0.5–1 cm long, 1/10–1/5 of the leaf blade length, densely stellate-pubescent with trichomes like those of the stems and leaves. Inflorescences 1–3 cm long, internodal and lateral, un-branched, with ca. 5–10 flowers, only a few flowers open at any one time, densely stellate pubescent with porrect-stellate trichomes like those of the stems; peduncle 0.4–2.5 cm long; pedicels 4–9 mm long, 0.7–1 mm in diameter at the base, 1.3–1.5 mm in diameter at the apex, erect to recurved, densely stellate-pubescent with porrect stellate-trichomes like those of the axes, articulated at the base; pedicel scars spaced 0.5–2 mm apart. Buds elongate-ovoid, pointed at the tip, the corolla ca. halfway exserted from
<figure><img src="image_80.png" /><figcaption>**Figure 65.** *Solanum robinsonii* Bonati **A** herbarium specimen (holotype) collected in Vietnam in 1911 (Robinson 1082, P00054128) **B** habit (Nuraliev 3031, Vietnam) **C** detail of the abaxial surface of a leaf blade (Nuraliev 3031, Vietnam) **D** inflorescence and detail view of an open flower (Nuraliev 3031, Vietnam) **E** detail view of a fruit (Nuraliev 3031, Vietnam). Photograph credits: **A** CC-BY, Muséum national d'Histoire naturelle, Paris **B–E** M. Nuraliev.</figcaption></figure>

the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 1.4–2 mm long, campanulate, the lobes 2–3 mm long, 0.6–0.8 mm wide, narrowly deltate and narrowing to an elongate acumen, the acumen 2/3–3/4 the total lobe length, densely stellate-pubescent with porrect stellate-trichomes like those of the
pedicels. Corolla 1–2 cm in diameter, blue to purple, stellate, lobed ca. 1/2–2/3 of the way to the base, the lobes 4–5 mm long, 2–2.5 mm wide, deltate, spreading at anthesis, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal or very slightly unequal with 2 longer than the rest; anthers 5–6 mm long, 0.5–0.75 mm wide, if unequal then 2 ca. 0.1 mm longer than the rest, tapering, connivent, yellow, glabrous, poricidal at the tips, the pores not lengthening to slits with age; filament tube <0.5 mm long; free portion of the filaments 0.4–0.75 mm, glabrous. Ovary conical, with minute glandular hairs at the top; style 7.5–9 mm long, slender, curved at the apex, glabrous; stigma capitate, minutely papillate. Fruit a globose berry, 1–2 per infructescence, 0.7–1.2 cm in diameter, the pericarp smooth, red when mature, glabrous; fruiting pedicels 1.2–1.6 cm long, ca. 0.5 mm in diameter at the base, 1.5–1.8 mm in diameter at the apex, woody, erect to recurved; fruiting calyx lobes slightly expanding to 4–5 mm long, 1/2–2/3 the length of the mature fruit, narrowly deltate, appressed to the berry, ending with a long acumen. Seeds ca. 10–15 per berry, 3–4 mm long, 2.5–3 mm wide, flattened-reniform, dull yellow, the surface minutely pitted, the testal cells pentagonal in outline. Chromosome number: not known.

**Distribution (Fig. 66).** *Solanum robinsonii* is endemic to Vietnam; the few known collections are restricted to Khánh Hòa and Ninh Thuân provinces of southeast Vietnam.

<figure><img src="image_81.png" /><figcaption>**Figure 66.** Distribution of *S. robinsonii*.</figcaption></figure>
**Ecology and habitat.** *Solanum robinsonii* grows on hillsides in deciduous forests, at ca. 200 m elevation.

**Common names and uses.** None recorded.

**Preliminary conservation status (IUCN 2019).** Endangered (EN [B1,2abii,iii]). EOO (610 km², EN); AOO (20 km², EN). *Solanum robinsonii* has a narrow distribution and grows in coastal forests that are subject to much anthropogenic disturbance. The species is of concern due to extreme habitat alteration (Wikramanayake et al. 2002).

**Discussion.** *Solanum robinsonii* is superficially similar to *S. putii* of Thailand and *S. nienkui* of Hainan Island, China. The three taxa share a spindly shrubby habit, long, thin anthers and small red berries. *Solanum robinsonii* can be distinguished from *S. putii* in its more narrowly elliptic leaves that are more strongly discolorous when dry, and from *S. nienkui* in its slightly larger corollas (to 2 cm in diameter versus to 1.3 cm in diameter) that are less deeply stellate, and in its longer style (7.5–9 mm long versus 5.5–6 mm long).

The strongly discolorous leaves of *S. robinsonii* are reminiscent to those of *S. gигаnteum*, but that species has highly branched inflorescences and stout, prickly stems. Specimens of *S. robinsonii* have been considered to look like *S. elaeagnifolium* (annotation by D.E. Symon on the isolectotype specimen at Kew); this is due to the bulbous bases of the midpoints that make the trichomes look slightly lepidote. *Solanum elaeagnifolium*, however, has true lepidote trichomes that are more shield-like in structure (see Knapp et al. 2017).

Hul and Dy Phon (2014) erroneously cited the specimen in P as the holotype of *S. robinsonii*. Although Bonati worked in Paris, he did not cite any herbarium in the protologue or in the introductory part of his treatment (Bonati 1914), so lectotypification is necessary. We here select the Paris sheet (P00054128) annotated by Hul and Dy Phon as “holotype” as the lectotype for *S. robinsonii*.

**Specimens examined.** See Suppl. materials 1–3.

**39. Solanum robustum H.L.Wendl., Flora 27: 784. 1844.**

Fig. 41C–E

No names based on tropical Asian types have been published; for complete synonymy see Vorontsova and Knapp 2016.

**Type.** Cultivated in Hanover, Germany, from seeds sent from Brazil, *Anonymous s.n.* (no herbarium specimens cited, none found).

**Description.** Vorontsova and Knapp (2016: 280–284); http://www.solanaceae-source.org/solanaceae/solanum-robustum.

**Distribution.** *Solanum robustum* has been collected in India (state of Tamil Nadu) and on the island of Réunion (two collections from the 1960s); it is native to southern South America. Outside of its native range it is apparently used as an ornamental (see Vorontsova and Knapp 2016).
**Discussion.** *Solanum robustum* is a distinctive species with strongly winged stems and small, pubescent berries. Even vegetatively it is not easily confused with any other species in the region. It has the potential to become a serious pest; in Hawai'i it is clas- sified as a noxious weed (Wagner et al. 1999).

*Solanum robustum* was described without reference to herbaria or specific collections; we, like Vorontsova and Knapp (2016), leave neotypification of this name for a mon- ographer of the group to which this distinctive species belongs (Erythrotrichum clade).

**Specimens examined.** See Suppl. materials 1–3.

**40. *Solanum schefferi* F.Muell., Descr. Notes Papuan Pl. 1: 44. 1876.**
Fig. 67

*Solanum incanum* Scheff., Ann. Jard. Bot. Buitenzorg 1: 39. 1876, nom. illeg., non *Solanum incanum* L., 1753. Type. Indonesia. West Papua: Andai (“Andaj”), sin. dat., *G. Teijsmann s.n.* (lectotype, designated here: BO [acc. # 1324395]; isolectotype: BO [acc. # 1323468]).

*Solanum lianoides* Elmer, Leaflets Philipp. Bot. 2: 733. 1910. Type. Philippines. MIMAROPA: Luzon, Romblon Province, Sibuyan Island, May 1910, A.D.E. Elmer 10752 (lectotype, designated here: BM [BM000778113]; isolectotypes: A [00077845], BISH [BISH1005079], CAL [acc. # 316476], E [E00273864], F [v0073460F, acc. # 290900], G [G00442946], GH [GH00077844], HBG [HBG511454], K [K000196018], LE [LE00016968, LE00016969], MO [MO-503846, acc. # 3783069], NY [00172284], US [00027651, acc. # 779390]).

*Solanum smilacocladum* Bitter, Bot. Jahrb. Syst. 55: 79. 1919. Type. Indonesia. Sulawesi: [Sulawesi Utara] “Celebes (Minahassa): Bojong, N. Celebes”, 1888, *O. Warburg 15072* (lectotype, designated here: NY [00804057]).

**Type.** Based on (replacement name for) *Solanum incanum* Scheff., non L.

**Description.** Scrambling shrubs or woody lianas to 10 m long, armed. Stems flexuous, terete, moderately prickly and sparsely to densely stellate-pubescent; prickles 2–4 mm long, 2–2.5 mm wide at the base, strongly recurved (hooked), pale yellowish tan; pubescence of stalked porrect-stellate trichomes, the stalks to 0.5 mm long, multiseriate, the rays 4–8, 0.2–0.4 mm long, the midpoints absent or slightly longer than the rays; new growth moderately to densely stellate-pubescent, the trichomes sessile to short-stalked, the stalks 0.2–0.4 mm, the rays 4–8, 0.2–0.5 mm long, the midpoints ca. 0.4 mm long, equal to the rays; bark of older stems reddish brown and somewhat shiny. Sympodial units unifoliate, the leaves not geminate. Leaves simple, not lobed, the blades 2.5–15 cm long, 1.5–7.5 cm wide, ca. 2 times longer than wide, elliptic-ovate to ovate, widest in the middle or the lower third, chartaceous, somewhat discolorous, armed or unarmed; adaxial surface evenly and sparsely to densely pubescent with mixed sessile and short-stalked porrect-stellate trichomes, the multiseriate stalks to 0.2 mm long and conical, the rays 4–6, 0.1–0.2 mm long, sometimes ascending
<figure><img src="image_82.png" /><figcaption>**Figure 67.** *Solanum schefferi* F.Muell. Herbarium specimen collected the Philippines in 1927 (*Ramos & Edaño 49324*, P00379535). Photograph credit: CC-BY, Muséum national d'Histoire naturelle, Paris.</figcaption></figure>
(pointing upwards from the stalk), the midpoints absent or very short; abaxial surface moderately to densely pubescent with mixed stalked and sessile porrect-stellate trichomes like the adaxial surface, but the pubescence denser, especially along the principal veins; major veins 4–6 pairs, impressed above, densely pubescent especially abaxially; base acute, usually strongly oblique; margins entire; apex acute; petioles 1–1.2 cm long, 1/10–1/5 as long as the leaf blades, densely pubescent with porrect-stellate trichomes like those of the stems, unarmed or armed with 0–5 hooked prickles. Inflorescences 3–10 cm long, internodal and lateral, unbranched or more commonly many times branched, with (2–)10–40 flowers, only a few flowers open at any one time, pubescent with porrect-stellate trichomes like those of the stems, occasionally with a few minute hooked prickles in larger inflorescences; peduncle 0.5–3 cm long; pedicels 0.8–1 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading, sparsely to moderately stellate-pubescent with porrect-stellate trichomes like the inflorescence axes, articulated above the base leaving a small peg on the inflorescence axis, the pegs 0.5–1 mm long, more glabrous than either the pedicels or the inflorescence axis, drying dark brown to black; pedicel scars irregularly spaced 1–3 mm apart, often raised as short pegs ca. 1 mm long. Buds stout and strongly tapering, strongly exserted from the calyx before anthesis. Flowers 5-merous, heterostylous with the distal flowers on each inflorescence branch short-styled and the plants weakly andromonoecious. Calyx with the tube 2.5–3 mm long, conical, in bud a hyaline cup with darker acumens from the rim, tearing irregularly at anthesis, the lobes 1–2 mm long including the acumens, ca. 1 mm wide, deltate with a subulate tip to 1 mm long, sparsely to moderately stellate-pubescent with porrect-stellate trichomes like those of the pedicels, unarmed. Corolla 2–3 cm in diameter, violet to dark purple, deeply stellate, lobed nearly to the base, interpretalar tissue scarce, but the petal margins thin and somewhat “ruffly”, the lobes 10–14 mm long, 3.5–5 mm wide, spreading at anthesis, mostly glabrous adaxially but in more pubescent individuals with porrect-stellate trichomes on the petal surface, densely stellate-pubescent abaxially with densely tangled stalked porrect-stellate trichomes where exposed in bud, the trichomes with 5–8 rays ca. 0.5 mm long and midpoints equal to the rays, sometimes tinged with purple, the petal margins and tips densely papillate, the tips cucullate. Stamens equal or very slightly unequal; anthers 6–8 mm long, ca. 1.5 mm wide in the lower third, strongly tapering, yellow, 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, sparsely pubescent with simple glandular trichomes and papillae; style in long-styled flowers 9–10 mm long, strongly hooked at the tip, minutely glandular papillate in the lower 1/4, otherwise glabrous, in short-styled flowers ca. 1 mm long, straight, glabrous; stigma slightly lobed, borne on the inner surface of the hooked style tip, the surfaces minutely papillose. Fruit a globose or ovoid berry, 2–3 cm long, ca. 3 cm in diameter (fide Symon 1985: 145; 3–5 cm long fide Elmer 1910: 734), green, the pericarp leathery and shiny, glabrous; fruiting pedicels 1.8–3 cm long, ca. 1 mm in diameter at the base, tapering to an apex ca. 3 mm in diameter, woody, spreading or pendent from weight of berries; fruiting
<figure><img src="image_83.png" /><figcaption>**Figure 68.** Distribution of *S. schefferi*.</figcaption></figure>

calyx slightly accrescent, the lobes ca. 5 mm long, ca. 2 mm wide, spreading. Seeds more than 100 per berry, ca. 2 mm long, ca. 1.5 mm wide, reddish brown or tan, the surfaces very minutely pitted or smooth, the testal cells not visible. Chromosome number: not known.

**Distribution (Fig. 68).** *Solanum schefferi* occurs from the Philippines to northern New Guinea and the Solomon Islands (Malaita Island) to northern Sulawesi and the Malakus in Indonesia.

**Ecology and habitat.** *Solanum schefferi* has been collected growing in open areas near abandoned fields or at forest edges from approximately sea level to 1,500 m elevation. It appears to grow on limestone, a soil type unusual for *Solanum*.

**Common names and uses.** None recorded.

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (1,012,899 km², LC); AOO (32 km², EN). In the area of tropical Asia treated here *S. schefferi* is at the northwestern part of its range; it is more widely distributed on New Guinea. The small AOO is common in vines of forest canopies, these types of plants are notoriously under-collected but are also often rare where they occur.

**Discussion.** *Solanum schefferi* is a species with most of its distribution on the island of New Guinea, but several collections are from the Malaku islands and northern Sulawesi. Symon (1985) suggested this distribution and its distinct differences from other New Guinea species suggested it was an introduction from the Americas. Molecular phylogenetic study, however, shows that *S. schefferi* is clearly a member of the Eastern Hemisphere clade of spiny solanums (Aubriot et al. 2016a) and is member of a small monophyletic group containing *S. athenae* Symon and *S. leptacanthum* Merr. & L.M.Perry, both New Guinea endemics (but see below).
*Solanum schefferi* is a striking species with large, showy (fide *Brass 24330*) flowers and large fruits. The plants climb with their hooked prickles (like rose thorns) and unlike many other species of New Guinean spiny solanums, the prickles on all parts are hooked. The berries are ovoid when young and grow to a considerable size (fide Elmer 1910: 734; Symon 1985: 145); smaller berries appear to be globose, elongating to ovoid with maturity.

*Solanum lianoides* has not previously been considered a synonym of *S. schefferi* (Symon 1985), but examination of the pubescence and flowers leads us to consider them as be-longing to the same taxon. Molecular results of Aubriot et al. (2016a) place *S. lianoides* in the larger Sahul-Pacific clade and forming a group with *S. dunalianum* and *S. gracili-florum*, not closely related to *S. schefferi*. These results remain to be revisited to ascertain if contamination of one or other of these samples influenced the results. It is clear, however, that *S. schefferi* is native to tropical Asia, and is not an introduction from the Americas.

The protologue of *S. lianoides* cites a single collection (Elmer 10752) but does not mention a herbarium (Elmer 1910). We have selected the only one of the many dupli-cates of this collection we have seen that has both flowers and fruits (BM000778113) as the lectotype for this name.

In coining the replacement name *S. smilacocladum* Bitter (1919) cited the same sheets in BO as those used by Scheffer in his description of “S. incanum” (“Herb. Buitenzorg 9853! Sub nom. *S. incanum* Scheff.”), as well as additional syntypes from Berlin (*Schlechter 17627*, *Warburg 15072*, *Warburg 21246*) and Bogor (*Koorders 18033B*, *Koorders 18046B*). We have lectotypified *S. smilacocladum* with *Warburg 15072* from northern Sulawesi (NY 00804057), to not make the name homotypic with *S. schefferi*.

**Specimens examined.** See Suppl. materials 1–3.

41. *Solanum sisymbriifolium* Lam., Tabl. Encycl. 2: 25. 1794, as "*sisymbrifolium*".
Fig. 41F–H

No names based on tropical Asian types have been published; for complete synonymy
see Vorontsova and Knapp 2016.

**Type.** ARGENTINA. Buenos Aires: Buenos Aires, *P. Commerson s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 307: P-LA [P00357630, lower plant fragment, Morton neg. 8391]); isolectotypes: P [P00371604, P00371605, P00371606].

**Description.** Vorontsova and Knapp (2016: 307–312); http://www.solanaceae-source.org/solanaceae/solanum-sisymbriifolium.

**Distribution.** *Solanum sisymbriifolium* has been collected in tropical Asia in Bangladesh, China, and throughout India (see Saha and Datta 2013); it is native to South America but is widely adventive and somewhat invasive, so it is to be expected throughout the region in highly disturbed areas.

**Common names.** China. suan jie qie (Zhang et al. 1994). India. mulathurivan [Mikir] (Jain and Borthakur 1986).
**Discussion.** *Solanum sisymbriifolium* is the only spiny solanum species in tropical Asia with deeply pinnatifid to bipinnatifid leaves coupled with accrescent calyces. *Solanum multiflorum* of the Western Ghats sometimes has deeply pinnatifid leaves, but the calyces are not accrescent and it is a shrub with much smaller flowers (1.3–1.5 cm versus 2–3 cm in diameter). The prickly, accrescent calyx lobes turn back at fruit maturity to reveal the sticky red berries (often cultivated as fruit and called vila-vila in other parts of the world). Some specimens from the Americas have less deeply divided leaves, but all those we have seen from tropical Asia are deeply pinnatifid. Saha and Datta (2013) record it as very common in Tripura.

*Solanum sisymbriifolium* was introduced very early to European botanical gardens, from where it perhaps was introduced as colonisation expanded. It is a very weedy species, even in its native range; overgrazed pastures can become overgrown with *S. sisymbriifolium* very quickly. In some parts of Europe, it is planted in fallow potato fields as a trap crop for cyst nematodes; the nematodes lay eggs in the roots of *S. sisymbriifolium*, but the plants are destroyed before the eggs hatch (Timmermans et al. 2007 a, b).

**Specimens examined.** See Suppl. materials 1–3.

**42. Solanum sulawesi X.Aubriot & S.Knapp, sp. nov.**

urn:lsid:ipni.org:names:77298805-1
Fig. 69

**Diagnosis.** Like *Solanum involucratum* Blume, but differing in its lack of dense stellate pubescence on leaf blades and new growth, attenuate leaf bases, and fruiting calyces that are not accrescent.

**Type.** INDONESIA. Sulawesi: Sulawesi Utara, 220 km W. of Manado, 50 km inland from Pangi, on tributary of Sungai Ilanga, 350–750 m, 3 Mar 1990, *J.S. Burley et al.* 3625 (holotype: L [L.2882218]; isotypes: A, BO [n.v.], K [K000014635]).

**Description.** Shrubs to 1.5 m tall, strongly armed. Stems erect, terete, densely prickly, glabrous, purple (fide *Burley et al.* 3625); prickles 0.3–1 cm long, 0.1–0.25 cm in diameter at the base, straight, of many different sizes, pale yellowish tan; pubescence absent; new growth densely prickly and moderately stellate-pubescent with short-stalked porrect-stellate trichomes, the stalks ca. 0.2 mm long, the rays 4–6, ca. 0.5 mm long, the midpoints ca. 0.5 mm long, equal to the rays, the trichomes soon deciduous and the stems glabrate; bark of older stems pale tan. Sympodial units difoliate, the leaves not geminate. Leaves simple, shallowly lobed, the blades (5–)9–17 cm long, (4–)7.5–12 cm wide, 1–1.5 times longer than wide, broadly elliptic, chartaceous, concolorous, strongly armed along the midrib and major veins with straight prickles; adaxial surface glabrate, sparsely pubescent along the veins with scattered sessile porrect-stellate trichomes, the rays 4, 0.1–0.2 mm long, the midpoints 1–1.5 mm long, much longer than the rays; abaxial surface sparsely pubescent with multangulate stalked trichomes, the stalks to 0.5 mm long, the rays 8–10, to 0.5 mm long, irregular in size on a single trichome; major veins 5–6 pairs, densely prickly on both surfaces,
<figure><img src="image_84.png" /><figcaption>**Figure 69.** *Solanum sulawesi* X.Aubriot & S.Knapp - Herbarium specimen (holotype) collected in Indonesia in 1990 (Burley et al. 3625, L.2882218). Photograph credit: Naturalis Biodiversity Center.</figcaption></figure>
the prickles ca. 40 per face, 0.3-1 cm long, straight; base attenuate; margins shallowly lobed, the lobes 5-6 on each side, 1-3 cm long, deltate, acute at the tips and occasionally somewhat bilobed, the sinuses ca. halfway to the midrib; apex acute to acuminate; petioles 2-4 cm long, 1/4-1/3 as long as the leaf blades, densely prickly and sparsely pubescent, the prickles like those of the stems and leaves, the pubescence of scattered stalked porrect-stellate-trichomes, the stalks to 0.5 mm, the rays 6-8, ca. 0.3 mm, the midpoints equalling the rays in length, soon deciduous and the petioles of older leaves glabrate. Inflorescences 0.5-3 cm long, internodal and lateral, unbranched, with 4-8 flowers, only 1 or 2 flowers open at any one time, glabrous except at the very tips, densely prickly, the prickles 2-3 mm long, straight; peduncle absent to 0.9 cm long, densely prickly; pedicels 0.5-0.6 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading and slightly nodding at anthesis, densely prickly and sparsely stellate-pubescent like the leaf adaxial surfaces, articulated at the base, the trichomes porrect-stellate, the rays 4-8, 0.2-0.5 mm long, midpoints not clearly differentiated from the rays; pedicel scars tightly packed and almost overlapping. Buds broadly ellipsoid and somewhat tapering, included in the calyx lobes until just 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, cup-shaped to slightly urceolate, abruptly narrowing to the pedicel and densely prickly, the lobes 4-6 mm long, ca. 2 mm wide, narrowly deltate, densely prickly on the midveins and sparsely stellate-pubescent abaxially with porrect-stellate trichomes like those of the pedicels. Corolla 1.4-1.8 cm in diameter, white, stellate, lobed 2/3 of the way to the base, interpetalar tissue minimal, the lobes 5-6 mm long, 2.5-3.5 mm wide, narrowly deltate, spreading at anthesis, mostly glabrous adaxially but with a few stellate trichomes at the tips, sparsely stellate-pubescent abaxially with densely tangled sessile trichomes where exposed in bud, these densest at the tips, a few tiny prickles along the petal midvein. Stamens equal; anthers 5-5.5 mm long, ca. 1.5 mm wide, tapering, yellow, densely papillate in the lower half, 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, densely pubescent with sessile porrect-stellate trichomes with tiny rays and elongate midpoints; style 6-6.5 mm long, glabrous; stigma strongly bilobed, the surfaces minutely papillose. Fruit a globose berry, several per infructescence, 1.5-2 cm in diameter, green (immature?), the pericarp thin and slightly shiny, evenly pubescent with porrect-stellate trichomes with 4-6 minute rays less than 0.1 mm long, the midpoints 3+-celled 1-2.5(-3) mm long; fruiting pedicels 0.8-1.1 cm long, ca. 2 mm in diameter at the base, abruptly enlarged at the apex to ca. 4 mm in diameter, woody, spreading to more or less deflexed from the weight of the berry; fruiting calyx accrescent, the lobes expanding to 7-8 mm long, reflexed and often breaking off. Seeds 50-100 per berry, ca. 2.5 mm long, ca. 2 mm wide, flattened reniform, yellowish tan or brown, the surfaces minutely pitted, the testal cells with straight or slightly sinuate margins. Chromosome number: not known.

**Etymology.** *Solanum sulawesi* is named for the island on which it appears to be endemic, Sulawesi, which is of an extraordinary composite geologic nature at the collision zone of the Asian, Pacific, and Australian tectonic plates (Villeneuve et al. 2002).
<figure><img src="image_85.png" /><figcaption>**Figure 70.** Distribution of *S. sulawesi*.</figcaption></figure>

**Distribution (Fig. 70).** *Solanum sulawesi* is endemic to the Indonesian island of Sulawesi (previously known as Celebes under Portuguese colonial occupation); two collections are from the northern peninsula and the third from the southwestern peninsula of the island.

**Ecology and habitat.** *Solanum sulawesi* has been collected in open areas in lowland tropical rainforest, from 350 to 750 elevation.

**Common names and uses.** None recorded.

**Preliminary conservation status (IUCN 2019).** Data Deficient (DD). The few collections, coupled with the possibility of more being uncovered as more specimens of *S. involucratum* are examined carefully, suggests that it is too early to assign a preliminary conservation status for this species.

**Discussion.** *Solanum sulawesi* is morphologically similar to both *S. involucratum* and *S. lasiocarpum*, both of which have large, repand leaves and pubescent berries. It differs from *S. involucratum* in its calyx that is not accrescent in fruit, its glabrous stems and extremely sparsely pubescent leaves that are cuneate to attenuate and widest near the middle (versus truncate and widest near the base). *Solanum lasiocarpum* is a much more densely pubescent plant and is often completely unarmed. The leaves are more ovate in outline and more densely pubescent. The midpoints of the fruit trichomes in *S. sulawesi* are unusual in being 2–3-celled; those of both *S. lasiocarpum* and *S. involucratum* are single-celled. Fruits of *S. sulawesi* are smaller than those of *S. lasiocarpum*, and less densely pubescent.

We would have preferred to place the holotype for this species in a herbarium in Indonesia, but extensive searches in the herbarium at Bogor (BO; A. Kartonegoro, pers. comm.) failed to locate a duplicate of *Burley et al. 3625*, although one should be located there.
Material of *S. sulawesi* has not been included in molecular analyses but we suspect it will prove a member of a clade comprised of southeast Asian species and not a member of the largely American Lasiocarpa clade.

We have seen very few collections of *S. sulawesi*, but specimens of this species may have been previously identified as *S. involucratum* or *S. lasiocarpum*. These and unidentified collections in BO and other Indonesian herbaria are of particular interest and are a priority for examination.

**Paratypes.** [also see Suppl. materials 1–3] **INDONESIA. Sulawesi:** Sulawesi Tenggara, Southeast Celebes. Ladongi - Tirawuta - Kolaka [Kolaka Timur], 19 Oct 1978, *Prawiroatmodjo & Maskuri 1356* (K, L); Sulawesi Utara, “Ins. Celebes: (Minahassa): N Celebes”, 1888, *Warburg 15844* (NY).

**43. Solanum torvoideum Merr. & L.M.Perry, J. Arnold Arb. 30: 47. 1949.**
Fig. 71

**Type.** PAPUA NEW GUINEA. Central: Mafulu, Sep-Nov 1933, *L.J. Brass* 5411 (lectotype designated by Symon 1985, pg. 150, as “holotype”: A [00077838]; isolectotypes: BM [BM000886267], BRI [BRI-AQ0080383], L [L.0003661], NY [00172294]).

**Description.** Shrubs to small trees, 1–5 m tall, unarmed or rarely sparsely armed. Stems erect, terete, usually unarmed or with a few broad-based straight prickles, stellate-pubescent; prickles, if present, 2–3 mm long, to 1.5 mm wide at the base, very sparse, straight, pale yellowish tan; pubescence of sessile to very short-stalked porrect-stellate trichomes, the stalks to 1 mm long, the rays 6–8, ca. 0.5 mm long, the midpoints absent or to 1 mm long (New Guinea only); new growth densely stellate-pubescent, the trichomes tangled, somewhat golden in dry material; bark of older stems grey to dark brown, glabrescent. Sympodial units difoliate, the leaves geminate, the leaves of pair equal in size and shape or one leaf slightly smaller. Leaves simple, shallowly lobed, the blades (7–)12–25 cm long, (2–)3–11 cm wide, 2.5–4 times longer than wide, elliptic to narrowly elliptic, chartaceous, discolorous, unarmed or sparsely armed along the midrib and major veins with small straight prickles; adaxial surface dark green, evenly and moderately to densely pubescent with golden short-stalked porrect-stellate trichomes, the stalks to 0.5 mm long, the rays 4–6, ca. 0.5 mm long, the midpoints absent or to 0.5 mm (in some New Guinea populations the midpoints glandular); abaxial surface with similar short-stalked porrect-stellate trichomes, but with more numerous rays (up to 10–12); major veins 4–7 pairs, densely pubescent especially abaxially; base acute, somewhat oblique; margins shallowly lobed, the lobes 4–7, 0.5–3 cm long, deltate to triangular, apically acute, the sinuses ca. one third of the way to the midrib; apex acute; petioles 1.5–3(–4+) cm long, ca. 1/4 as long as the leaf blades, sparsely prickly and densely to moderately pubescent with short-stalked stellate-trichomes like those of the stems, unarmed or prickly with 1–3 prickles like those of the stems. Inflorescences 4–9 cm long, internodal and lateral, forked to several times branched, with 50–70+ flowers, several flowers open at any one time,
<figure><img src="image_86.png" /><figcaption>**Figure 71.** *Solanum torvoideum* Merr. & L.M.Perry **A** herbarium specimen collected in the Philippines in 1954 (*Frake 20323*, L.2771110) **B** habit (field photograph, unvouchered, Philippines) **C** detail of the inflorescence (field photograph, unvouchered, Philippines). Photograph credits: **A** Naturalis Biodiversity Center **B**, **C** P.B. Pelser.</figcaption></figure>

densely pubescent with short-stalked stellate-porrect trichomes like those of the stems, with 6–8 rays ca. 0.5 mm long and midpoints to 1 mm long, unarmed; peduncle absent and the first inflorescence branches appearing to arise directly from the stem, or 0.5 cm long, unarmed; pedicels 1–1.2 cm long, ca. 1 mm in diameter at the base and apex, spreading to erect at anthesis, densely stellate-pubescent like the inflorescence axes, articulated at the base, unarmed; pedicel scars irregularly spaced 1–2 mm apart. Buds elongate ellipsoid and somewhat tapering, strongly exerted from the calyx before anthesis. Flowers 5-merous, apparently all perfect, but some distal flowers may be short-styled. Calyx with the tube 2–3 mm long, conical, the lobes 2–5 mm long, 1–2 mm wide, deltate with a subulate acumen 1.5–2 mm long, densely stellate-pubescent with porrect-stellate trichomes like those of the pedicels, the acumen sometimes more
glabrous and drying dark. Corolla 2-2.6 cm in diameter, white, shallowly stellate, lobed ca. halfway to the base, interpretalar tissue present and abundant, the lobes 8-10 mm long, 4-7 mm wide, spreading at anthesis, mostly glabrous adaxially but with a few stellate trichomes at the tips, densely stellate-pubescent abaxially with densely tanged sessile trichomes where exposed in bud, these densest at the tips, the interpretalar tissue glabrous. Stamens equal; anthers 5-6.5 mm long, ca. 1 mm wide, connivent or somewhat spreading, 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 0.5-1 mm long, glabrous. Ovary conical, glabrous; style 6-9 mm long, glabrous; stigma capitate, the surfaces minutely papillose. Fruit a globose berry, several to many per infructescence, 1-1.4 cm in diameter, the pericarp thin and shiny, yellow to orange-yellow when ripe, aging to brown or almost black (fide Symon 1985), glabrous; fruiting pedicels 1.5-2 cm long, ca. 1.5 mm in diameter at the base, 3-3.5 mm in diameter at the apex, woody, erect or somewhat spreading from weight of berries; fruiting calyx not accrescent, the lobes often breaking off. Seeds 60-100 per berry, 2-3.5 mm long, ca. 2 mm wide, flattened reniform, pale tan or yellowish brown, the surfaces minutely pitted, the testal cells with sinuate margins. Chromosome number: n = 12 (Symon 1985; voucher *Symon 10669* from Papua New Guinea).

**Distribution (Fig. 72).** *Solanum torvoideum* occurs from the Sunda Islands in Indonesia and Timor Leste east to the Philippines and the island of New Guinea.

**Ecology and habitat.** *Solanum torvoideum* is a species of disturbed areas such as old garden sites, treefalls or roadsides; from 300 to 2,000 m elevation (higher elevations from New Guinea specimens). Symon (1985) records it as occurring in fagaceous forest and on gravelly stream beds on New Guinea.

<figure><img src="image_87.png" /><figcaption>**Figure 72.** Distribution of *S. torvoideum*.</figcaption></figure>
**Common names and uses.** Philippines. Mindanao: bokul [Bukid] (Sulit 9868), tan-gutaungan (Zwickey 52), gebubruca (Frake 38012); Luzon: tanutong (Maliwanang 229).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (2,372,285 km², LC); AOO (212 km², EN). As a specialist of disturbed areas with a wide distribution when the island of New Guinea is taken into account, *S. torvoideum* is not of conservation concern.

**Discussion.** *Solanum torvoideum* is a member of the Torva clade (see Aubriot et al. 2016a) and like others in the group has difoliate sympodia with usually geminate leaves, highly branched inflorescences, straight or only slightly curved stem prickles, and sticky leathery berries. The Asian species of “torvoids” differ from their American relatives in having red, rather than green berries at maturity. In the area treated here *S. torvoideum* is most similar to the introduced and widespread *S. torvum*, from which it differs in its denser, more reddish gold pubescence, eglandular inflorescences, and red berries. In general, plants of *S. torvoideum* are more robust-looking than those of *S. torvum*. *Solanum torvoideum* is superficially similar to another introduced species, *S. chrysotrichum* in its pubescence, but is not sympatric with it, and can be distinguished by fruit colour. It can be distinguished from *S. poka* and *S. pseudosapona-ceum* in its more branched inflorescences with denser pubescence and larger flowers (to 2.6 cm in diameter versus to 2 cm or 1.5 cm in diameter).

On the island of New Guinea, *S. torvoideum* is most similar to the sympatric *S. dammerianum* Lauterb. & Schum. (Symon 1985). *Solanum torvoideum* differs from *S. dammerianum* in is rusty (versus drab) leaf pubescence, its more compact and fewer branched inflorescence and its larger mature berries (to 2 cm in diameter versus to 1.5 cm in diameter). The type of *S. dammerianum* has been lost (Symon 1985) so further intensive study of New Guinea populations may reveal additional differences.

The Torva clade species occurring in tropical Asia are all each other's closest relatives, suggesting a single dispersal event from the Americas (Aubriot et al. 2016a).

**Specimens examined.** See Suppl. materials 1–3.

**44. Solanum torvum Sw., Prodr. [O.P. Swartz] 47. 1788.**

Figs 3G, 73A, B

Names based on tropical Asian types only; for complete synonymy see Vorontsova and Knapp 2016.

*Solanum macaoense* Dunal, Prodr. [A.P. de Candolle] 13(1): 264. 1852. Type. China. Guangdong: “Macao”, *S. Callery 28/29* (holotype: P [P00055495]).

**Type.** “Indiae occidentalis” [“Provenit in sepibus Jamaicae, Hispaniolae, Insulis Bermudensibus” Swartz 1797: 456], *O. Swartz s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 351: S [S-R-5814]).

**Description.** Vorontsova and Knapp (2016: 351–355); http://www.solanaceae-source.org/solanaceae/solanum-torvum-1.
<figure><img src="image_88.png" /><figcaption>**Figure 73.** Introduced species of *Solanum*. *Solanum torvum* Sw. **A** habit (Meeboonya et al. RM 267, Thailand) **B** detail view of an inflorescence with flowers and fruits (Meeboonya et al. RM 267, Thailand). *Solanum viarum* Dunal **C** habit (Sampath Kumar et al. 126976, India) **D** detail view of a flower (Sampath Kumar et al. 126923, India) **E** detail view of a fruit (Sampath Kumar et al. 126923, India). *Solanum wrightii* Benth. **F** habit (Meeboonya et al. RM 271, Thailand) **G** detail view of flowers (Meeboonya et al. RM 271, Thailand) **H** detail view of fruits (Meeboonya et al. RM 271, Thailand). Photograph credits: X. Aubriot.</figcaption></figure>
**Distribution.** *Solanum torvum* is naturalised throughout tropical Asia; it is native to Central America and the Caribbean but has achieved a worldwide distribution in tropical and subtropical areas.

**Common names.** Bangladesh. tith begum (*Rahman's collector* 1124). Brunei Darussalam. terong pipit (*Norul Rozimah Pg Hj Seruji 18-b-3000*); tarong cit, tarong lowow (*Yati* 412). Cambodia. trâp put num nhong, trâp rom nhong (Hul and Dy Phon 2014). China. shui qie (Zhang et al. 1994). India. Assam: hati bhekur (*Marriot* 86); Bihar: jangali baigan (Varma 1981); Chhattisgarh: ban bhata, kutmi (*Mooney* 120); Kerala: anachunda (Mohanan and Henry 1994); Maharashtra: adhvi badnae [Kanara] (*Fernandes* 1377), ran vanghae (Marathi, *Fernandes* 1377); Odisha: tutugana (*Mooney* 688); Rajasthan: bhurat [Hindi] (Singh 1991); Tamil Nadu: sundai [Tamil] (Matthew 1983); sundaikai [Tamil] (Henry et al. 1987); sundaikkaai (*Sampath Kumar et al. 126940*). Indonesia. Borneo: terung pipit (*Wiriadianata* 1196); Sumatra: têrong pipit (Burkill 1935). Laos. kh'èngz do:n, kh'èngz hna:m hlwàng, kh'èngz khâm, kh'èngz saph'ao (Hul and Dy Phon 2014). Malaysia. Malacca: trong naya bang (*Anonymous* 418); terong manggor (*Anonymous s.n.* K001153289). Malaysia/Singapore. têrong pipit puteh, têrong rembang (Burkill 1935, but care to be taken with these names, they include other Torva clade taxa). Mauritius. Bringelle marron (*Drennen* 24). Sri Lanka. gona batu [Sinhala] (*Hepper & Silva* 4675, *Hepper* 4411, *Hepper* 4415); tibatu [Sinhala] (*Veldkamp* 7852). Thailand. weng (387); sambalan (*Hoed & Kostermans* 962). Vietnam. cà có, cà rùng, cà gai, cà pháo, trong phang, trong phet (Hul and Dy Phon 2014).

**Discussion.** *Solanum torvum* is by far the most commonly collected spiny solanum in tropical Asia; most recent collections sent to us for identification are this species. It may be becoming more common with anthropogenic change or with spread due to human use. *Solanum torvum* is ubiquitous and fully naturalised in the region and has become culturally important in many countries (see multiplicity of common names above). The fruits are used in cooking in China and Indochina; they are exported to Europe under the name pea eggplant. Hul and Dy Phon (2014) record the use of seeds in traditional medicine in Indochina, and in India extracts of the plant are used for relief from bites of venomous insects or snakes. In Bhutan the roots and leaves are used medicinally and the fruits are eaten (Mill 2001).

Vorontsova and Knapp (2016) incorrectly placed *S. pseudosaponaceum* and its synonym *S. macaoense* in the synonymy of *S. torvum*, our examination of more extensive material from tropical Asia clearly shows that *S. pseudosaponaceum* is a distinct taxon. *Solanum torvum* is a member of the Torvum clade and can be distinguished from all other members of that group in Asia (*S. chrysotrichum*, *S. comitis*, *S. kachinense*, *S. peikuoense*, *S. poka*, *S. pseudosaponaceum*, *S. torvoideum*) by its densely glandular inflorescences with tiny simple glandular trichomes, in addition to the stellate trichomes found in the rest of the plant; the rest of the plant is eglandular.

**Specimens examined.** See Suppl. materials 1–3.
45. *Solanum trilobatum* L., Sp. Pl. 188. 1753.

Figs 1B, 2F, 4F, 74

*Solanum acetosifolium* Lam., Tabl. Encycl. 2: 24. 1794, as "*acetosaefolium*". Type. India. Sin loc., *P. Sonnerat s.n.* (lectotype, designated here: P [P00055514]).

*Solanum canaranum* Miq., Hohen. Pl. Ind. Or. (Terr. Canara & Confin.) Exsicc. No. 740. 1847. Type. India. Karnakata: "prope Bettighery", Feb., *R.F. Hohenacker 740* (lectotype, designated here: U [U.1740185]; isolectotypes: BM [BM000900314], K [K000014877]).

*Solanum trilobatum* L. var. *griffithii* C.B.Clarke, Fl. Brit. India [J. D. Hooker] 4: 237. 1883. Type. "Birma and Malay Peninsula", *W. Griffith 5915* (lectotype, designated here: K [K000196631]).

*Solanum griffithii* (C.B.Clarke) Kuntze, Revis. Gen. Pl. 2: 454. 1891. Type. Based on *Solanum trilobatum* L. var. *griffithii* C.B.Clarke

**Type.** "Habitat in India", *Anonymous s.n.* (lectotype, designated by Deb 1980, pg. 51: LINN [acc. # 248.44]).

**Description.** Scandent herbs or shrubs, to 2.5 m tall, armed. Stems erect or spreading, terete, moderately to densely prickly, glabrous or with a few scattered stellate trichomes; prickles to 6 mm long, to 5 mm wide at the base, deltate, markedly hooked, laterally flattened, pale yellow to brown, glabrous; pubescence of sessile to very short-stalked porrect-stellate trichomes, the rays 2-5, 0.1-0.25 mm long, the midpoints absent or to 0.25 mm long; new growth sparsely stellate-pubescent, glabrescent, light yellowish green in dry material; bark of older stems brownish, glabrous. Sympodial units apparently difoliate, the leaves geminate, usually similar in size. Leaves simple, more or less deeply lobed, the blades 2.5-6.5 cm long, 2-7.5 cm wide, ca. 1-1.5 times longer than wide, ovate to broadly ovate, somewhat fleshy, slightly discolorous, unarmed or with a few hooked prickles like those of the stems, per face; adaxial surface green, glabrous or with a few scattered porrect-stellate trichomes, the trichomes sessile to sub sessile, the rays 4-8, 0.1-0.25 mm long, the midpoints to 0.1 mm long; abaxial surface light-green, glabrescent, sometimes with a few (i.e., < 10) porrect-stellate trichomes like those of the adaxial surface; major veins 3-5 pairs drying brown; base truncate to shortly attenuate; margins shallowly to deeply lobed, the lobes 1-2 on each side, 0.5-0.7 cm long, deltate, apically rounded, the sinuses extending up to 3/5 of the distance to the midvein; apex rounded; petioles 1.2-3 cm long, 1/2-2/3 of the leaf blade length, glabrescent, unarmed or with a few prickles like those of the stems. Inflorescences 1-6 cm long, apparently lateral, unbranched or forked, with ca. 3-11 flowers, only a few flowers open at any one time, glabrescent, unarmed; peduncle ca. 0.5 cm long, unarmed or a few prickles; pedicels 0.8-1.5 cm long, 0.8-1 mm in diameter at the base, 1.2-1.5 mm in diameter at the apex, erect, unarmed or with a few hooked prickles especially on the lower flowers, glabrescent, articulated at the base; pedicel scars spaced 0.4-1 cm apart. Buds ellipsoid, the corolla strongly exerted from
<figure><img src="image_89.png" /><figcaption>**Figure 74.** *Solanum trilobatum* L. **A** herbarium specimen collected in Thailand in 1966 (*Larsen 1614*, P00055524) **B** habit (*Meeboonya et al. RM 243*, Thailand) **C** detail of the adaxial surface of a leaf blade (*Meeboonya et al. RM 301*, Thailand) **D** detail view of an open flower (*Meeboonya et al. RM 245*, Thailand) **E** detail view of an infructescence (field photograph, unvouchered, India). Photograph credits: **A** CC-BY, Muséum national d'Histoire naturelle, Paris **B–E X. Aubriot**.</figcaption></figure>

the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 1.5–2 mm long, cup-shaped and somewhat keeled, the lobes 1–2 mm long, ca. 1 mm wide, narrowly deltate, apically acute, unarmed or with a few hooked to straight
prickles, glabrous to sparsely stellate-pubescent with sessile porrect-stellate trichomes like those of the stems. Corolla 2–3 cm in diameter, purplish blue, stellate, lobed ca. 3/5–4/5 of the way to the base, the lobes 8–12 mm long, 2.5–7 mm wide, deltate to narrowly deltate, spreading at anthesis, sparsely to moderately stellate-pubescent abaxially on parts exposed in bud. Stamens equal; anthers 7–9 mm long, 1–1.5 mm wide, connivent, not tapering at anthesis, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying. Ovary globose, with minute glandular hairs; style 1.4–1.8 cm long, slender, curved at the apex, glabrous; stigma capitate, minutely papillate. Fruit a globose berry, 1 to several per infructescence, 0.7–1.2(–1.7) cm in diameter, red at maturity, the pericarp thin and shiny, glabrous; fruiting pedicels 1.2–2.6 cm long, ca. 1 mm in diameter at the base, 2–2.5 mm in diameter at the apex, erect to recurved, not markedly woody; fruiting calyx lobes slightly expanding to ca. 3 mm long, 1/5–1/3 the length of the mature fruit, deltate, not reflexed. Seeds ca. 16–47 per berry, 3–4 mm long, 2.5–3 mm wide, flattened-reniform, dull yellow, the surface minutely pitted, the testal cells pentagonal in outline. Chromosome number: n = 12, 2n = 24 (Madhavadian 1968).

**Distribution** (Fig. 75). *Solanum trilobatum* is widely distributed in tropical Asia; from South India and Sri Lanka to Indochina, and Java. The single collection from the Philippines (*Loher s.n.*) is from the railway station in Manila and is probably cultivated.

<figure><img src="image_90.png" /><figcaption>**Figure 75.** Distribution of *S. trilobatum*.</figcaption></figure>
**Ecology and habitat.** *Solanum trilobatum* grows in open areas or forest edges, on sand or limestone, close to coastal areas; often in open field or mangrove forest; from around sea level to 700 m elevation.

**Common names and uses.** Cambodia. traob (Khmer, *Hul 4774*); India. Andhra Pradesh: uchinta (*Venkanna 5562*), usti (*Raji 291*); Karnataka: ullul (Singh 1988); Odisha: ankaranti (*Panigrahi EC-23792*); Tamil Nadu: toundou valay [Tamul] (*Com- merson s.n.*), thoothuvalai [Tamul] (*Matthew 8323*), tudulai [Tamul] (*Mehen-Homji s.n.*), thoodhuvalai [Tamul] (*Narasimhan 624*, *Sampath Kumar et al. SC-126970*), thudhuvaelai [Tamul] (Matthew 1983), tuduvali [Tamul] (Henry et al. 1987); West Bengal: kantikuri (*Tribedi CNH-1435*); Malaysia. priapantie (*Curtis 858*).

It is reported that in Thailand the plant is boiled to serve as a decoction for ulcerated throat (*Collins 1380*). Indian collectors reported that the fruits are edible and used as medicinal (*Raji 291*, *Sengupta 145*). In Penang (Malaysia) the root is used medicinally (*Curtis 858*). For the many uses in traditional medicine in India see section on Uses (pg. 18–20).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (2,326,480 km², LC); AOO (528 km², VU). Widely distributed and likely to be protected by people as a plant of considerable medicinal importance, *S. trilobatum* is not of immediate conservation concern. Methods of collection for medicinal use, however, have not been assessed and if these are destructive this could be cause for concern.

**Discussion.** *Solanum trilobatum* is widely distributed through India and Indonesia. It was one of the first Asian spiny solanums to be described (Linnaeus 1753) and was brought into cultivation in European botanical gardens in the late 18th century. It is a distinctive species with its rhomboidal to triangular leaves that are usually widest in the lower third, but has been confused with *S. procumbens*, a similarly scandent plant. Flowers of *S. trilobatum* are much larger than those of *S. procumbens* (2–3 cm in diameter versus 1–1.5 cm in diameter), with abundant interpetalar tissue; *S. procumbens* flowers are deeply stellate with narrow corolla lobes. The two species do not overlap in distribution.

No specimen labelled as *S. acetosifolium* has been found in the Lamarck herbarium (P-LAM), however in the French language column Lamarck (1794) cited “Indes Orient, Sonnerat” clearly indicating a collector. We have lectotypified *S. acetosifolium* with a specimen collected by Sonnerat labelled “Inde” in the general collection at Paris (P00055514).

In describing *S. trilobatum* var. *griffithii*, Clarke (1883) cited only “Griffith 5915”; we have selected the single specimen thus labelled (K000196631) as the lectotype.

The name *S. canaranum* was effectively published by being printed on herbarium labels that were distributed to herbaria before 1953 (Turland et al. 2018, Art. 30.8); the name is also accompanied by a diagnosis attributed to Miquel. We have chosen the duplicate in Utrecht [U.1740185], where Friedrich Miquel worked (Stafleu 1966), as the lectotype for this name.

**Specimens examined.** See Suppl. materials 1–3.
46. *Solanum vagum* Nees, Trans. Linn. Soc. London 17(1): 48 1834.
Fig. 76

**Type.** INDIA. Sin. loc., "S. corymbosum Pers?., hb. Wight", *Herb. Wight s.n.* (Walter cat. 2624b) (lectotype, designated here: GZU [GZU000255872]; isolectotypes: K [K001152841], K-W [K001116639]).

**Description.** Erect shrubs 0.5-2.5 m tall, unarmed. Stems erect, terete, sparsely to moderately stellate-pubescent, soon glabrescent; pubescence of sessile to short-stalked porrect-stellate to somewhat multangulate trichomes, the stalks 0.1-0.2 mm long, the rays 4-12, 0.1-0.2 mm long, the midpoints shorter than or as long as the rays; new growth densely stellate-pubescent, the trichomes porrect-stellate, sessile or short-stalked, the stalks 0.1-0.2 mm long, the rays 6-8, 0.2-0.5 mm long, the midpoints equal to the rays; bark of older stems glabrescent, usually with only a few scattered stellate trichomes, greyish brown and somewhat shiny. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, entire, the blades 5-15 cm long, 1.5-7 cm wide, lanceolate to narrowly elliptic to elliptic, membranous, discolorous; adaxial surface drying dark, glabrous or with a few scattered sessile porrect-stellate trichomes on the veins and lamina, the trichomes more delicate than those of the abaxial surfaces, with 4-8 rays 0.1-0.2 mm long and midpoints equal to the rays; abaxial surface paler, almost glabrous to moderately pubescent, the trichomes sessile or very short-stalked, the rays 4-8, 0.3-0.5 mm long, the midpoints equal to the rays; major veins 6-8 pairs, prominent and drying yellowish or darker than the lamina abaxially; base acute, somewhat oblique, but not markedly so; margins entire or slightly sinuate and irregularly lobed; apex acute to somewhat acuminate; petiole 2-6 cm long, 1/2 or more of the leaf blade length, glabrous to sparsely stellate-pubescent with porrect-stellate stellate trichomes like the stems. Inflorescences 3-8 cm long, apparently terminal or lateral, usually unbranched but occasional forked, with 4-10 flowers, 1-2 flowers open at any one time, glabrous to very sparsely pubescent with sessile porrect-stellate trichomes like those of the stems; peduncle 1-3 cm long; pedicels 1-1.2 cm long, 1-1.2 mm in diameter at the base, 1.5-1.8 mm in diameter at the apex, erect at anthesis, glabrous or sparsely stellate-pubescent like the inflorescence axes, articulated at the base or just above and leaving a minute peg on the inflorescence axis; pedicel scars spaced 1-3 mm apart. Buds ovoid, not markedly tapering, densely stellate-pubescent, strongly exerted from the calyx tube before anthesis, but the calyx acumens nearly to the tips of the buds. Flowers 5-merous, apparently all perfect. Calyx with the tube 1.5-2 mm long, conical, the lobes 1-2 mm long, 1-1.2 mm wide, narrowly deltate with a long subulate acumen to 2.5 mm long, glabrous or very sparsely pubescent abaxially with sessile porrect-stellate trichomes like those of the pedicels. Corolla 2-3 cm in diameter, white, stellate, lobed nearly to the base, the lobes 8-10 mm long, 2.5-3 mm wide, narrowly deltate, spreading at anthesis, glabrous adaxially, moderately pubescent abaxially with sessile stellate porrect trichomes, the rays 3-10, ca. 0.1 mm long, the midpoints equal to the rays, the tips somewhat cucullate. Stamens equal in bud, unequal at anthesis, with 4 short and one long and curved; long anther 7-9 mm long and strongly curved upwards, 1-1.5 mm wide, the short anthers 3-6 mm long, 1-1.5 mm wide, elliptic but the longer one somewhat tapering, all yellow, glabrous, poricidal at the tips, the pores
<figure><img src="image_91.png" /><figcaption>**Figure 76.** *Solanum vagum* B.Heyne ex Nees A herbarium specimen collected in India in 1818 (*Herb. Wight s.n.* (Wallich cat. 2624A), K001116638) B habit (field photograph, unvouchered, India) C detail view of an infructescence (field photograph, unvouchered, India). Photograph credits: A © copyright of the Board of Trustees of the Royal Botanic Gardens, Kew B, C R. Vellingiri.</figcaption></figure>

directed distally and not elongating with age; filament tube minute, glabrous; free portion of all filaments ca. 0.5 mm long, glabrous; Ovary conical, glabrous or with a few porrect-stellate trichomes at the apex; style 9–11 mm long, slender, strongly curved in same direction as the long anther, glabrous or with a few porrect-stellate trichomes at the very base; stigma capitate, minutely papillate. Fruit a globose berry, 2–4 per infructescence, 0.8–1 cm in diameter, red at maturity, the pericarp thin and shiny, glabrous; fruiting pedicels 1.5–1.8 cm long, ca. 1 mm in diameter at the base, 2–2.5 mm in diameter at the apex, woody and erect; fruiting calyx lobes not markedly accrescent, splitting to the base and expanding to 4–6 mm long including the acumen which often breaks off in dry specimens. Seeds 10–20 per berry, 4–4.5 mm long, 3–4 mm wide, flattened-reniform, pale tan, the surfaces smooth in the seed centre, minutely pitted on the thickened margins, the testal cells somewhat sinuate in outline. Chromosome number; n = 12 (Madhavadian 1968).
<figure><img src="image_92.png" /><figcaption>**Figure 77.** Distribution of *Solanum vagum*.</figcaption></figure>

**Distribution (Fig. 77).** *Solanum vagum* occurs in southern India (states of Kerala and Tamil Nadu) and in Sri Lanka.

**Ecology and habitat.** *Solanum vagum* grows in dry open woodland, degraded vegetation, grasslands and open areas; occurring from sea level to 750 m elevation.

**Common names and uses.** None recorded.

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (75,338 km², LC); AOO (76 km², EN). *Solanum vagum* is found both in southern India and Sri Lanka, and where it occurs it is a plant of open, disturbed areas, suggesting that it is not of immediate conservation concern.

**Discussion.** *Solanum vagum* is morphologically similar and probably closely related to *S. pubescens*, sharing with that species zygomorphic flowers at anthesis, heteromorphic anthers and shiny berries on erect or slightly pendulous pedicels. Hepper (1987) treated material of *S. vagum* from Sri Lanka as *S. pubescens* in the "Revised Handbook to the Flora of Ceylon". It differs from *S. pubescens* in its complete lack of glandular pubescence, its narrowly elliptic leaves with attenuate bases and slightly larger, white (rather than violet) flowers. The pubescence of adaxial leaf surfaces of *S. vagum* is of very sparse sessile trichomes with midpoints usually equal to the rays, and the lamina is clearly visible, while in *S. pubescens* the lamina of young leaves is obscured by the dense covering of glandular stellate to multangulate stalked trichomes with elongate
midpoints. The two species apparently co-occur in southern India, while all specimens we have seen identified as *S. pubescens* from Sri Lanka are referable to *S. vagum*.

The long anther in the flowers of *S. vagum* appears to expand after anthesis; specimens with buds or very young flowers have the anthers all of equal size, while older flowers have one anther markedly longer than the rest. Post-anthesis anther expansion occurs in the unrelated *S. turneroides* Chodat (Brevantherum clade, see Stern et al. 2013) of southern South America. Buds of *S. pubescens* and the African species *S. somalense* Franch., which greatly resemble those of *S. vagum*, also have the anthers of more or less equal length and buds that are not markedly curved.

*Solanum vagum* is probably a member of the Giganteum clade (sensu Vorontsova et al. 2013; Aubriot et al. 2016a) based on similarity with *S. pubescens* and the African/ Middle Eastern species *S. somalense*. These taxa can be distinguished from *S. giganteum*, the only other member of the group occurring in tropical Asia, by their zygomorphic flowers and calyx lobes with long subulate tips that are especially visible in bud.

The species has been indicated as endemic to the southern part of the Western Ghats biodiversity hotspot (https://indiabiodiversity.org/biodiv/species/show/264048) but also occurs in Sri Lanka.

Nees van Esenbeck (1834) validated the herbarium name previously listed as Wallich cat. 2624 citing two collections; “Solanum vagum. Herb. Heyn., Wall. l.c.” (Wallich cat. 2624a) and “Solanum corymbosum? Herb. Wight” (Wallich cat. 2624b). We have selected the specimen at GZU that comes from Nees van Esenbeck’s own herbarium (Stafleu and Cowan 1981) as the lectotype for *S. vagum* because it is clear that it was used by Nees, comes from one of the original collections in the Wallich herbarium and has both mature flowers and fruits. This collection from the Wight herbarium (Wallich cat. 2624b) also has more widely distributed duplicates than the other syntype from the Heyne herbarium (Wallich cat. 2624a). Although a specimen in BM (BM000900081) is labelled “Wall. cat. 2624b” it is also labelled as being from “herb. Heyne” and so is here treated as a syntype, not as an isolectotype.

**Specimens examined.** See Suppl. materials 1–3.

**47. Solanum viarum Dunal, Prodr. [A.P. de Candolle] 13(1): 240. 1852.**

Figs 3H, 73C–E

Names based on tropical Asian types only; for complete synonymy see Nee 1979; Vorontsova and Knapp 2016.

*Solanum khasianum* C.B.Clarke var. *chatterjeeanum* Sengupta, Bull. Bot. Surv. India 3: 413. 1961. Type. India. Tamil Nadu: “Madras, Nilgiris district”, 1075 m, 17 Jul 1960, *K. Subramanyam 10413* (holotype: CAL [CAL-10413A]; isotypes: CAL [CAL-10413B, CAL-10413C]).

**Type.** BRAZIL. São Paulo: sin. loc., *P. Lund* 799 (holotype: G-DC [G00145818, F neg. 6816, IDC microfiche 800-61.2080:I.1]).
**Description.** Vorontsova and Knapp (2016: 365–368); http://www.solanaceae-source.org/solanaceae/solanum-viarum.

**Distribution.** *Solanum viarum* is widely distributed in tropical Asia, particularly in China and northern India; native to Brazil, this species is widely adventive and can be invasive (see below). *Solanum viarum* has been formally assessed as of Least Concern (LC) on the IUCN Red List (Knapp 2020).

**Common names.** China. mao guo qie (Zhang et al. 1994). India. Andaman and Nicobar Islands: jangli baigun (Basu ANC-6866).

**Discussion.** *Solanum viarum* is a member of the Acanthophora clade (sensu Stern et al. 2011; Gagnon et al. 2022), along with *S. aculeatissimum*, *S. capsicoides*, and *S. mammosum*. All members of the clade can be distinguished by their pubescence of simple trichomes (derived from stellate midpoints, see Whalen 1984) on upper leaf surfaces. It can be distinguished from the rest of the members of the clade in tropical Asia in its dense and fine glandular pubescence on all parts, relatively small fruits, unwinged seeds, and small, pale greenish cream flowers. It is most similar to *S. aculeatissimum*, which has similarly small berries, but differs from that species in its deltate, rather than long acuminate, calyx lobes and puberulent, rather than stipitate-glandular, ovary. The prickles of *S. viarum* are often slightly curved and of uniform length, while those of *S. aculeatissimum* are straight and of varying lengths.

*Solanum viarum* and *S. aculeatissimum* were both previously known as *S. khasiense* (see Babu and Hepper 1979), but subsequent monographic work across the native range of these taxa (Nee 1979) clarified that the two were distinct.

*Solanum viarum* has potent antioxidant compounds similar to those used medically from other species in tropical Asia (e.g., *S. insanum*, see Uses, pg. 18–20) and has potential for medicinal or other use in the area (Wu et al. 2012). In Bhutan it has been recorded as being grown as a source of the alkaloid solasodine (Mill 2001).

*Solanum viarum* (under the common name soda apple or tropical soda apple) has been classified as a noxious weed in the southern United States and Australia (https://www.ars.usda.gov/news-events/news/research-news/1999/digging-burning-thwart-soda-apple-weed/; https://weeds.dpi.nsw.gov.au/Weeds/Details/186). Where we have seen it in tropical Asia it is similarly aggressive and can take over fields and pastures.

**Specimens examined.** See Suppl. materials 1–3.

48. *Solanum violaceum* Ortega, Nov. Pl. Descr. Dec. 56. 1798.
Figs 3A, 78

*Solanum chinense* Dunal, Hist. Nat. Solanum 240. 1813. Type. China. Sin. loc. (lectotype, designated here: [illustration] Plukenet, Phytographia, pars altera, tab. 62, fig. 1. 1691).

*Solanum heynei* Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 669. 1819, as "Heynii". Type. "H. in India orientali. B. Heyne", *B. Heyne s.n.* (lectotype, designated by Turner 2021, pg. 418: L [L 0403734]).
*Solanum pinnatifidum* Roth, Nov. Pl. Sp. 130. 1821, nom. illeg. non *S. pinnatifidum* Lam., 1794. Type. Based on same material and homotypic with *S. heynei* Roem. & Schult.

*Solanum indicum* Nees var. sinuato-lobatum Dunal, Prodr. [A. P. de Candolle] 13(1): 309. 1852. Type. Indonesia. “Malacca”, *H. Cuming* 2261 (lectotype, designated here: G [G00442779]; isolectotypes: BM [BM000886164, BM000886188], E [E00526948], LE, K [K000014604, K000014606], P [P00055666]).

*Solanum indicum* Nees var. eroso-pinnatifidum Dunal, Prodr. [A. P. de Candolle] 13(1): 310. 1852. Type. India. Sin. loc., “Bengala inferior”, 1817, *N. Wallich s.n.* [Wallich Catal. 2626e] (lectotype, designated here: G-DC [G00130375]; isolectotypes: GZU [GZU000255428], K-W [K001116646 pro parte, bottom L hand fragment only]).

*Solanum indicum* Nees var. parvifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 310. 1852. Type. Mauritius. “Isle Maurice”, 1839, *L. Bouton s.n.* (lectotype, designated here: G-DC [G00130374]).

*Solanum junghuhnii* Miq., Fl. Ned. Ind. 2: 649. 1857. Type. Indonesia. Java: Central Java, “op Tjilatjap” [Cilacap], *H. Gesker s.n.* [195] (lectotype, designated here: L [L0003638]).

*Solanum indicum* L. var. inerme Van Heurck & Müll.Arg., Observ. Bot. (Van Heurck) 2: 133. 1871. Type. India. Assam: Sin. loc., *W. Griffith 1000* (holotype: BR [AWH10071564]; isotype: BM [BM000900299]).

*Solanum pubescens* Willd. var. lobatum C.B.Clarke, Fl. Brit. India [J. D. Hooker] 4: 231. 1883. Type. India. Meghalaya: “Assam, Khasi Hills” [Khasia Hills], *S. Kurz s.n.* (lectotype, designated here: CAL [CAL0000018697]).

*Solanum kurzii* Brace ex Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 65: 541. 1896. Type. India. Sikkim: Sin. loc., 8 May 1874, *G. King s.n.* (lectotype, designated here: K [K000441390]).

*Solanum indicum* L. var. mesarchon Bitter, Repert. Spec. Nov. Regni Veg. Beih. 16: 9. 1923. Type. Mauritius. Pamplemousses District: “Umgebung von Pamplemousse” [surroundings of Pamplemousse], May 1887, *S. Paulay s.n.* (holotype: W [acc # 1887-0010053]).

*Solanum sanitwongsei* Craib, Kew Bull. 1928: 246. 1928. Type. Thailand. Bangkok: “cultivated”, 20 Nov 1927, *A.F.G. Kerr s.n.* (lectotype, designated here: K [K000922035]).

*Solanum nivalomontanum* C.Y.Wu & S.C.Huang, Acta Phytotax. Sin. 16(2): 74. 1978. Type. China. Yunnan: Fengqing, “Shunning, Snowrange [transl. from the protologue]”, 25 May 1938, *T.T. Yu* 15962 (lectotype, designated here: PE [PE00031398]); isolectotype: KUN [KUN183591], PE [PE00031399]).

*Solanum indicum* L. forma album C.Y.Wu & S.C.Huang, Fl. Yunnanica 2: 580. 1979. Type. China. Yunnan: Yanshan, “Yan-shan-hsien, Bar-garh [transl. from the protologue]”, 16 Nov 1939, *C.W. Wang* 85018 (lectotype, designated here: PE [PE00031401]); isolectotype: KUN [KUN183578]).

**Type.** INDIA. "*Habitat* en Bahia Botanica. *Floret* Octobri et Novembri in Reg. Hort. Matrit. è seminibus Londino missis per *Exc. D. Marchionissam de Bute*" (neotype, designated by Knapp 2013b, pg. 59: MA [MA307449]).
<figure><img src="image_93.png" /><figcaption>**Figure 78.** *Solanum violaceum* Ortega **A** herbarium specimen collected in Thailand in 1920 (Kerr s.n., BM000886106) **B** habit (field photograph, unvouchered, Thailand) **C** detailed view of a white flower with immature fruits (*Sampath Kumar et al. 126947*, India) **D** detailed view of a purple flower (field photograph, unvouchered, Thailand) **E** detail view of an infructescence (*Sampath Kumar et al. 126945*, India). Photograph credits: **A** CC-BY, Muséum national d'Histoire naturelle, Paris **B–E** X. Aubriot.</figcaption></figure>

**Description.** Erect shrub, to 3 m tall, armed or unarmed. Stems erect, flattened to terete, prickly or less often unarmed, moderately to densely stellate-pubescent; prickles to 10 mm long, to 7 mm wide at the base, usually curved, sometimes straight, rounded or flattened, straw-yellow to orange-brown, glabrous or sparsely pubescent
in the lower 1/3; pubescence of sessile or shortly stalked porrect-stellate trichomes, the stalks less than 0.1 mm long, the rays 6–8, 0.1–0.2 mm long, the midpoints same length as the rays or to 1.5 mm long; new growth densely stellate-pubescent, the trichomes like those of the stems; bark of older stems brown, glabrescent. Sympodial units difoliate, the leaves geminate. Leaves simple, more or less deeply lobed, the blades 4–12 cm long, 2.5–11 cm wide, 1.5–2 times longer than wide, usually ovate in outline, sometimes elliptic, chartaceous, strongly discolorous but sometimes concolorous on dry material, unarmed or with several straight prickles to 10 mm long on veins of both surfaces; adaxial surface evenly and moderately to densely stellate-pubescent with sessile or short-stalked trichomes, the stalks to 0.5 mm, the rays 4–10, ca. 0.5 mm long, the midpoints shorter than the rays or to 2 mm long; abaxial surface densely stellate-pubescent with trichomes like those of the adaxial surface but with longer rays and midpoints, the lamina not usually visible; principal veins 3–5 pairs; base usually truncate, often oblique; margins lobed, the lobes (2-3)(-4) on each side, 1–4.5 cm long, broadly deltate to oblong or obovate, often with secondary lobes, apically obtuse, sometimes acute or rounded, the sinuses extending to 2/3 of the way to the midrib; apex acute; petiole 1–6 cm long, 1/5–3/5 of the leaf blade length, densely stellate-pubescent, unarmed or with a few straight or slightly curved prickles. Inflorescences, 2.5–8(–10) cm long, lateral or leaf-opposed, unbranched or sometimes forked, with 5–15(–30) flowers, 2–3 flowers open at any one time, moderately to densely pubescent with stellate-porrect trichomes like those of the stems, unarmed or with a few straight prickles; peduncle 0.1–1(–2.5) cm long, usually unarmed; pedicels 0.8–1.7 cm long, 0.7–1 mm in diameter at the base, 1.6–2 mm in diameter at the apex, straight and slender, unarmed, densely stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars spaced 2–10 mm apart. Buds ovoid, the corolla strongly exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 1.5–2 mm long, obconical to cup-shaped, the lobes 1.5–4 mm long, 1.5–2.5 mm wide, deltate, apically apiculate to acute, rarely long-acuminate, with no venation visible, unarmed or with up to 10 straight prickles, densely stellate-pubescent with sessile porrect-stellate trichomes like those of the rest of the inflorescence. Corolla 1.3–3 cm in diameter, pale blue to purple or more rarely white, stellate, lobed 1/2–2/3 of the way to the base, the lobes 4.5–8 mm long, 3–5 mm wide, deltate, spreading to slightly reflexed at anthesis, mostly glabrous adaxially, moderately to densely stellate-pubescent abaxially, the trichomes porrect, sessile to shortly stalked, the stalks up to 0.1 mm, the rays 6–8, 0.1–0.2 mm, the midpoints shorter than the rays, but lengthening towards corolla lobe apices. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, glabrous; anthers 4.5–8.5 mm long, 1.2–1.5 mm wide, yellow-orange or yellow, connivent, tapering, glabrous but sometimes with a few stellate trichomes, poricidal at the tips, the pores directed distally and lengthening to slits with age. Ovary ovoid, glabrous but with a few stellate trichomes towards the apex; style 7–11(15) mm long, filiform, straight to gently curved, stellate-pubescent in the lower 1/3–2/3; stigma clavate to capitate, the surface minutely papillose. Fruit a globose berry, several per infructescence, 0.7–0.9 cm in diameter, yellow to orange when ripe, the pericarp
thin and shiny, glabrous; fruiting pedicels 1.2–1.8 cm long, 0.5–1.2 mm wide at the base, 3–3.5 mm in diameter at the apex, somewhat woody, straight, spreading, unarm ed or with a few straight prickles; fruiting calyx lobes elongating to 3.5–6(–8) mm long, 1/4–1/2 of the length of the mature fruit, spreading and usually not reflexed, unarm ed or with up to 10 straight prickles. Seeds ca. 10–15 per berry, 2.2–3.5 mm long, 1.8–3 mm wide, flattened-reniform, yellow to orange-brown, the surfaces minutely pitted, the testal cells with thick, strongly sinuate margins. Chromosome number: n = 12 (Karihaloo 1991), 2n = 24 (Karihaloo 1991; Das and Borah 2015, as S. *kurzii*).

**Distribution** (Fig. 79). *Solanum violaceum* occurs across Asia, from India to China, Vietnam and Malaysia, from the coasts to inland mountains between ca. 3–27° latitude. It is very common in India, southern China, and Thailand. The westernmost recorded occurrences are on the islands of Mauritius, Réunion and Rodrigues. Chinese populations are restricted to the more southern regions of Fujian, Guangdong, Guangxi, Hainan, Hong Kong, Sichuan Taiwan, and Yunnan (Zhang et al. 1994). There are occasional collections from Sumatra and Java, but it is not clear whether these are from the wild or cultivated. *Solanum violaceum* does not occur wild in the Philippines, New Guinea, or Australia.

<figure><img src="image_94.png" /><figcaption>**Figure 79.** Distribution of *Solanum violaceum*.</figcaption></figure>

**Ecology and habitat.** *Solanum violaceum* is usually found growing in open places, abandoned cultivation, and roadsides, in a variety of forest types; between sea level and 2,000 m elevation.

**Common names and uses.** China. ci tian qie (Zhang et al. 1994); Yunnan: xue shan qie (eggplant from the snow mountain; *Yu 15962*). India. Bihar: jangali baigan (Varma 1981, as *S. indicum*); Goa: ringani, inoti-ringani, badane, dorli (darli),
motaring (Naithani et al. 1997); Karnataka: mullsunde, kad badne, ustikai (Singh 1988); Kerala: cheruchunda (Mohanan and Henry 1994, as *S. anguivi* Lam.); Tamil Nadu: naaimulli, mulluchundai [Tamil] (Matthew 1983); cherukinda, cheruvazhuthanai [Malayalam], ciruvaludalai, kondal, karimulli [Tamil] (Nair and Nayar 1987, as *S. indicum*), mulli, pappara-mulli, karimulli [Tamil] (Henry et al. 1987, as *S. anguivi*). Malaysia/Singapore: těrong pipit puteh, těrong pipit hijau (Burkill 1935), tèrong peuheur [Sundanese] (Burkill 1935).

The roots of *S. violaceum* (as *S. indicum*) are one of the ingredients of Dashamoola of Ayurvedic medicine used to treat inflammatory conditions of all kinds (Naithani et al. 1997; Jain et al. 2000). It is regarded as a diuretic, is used for treatment of dropsy, and as an asthma and catarrhal expectorant (Jain et al. 2000). Unripe fruits are used in the preparation of curries (Naithani et al. 1997).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (10,078,363 km², LC); AOO (988 km², VU). *Solanum violaceum* is one of the most widely distributed spiny solanums of tropical Asia and grows in a wide variety of disturbed habitats, often as large populations of individuals.

**Discussion.** *Solanum violaceum* is very widespread and is common where it occurs. In the older literature the name *Solanum indicum* was often used for this taxon, but considerable confusion over its application led to its suppression (Hepper 1978; see Doubtful and excluded names).

*Solanum violaceum* is a plant of disturbed areas, often growing along roadsides and in stream beds. It has been confused with *S. anguivi* Lam., an African taxon, because of its small orange to red berries and small, hermaphroditic flowers (e.g., Hepper 1978; Singh 1991), but is not directly related to that species. It differs from *S. anguivi* in its orange, rather than red, berries, its usually violet or pale violet, rather than white flowers, and in its straight, spreading pedicels in fruit, rather than slightly curved pedicels.

In the Flora of Bhutan (Mill 2001) *S. violaceum* was treated as *S. kurzii* (specimens with few prickles) and *S. anguivi* (prickly plants).

*Solanum violaceum* is sympatric with its close relatives *S. deflexicarpum* of southern China, *S. hovei*, and *S. multiflorum*, the latter two endemic to India. All of these species have strongly deflexed pedicels in fruit, while those of *S. violaceum* are broadly spreading and usually longer. See those species descriptions for more details. *Solanum violaceum* is quite variable across its range in leaf shape and prickliness; populations from Sri Lanka and Thailand lacking prickles have been called *S. kurzii* and *S. sanitwongsei*, respectively. Similar unarmed plants, however, are found across the range of *S. violaceum* and even within populations.

*Solanum chinense* was described as a species of uncertain status related to *S. violaceum* (Dunal 1813) referring only to Tournefort (1700) and an illustration from Pluke-net (1691); we have found no other authentic original material, and so lectotypify the name using the Plukenet illustration which matches the protologue.

The varieties of *S. indicum* sensu Nees described by Dunal (1852), were based on the polynomial varieties from Nees van Esenbeck (1834), which themselves were based entirely on specimens from Wallich's herbarium that he saw in London (De
Candolle and Radcliffe-Smith 1981). Dunal (1852) cited a number of specimens and concepts in synonymy, making typification difficult, but in lectotypifying these in-
fraspecific names we have used specimens at G-DC that Dunal definitely would have seen during his preparation of the *Prodromus* treatment. For var. *sinuatolobatum* we have selected as the lectotype a collection specifically mentioned in the protologue "Cuming 2261, hb. Boiss." (G00442779) that is widely duplicated. For var. *parvi-
folium* we have selected the *Bouton s.n.* specimen from Mauritius in G-DC cited in the protologue (G00130374) because it is an unambiguous, well-preserved specimen. Var. *eroso-pinnatifidum* cited "S. pinnatifidum *Roth*, S. heynii *R. & S.*, *Solanum indicum* *Wall. cat.* 2626 D,E" as material; we have selected the G-DC specimen of "Wallich cat. 2626E" (G00130375) as the lectotype of this variety. The "duplicate" in the Wallich herbarium at Kew (K00116646) is a mixture of potentially three elements, only the lower L hand stem of which we regard as isolectotype material; this fragment has the letter "E" in pencil, as does the larger R hand fragment. This larger fragment appears to be a collection of *S. multiflorum*, and so we exclude it as type material for var. *eroso-
pinnatifidum*. Duplicates of all these varieties in Nees van Esenbeck's own herbarium at GZU all correspond to *S. violaceum* and are considered isolectotypes.

Miquel (1857) cited three specimens in the protologue of *S. junghuhnii: Gesker*
*s.n.* from "Tjiltajap", *Junghuhn s.n.* from "Awoe Awoe" and *Horsfield s.n.* from "Soera-
karta". We have selected the Leiden specimen (L 0003638) collected by F.W. Junghuhn
as the lectotype, it is labelled unambiguously as to locality and matches the protologue.

Clarke (1883) indicated his uncertainty over the identity of his *S. pubescens* var. *lobatum* by stating “Var. ? *lobata*” and “it resembles the unarmed form of *S. Melongena* but the flowers are too small.” He cited a specimen “from Herb. Calcutta, named *S. pu-bescens* by Kurz”. Several specimens in CAL are so labelled, we have selected the best of these (CAL0000018697) as the lectotype.

The protologue of *S. kurzii* (Prain 1896) cites three specimens, two from Sikkim (*Thomson s.n.* and *King s.n.*) and one from “Khasia” (*Mann s.n.*). Specimens collected from the effort towards the flora of British India (Endersby 2008) are usually un-numbered and have minimal to no locality information. Tracing them can be challenging. A sheet at Kew (K000441390) collected by George King in Sikkim in 1874 matches the protologue in being nearly without prickles and is here selected as the lectotype for *S. kurzii*.

The protologue of *S. santiwongsei* cites only an un-numbered collection of A.F.G.
Kerr from Bangkok, with no further details. Craib in his many papers of additions to
“Siamese plants” never cited herbaria, but in the final compilation of these lists (Craib
1928) stated that the distributions of all species were taken from specimens at Kew. We
have therefore selected a specimen collected by Kerr in Bangkok of a cultivated plant
annotated with the name *S. sanitwongsei* (K000922035), as the lectotype of this name.

The type of *S. nivalomontanum* is indicated as being in PE (Wu and Huang 1978), but two duplicates of the type gathering are housed there; we here select the better preserved of these (PE00031398) as the lectotype. The protologue of *S. indicum* forma *album* cites a gathering (Wang 85018; Wu and Huang 1979), but no herbarium
is mentioned. We have selected the duplicate preserved in PE (PE00031401) as the lectotype, because we have been able to access images of it and ascertain its identity.

**Specimens examined.** See Suppl. materials 1–3.

49. *Solanum virginianum* L., Sp. Pl. 187. 1753.

Figs 2D, 80

*Solanum surattense* Burm.f., Fl. Ind. (N. L. Burman) 57. 1768. Type. India. Gujarat: “Surat”, *L. Garcin s.n.* (lectotype, designated by Hepper and Jaeger 1986, pg. 434, as “type”; second step designated here: G-PREL [G00811278]; isolectotype: G-PREL [G00811279]).

*Solanum virginicum* L., Mant. 2: 340. 1771. Type. Based on (orthographic error for) *Solanum virginianum* L., not intended as a new name.

*Solanum armatum* Forssk., Fl. Aegypt.-Arab. 47. 1775. Type. Yemen. Al Hudaydah: Hays, “ad Haes” [from protologue], *P. Forsskål s.n.* (lectotype, designated here: KIEL [KIEL0005103]).

*Solanum xanthocarpum* Schrad. & J.C.Wendl., Sert. Hanov. 1: 8, tab. 2. 1795. Type. Based on *Solanum virginianum* L. (see Hepper and Jaeger 1986).

*Solanum jacquinii* Willd., Sp. Pl., ed. 4 [Willdenow] 1(2): 1041. 1798, nom. illeg. superfl. Type. Based on *Solanum virginianum* L. (cited indirectly in synonymy).

*Solanum arabicum* Dunal, Hist. Nat. Solanum 240. 1813, nom. illeg. superfl. Type. Based on *Solanum armatum* Forssk. (cited in synonymy).

*Solanum xanthocarpum* Schrad. & J.C.Wendl. var. *schraderi* Dunal, Prodr. [A. P. de Candolle] 13(1): 302. 1852. Type. Cultivated in the Calcutta Botanical Garden, “Hort Calc”, *N. Wallich s.n.* [Wallich Catal. 2612b] (lectotype, designated here: G-DC [G00130292]; isolectotypes: BM [BM000900258], K [K001080635], K-W [K001116573]).

*Solanum xanthocarpum* Schrad. & J.C.Wendl. var. *jacquinii* (Willd.) Dunal, Prodr. [A. P. de Candolle] 13(1): 303. 1852. Type. Based on *Solanum jacquinii* Willd.

*Solanum mairei* H.Lév., Repert. Spec. Nov. Regni Veg. 12: 531. 1913. Type. China. Yunnan: “Plaine de Kiao-Kia”, May 1912, *E.E. Maire s.n.* (lectotype, designated here: E [E00284473]).

*Solanum mccannii* Santapau, J. Bombay Nat. Hist. Soc. 47: 654. 1948. Type. India. Maharashtra: Pune, “Khandala”, 18 Oct 1943, *H. Santapau 2972* (lectotype, designated here: BLAT [acc. # 89719]).

**Type.** “America” (lectotype, designated by Hepper and Jaeger 1986, pg. 434: [illustration] “Solanum American. laciniatum spinossimum”, Dillenius, Hort. Eltham. 360, t. 267, f. 346. 1732).

**Description.** Prostrate shrubs, to 60 cm tall, heavily armed. Stems sprawling to erect, terete, densely prickly and sparsely stellate-pubescent; prickles to 2 cm long, broad-based, straight, straw-yellow in dry material, pale whitish green in live plants; pubescence of sessile porrect-stellate trichomes, the rays 4–7, 0.2–0.5 mm long, the
<figure><img src="image_95.png" /><figcaption>**Figure 80.** *Solanum virginianum* L. **A** herbarium specimen collected in India in 1958 (Stainton 2306, BM000900270) **B** habit (Sampath Kumar et al. 126968, India) **C** detailed view of a flower (Sampath Kumar et al. 126968, India) **D** detailed view of immature fruits (Sampath Kumar et al. 126968, India) **E** detailed view of mature fruits (field photograph, unvouchered, India). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B–D** X. Aubriot **E** S. Knapp.</figcaption></figure>

midpoints more or less equal in length to the rays; new growth densely stellate-pubescent with a dense covering of papillose glandular unicellular trichomes, yellowish green or purplish green; bark of older stems brown or purplish brown, glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, deeply lobed, the
blades (1–)3–11 cm long, 2–4 cm wide, 1–2.5 times longer than wide, elliptic on
outline, membranous to somewhat fleshy, concolorous or slightly discolorous, densely
armed on both surfaces along the midrib and major veins with prickles like those of
the stems; adaxial surface dark green and shiny, sparsely pubescent with sessile porrect-
stellate trichomes, the rays 4–7, 0.2–0.5 mm long, the midpoints longer than the rays,
moderately papillose with minute unicellular glandular papillae; abaxial surface lighter
and usually more densely stellate-pubescent with trichomes like those of the adaxial
surface; major veins 3–6 pairs, drying whitish yellow; base truncate, unevenly oblique
with one side basiscopically extended along the petiole; margins deeply lobed, the
lobes ca. 20 per side, 1.5–2 cm long, 1–1.5 cm wide, narrowly deltate to triangular,
often with secondary lobing, apically acute, the sinuses extending up to halfway to the
midrib; apex acute; petiole 1.5–5 cm long, 1/2–3/4 of the leaf blade length, very spare-
ly stellate-pubescent with sessile porrect trichomes, densely prickly with more than
10 prickles like those of the stems. Inflorescences 1.5–7 cm long, internodal or occa-
sionally almost opposite the leaves, usually unbranched, but occasionally forked, with
4–8(–10) flowers, only a few open at any one time, glabrous or with a few scattered
porrect-stellate trichomes, moderately prickly with straight, straw-coloured prickles to
1.5 cm long; peduncle 1.4–2.5 cm long, moderately prickly with prickles to 1.5 cm
long; pedicels 0.6–0.8(–1) cm long, ca. 1 mm in diameter at the base, 1–1.5 mm in
diameter at the apex, erect, glabrous or sparsely pubescent with porrect-stellate tri-
chomes like those of the stems articulated at the base; pedicel scars spaced 0.7–2 cm
apart. Buds elongate-ovoid, pointed at the tip, strongly exserted from the calyx tube
before anthesis. Flowers 5-merous, heterostylous and the plants weakly andromono-
cious, with the lowermost flower(s) long-styled and hermaphrodite, the distal flowers
short-styled and staminate. Calyx with the tube 2–3 mm long, cup-shaped, the lobes
2–3 mm long, 1.5–2 mm wide, broadly deltate abruptly narrowing to a slender subu-
late acumen 1–1.5 mm long, apically acute, densely prickly with prickles to 0.7 cm
long, the prickles denser on long-styled flowers, glabrous or sparsely stellate pubescent
with porrect trichomes like those of the pedicels. Corolla 2.5–3 cm in diameter, deep
violet and fragrant, stellate to almost rotate with abundant interpretalar tissue, lobed
1/4–1/2 of the way to the base, the lobes 0.7–1 cm long, 0.5–0.9 cm wide, broad-
deltate, spreading or slightly reflexed, glabrous or sparsely stellate-pubescent along the
midveins adaxially, moderately stellate-pubescent abaxially especially on the parts of
the corolla exposed in bud and on the tips. Stamens equal; anthers 7–9 mm long,
1.7–2 mm wide, tapering, more or less connive, bright yellow, glabrous, poricidal
at the tips, the pores directed distally, not elongating to slits with age; filament tube
minute; free portion of the filaments 0.5–1 mm long, glabrous. Ovary conical, stel-
late-pubescent with white trichomes that are soon deciduous; style 12–15 mm long
and strongly recurved in long-styled flowers, 2–2.5 mm long in short-styled flowers,
glabrous or very sparsely pubescent with a few stellate trichomes near the base, the
trichomes weak and deciduous; stigma capitate, minutely papillose. Fruit a globose
berry, 1–3 per infructescence, 1.5–2.5 cm in diameter, marbled green and white when
young, bright yellow when mature, the pericarp thick and leathery, smooth, glabrous;
<figure><img src="image_96.png" /><figcaption>**Figure 81.** Distribution of *S. virginianum*.</figcaption></figure>

fruiting pedicels 1.2–1.5 cm long, 2–3 mm in diameter at the base, 3–5 mm in diameter at the apex, unarmed or with a few small straight prickles shorter than those of the calyx, thickened and slightly fleshy, strongly recurved, the apical quarter ridged; fruiting calyx lobes elongating to 4 mm long, ca. 1/4 the length of the mature fruit, thickened and slightly fleshy in live plants, appressed to the berry. Seeds 50–100 per berry, 2.5–4 mm long, 2–2.5 mm wide, reniform, not markedly flattened, pale yellowish tan, the surfaces minutely pitted, the testal cells sinuate in outline. Chromosome number: 2n = 24 (Rajasekaran 1971; Das and Borah 2015, both as *S. xanthocarpum*).

**Distribution (Fig. 81).** *Solanum virginianum* occurs from the Arabian Peninsula (Saudi Arabia, Yemen) to China and Myanmar.

**Ecology and habitat.** *Solanum virginianum* is a plant of disturbed places, such as roadsides, streambeds and edges of fields, and often occurs in large populations and forms impenetrable mats; from sea level to 3,100 m elevation (highest elevation from Afghanistan).

**Common names and uses.** China. mao guo qie (Zhang et al. 1994). India. Bihar: rengani, kataila (Varma 1981), as *S. surattense*); Haryana: kateli (Jain et al. 2000, as *S. surattense*); Kerala: kandankathiri (Mohanan and Henry 1994, as *S. surattense*); Rajasthan: baiga-kateli, bhurangi, dhaturi [Hindi] (Singh 1991); Tamil Nadu: kandankathiri [Tamil] (Matthew 1983); kandangattiri [Tamil] (Henry et al. 1987).
*Solanum virginianum* is widely used in medicine as one of the “dasa mulikas” of traditional Indian medicine; used in treatment of asthma, cough, fever etc. (Tadul-ingam and Venkatanarayana 1932; Jain et al. 2000). The whole plant is considered highly medicinal (Singh 1988).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (7,899,801 km², LC); AOO (336 km², EN). *Solanum virginianum* is a weedy species, forming dense patches in open disturbed areas and along dry riverbeds. Tropical Asia represents the easternmost part of its range.

**Discussion.** *Solanum virginianum* is a seriously mis-named species, Linnaeus (1753: 187) thought it originated in the then-English colony of Virginia (“Habitat in America” — taken from the polynomials cited in synonymy), while in fact it is not found there except as a recent introduction in ships ballast. With its almost glabrous leaves, prominent straw-coloured prickles, large, fragrant purple flowers and weedy habit it is easily recognised and unlikely to be confused with any other species in the region. In the field the flowers of *S. virginianum* are sweetly fragrant but little is known about the pollination of this, or any other spiny solanum in the region (except *S. insanum*, see Davidar et al. 2015).

The introduced *S. aculeatissimum* and *S. viarum* are similarly prickly, but the prickles are not as long and in those species the leaf pubescence is apparently simple, not stellate. The flowers of both these taxa are white or pale greenish white and the leaves are ovate to broadly elliptic rather than narrowly elliptic in outline.

*Solanum virginianum* is sister to the main set of African clades of spiny solanums (Aubriot et al. 2016a), albeit this relationship is only weakly supported and is an isolated lineage; no other species clustered with it in the molecular analyses. Its affinities apparently lie with African rather than tropical Asian lineages such as the ‘Sahul-Pacific clade’.

Hepper and Jaeger (1986) inadvertently and effectively lectotypified *S. surattense* with citation of “Type.: Pakistan, Surat, [....] *Garcin* in Hb. *Burmann*” (G-Herb. Delessert!). The Burmann herbarium is held in G-PREL; pre-Linnaean collections were extracted from the general herbarium after 1992, and are no longer part of the Delessert herbarium. Two sheets of *Garcin s.n.* are housed in G-PREL necessitating a second lectotypification step. We have chosen the more complete of these sheets (G00811278) as the lectotype for *S. surattense*.

We have lectotypified *S. armatum* using the only authentic Forsskål material we have found in the Kiel University herbarium (KIEL0005103), although the specimen does not have an indication of the locality.

*Solanum jacquinii* is an illegitimate superfluous name because *S. virginianum* was cited in synonymy indirectly by citation of Jacquin’s (1786–1793) illustration, which itself cited Linnaeus. The specimen in the Willdenow herbarium at Berlin (B-W-04384-00 0) bears a copy of Willdenow’s short description on the cover and is certainly that used in the description.

In the description of *S. xanthocarpum* var. *schraderi* Dunal (1852) cited a number of names and elements, including *S. xanthocarpum* itself, suggesting he was considering this the typical variety. We have elected to typify the name however, since the
citations are not clear, and choose as the lectotype the G-DC duplicate of *Wallich cat. 2618b* (G00130292) that is specifically cited in the protologue.

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. We have selected the specimen at E (E00284473) that corresponds to the description, collector and locality (Léveillé 1913), as the lectotype for *S. mairei*.

**Specimens examined.** See Suppl. materials 1–3.

50. *Solanum wightii* Nees, Trans. Linn. Soc. London 17(1): 51. 1834.
Fig. 82

*Solanum hohenackeri* Van Heurck & Müll.Arg., Observ. Bot. (Van Heurck) 87. 1870. Type. India. Tamil Nadu: Nilgiris, "in montibus Nilagiri", *R.F. Hohenacker 1076* (lectotype, designated here: BR [AWH10071212]; isotypes: BM [BM000778309], G [G00442609, G00442943, G00442944], HAL [HAL0010834], K [K000441384], L [0403690], LE [2 sheets], MEL [MEL2446521], P [P00055713, P00055714], W [acc. # 0000605]).

*Solanum pulneyensis* Soosairaj, Adansonia sér. 3, 43(21): 236. 2021. Type. India. Tamil Nadu: Dindigul district, Palani Hills National ark, Thonimalai, ca. 1300 m, 29 Jan 2018. *S. Soosairaj* 2514 (holotype: RHT [acc. # 076723]; isotype: MH).

**Type.** INDIA. Sin. loc., "Peninsula Ind. orientalis", *R. Wight 1576/126* (lectotype, designated here: GZU [GZU000255932]; isotypes: BM [BM000900156], E [E00179475, E00179476, E00179477], G [G00442945], K [K000441376, K000441378, K000441379], LE [LE00017069]).

**Description.** Scrambling herbs or shrubs to 2 m tall, armed or unarmed. Stems erect or spreading, terete, prickly and stellate-pubescent; prickles, if present, to 3 mm long, 1.5-2 mm wide at the base, slightly curved, pale yellowish tan; pubescence of sessile to very short-stalked porrect-stellate trichomes, the stalks, if present, to 0.2 mm, the rays 4-6, ca. 0.5 mm long, the midpoints 2-4 celled, to 2 mm long, glandular tipped, drying with violet tinge at cell junctions (trichomes described as black to reddish *Clarke 10793*); new growth densely stellate-pubescent, the trichomes tangled, soon deciduous and the stems glabrate; bark of older stems ashy white. Sympodial units difoliate, the leaves not geminate. Leaves simple, shallowly lobed, the blades 2-9 cm long, 1.7-6.8 cm wide, 1-1.3 times longer than wide, ovate to broadly triangular, widest in the lower third, chartaceous, somewhat discolorous, unarmed or very occasionally sparsely armed along the midrib and major veins with small prickles; adaxial surface evenly and densely pubescent with sessile and very short-stalked porrect-stellate trichomes, the rays 4-8, to 0.5 mm long, glandular at the tips, the midpoints 2-4-celled, to 2 mm long, glandular at the tips; abaxial surface with similar porrect-stellate trichomes, but these denser especially along the veins; major veins
<figure><img src="image_97.png" /><figcaption>**Figure 82.** *Solanum wightii* Nees A herbarium specimen collected in India in 1937 (*Vine* 176, BM000900152) B detailed view of a flower (field photograph, unvouchered, India) C detailed view of a fruit (field photograph, unvouchered, India). Photograph credits: A CC-BY, © copyright The Trustees of the Natural History Museum, London B, C G. Gnanasekaran.</figcaption></figure>

3–4 pairs, densely pubescent; base abruptly truncate to cordate, somewhat oblique; margins shallowly lobed, the lobes 3–4 on each side, to 0.5 cm long, broadly deltate, apically rounded, the sinuses less than halfway to the midrib; apex acute to obtuse; petioles 1–4 cm long, ca. half as long as the leaf blades, unarmed or with a few prickles, densely stellate pubescent like the stems. Inflorescences 0.3–1 cm long, internodal and lateral, unbranched, with 1–3 flowers, only 1 or 2 flowers open at any one time, pubescent with mixed sessile and short-stalked stellate-porrect trichomes like those of the stems, with multicellular midpoints to 2 mm long, unarmed; peduncle absent to 0.2 cm long; pedicels 2.5–4 cm long, ca. 0.7 mm in diameter at the base, ca. 0.7 mm in diameter at the apex, spreading and slightly nodding at anthesis, unarmed or with a few prickles, more sparsely stellate-pubescent than the inflorescence axes, articulated
at the base; pedicel scars tightly spaced ca. 1 mm apart. Buds elongate and tapering, curved, strongly exserted from the calyx before anthesis. Flowers 5-merous, heterostylenous and the plants andromonoecious, with the distal flower(s) short-styled and smaller than the hermaphroditic flowers. Calyx with the tube 3.5-4 mm long, conical, sparsely prickly, the lobes 6-8 mm long, ca. 1 mm wide, long-triangular to lanceolate, unarmed, densely stellate-pubescent with mixed sessile and short-stalked porrect-stellate trichomes with multicellular glandular midpoints like those of the pedicels, the pubescence denser than that of the pedicels. Corolla (2-3.5-5 cm in diameter, violet or deep purple, rotate-stellate, lobed to 1/3 of the way to the base, abundant interpretalar tissue present, the lobes (10-14-17 mm long, (11-14-18 mm wide, broaddeltate, spreading or somewhat campanulate at anthesis, mostly glabrous adaxially but with a few stellate trichomes along the petal midvein, densely stellate-pubescent abaxially with densely tangled sessile trichomes where exposed in bud, these densest at the tips, the interpretalar tissue glabrous. Stamens markedly unequal; anthers 3 long and 2 short, the long anthers 12-15 mm long, ca. 2 mm wide, strongly curved and tapering, the short anthers ca. 10 mm long, 0.6-0.7 mm wide, straight or slightly curved, all anthers 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 1-1.5 mm long, glabrous. Ovary conical, glabrous; style ca. 5 mm long in short-styled flowers, 12-15 mm long, in long-styled flowers, strongly curved, glabrous; stigma capitate, the surfaces minutely papillose. Fruit a globose berry, 1-2 per infructescence, 1.1-1.5 cm in diameter, completely enclosed in the accrescent calyx, yellowish brown when ripe, drying and breaking into 4 irregular valves when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 3-3.5 cm long, 1-1.5 mm in diameter at the base, 3-4.5 mm in diameter at the apex, unarmed, somewhat woody, sharply deflexed and pendent; fruiting calyx strongly accrescent, the lobes often breaking off. Seeds 20-30 per berry, 4-5 mm long, 3-3.5 mm wide, flattened reniform, pale tan or yellowish brown, the surfaces minutely pitted, the testal cells with straight walls, pentagonal in outline, the margins incrassate. Chromosome number: not known.

**Distribution (Fig. 83).** *Solanum wightii* is endemic to the mountains of southeastern India in the states of Tamil Nadu and Andhra Pradesh.

**Ecology and habitat.** *Solanum wightii* occurs in dry forests and forest margins, from 850 to 2,200 m elevation.

**Common names and uses.** None recorded.

**Preliminary conservation status (IUCN 2019).** Vulnerable (VU). EOO (32,261 km², VU); AOO (44 km², EN). *Solanum wightii* is rather narrowly distributed and occurs in open grassy areas subject to human disturbance; it occurs in the proposed Palani Hills National Park, so there is afforded some degree of protection.

**Discussion.** *Solanum wightii* is a beautiful, distinctive Indian endemic with large, zygomorphic flowers and berries enclosed in accrescent calyces that are borne on long, strongly deflexed pedicels. The only other species in India with purple zygomorphic flowers is *S. pubescens* which has sticky pubescence, smaller flowers with a single long stamen
<figure><img src="image_98.png" /><figcaption>**Figure 83.** Distribution of *S. wightii*.</figcaption></figure>

rather than three, and more fruit on each infructescence borne on shorter pedicels. The berry of *S. wightii* has been recorded as dry and dehiscent (breaks into four valves, *Anon. s.n.*, in MH; described as “subcapsular” in Soosairaj et al. 2021) like members of the Androceras clade (Whalen 1979) and the Elaeagnifolium clade (see Knapp et al. 2017). Field studies of dispersal in *S. wightii* are needed; this type of dry berry is also found in members of the Leptostemonum Clade from Australia (Symon 1981; Martine et al. 2019) where dispersal can be either via a censer (shaking) mechanism or as trample burrs.

Aubriot et al. (2016a) suggest *S. wightii* is sister to *S. praetermissum*, but the relationship is poorly supported, and additional analyses are necessary.

Ramachandran and Viswanathan (2010) reported specimens of *S. wightii* from Sethukadai in the Namakkal District of Tamil Nadu as *S. cordatum*. The illustration in their paper is clearly of *S. wightii* although the description appears to be a mixture of information from published works and the specimens they used. The recently described *S. pulneyensis* (Soosairaj et al. 2021) clearly falls within the range of variation of *S. wightii*, and the holotype specimen at RHT corresponds to *S. wightii*. The character used to distinguish the two species is the dry pseudo-capsular berry, something that is known to occur in *S. wightii* (see above).

We have selected the sheet of *Wight 1576/126* from Nees van Esenbeck’s personal herbarium (GZU000255932) that is annotated in his handwriting as “S. Wightii”
n.sp." as the lectotype of *S. wightii*; this collection is widely duplicated. After its initial description (Nees van Esenbeck 1834), Nees van Esenbeck (1836) published a more extensive treatment of *S. wightii*, in which he clarified his ideas about its relationships and illustrated the plant.

Two collections and two herbaria were cited in the protologue (Van Heurck 1870) of *S. hohenackeri* – “Hohenacker 1076, 1417! in hb. Van Heurck et hb. DC”. The van Heurck herbarium, previously held in Antwerp at AWH, has now been acquired by BR, but sheets are still barcoded with the AWH herbarium code. We have selected duplicate of *Hohenacker 1076* held in BR (AWH10071212) as the lectotype for *S. hohenackeri*; this specimen is well-preserved and duplicates of *Hohenacker 1076* are widely distributed.

**Specimens examined.** See Suppl. materials 1–3.

**51. Solanum wrightii Benth., Fl. Hongk. 243. 1861.**

Fig. 73F–H

Names based on tropical Asian types only; for complete synonymy see Vorontsova and Knapp 2016.

**Type.** CHINA. “Hongkong”, 1853–1856, *C. Wright* 489 (lectotype, designated here: K [K000545757]; isotypes: GH [00077826], US [acc. # 79466, barcode 00731429]).

**Description.** Vorontsova and Knapp (2016: 372–375); http://www.solanaceae-source.org/solanaceae/solanum-wrightii.

**Distribution.** *Solanum wrightii* has been recorded from several areas in tropical Asia in cultivation as a street tree; it is native to Bolivia but widely cultivated for ornament and as a shade tree in coffee plantations. *Solanum wrightii* has been formally assessed as of Least Concern (LC) for the IUCN Red List (Botanical Gardens Conservation International (BGCI) and IUCN SSC Global Tree Specialist Group 2018).

**Common names.** China. da hua qie (Zhang et al. 1994).

**Discussion.** Although first described from Hong Kong, *S. wrightii* is a plant of the eastern slopes of the Andes in Bolivia. It is a member of the Crinitum clade (sensu Stern et al. 2011; Gagnon et al. 2022), characterised by large flowers and fruits, and usually large, repand leaves. In tropical Asia it is widely planted as a street and shade tree, and also sometimes as living fencing. It is not known to naturalise and is rarely collected out of cultivation.

Bentham (1861) did not state where he had examined herbarium material for his "Flora of Hong-Kong", and subsequent authors have assumed the Kew sheet of *Wright 849* is the holotype for this name (e.g., Vorontsova and Knapp 2016). This is likely to have been the only material Bentham saw, as Wright's collections were distributed from the Smithsonian, but since he did not cite a single collection in a single herbarium it is necessary to designate a lectotype for this widely used name (McNeill 2014); it is surprising it has not been done inadvertently previously. We lectotypify *S. wrightii* here in conformity with previous usage, selecting the Kew duplicate of *Wright 849* as the lectotype (K000545757).

**Specimens examined.** See Suppl. materials 1–3.
<table><caption>**Table 3.** Preliminary threat status for native spiny solanums of tropical Asia (introduced species not assessed here, nor is the cultivated *S. melongena*). Also see individual species treatments for more details; *S. comitis*, *S. kachinense*, *S. putii* and *S. sulawesi* are known only from the types or too few collections to calculate a preliminary assessment and are therefore assessed as Data Deficient (DD). The highly fragmented island nature of these species' distributions means more detailed assessments should be undertaken in the future once local herbaria are consulted (e.g., Delves 2021).</caption><thead><tr><th>Species</th><th>EOO (km²)</th><th>AOO (km²)</th><th>Assessment</th></tr></thead><tbody><tr><td>Solanum arundo Mattei</td><td>1,277,240</td><td>216</td><td>LC</td></tr><tr><td>Solanum barbisetum Nees</td><td>1,244,077</td><td>248</td><td>LC</td></tr><tr><td>Solanum camranhense Dy Phon & Hul</td><td>555</td><td>16</td><td>EN</td></tr><tr><td>Solanum comitis Dunal</td><td>n/a</td><td>n/a</td><td>DD</td></tr><tr><td>Solanum cordatum Forssk.</td><td>Formally assessed for the IUCN Red List</td><td></td><td>LC (Knapp 2021)</td></tr><tr><td>Solanum cyanocarphium Blume</td><td>1,492,571</td><td>112</td><td>LC</td></tr><tr><td>Solanum deflexicarpum C.Y.Wu & S.C.Huang</td><td>65,775</td><td>16</td><td>NT</td></tr><tr><td>Solanum dunalianum Gaudich.</td><td>887,043</td><td>100</td><td>LC</td></tr><tr><td>Solanum forskalii Dunal</td><td>Formally assessed for the IUCN Red List</td><td></td><td>LC (Knapp 2021)</td></tr><tr><td>Solanum giganteum Jacq.</td><td>Formally assessed for the IUCN Red List</td><td></td><td>LC (BGCI and IUCN SSC 2020)</td></tr><tr><td>Solanum graciliflorum Dunal</td><td>131,936</td><td>16</td><td>NT</td></tr><tr><td>Solanum harmandii Bonati</td><td>n/a</td><td>n/a</td><td>DD</td></tr><tr><td>Solanum hovei Dunal</td><td>173,911</td><td>564</td><td>LC</td></tr><tr><td>Solanum insanum L.</td><td>10,914,652</td><td>1,052</td><td>LC</td></tr><tr><td>Solanum involucratum Blume</td><td>1,868,808</td><td>216</td><td>LC</td></tr><tr><td>Solanum kachinense X.Aubriot & S.Knapp</td><td>n/a</td><td>n/a</td><td>DD</td></tr><tr><td>Solanum lasiocarpum Dunal</td><td>6,950,615</td><td>1,032</td><td>LC</td></tr><tr><td>Solanum melongena L.</td><td>cultivated</td><td></td><td>Not assessed</td></tr><tr><td>Solanum miyakojimense T.Yamaz. & Takushi</td><td>641</td><td>40</td><td>EN</td></tr><tr><td>Solanum multiflorum Roth</td><td>669,122</td><td>324</td><td>LC</td></tr><tr><td>Solanum nienkui Merr. & Chun</td><td>64,999</td><td>72</td><td>NT</td></tr><tr><td>Solanum peikuense S.S.Ying</td><td>10,013</td><td>44</td><td>VU</td></tr><tr><td>Solanum poka Dunal</td><td>1,202,780</td><td>68</td><td>LC</td></tr><tr><td>Solanum praetermissimum Kerr ex Barnett</td><td>1,191,003</td><td>88</td><td>LC</td></tr><tr><td>Solanum procumbens Lour.</td><td>1,575,888</td><td>164</td><td>LC</td></tr><tr><td>Solanum pseudosaponaceum Blume</td><td>2,387,194</td><td>160</td><td>LC</td></tr><tr><td>Solanum pubescens Willd.</td><td>996,977</td><td>272</td><td>LC</td></tr><tr><td>Solanum putii Kerr ex Barnett</td><td>n/a</td><td>n/a</td><td>DD</td></tr><tr><td>Solanum retrorsum Elmer</td><td>115,321</td><td>48</td><td>NT</td></tr><tr><td>Solanum robinsonii Bonati</td><td>610</td><td>20</td><td>EN</td></tr><tr><td>Solanum schefferi F.Muell.</td><td>1,012,899</td><td>32</td><td>LC</td></tr><tr><td>Solanum sulawesi X.Aubriot & S.Knapp</td><td>n/a</td><td>n/a</td><td>DD</td></tr><tr><td>Solanum torvoideum Merr. & L.M.Perry</td><td>2,372,285</td><td>212</td><td>LC</td></tr><tr><td>Solanum trilobatum L.</td><td>2,326,480</td><td>528</td><td>LC</td></tr><tr><td>Solanum vagum Nees</td><td>75,338</td><td>76</td><td>LC</td></tr><tr><td>Solanum violaceum Ortega</td><td>10,078,363</td><td>988</td><td>LC</td></tr><tr><td>Solanum wightii Nees</td><td>7,899,801</td><td>336</td><td>VU</td></tr></tbody></table>

## Doubtful and suppressed names

*Solanum canescens* Blume, Bijdr. Fl. Ned. Ind. 13: 701. 1826. No specimens or locality cited; Blume cited no specimens or localities, and it is unclear if his description is a re-description of *S. incanum* sensu Forssk. (= *S. incanum* L., see Vorontsova and Knapp 2016), or if he based it on Indonesian specimens. The protologue states “(aff. *S. sancto*; *S. incanum* Forssk, forsan eadem species)” perhaps indicating this is a replacement name for the unpublished *S. incanum* sensu Forssk., a bibliographic reference to *S. incanum* L. and not intended as a new name. No material in Leiden
is annotated with this name, nor have we found any other material that Blume
used annotated as *S. canescens*. We are treating this name as doubtful.

*Solanum ferox* L., Sp. Pl. ed. 2: 267. 1762, nom. utique rej. (suppressed name; Knapp 2011, see Shenzhen Code Appendices at https://naturalhistory2.si.edu/botany/codes-proposals/index.cfm).

*Solanum indicum* L., Sp. Pl. 187. 1753, nom. utique rej. (suppressed name; Hepper 1978, see Shenzhen Code Appendices at https://naturalhistory2.si.edu/botany/codes-proposals/index.cfm).

*Solanum sodomeum* Dunal, Prodr. [A. P. de Candolle] 13(1): 365. 1852, nom. utique rej. (suppressed name; Hepper 1978, see Shenzhen Code Appendices at https://naturalhistory2.si.edu/botany/codes-proposals/index.cfm).

"Names" (designations) not validly published

*Solanum album* Noronha, Verh. Batav. Genootsch. Kunst. 5(Art. 4): 84. 1791, nomen nudum, name in list, no description or diagnosis (Art. 38.1). [Page 84 in 1827 exact reprint, read 1790, printed 1791] = *S. melongena* L.

*Solanum blattaroïoides* Koen. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 127. 1852, pro syn. *Solanum pubescens* Willd. = *S. pubescens* Willd. (herbarium name on König s.n. specimen of *S. pubescens* [BM000900094]).

*Solanum canaranum* Miq. ex C.B.Clarke, Fl. Brit. India [J. D. Hooker] 4(10): 236. 1883, pro syn. *Solanum trilobatum* L. = *S. trilobatum* L.

*Solanum coccineum* Dunal, Prodr. [A. P. de Candolle] 13(1): 310. 1852, pro syn. *Solanum indicum* Nees var. *sinuato-lobatum* Dunal = *S. violaceum* Ortega

*Solanum diffusum* Roxb. ex Wall., Hort. Bengal. 17. 1814, nomen nudum, no description or diagnosis (Art. 38.1) = *Solanum virginianum* L. (i.e., K001116578)

*Solanum ferox* Jungh. ex Miq., Fl. Ned. Ind. 2: 650. 1857, pro syn. *Solanum junghuhnii*
Miq. = *S. violaceum* Ortega

*Solanum fuscum* B.Heyne ex Wall., Numer. List [Wallich] no. 2622b. 1831, nomen nudum [probably from Heyne's herbarium], no description or diagnosis (Art. 38.1) = *S. trilobatum* L.

*Solanum indicum* Roxb., Hort. Bengal. 17. 1814, nomen nudum, no description or diagnosis (Art. 38.1); identity uncertain, could apply to any number of species from the region; probably a reference to *S. indicum* L. (nom. utique rej.)

*Solanum intermedium* Dunal, Prodr. [A. P. de Candolle] 13(1): 311. 1852, pro syn. *Solanum hovei* Dunal (herbarium name on holotype specimen of *S. hovei*, *Hove s.n.* [BM000900293]) = *S. hovei* Dunal

*Solanum involucratum* Kurz, Forest Fl. Burma 2: 224. 1877, nomen nudum, no description or diagnosis (Art. 38.1), probably not intended as a new name; probably = *S. barbisetum* Nees

*Solanum lividum* Willd. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 310. 1852, pro syn. *Solanum indicum* Nees var. *sinuato-lobatum* Dunal = *S. violaceum* Ortega

*Solanum longum* Roxb., Hort. Bengal. 16. 1814, nomen nudum, name in list, no description or diagnosis (Art. 38.1) = *Solanum melongena* L.
*Solanum melongena* L. subsp. *agrestis* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 318. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1) = *S. melongena* L.

*Solanum melongena* L. var. *agrestis* Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984. Not validly published, no description, diagnosis or type (seems to be homotypic with his subsp. *agrestis*) = *S. melongena* L.

*Solanum melongena* L. var. *americanum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 317. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *meridionale* Filov var. *americanum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *arabicum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 318. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *subspontaneum* Filov var. *arabicum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *azerbaijanicum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 313. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *occidentale* Haz. var. *azerbaijanicum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *bulgaricum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 313. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *occidentale* Haz. var. *bulgaricum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *europaeum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 314. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *occidentale* Haz. var. *europaeum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *giganteum* (Alef.) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as “*gigantea*”; not a new name, re-publication of Alefeld’s name at the same rank. = *S. melongena* L.

*Solanum melongena* L. var. *insanum* (L.) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 321. 1958, not validly published, no full and direct references to original place of publication (Art. 41.5); also nom. illeg., later homonym as *Solanum melongena* L. subsp. *agrestis* Filov var. *insanum* (L.) Filov = *S. insanum* L.

*Solanum melongena* L. var. *leucoum* (Alef.) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984; not a new name, re-publication of Alefeld’s name at the same rank. = *S. melongena* L.

*Solanum melongena* L. subsp. *meridionale* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 315. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1) [cites as syn. var. *insanum* Nees, var. *esculentum* Bailey] = *S. melongena* L.

*Solanum melongena* L. subsp. *orientale* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 306. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1) = *S. melongena* L.

*Solanum melongena* L. var. *ovigera* Lam., Tabl. Encycl. 2: 19. 1794, the name that occurs in the agricultural and some other indices, is not presented as a binomial in Lamarck (1794). The protologue reads “β Idem fructu ovato, alb. Melongena ovigera. Ex Hort. Paris”. We do not consider this to be the coining of a new name (Art. 23). A specimen in the Lamarck herbarium at P (P00357638) labelled “Solanum melongena ovigera/Dict. No. 43” in Lamarck’s hand is probably original material associated with this polynomial. = *S. melongena* L.

*Solanum melongena* L. var. *ovigera* Pers., Syn. Pl. [Persoon] 1: 221. 1805, this name that occurs in some indices was not published as a binomial, the protologue reads
"β Melongena (ovigera)", along with citations of Tournefort (1700), Miller (1768) and Lamarck ("encl.") We do not consider this name to be validly published (Art. 23). = *S. melongena* L.

*Solanum melongena* L. var. *pekinense* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 310. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *orientale* Filov var. *pekinense* Filov= *S. melongena* L.

*Solanum melongena* L. var. *palestinicum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 316. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *meridionale* Filov var. *palestinicum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *racemosum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 320. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1); as *Solanum melongena* L. subsp. *agrestis* Filov var. *racemosum* Filov = *S. melongena* L.

*Solanum melongena* L. var. *stenoleucum* (Alef.) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 104. 1984, as "*stenoleuca*"; not a new name, re-publication of Alefeld's name at the same rank.

*Solanum melongena* L. subsp. *subspontaneum* Filov, Kult. Fl. SSSR (Zhukovskii) 10: 318. 1958., not validly published, no description or diagnosis in Latin (Art. 39.1) [cites in synonymy subsp. *italo-arabicum* Filov, perhaps a herbarium name?] = *S. melongena* L.

*Solanum melongena* L. var. *variegatum* (Alef.) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "*variegata*"; not a new name, re-publication of Alefeld's name with spelling correction. = *S. melongena* L.

*Solanum melongena* L. var. *violaceum* (Alef.) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "*violacea*"; not a new name, re-publication of Alefeld's name at the same rank. = *S. melongena* L.

*Solanum nelsonii* Zipp. ex Span., Linnaea 15: 337. 1841, pro syn. *Solanum indicum* L. = *S. violaceum* Ortega.

*Solanum ovigerum* Dunal var. *album* Sweet, Hort. Brit., ed. 2: 387. 1830, nomen nudum, listed in table, no description or diagnosis (Art. 38.1) = *S. melongena* L.

*Solanum ovigerum* Dunal var. *luteum* Sweet, Hort. Brit., ed. 2: 387. 1830, nomen nudum, listed in table, no description or diagnosis (Art. 38.1) = *S. melongena* L.

*Solanum ovigerum* Dunal var. *ruber* Sweet, Hort. Brit., ed. 2: 387. 1830, nomen nudum, listed in table, no description or diagnosis (Art. 38.1) probably = *S. aethiopicum* L.

*Solanum ovigerum* Dunal var. *violaceum* Sweet, Hort. Brit., ed. 2: 387. 1830, nomen nudum, listed in table, no description or diagnosis (Art. 38.1) = *S. melongena* L.

*Solanum ovoideum* Zipp. ex Miq., Fl. Ned. Ind. 2: 647. 1857, pro syn. *Solanum ferox* L. var. *involucratum* (Blume) Miq. = *S. involucratum* Blume

*Solanum peikuoense* S.S.Ying, Quarterly J. Exp. For. Natl. Taiwan Univ. 11(1): 14. 1997, as "peikuoensis", name not validly published; the words holotype or the equivalent were not used contrary to Art. 40.6. = *S. peikuoense* S.S.Ying

*Solanum prostratum* Raeusch., Nomencl. Bot. 60. 1797, nomen nudum, no description or diagnosis (Art. 38.1); identity uncertain.

*Solanum pubescens* B.Heyne ex Walp., Repert. Bot. Syst. (Walpers) 3: 83. 1844, pro syn. *Solanum indicum* L. var. δ = *S. pubescens* Willd.
*Solanum quercifolium* Banks ex Dunal, Prodr. [A. P. de Candolle] 13(1): 345. 1852, pro syn. *Solanum poka* Dunal = *S. pseudosaponaceum* Blume (herbarium name on Banks & Solander s.n. specimen of *S. pseudosaponaceum* [BM000886238]).

*Solanum racemosum* Noronha, Verh. Batav. Genootsch. Kunst. 5(Art. 4): 26. 1790, nomen nudum, no description or diagnosis (Art. 38.1); identity uncertain, probably misuse of *S. racemosum* Jacq., a Caribbean species that is a synonym of *S. bahamense* L.

*Solanum reflexispinosum* Merr., Fl. Arfak Mts. [Gibbs] 178. 1917, pro syn. *Solanum retrorsum* Elmer (herbarium name on a specimen of *S. retrorsum*, Williams 1074 [K000195907]) = *S. retrorsum* Elmer

*Solanum serpentinum* Noronha, Verh. Batav. Genootsch. Kunst. 5(Art. 4): 84. 1791, nomen nudum, name in list, no description or diagnosis (Art. 38.1). = *S. melongena* L.

*Solanum serpentinum* Desf., Cat. Pl. Horti Paris 397. 1829, nomen nudum; on page 115 in list as "serpentinum H."; on page 397 as "Solanum serpentinum/ Fructus elongatus serpentinus. An varietas S. Melongenae?"; refers to a horticultural name and not with the intention of coining a new name, this description is much less detailed than those in which Desfontaines did intend new names in this work. = *S. melongena* L.

*Solanum surattense* Burm.f. var. *awanicum* Yousef, Mir A.Khan & Shinwari, Pakistan J. Bot 41(5) 2099. 2009, not validly published, no description or diagnosis in Latin (Art. 39.1). = *S. virginianum* L.

*Solanum tomentosum* Herb. Hasselt. ex Miq., Fl. Ned. Ind. 2: 654. 1857, pro syn. *Solanum undatum* Lam. = *S. insanum* L.

*Solanum undatum* Poir., Encycl. (Lamarck) 4: 301. 1797, not a new name, Poiret makes clear reference to Lamarck's protologue.

*Solanum vagum* B.Heyne ex Wall., Numer. List [Wallich] n. 2624. 1831, nomen nudum, no description or diagnosis (Art. 38.1) = *Solanum vagum* Nees.

*Solanum violaceum* DC. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 359. 1852, pro syn. *Solanum undatum* Lam. var. *violaceum* Dunal = *S. insanum* L.

*Solanum vincentii* Delile ex Dunal, Prodr. [A. P. de Candolle] 13(1): 310. 1852, pro syn. *Solanum indicum* Nees var. *sinuato-lobatum* Dunal = *S. violaceum* Ortega

*Solanum virginianum* Jacq., Icon. Pl. Rar. [Jacquin] 2: 11, t. 332. 1792, not a new name, Jacquin (1792) makes clear reference to the Linnaean protologue.

## Acknowledgements

Many people have provided invaluable assistance over the course of preparation this revision. First and foremost, we would like to thank the curators of the herbaria mentioned in the text and in the Suppl. materials 1–3 for loan of or permission to examine specimens in their care. We particularly thank Sovanmoly Hul (P), Shuichi Nemoto (TI), Kuo-Fang Chung (HAST), Abdulrokhman Kartonegoro (BO), Caroline Loup (MPU), Dietrich Ober (KIEL), Sarah Phillips (K), Roxali Bijmoer (L), Anand Kumar (CAL) and Nicolas Fumeaux (G) for providing specimen scans and information not available online; Tiina Särkinen (E) kindly prepared the maps. We thank the following herbaria and their institutions for permission to publish photographs of herbarium specimens from
their collections: BM (Natural History Museum), CAL (The Director, Botanical Survey of India, Kolkata), E (Royal Botanic Garden Edinburgh), K (Royal Botanic Gardens Kew), L (Naturalis Biodiversity Center), P (Muséum national d'Histoire naturelle), TI (University of Tokyo) and W (Naturhistorisches Museum, Wien); details of copyright given in the individual figures. Permission to use photographs of living plants was kindly granted by Prashant Awale (India), Ehoarn Bidault (MO), Indiana Coronado González (via MO), Gunadayalan Gnanasekaran (Madras Christian College), Sovanmoly Hul (P), Prabhu Kumar (India), Olga Martha Montiel (MO), Sushant More (India), Maxim Nuraliev (Lomonosov Moscow State University, Vietnam-Russian Tropical Center), Dennis Pedersen (Aarhus University), Pieter B. Pelser (University of Canterbury, New Zealand), Stephen Stern (Colorado Mesa University), D. Tandang (National Museum of the Philippines), Ravichandran Vellingiri (Botanical Survey of India), Maria S. Vorontsova (K) and Ming-I Weng (Taiwan). Mark Watson (E), Jacek Wajer (BM), and Henry Noltie (E) provided assistance with historical material from India; Franco Chiarini (CORD) generously shared chromosome data for many of these species; Ranee Prakash (BM) kindly facilitated verification of many specimens in Indian herbaria; Jin Xiu Wang and Tan Gang Gao (both PE) facilitated SK's fieldwork in China; Henrik Balslev hosted XA in the field in Thailand; staff of the Bangkok Forest Herbarium (BKF) and of Khon Kaen University (KKU) kindly assisted XA in the field in Thailand; staff of the Botanical Survey of India (BSI) centres in Coimbatore and the Blatter Herbarium (BLAT) assisted XA in the field in southern India; staff of the BSI Centres in Dehradun, Kolkata, Sikkim, and Port Blair kindly assisted SK in the field in northern India; Ranee Prakash (BM) accompanied SK in the field in northern India, and with Jacek Wajer (BM), XA in southern India. Many others have contributed to this study in ways too numerous to mention – thank you all. This work was funded by the National Science Foundation's Planetary Biodiversity Inventory programme (DEB-0316614 'PBI Solanum' – a worldwide treatment) http://www.solanaceae source.org to SK; the SYNTHESYS Project http://www.synthesys.info/ which was financed by European Community Research Infrastructure Actions under the FP6 and FP7 "Structuring the European Research Area" Programme financed herbarium visits; NHM Special Funds provided support for collecting in India (XA) and China (SK); the UK Government Department of Business, Energy and Industrial Strategy (BEIS) Rutherford Fund provided support for travel to India (SK). Stephen Stern, Donald McClelland, and Leandro Giacomin greatly improved the manuscript with their reviews.

## References

Al Wadi HM (2002) Cytomorphology of three *Solanum* species in the Aseer Region, South-West Saudi Arabia. Online Journal of Biological Sciences 2(2): 106–108. https://doi.org/10.3923/jbs.2002.106.108

Alefeld FGC (1866) Landwirthschaftliche Flora. Weigandt & Hempel, Berlin.

Anderson GJ, Symon DE (1989) Functional dioecy and andromonoecy in *Solanum*. Evolution; International Journal of Organic Evolution 43(1): 204–219. https://doi.org/10.1111/j.1558-5646.1989.tb04218.x
Anderson GJ, Bernardello G, Bohs L, Weese T, Santos-Guerra A (2007) Phylogeny and bio-
geography of the Canarian *Solanum vespertilio* and *S. lidii* (Solanaceae). Anales del Jardin
Botanico de Madrid 63: 159–167. https://doi.org/10.3989/ajbm.2006.v63.i2.5

Anderson GJ, Anderson MKJ, Patel N (2015) The ecology, evolution, and biogeography of
dioecy in the genus *Solanum*: With paradigms from the strong dioecy in *Solanum polyg-
mum*, to the unsuspected and cryptic dioecy in *Solanum conocarpum*. American Journal of
Botany 102(3): 471–486. https://doi.org/10.3732/ajb.1400486

Aparna S, Ved DK, Lalitha S, Venkatasubramanain P (2012) Botanical identity of plant sources of Dašūamūla drugs through an analysis of published literature. Ancient Science of Life 32(1): 3–10. https://doi.org/10.4103/0257-7941.113790

Arousseau L (1922) Jules Harmand. In: École française d'Extrême-Orient (Ed.) Bulletin de l'École française d'Extrême-Orient. F.-H. Schneider, Hanoi, 22, 402–404. https://doi.org/10.3406/befeo.1922.2980

Aubriot X, Singh P, Knapp S (2016a) Tropical Asian species show the Old World clade of “spiny solanums” (subgenus *Leptostemonum* pro parte: Solanaceae) is not monophyletic. Botanical Journal of the Linnean Society 181(2): 199–223. https://doi.org/10.1111/boj.12412

Aubriot X, Loup C, Knapp S (2016b) Confirming the identity of two enigmatic "spiny solanums" (*Solanum* subgenus *Leptostemonum*, Solanaceae) collected by Jean-Baptiste Le-schenault in Java. PhytoKeys 70: 97–110. https://doi.org/10.3897/phytokeys.70.9758

Aubriot X, Knapp S, Syfert MM, Poczai P, Buerki S (2018) Shedding new light on the origin and spread of the brinjal eggplant (*Solanum melongena* L.) and its wild relatives. American Journal of Botany 105(7): 1175–1187. https://doi.org/10.1002/ajb2.1133

Babu CR, Hepper FN (1979) Taxonomy and nomenclature of *Solanum khasianum* and some of its relatives. Kew Bulletin 34(2): 407–411. https://doi.org/10.2307/4110004

Barchi L, Pietrella M, Avanzato C, Venturini L, Minio A, Topppino L, Acquadro A, Andolfo G, Aprea G, Avanzato C, Bassolino L, Comino C, Dal Molin A, Ferrarin A, Maor LC, Portis E, Reyes-Chin Wo S, Rinaldi R, Sala T, Scaglione D, Sonawane P, Tononi P, Almekias-Siegl E, Zago E, Lanteri S, Rotino GL (2019) A chromosome-anchored eggplant genome sequence reveals key events in Solanaceae evolution. Scientific Reports 9(1): e11769. https://doi.org/10.1038/s41598-019-47985-w

Barnett EC (1963) Contributions to the Flora of Thailand: LVI: Descriptions of new species from Dr. Kerr's manuscript. Kew Bulletin 16(3): 485–490. https://doi.org/10.2307/4114708

Bauhin J (1651) Historia plantarum universalis, nova et absolutissima. Vol. 3. Yverdon.

Bean AR (2004) The taxonomy and ecology of *Solanum* subg. *Leptostemonum* (Dunal) Bitter (Solanaceae) in Queensland and far north-eastern New South Wales, Australia. Austrobai-
leya 6(4): 639–816.

Bean AR (2016) Taxonomic novelties in the *Solanum ferocissimum* group (Solanaceae: *Solanum* subg. *Leptostemonum*) from New Guinea. Austrobaileya 9(4): 560–599.

Bentham G (1861) Flora hongkongensis. L. Reeve, London.

Bitter G (1919) Die papuasischen Arten von *Solanum*. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 55: 59–112.

Bitter G (1923) Solana africana IV. Repertorium Specierum Novarum Regni Vegetabilis 16: 1–320.

Blanco CE (1837) Flora de Filipinas. Vol. 1. 3rd edn. Sto. Thomas por D. Candido López, Manila.
Blume CL (1826) *Solanum*. In: Bijdragen tot de flora van Nederlandsch Indië. Ter Lands Druk-
kerij, Batavia [Djakarta], 694–702. https://doi.org/10.5962/bhl.title.115427

Bohs L (2004) A chloroplast phylogeny of *Solanum* section *Lasiocarpa*. Systematic Botany 29(1): 177–187. https://doi.org/10.1600/036364404772974310

Bohs L (2005) Major clades in *Solanum* based on ndhF sequences. In: Keating RC, Hollowell VC, Croat TB (Eds) A festschrift for William G. D'Arcy: the legacy of a taxonomist, Monographs in Systematic Botany from the Missouri Botanical Garden, Vol. 104. Missouri Botanical Garden Press, St. Louis, 27–49.

Bohs L, Weese T, Myers N, Lefgren V, Thomas N, Stern S (2007) Zygomorphy and heterandry in *Solanum* in a phylogenetic context. In: Bohs L, Giovannoni J, Olmstead R, Shibata D, Spooner DM (Eds) Solanaceae VI: genomics meets biodiversity [Actae Horticulturae 745] International Society for Horticultural Science, Leuven, 201–224. https://doi.org/10.17660/ActaHortic.2007.745.8

Bonati G (1914) Primulacées, Solanacées et Scrofulariacées nouvelles de la Chine, de l'Indochine et du Turkestan. Bulletin de la Société Botanique de Genève, sér., 2 5: 297–316.

Bonati G (1915) Solanacées. In: Flore generale d'Indo-Chine 4: 313–336.

Botanic Gardens Conservation International (BGCI), IUCN SSC Global Tree Special-
ist Group (2018) *Solanum wrightii*. The IUCN Red List of Threatened Species 2018:
e.T134512852A135681297. https://doi.org/10.2305/IUCN.UK.2018-2.RLTS.
T134512852A135681297.en [downloaded 22 November 2021]

Botanic Gardens Conservation International (BGCI), IUCN SSC Global Tree Special-
ist Group (2020) *Solanum giganteum*. The IUCN Red List of Threatened Species 2020:
e.T146459232A146459234. https://doi.org/10.2305/IUCN.UK.2020-2.RLTS.
T146459232A146459234.en [downloaded 22 November 2021]

Bowers KAW (1975) The pollination ecology of *Solanum rostratum* (Solanaceae). American Journal of Botany 62(6): 633–638. https://doi.org/10.1002/j.1537-2197.1975.tb14094.x

Bridson GDR (2004) BPH-2: Periodicals with Botanical Content. Hunt Institute for Botani-
cal Documentation, Pittsburgh. [Available and searchable online at] http://fmhibd.library.cmu.edu/fmi/iwp/cgi?-db=BPH_Online&-loadframes

Bruneau A, Dickson EE, Knapp S (1995) Congruence of chloroplast DNA restriction site char-
acters with morphological and isozyme data in *Solanum* sect. *Lasiocarpa*. Canadian Journal
of Botany 73(8): 1151–1167. https://doi.org/10.1139/b95-125

Buchanan-Hamilton F (1824) A commentary on the second part of the *Hortus Ma-labaricus*. Transactions of the Linnean Society of London 14(2): 171–312. https://doi.org/10.1111/j.1095-8339.1823.tb00088.x

Buchmann SL, Jones CE, Colin LJ (1977) Vibratile Pollination of *Solanum douglassi* and *S. xanti* (Solanaceae) in Southern California. Wasmann Journal of Biology 35: 1–25.

Burkill IH (1935) A dictionary of the economic products of the Malay Peninsula. Vol. 2: I-Z.
Crown Agents for the Colonies, London.

Caguiat XGI, Hautea DM (2014) Genetic diversity analysis of eggplant (*Solanum melongena* L.) and related wild species in the Philippines using morphological and SSR markers. SA-BRAO Journal of Breeding and Genetics 46(2): 183–201.

Cámara-Leret R, Frodin DC, Adema F, Anderson C, Appelhans MS, Argent G, Arias Guer-
rero S, Ashton P, Baker W, Barfod AS, Barrington D, Borosova R, Bramley GLC, Briggs
M, Buerki S, Cahen D, Callmander MW, Cheek M, Chen C-W, Conn B, Coode MJE, Darbyshire I, Dawson S, Dransfield J, Drinkell C, Duyjfies B, Ebihara A, Ezedin Z, Fu L-F, Heatubun CD, Hind DJN, Hoch P, Hovenkamp P, Hughes M, Jebb M, Jennings L, Jimbo T, Kessler M, Kiew R, Knapp S, Lehnert M, Lewis GP, Linder HP, Lindsay S, Low YW, Lucas E, Mancera JP, Monro AK, Moore A, Middleton DJ, Nagamusu H, Newman MF, Nic Lughadha E, Melo PHA, Ohlsen D, Pannell CM, Parris B, Pearce L, Penneys D, Perrie LR, Petoe P, Poulsen AD, Prance GT, Quakenbush JP, Raes N, Rodda M, Rogers ZS, Schuiteman A, Schwartsburd P, Scotland RW, Simmons MP, Simpson DA, Stevens PF, Sundue M, Testo W, Trias-Blasi A, Turner I, Utteridge T, Walsingham L, Webber BL, Wei R, Weiblen G, Weigend M, Weston P, de Wilde W, Wilkie P, Wilmot-Dear CM, Wilson HP, Wood JRI, Zhang L-B, van Welzen P (2020) New Guinea has the world's richest island flora. Nature 584(7822): 579–583. https://doi.org/10.1038/s41586-020-2549-5

Cericola F, Portis E, Toppino L, Barchi L, Acciarri N, Ciriaci T, Sala T, Rotino GL, Lanteri S (2013) The population structure and diversity of eggplant from Asia and the Mediterranean Basin. PLoS ONE 8(9): e0073702. https://doi.org/10.1371/journal.pone.0073702

Chapman MA [Ed.] (2019) The eggplant genome. In: Kole C (series Ed.) Compendium of plant genomes. Springer International, Switzerland. https://doi.org/10.1007/978-3-319-99208-2

Chatterjee SA, van Andel T (2019) Lost grains and forgotten vegetables from Japan: The Seikei Zusetsu Agricultural Catalog (1793–1804). Economic Botany 73(3): 375–389. https://doi.org/10.1007/s12231-019-09466-z

Chennaveeraiah MS, Krishnappa DG (1971) Karyomorphological studies of nonspinaceous species of *Solanum*. Journal of Cytology and Genetics 5: 89–94.

Child A, Lester RN (1991) Life form and branching with the Solanaceae. In: Hawkes JG, Lester RN, Nee M, Estrada N (Eds) Solanaceae III: taxonomy, chemistry, evolution. Royal Botanic Gardens Kew, Richmond, 151–159.

Child A, Lester RN (2001) Synopsis of the genus *Solanum* L. and its infrageneric taxa. In: van den Berg RG, Barendse, GWM, van der Weerden G, Mariani C (Eds) Solanaceae V: advances in taxonomy and utilization. Nijmegen University Press, Nijmegen, 39–52.

Clark JL, Nee M, Bohs L, Knapp S (2015) A revision of *Solanum* section *Aculeigerum* (the *Solanum wendlandii* group, Solanaceae). Systematic Botany 40(4): [1–36] 1102–1136. https://doi.org/10.1600/036364415X690148

Clarke CB (1883) Solanaceae. In: Hooker JD (Ed.) Flora of British India. Vol. 4. Reeve & Co., London, 228–246.

Cracraft J (1989) Speciation and its ontogeny: the empirical consequences of alternative species concepts for understanding patterns of differentiation. In: Otte D, Endler JA (Eds) Speciation and its consequences. Sinauer and Associates, New York, 28–59.

Craib WG (1928) Contributions to the flora of Siam. Additamentum XXV. Kew Bulletin 1928: 234–247. https://doi.org/10.2307/4107694

D'Arcy WG (1972) Solanaceae studies II: Typification of subdivisions of *Solanum*. Annals of the Missouri Botanical Garden 59(2): 262–278. https://doi.org/10.2307/2394758

D'Arcy WG (1974a) [1973] Solanaceae, Flora of Panama. Part 9. Annals of the Missouri Botanical Garden 60: 573–780. https://doi.org/10.2307/2395139

D'Arcy WG (1974b) *Solanum* and its close relatives in Florida. Annals of the Missouri Botanical Garden 61(3): 819–867. https://doi.org/10.2307/2395032
D'Arcy WG (1986) The calyx in *Lycianthes* and some other genera. Annals of the Missouri Botanical Garden 73(1): 117-217. https://doi.org/10.2307/2399143

D'Arcy WG, Keating RC, Buchmann SL (1996) The calcium oxalate package or so-called resorption tissue in some angiosperm anthers. In: D'Arcy WG, Keating RC (Eds) The anther: form, function and phylogeny. Cambridge University Pres, Cambridge, 159-191.

Danert S (1958) Die Verzweigung der Solanaceen im reproduktiven Bereich. Abhandlungen der Deutschen Akademie der Wissenschaften zu Berlin, Klasse für Chemie, Geologie und Biologie 1957: 1-183.

Danert S (1967) Die Verzweigung als infragenerisches Grüppenmerkmal in der Gattung *Solanum*. L. Die Kulturpflanze 15(1): 275-292. https://doi.org/10.1007/BF02095719

Danert S (1970) Infragenerische Taxa der Gattung *Solanum*. L. Die Kulturpflanze 18(1): 253-297. https://doi.org/10.1007/BF02095597

Das AK, Borah SP (2015) Karyomorphological characterization of some wild species of *Solanum*. International Journal of Pharma and Bio Sciences 6(1): B1196-B1209.

Daunay M-C, Hazra P (2012) Eggplant. In: Peter KV, Hazra P (Eds) Handbook of vegetables. Studium Press, Houston, 257-322.

Daunay M-C, Janick J (2007) History and iconography of eggplant. Chronica Horticulturae 47: 16-22.

Davidar P, Snow AA, Rajkumar M, Pasquet R, Daunay M-C, Mutegi E (2015) The potential for crop to wild hybridization in eggplant (*Solanum melongena*; Solanaceae) in southern India. American Journal of Botany 102(1): 128-138. https://doi.org/10.3732/ajb.1400404

Davis JI (1997) Evolution, evidence, and the role of species concepts in phylogenetics. Systematic Botany 22(2): 373-403. https://doi.org/10.2307/2419463

Davis PH, Heywood VH (1963) Principles of angiosperm taxonomy. Van Nostrand, New York.

De Candolle R, Radcliffe-Smith A (1981) Nathaniel Wallich, MD, PhD, FRS, FLS, FRGS, (1786-1854) and the Herbarium of the Honourable East Indian Comapany, and their relation to the de Candolles of Geneva and the great *Prodromus*. Botanical Journal of the Linnean Society 83(4): 325-348. https://doi.org/10.1111/j.1095-8339.1981.tb00355.x

Dean EA (2001) The post-anthesis floral biology of *Lycianthes* series *Meizonodontae* (*Solanaceae*): variation in filament elongation, anther dehiscence, floral movement and corolla growth. In: van den Berg RG, Barendse GWM, van der Weerden GM, Mariani C (Eds) Solanaceae V; advances in taxonomy and utilization. Nijmegen University Press, Nijmegen, 137-151.

Deb DB (1980) Enumeration, synonymy and distribution of the Solanaceae in India. Journal of Economic and Taxonomic Botany 1: 33-54.

Dempewolf H, Eastwood RJ, Guarino L, Khoury CK, Müller JV, Toll J (2014) adapting agriculture to climate change: A global initiative to collect, conserve, and use crop wild relatives. Agroecology and Sustainable Food Systems 38(4): 369-377. https://doi.org/10.1080/021683565.2013.870629

Diaz R, Overholt WA, Langeland K (2008) Jamaican nightshade (*Solanum jamaicense*): A threat to Florida's hammocks. Invasive Plant Science and Management 1(4): 422-425. https://doi.org/10.1614/IPSM-08-079.1

Diggle P, Miller J (2004) Architectural effects mimic floral sexual dimorphism in *Solanum* (*Solanaceae*). American Journal of Botany 91(12): 20-30. https://doi.org/10.3732/ajb.91.12.2030
Dikii SP (1984) Infraspecific taxonomy of *Solanum melongena* L. Trudy Prikladnoi Botanike i Seleksii 88: 103–106. [in Russian] [Bulletin of Applied Botany and Selection]

Doganlar S, Frary A, Daunay M-C, Lester RN, Tanksley SD (2002) a comparative genetic linkage map of eggplant (*Solanum melongena* L.) and its implications for genome evolution in the Solanaceae. Genetics 161(4): 1697–1711. https://doi.org/10.1093/genetics/161.4.1697

Dulberger R, Levy A, Palevitch D (1951) Andromonoecy in *Solanum marginatum*. Botanical Gazette (Chicago, Ill.) 142(2): 259–266. https://doi.org/10.1086/337221

Dunal M-F (1813) Histoire naturelle, médicale et économique des *Solanum* et des genres qui ont été confundus avec eux. Renaud, Montpellier. https://doi.org/10.5962/bhl title.164866

Dunal M-F (1814) Morelle, *Solanum*. In: Poiret JLM (Ed.) Encyclopédie méthodique. Botanique, Supplément 3: 738–780. [H. Agasse, Paris]

Dunal M-F (1816) Solanorum generumque affinium synopsis. Renaud, Montpellier.

Dunal M-F (1852) Solanaceae. In: Candolle AP de (Ed.) Prodromus systematis naturalis regni vegetabilis 13(1): 1–690. [V. Masson, Paris]

Dupin J, Matzke NJ, Särkinen T, Knapp S, Olmstead RG, Bohs L, Smith SD (2016) Bayesian estimation of the global biogeographic history of the Solanaceae. Journal of Biogeography 44(4): 887–899. https://doi.org/10.1111/jbi.12898

Echeverría-Londoño S, Särkinen T, Fenton I, Knapp S, Purvis A (2020) Dynamism and context dependency in the diversification of the megadiverse plant genus *Solanum* (Solanaceae). Journal of Systematics and Evolution 58(6): 767–782. [first published online 19 May 2020] https://doi.org/10.1111/jse.12638

Edmonds JM (2012) *Solanum giganteum*. In: Beentje H (Ed.) Flora of Tropical East Africa. Solanaceae. Royal Botanic Gardens Kew, 133–135.

Elmer ADE (1910) The genus *Solanum* from Mt. Apo. Leaflets of Philippine Botany 2: 729–734.

Elmer ADE (1913) Four score of new plants. Leaflets of Philippine Botany 5: 1791–1854.

Endersby J (2008) Joseph Hooker: Philosophical botanist. Journal of Biosciences 33(2): 163–169. https://doi.org/10.1007/s12038-008-0033-8

Filov AI (1958) *Solanum melongena* L. [in Russian]. In: Brezhnev DD (Ed.) Vegetable Plants. Fam. Solanaceae. In: Zhukovsky PM (Ed. gen). Cultivated Flora USSR 10: 292–370.

Frodin DG (2004) History and concepts of big plant genera. Taxon 53(3): 753–776. https://doi.org/10.2307/4135449

Gagnon E, Hilgenhof R, Orejuela A, McDonnell A, Sablok G, Aubriot X, Giacomin L, Gouvèa Y, Bragonis T, Stehmann JR, Bohs L, Dodsworth S, Martine C, Poczai P, Knapp S, Särkinen T (2022) Phylogenomic discordance suggests polytomies along the backbone of the large genus *Solanum*. American Journal of Botany 109(4): 580–601. [online 16 Feb 2022] https://doi.org/10.1002/ajb2.1827

García-Fortea E, Gramazio P, Vilanova S, Fita A, Mangino G, Villanueva G, Arrones A, Knapp SJ, Prohens J, Plazas M (2019) First successful backcrossing towards eggplant (*Solanum melongena*) of a New World species, the silverleaf nightshade (*Solanum elaeagnifolium*), and characterization of interspecific hybrids and backcrosses. Scientia Horticulturae 246: 563–573. https://doi.org/10.1016/j.scienta.2018.11.018

Govindan S, Viswanathan S, Vijayaesekaran V, Alagappan R (1999) A pilot study on the clinical efficacy of *Solanum xanthocarpum* and *Solanum trilobatum* in bronchial asthma. Journal of Ethnopharmacology 66(2): 205–210. https://doi.org/10.1016/S0378-8741(98)00160-3
Govindan S, Viswanathan S, Vijayaesekaran V, Alagappan R (2004) Further studies on the clinical efficacy of *Solanum trilobatum* in bronchial asthma. Phytotherapy Research 18: 805-809. https://doi.org/10.1002/ptr.1555

Gramazio P, Prohens J, Plazas M, Andjar I, Herraiz FJ, Castillo E, Knapp S, Meyer RS, Vilanova S (2014) Location of chlorogenic acid biosynthesis pathway and polyphenol oxidase genes in a new interspecific anchored linkage map of eggplant. BMC Plant Biology 14(1): e350. https://doi.org/10.1186/s12870-014-0350-z

Gramazio P, Prohens J, Plazas M, Mangino G, Herraiz FJ, Vilanova S (2017) Development and genetic characterization of advanced backcross materials and an introgression line population of *Solanum incanum* in a *S. melongena* background. Frontiers in Plant Science 8: e1477. https://doi.org/10.3389/fpls.2017.01477

Gramazio P, Yan H, Hasing T, Vilanova S, Prohens J, Bombarely A (2019) Whole-genome resequencing of seven eggplant (*Solanum melongena*) and one wild relative (*Solanum incanum*) accessions provides new insights and breeding tools for eggplant enhancement. Frontiers in Plant Science 10: e1220. https://doi.org/10.3389/fpls.2019.01220

Hance HF (1868) Sertulum Chinense tertium: A third decade of new Chinese plants. Le Journal de Botanique 6: 322-335.

Heiser Jr CB (1986) A new domesticated variety and relationships of *Solanum lasiocarpum*. In: D'Arcy WG (Ed.) Solanaceae: biology and systematics. Columbia University Press, New York, 412-415.

Heiser Jr CB (1987) Origins of *Solanum lasiocarpum* and *S. repandum*. American Journal of Botany 74(7): 1045-1048. https://doi.org/10.1002/j.1537-2197.1987.tb08713.x

Heiser Jr CB (1996) Reappraisal of *Solanum ferox*, *S. lasiocarpum* and *S. repandum*. Solanaceae Newsletter 4: 44-50.

Heiser Jr CB (2001) Is *Solanum ferox* var. *ferox* (Solanaceae) extinct? Flora Malesiana Bulletin 12(7/8): 392-392.

Henry AN, Kumari GR, Chithra V (1987) Flora of Tamil Nadu, India. Series I: Analysis. Botanical Survey of India, Coimbatore.

Hepper FN (1978) (456, 457) Proposals to list *Solanum indicum* L. and *Solanum sodumeum* L. as rejected names under Article 69 of the ICBN. Taxon 27(5/6): 555-555. https://doi.org/10.2307/1219929

Hepper FN (1987) Solanaceae. In: Dassanayake MD (Ed.) A revised handbook to the flora of Ceylon, Vol. VI. Smithsonian Institution and Amerind Publishing, Washington DC and New Delhi, 365-409.

Hepper FN, Jaeger PML (1985) The typification of six Linnaean names in *Solanum*. Kew Bulletin 40(2): 387-391. https://doi.org/10.2307/4108264

Hepper FN, Jaeger PML (1986) Name changes for two Old World *Solanum* species. Kew Bulletin 41(2): 433-435. https://doi.org/10.2307/4102956

Hill J (1765) The vegetable system. Vol. 9. Published by the author, London.

Hsu T-W, Peng C-I, Peng J-J, Chiang T-Y (2007) Confirmation of the distribution of *Solanum miyakojimense* Yamazaki & Takushi (Solanaceae) in Taiwan. Taiwania 52(2): 190-193.

Hul S, Dy Phon P (2014) Solanaceae. In: Aubréville A, Leroy J, Morat P (Eds) Flore du Cambodge, du Laos et du Vietnam. Muséum National d'Histoire Naturelle & Royal Botanic Garden Edinburgh, Paris & Edinburgh, 1-93.
Hurtado M, Vilanova S, Plazas M, Gramazio P, Fonseka HH, Fonseka R, Prohens J (2012) Diversity and relationships of eggplants from three geographically distant secondary centers of diversity. PLoS ONE 7(7): e41748. https://doi.org/10.1371/journal.pone.0041748

Husnudin BA, Suharyanto, Daryono BS, Purnomo (2019) Variation and non-formal classification of Indonesian eggplant (*Solanum melongena* L.) accessions based on macro- and micro-morphological characters. AGRIVITA Journal of Agricultural Science 41(3): 544–560. https://doi.org/10.17503/agrivita.v41i3.2039

Hutchings A, Scott AH, Lewis G, Cunningham AB (1996) Zulu medicinal plants: an inventory. University of Natal Press, Pietermaritzburg.

IUCN (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee of the IUCN Species Survival Commission. http://www.iucnredlist.org/documents/RedListGuidelines.pdf

Jacquin NJ (1792) Icones plantarum rariorum. Vol 2. C.F. Wappler, Vienna.

Jain SK, Borthakur SK (1986) Solanaceae in Indian tradition, folklore and medicine. In: D'Arcy WG (Ed.) Solanaceae: biology and systematics. Columbia University Press, New York, 575–583.

Jain SP, Verma DM, Singh SC, Singh JS, Kumar S (2000) Flora of Haryana. Central Institute of Medicinal Plants, Lucknow.

Jaiswal BS (2012) *Solanum torvum*: A review of its traditional uses, phytochemistry and pharmacology. International Journal of Pharma and Bio Sciences 3(4): 104–111.

Kandoliya UK, Bajaniya VK, Bhadja NK, Bodar NP, Golakiya BA (2015) And antioxidant and nutritional components of Egg plant (*Solanum melongena* L.) fruit grown in Saurastra region. International Journal of Current Microbiology and Applied Sciences 4(2): 806–813.

Karihaloo JL (1991) Desynapsis due to temperature stress in three species of *Solanum*. Cytologia 56(4): 603–611. https://doi.org/10.1508/cytologia.56.603

Kaushik P, Prohens J, Vilanova S, Gramazio P, Plazas M (2016) Phenotyping of eggplant wild relatives and interspecific hybrids with conventional and phenomics descriptors provides insight for the potential utilization in breeding. Frontiers in Plant Science 7: e677. https://doi.org/10.3389/fpls.2016.00677

Khatoon S, Ali SI (1993) Chromosome atlas of the angiosperms of Pakistan. University of Karachi, Karachi.

Kingdon-Ward F (1956) Return to the Irrawaddy &c. Andrew Melrose, London.

Kirti PB, Rao BGS (1983) The cytogenetic basis of sterility in spinous *Solanum* hybrids. Caryologia 36(2): 155–164. https://doi.org/10.1080/00087114.1983.10797655

Knapp S (2001) Is morphology dead in *Solanum* taxonomy? In: van den Berg RG, Barendse GWM, van der Weerden GM, Mariani C (Eds) Solanaceae V: advances in taxonomy and utilization. Nijmegen University Press, Nijmegen, 23–38.

Knapp S (2002a) Assessing patterns of plant endemism in Neotropical uplands. Botanical Review 68(1): 22–37. https://doi.org/10.1663/0006-8101(2002)068[0022:APOPEI]2.0.CO;2

Knapp S (2002b) *Solanum* section *Geminata* (G. Don) Walpers (Solanaceae). Flora Neotropica 84: 1–405.

Knapp S (2007) Lectotypification of Cavanilles' names in *Solanum* (Solanaceae). Anales del Jar-din Botanico de Madrid 64(2): 195–203. https://doi.org/10.3989/ajbm.2007.v64.i2.175
Knapp S (2008) Species concepts and floras: What are species for? Biological Journal of the Linnean Society. Linnean Society of London 95(1): 17–25. https://doi.org/10.1111/j.1095-8312.2008.01090.x

Knapp S (2011) (2042–2043) Proposals to reject the names *Solanum ferox* and *S. fuscatum* (*Solanaceae*). Taxon 60(6): 1782–1783. https://doi.org/10.1002/tax.606032

Knapp S (2013a) A revision of the Dulcamaroid clade of *Solanum* L. (Solanaceae). PhytoKeys 22(0): 1–432. https://doi.org/10.3897/phytokeys.22.4041

Knapp S (2013b) Typification of *Solanum* species (Solanaceae) described by Casimiro Gómez Ortega. Anales del Jardín Botánico de Madrid 70(1): 56–61. https://doi.org/10.3989/ajbm.2340

Knapp S (2020) *Solanum viarum*. The IUCN Red List of Threatened Species 2020: e.T101534565A101534580. https://doi.org/10.2305/IUCN.UK.2020-3.RLTS. T101534565A101534580.en [accessed 22 November 2021]

Knapp S (2021a) *Solanum cordatum*. The IUCN Red List of Threatened Species 2021: e.T186619213A186619254. https://doi.org/10.2305/IUCN.UK.2021-1.RLTS. T186619213A186619254.en [accessed 22 November 2021]

Knapp S (2021b) *Solanum forskalii*. The IUCN Red List of Threatened Species 2021: e.T101526207A101526210. https://doi.org/10.2305/IUCN.UK.2021-1.RLTS. T101526207A101526210.en [accessed 22 November 2021]

Knapp S, Jarvis CE (1990) The typification of the names of New World *Solanum* species described by Linnaeus. Botanical Journal of the Linnean Society 104(4): 113–139. https://doi.org/10.1111/j.1095-8339.1990.tb02227.x

Knapp S, Vorontsova MS (2016) A revision of the “African Non-Spiny” Clade of *Solanum* L. (*Solanum* sections *Afrosolanum* Bitter, *Benderianum* Bitter, *Lemurisolanum* Bitter, *Lycioso-lanum* Bitter, *Macronesiotes* Bitter, and *Quadrangulare* Bitter: *Solanaceae*). PhytoKeys 66: 1–142. https://doi.org/10.3897/phytokeys.66.8457

Knapp S, Persson V, Blackmore S (1998) Pollen morphology and functional dioecy in *Solanum* (Solanaceae). Plant Systematics and Evolution 210(1–2): 113–139. https://doi.org/10.1007/BF00984731

Knapp S, Vorontsova MS, Prohens J (2013) Wild relatives of the eggplant (*Solanum melongena* L.: Solanaceae): new understanding of species names in a complex group. PLoS ONE 8(2): e57039. https://doi.org/10.1371/journal.pone.0057039

Knapp S, Sagona E, Carbonell AKZ, Chiarini F (2017) A revision of the *Solanum elaeagnifolium* clade (Elaeagnifolium clade, subgenus Leptostemonum, Solanaceae). PhytoKeys 84: 1–104. https://doi.org/10.3897/phytokeys.84.12695

Knapp S, Barboza GE, Bohs L, Särkinen T (2019a) A revision of the Morelloid Clade of *Solanum* L. (Solanaceae) in the Caribbean and North and Central America. PhytoKeys 123: 1–174. https://doi.org/10.3897/phytokeys.123.31738

Knapp S, Aubriot X, Prohens J (2019b) Eggplant (*Solanum melongena* L.): taxonomy and relationships. In: Chapman MA (Ed.) The eggplant genome. Springer International Publishing AG, New York & Basel, 11–22. https://doi.org/10.1007/978-3-319-99208-2_2

Kouassi B, Prohens J, Gramazio P, Kouassi AB, Vilanova S, Galán-Ávila A, Herrraiz FJ, Kouassi A, Seguí-Simarro JM, Plazas M (2016) Development of backcross generations and new interspecific hybrid combinations for introgression breeding in eggplant (*Solanum melongena* L.). Scientia Horticulturae 213: 199–207. https://doi.org/10.1016/j.scienta.2016.10.039
Kumar V, Subramaniam B (1987) Chromosome atlas of flowering plants of the Indian subcontinent. Botanical Survey of India, New Delhi.

Lamarck JB (1794) Tableau encyclopédique et méthodique, Botanique. Vol. 2. Pancoucke, Paris.

Latha PS, Kannabiran K (2006) Antimicrobial activity and phytochemicals of *Solanum trilobatum* Linn. African Journal of Biotechnology 5(23): 2402–2404.

Ledermann C (1919) Einiges von der Kaiserin-Augusta-Fluß-Expedition. Botanische Jährbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 55: 33–44.

Lester RN (1997) Typification of nineteen names of African *Solanum* species described by A. Richard and others, including *S. campylacanthum* and *S. panduriforme*. Botanical Journal of the Linnean Society 125(4): 273–293. https://doi.org/10.1111/j.1095-8339.1997.tb02258.x

Lester RN, Durrands P (1984) Enzyme treatment as an aid in the study of seed surface structures of *Solanum* species. Annals of Botany 53(1): 129–131. https://doi.org/10.1093/oxfordjournals.aob.a086662

Lester RN, Hasan SMZ (1991) Origin and domestication of the brinjal eggplant, *Solanum melongena*, from *Solanum incanum*, in Africa and Asia. In: Hawkes JG, Lester RN, Nee M, Estrada E (Eds) Solanaceae III: taxonomy, chemistry and evolution, Royal Botanic Gardens Kew. Richmond, 369–387.

Lester RN, Hawkes JG (2001) Solanaceae. Mansfeld’s encyclopedia of agricultural and horticultural crops (except ornamentals). Vol. 4. Springer Verlag, Berlin, Heidelberg, New York, 1790–1857.

Lester RN, Francisco-Ortega J, Al-Ani M (1999) Convergent evolution of heterandry (unequal stamens) in *Solanum*, proved by spermoderm SEM. In: Nee M, Symon DE, Lester RN, Jessop JP (Eds) Solanaceae IV: advances in biology and utilization, Royal Botanic Gardens Kew, Richmond, 51–69.

Léveillé AAH (1908) *Solanum* et *Physalis* de Chine. Bulletin de la Société Botanique de France 55(3): 202–209. https://doi.org/10.1080/00378941.1908.10831958

Léveillé AAH (1913) CIII. Decades plantarum novarum. CXXVII–CXXXI. Repertorium Specierum Novarum Regni Vegetabilis 12(33–36): 531–538. https://doi.org/10.1002/fedr.19130123306

Levin RA, Watson K, Bohs L (2005) A four-gene study of evolutionary relationships in *Solanum* section *Acanthophora*. American Journal of Botany 92(4): 603–612. https://doi.org/10.3732/ajb.92.4.603

Levin RA, Myers NR, Bohs L (2006) Phylogenetic relationships among the “spiny solanums” (*Solanum* subgenus *Leptostemonum*, Solanaceae). American Journal of Botany 93(1): 157–169. https://doi.org/10.3732/ajb.93.1.157

Lim TK (2012) *Solanum lasiocarpum*. In: Lim TK (Ed.) Edible and medicinal and non-medicinal plants. Vol 6. Fruits. Springer, Dordrecht, 333–335. https://doi.org/10.1007/978-94-007-5628-1_39

Linnaeus C (1753) Species plantarum. L. Salvius, Stockholm.

Linnaeus C (1767) Mantissa plantarum. L. Salvius, Stockholm.

Lippman ZB, Cohen O, Alvarez IP, Abu-Abied M, Pekker I, Prana I, Eshed Y, Zamir D (2008) The making of a compound inflorescence in tomato and related nightshades. PLoS Biology 6(11): e288. https://doi.org/10.1371/journal.pbio.0060288
Loureiro J (1790) Flora cochinchinensis. Royal Portuguese Academy of Sciences, Lisbon.

Luckow M (1995) Species concepts: Assumptions, methods and applications. Systematic Botany 20(4): 589–605. https://doi.org/10.2307/2419812

Madhavadian P (1968) Chromosome numbers in south Indian Solanaceae. Caryologia 21(4): 343–347. https://doi.org/10.1080/00087114.1968.10796312

Mallet J (1995) A species definition for the modern synthesis. Trends in Ecology & Evolution 10(7): 294–299. https://doi.org/10.1016/0169-5347(95)00031-4

Margono BA, Potapov PV, Turubanova S, Stolle F, Hansen MC (2014) Primary forest cover loss in Indonesia over 2000–2012. Nature Climate Change 4(8): 1–6. https://doi.org/10.1038/nclimate2277

Martine CT, Anderson GJ, Les DH (2009) Gender-bending aubergines: Molecular phylogenetics of cryptically dioecious *Solanum* in Australia. Australian Systematic Botany 22(2): 107–120. https://doi.org/10.1071/SB07039

Martine CT, Jordan-Thaler IE, McDonnell AJ, Cantley JT, Hayes DS, Roche MD, Frawley ES, Gilman IS, Tank DC (2019) Phylogeny of the Australian *Solanum dioicum* group using seven nuclear genes, with consideration of Symon's fruit and seed dispersal hypothesis. PLoS ONE 14(4): e0207564. https://doi.org/10.1371/journal.pone.0207564

Matthew KM (1983) Flora of the Tamilnadu Carnatic. Volume 3. Dioscesan Press, Madras [Chennai].

Mayr E (1982) The growth of biological thought. Harvard University Press, Cambridge.

McClelland DHR (2012) Systematics and taxonomy of *Solanum* sections *Dunaliana* and *Irenosolanum* (Solanaceae). Doctoral dissertation presented to the City University of New York, New York.

McClelland DHR, Nee M, Knapp S (2020) new names and status for Pacific spiny species of *Solanum* (Solanaceae, subgenus *Leptostemonum* Bitter; the *Leptostemonum* Clade). PhytoKeys 145: 1–36. https://doi.org/10.3897/phytokeys.145.48531

McDonnell AJ, Wetreich HB, Cantley JT, Jobson P, Martine CT (2019) *Solanum plastisexum*, an enigmatic new bush tomato from the Australian Monsoon Tropics exhibiting breeding system fluidity. PhytoKeys 124: 39–55. https://doi.org/10.3897/phytokeys.124.33526

McNeill J (2014) Holotype specimens and type citations: General issues. Taxon 63(5): 1112–1113. https://doi.org/10.12705/635.7

Merrill ED (1906) New or noteworthy Philippine plants V. Philippine Journal of Science 1(suppl.): 169–246.

Merrill ED (1917) The interpretation of Rumphius's Herbarium amboinense. Bureau of Printing, Manila. https://doi.org/10.5962/bhl.title.79163

Merrill ED (1918a) Species Blancoanae, a critical revision of the Philippine species of plants described by Blanco and Llanos. Bureau of Printing, Manila. https://doi.org/10.5962/bhl.title.2116

Merrill ED (1918b) New or noteworthy Philippine plants XIII. The Philippine Journal of Science, C. Botany 13: 1–66.

Merrill ED (1923) Solanaceae. Enumeration of Philippine flowering plants. Vol. 3. Bureau of Printing, Manila, 423–431. https://doi.org/10.5962/bhl.title.49412

Merrill ED (1935) Loureiro and his botanical work. Proceedings of the American Philosophical Society 72(4): 229–239.
Meyer RS, Karol KG, Little DP, Nee MH, Litt A (2012) Phylogeographic relationships among Asian eggplants and new perspectives on eggplant domestication. Molecular Phylogenetics and Evolution 63(3): 685–701. https://doi.org/10.1016/j.ympev.2012.02.006

Meyer RS, Bamshad M, Fuller DQ, Litt A (2014) Comparing medicinal uses of eggplant and related Solanaceae in China, India and the Philippines suggests the independent development of uses, cultural diffusions, and recent species substitutions. Economic Botany 68(2): 137–152. https://doi.org/10.1007/s12231-014-9267-6

Mill RR (2001) Solanaceae. In: Grierson AJC, Long DG, Springate LS (Eds) Flora of Bhutan, including a record of plants from Sikkim and Darjeeling Volume 2, part 3. Royal Botanic Garden Edinburgh, Edinburgh, 1037–1078.

Miller P (1768) The gardeners' dictionary, 8th edn. Printed for the author, London.

Miller J, Diggle P (2003) Diversification of andromonoecy in *Solanum* section *Lasiocarpa* (*Solanaceae*): The roles of phenotypic plasticity and architecture. American Journal of Botany 90(5): 707–715. https://doi.org/10.3732/ajb.90.5.707

Miquel FAW (1857) Ordo CLXIV. Solanaeae Juss. Flora Nederlandsch Indië. Vol. 2. CG van der Post, Amsterdam, 633–672.

Moat J (2007) Conservation assessment tools extension for ArcView 3.x, version 1.2. GIS Unit, Royal Botanic Gardens, Kew. http://www.rbgkew.org.uk/cats

Mohanan M, Henry AN (1994) Flora of Thiruvananthapuram, Kerala. Botanical Survey of India, Calcutta.

Mohanan PV, Rao JM, Kutti MAS, Devi SK (1998) Cytotoxicity of extracts of *Solanum trilo-batum* and anticarcinogenic activity of sobatum. Biomedicine 18: 106–111.

Muñoz-Falcón JE, Prohens J, Vilanova S, Nuez F (2009) Diversity in commercial varieties and landraces of black eggplant and implications for broadening the breeder's gene pool. Annals of Applied Biology 154(3): 453–465. https://doi.org/10.1111/j.1744-7348.2009.00314.x

Mutegi E, Snow AA, Rajkumar M, Pasquet R, Ponniah H, Daunay M-C, Davidar P (2015) Genetic diversity and population structure of wild/weedy eggplant (*Solanum insanum*, Solanaceae) in southern India: Implication for conservation. American Journal of Botany 102(1): 139–147. https://doi.org/10.3732/ajb.1400403

Naeem MY, Ugur S (2020) Nutritional content and health benefits of eggplant. Turkish Journal of Agriculture – Food Science and Technology (Campinas) 7: 31–36.

Nair KKN, Nayar M (1987) Flora of Courtallum. Vol. 2. Botanical Survey of India, New Delhi.

Naithani HB, Sahni KC, Bennett SSR (1997) Forest flora of Goa. International Book distributors, Dehra Dun.

Nee M (1979) A revision of *Solanum* section *Acanthophora*. PhD dissertation, University of Wisconsin, Madison.

Nee M (1993) Solanaceae II: *Solanum*. In: Sosa V (Ed.) Flora de Veracruz, Fascículo 72. Instituto de Ecología, Xalapa, 1–158.

Nees van Esenbeck CG (1834) Monograph of East Indian Solanaeae. Transactions of the Linnean Society of London 17(1): 37–82. https://doi.org/10.1111/j.1095-8339.1834.tb00017.x

Nees van Esenbeck CG (1836) Excursus de *Solano wightii*, quem Arnottii collegae observationibus addidit. Nova Acta Physico-Medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosum 18: 357–360. [Tab. XVIII]
Page A, Gibson J, Meyer RS, Chapman MA (2019) Eggplant domestication: Pervasive gene flow, feralization, and transcriptomic divergence. Molecular Biology and Evolution 36(7): 1359–1372. https://doi.org/10.1093/molbev/msz062

Paran I, van der Knaap E (2007) Genetic and molecular regulation of fruit and plant domestication traits in tomato and pepper. Journal of Experimental Botany 58(14): 3841–3852. https://doi.org/10.1093/jxb/erm257

Peralta IE, Spooner DM, Knapp S (2008) Taxonomy of wild tomatoes and their relatives (Solanum sections Lycopersicoides, Juglandifolia, Lycopersicon; Solanaceae). Systematic Botany Monographs 84: 1–186.

Plukenet L (1691) Phytographia, pars altera. London.

Plukenet L (1692) Phytographia, pars tertia. London.

Poiret JLM (1797) Morelle, *Solanum*. In: Lamarck JB (Ed.) Encyclopédie Methodique, Botanique. Vol. 4. H Agasse, Paris, 279–309.

Plazas M, Vilanova S, Gramazio P, Rodríguez-Burruezo R, Fita A, Herraiz FJ, Ranil R, Fonseka R, Niran L, Fonseka H, Kouassi B, Kouassi A, Houassi A, Prohens J (2016) Interspecific hybridization between eggplant and wild relatives in different genepools. Journal of the American Society for Horticultural Science 141(1): 34–44. https://doi.org/10.21273/JASHS.141.1.34

Prado J, Hirai RY, Moran RC (2015) (046–048) Proposals concerning inadvertent lectotypifications (and neotypifications). Taxon 64(3): 651–651. https://doi.org/10.12705/643.29

Prain D (1896) Noviciae Indiae XIV. Some additional Solanaceae. Journal of the Asiatic Society of Bengal 65(2): 541–543.

Prohens J, Plazas M, Raigón MD, Seguí-Simarro JM, Stommel JR, Vilanova S (2012) Characterization of interspecific hybrids and first backcross generations from crosses between two cultivated eggplants (*Solanum melongena* and *S. aethiopicum* Kumba Group) and implications for eggplant breeding. Euphytica 186(2): 517–538. https://doi.org/10.1007/s10681-012-0652-x

Prohens J, Whitaker BD, Plazas M, Vilanova S, Hurtado M, Blasco M, Gramazio P, Stommel JR (2013) Genetic diversity in morphological characters and phenolic acids content resulting from an interspecific cross between eggplant, *Solanum melongena*, and its wild ancestor (*S. incanum*). Annals of Applied Biology 162(2): 242–257. https://doi.org/10.1111/aab.12017

Prohens J, Gramazio P, Plazas M, Dempewolf H, Kilian B, Diéz MJ, Fita A, Herraiz FJ, Rodríguez-Burruezo A, Soler S, Knapp S, Vilanova S (2017) Introgressiomics: a new approach for using crop wild relatives in breeding for adaptation for climate change. Euphytica 213: e158. https://doi.org/10.1007/s10681-017-1938-9

Rajasekaran S (1971) Cytological studies on the F1 hybrid (*Solanum xanthocarpum* Schrad. and Wendl. × *S. melongena* L.) and its amphidiploid. Caryologia 24(3): 261–267. https://doi.org/10.1080/00087114.1971.10796434

Ramachandran A, Viswanathan MB (2010) *Solanum cordatum* (Solanaceae) – a new record to southern India. Rheedea 20: 56–58.

Ranil RHG, Prohens J, Aubriot X, Niran HML, Plazas M, Fonseka RM, Vilanova S, Fonseka HH, Gramazio P, Knapp S (2016) *Solanum insanum* L. (subgenus *Leptostemonum* Bitter, Solanaceae), the neglected wild progenitor of eggplant (*S. melongena* L): a review of taxonomy, characteristics and uses aimed at its enhancement for improved eggplant breeding.
Genetic Resources and Crop Evolution 64(7): 1707–1722. [e-published 22 Nov 2016, in hard copy 18 Sep 2017] https://doi.org/10.1007/s10722-016-0467-z

Rao YV, Rao BG (1987) Cytomorphological studies of hybrids between *Solanum pubescens*, *Solanum incanum* and *Solanum indicum*. Proceedings of the Indiana Academy of Sciences 97(1): 49–53. https://doi.org/10.1007/BF03053339

Rheede von tot Draakestein H (1689) Hortus indicus malabaricus. J. van Someren & J. van Dyk, Amsterdam.

Roe KE (1966) Juvenile forms in *Solanum mitlense* and *S. blodgettii* (Solanaceae) and their importance in taxonomy. Sida 2: 381–385.

Roe KE (1971) Terminology of hairs in the genus *Solanum*. Taxon 20(4): 501–508. https://doi.org/10.2307/1218251

Roemer JJ, Schultes JA (1819) Systema vegetabilium, 15th edn. [bis]. Vol. 4. J.G. Cotta, Stuttgart.

Roth AW (1821) Novae plantarum species praesetim Indiae orientalis ex collections Doct. Benj. Heynii. H. Vogler, Halberstadt. https://doi.org/10.5962/bhl title.10723

Roxburgh W (1814) Hortus Bengalensis, or a catalogue of the plants growing in the Honourable East India Company's Botanic Garden at Calcutta. Mission Press, Serampore.

Roxburgh W (1824) Flora indica, or descriptions of Indian plants. Vol. 2 [Wallich N, Carey W, eds]. Mission Press, Serampore.

Rumphius [Rumpf] GE (1747) Herbarium amboinense. Vol. 5. F. Changuion & H. Utewerf, Amsterdam.

Saha M, Datta BK (2013) New distributional record of *Solanum sisymbriifolium* Lamarck (Solanaceae) from Tripura, India. Pleione 7(2): 579–582.

Särkinen T, Olmstead RG, Bohs L, Knapp S (2013) A phylogenetic framework for evolutionary study of the nightshades (Solanaceae): A dated 1000-tip tree. BMC Evolutionary Biology 13(1): e214. https://doi.org/10.1186/1471-2148-13-214

Särkinen TE, Barboza GE, Knapp S (2015) True Black nightshades: Phylogeny and delimitation of the Morelloid clade of *Solanum*. Taxon 64(5): 945–958. https://doi.org/10.12705/645.5

Särkinen T, Poczai P, Barboza GE, van der Weerden GM, Baden M, Knapp S (2018) A revision of the Old World black nightshades (Morelloid Clade of *Solanum* L., Solanaceae). PhytoKeys 106: 1–223. https://doi.org/10.3897/phytokeys.106.21991

Scaldaferro M, Chiarini F, Santiñaque FF, Bernardello G, Moscone EA (2012) Geographical pattern and ploidy levels of the weed *Solanum elaeagnifolium* (Solanaceae) from Argentina. Genetic Resources and Crop Evolution 59(8): 1833–1847. https://doi.org/10.1007/s10722-012-9807-9

Schönbeck-Temesy E (1972) *Solanum* (Solanaceae). In: Rechinger KH (Ed.) Flora Iranica. 100:1–82. Verlag des Naturhistorischen Museums, Vienna.

Schumacher HCF (1827) Beskrivelse af Guineske planter: som ere fundne af Danske botanikere, isoer af etatsraad Thonning.F. Popps, Copenhagen. https://doi.org/10.5962/bhl title.51454

Scopoli JA (1786) Deliciae florae et faunae Insubricae. Monasterii S. Salvatoris, Tocino [Turin].

Seithe A (1962) Die Haararten der Gattung *Solanum* L. und ihre taxonomische Verwertung. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 81: 261–336.

Seithe A (1979) Hair types as taxonomic characters in *Solanum*. In: Hawkes JG, Lester RN, Skelding AD (Eds) The biology and taxonomy of the Solanaceae, Academic Press, London, 307–319.
Sen Gupta G (1961) A taxonomic note on *Solanum khasianum* Clarke and description of a new variety under it. Bulletin of the Botanical Survey of India 3: 411–415.

Sen Gupta G (1963) *Solanum arundo* Mattei, a new record for India. Indian Forester 89: 481–482. [1 fig.]

Sen Gupta G (1964) A rare and interesting Solanum (S. hovei Dun.) from western India. Bulletin of the Botanical Society of Bengal 17: 3–5.

Singh NP (1988) Solanaceae. In: Flora of eastern Karnataka. Vol. 2. Mittal Publications, Delhi, 455–462.

Singh V (1991) Solanaceae. In: Shetty BV, Singh V (Eds) Flora of Rajasthan Vol. 2. Botanical Survey of India, Calcutta, 562–579.

Soosairaj S, Raja P, Balaguru B, Tagore JK (2021) *Solanum pulneyensis* Soosairaj, sp. nov. (Solanaceae) from Palani Hills National Park of Tamil Nadu, India. Adansonia, sér. 3, 43(21): 235–240. https://doi.org/10.5252/adansonia2021v43a21

Spooner DM, Ghislain M, Simon R, Jansky SH, Gavrilenko T (2014) Systematics, diversity, genetics, and evolution of wild and cultivated potatoes. Botanical Review 80(4): 283–383. https://doi.org/10.1007/s12229-014-9146-y

Stafleu FA (1966) F.A.W. Miquel, Netherlands botanist. Mededelingen van het Botanisch Museum en Herbarium van de Rijksuniversiteit te Utrecht (Wentia 16(52): 1–95) 220: 1–95. https://doi.org/10.1111/j.1438-8677.1966.tb00026.x

Stafleu FA, Cowan RS (1981) Taxonomic Literature II. Vol 3: Authors Lh-O, ed. 2. Bonn, Scheltema & Holkema, Utrecht. https://www.sil.si.edu/DigitalCollections/tl-2/browse.cfm?vol=3#page/721

Stern SR, Agra M de F, Bohs L (2011) Molecular delimitation of clades within New World species of the "spiny solanums" (*Solanum* subgenus *Leptostemonum*). Taxon 60(5): 1429–1441. https://doi.org/10.1002/tax.605018

Stern S, Giacomin LL, Stehmann JR, Knapp S (2013) A revision of *Solanum* section *Gonatotrichum* Bitter (Solanaceae). Systematic Botany 38: 471–496. https://doi.org/10.1600/036364413X666624

Swartz O (1797) Flora indiae occidentalis. Vol. 1. J. Palm, Erlangen.

Syfert MM, Castañeda-Álvarez NP, Khoury CK, Särkinen T, Sosa CC, Achicanoy HA, Bernau V, Prohens J, Daunay M-C, Knapp S (2016) Crop wild relatives of the brinjal eggplant (*Solanum melongena*; Solanaceae): Poorly represented in genebanks and many species at risk of extinction. American Journal of Botany 103(4): 635–651. https://doi.org/10.3732/ajb.1500539

Symon DE (1970) *Dioecious Solanums*. Taxon 19(6): 909–910. https://doi.org/10.2307/1218308

Symon DE (1979) Sex forms in *Solanum* (Solanaceae) and the role of pollen collecting insects. In: Hawkes JD, Lester RN, Skelding AD (Eds) The biology and taxonomy of the Solanaceae. Academic Press, London, 385–397.

Symon DE (1981) A revision of *Solanum* in Australia. Journal of the Adelaide Botanic Gardens 4: 1–367.

Symon DE (1985) The Solanaceae of New Guinea. Journal of the Adelaide Botanic Gardens 8: 1–171.

Tadulingam C, Venkatanarayana G (1932) A handbook of South Indian weeds. Government Press, Madras.
Taher D, Solberg SØ, Prohens J, Chou Y-Y, Rakhla M, Wu T-H (2017) World Vegetable Center eggplant collection: Origin, composition, seed dissemination and utilization in breeding. *Frontiers in Plant Science* 8: e1484. https://doi.org/10.3389/fpls.2017.01484

Takeuchi S, Golman M (2002) The present status of Ledermann’s April River localities in Papua New Guinea. *Sida* 20: 55–70.

Tepe EJ, Bohs L (2011) A revision of *Solanum* section *Herpystichum*. *Systematic Botany* 36(4): 1068–1087. https://doi.org/10.1600/036364411X605074

Timmermans BGH, Vos J, Stomph TJ, Van Nieuwberg J, Van der Putten PEL (2007a) Field performance of *Solanum sisymbriifolium*, a trap crop for potato cyst nematodes. *I Dry matter accumulation in relation to sowing time, location, season, and plant density. Annals of Applied Biology* 150: 99–106. https://doi.org/10.1111/j.1744-7348.2006.00113.x

Timmermans BGH, Vos J, Stomph TJ, Van Nieuwberg J, Van der Putten PEL (2007b) Field performance of *Solanum sisymbriifolium*, a trap crop for potato cyst nematodes. *II Root charcateristics. Annals of Applied Biology* 150: 89–97. https://doi.org/10.1111/j.1744-7348.2006.00112.x

Tournefort JP de (1700) *Institutiones rei herbariae. Vol. 1. Typographia Regia, Paris*. https://doi.org/10.5962/bhl title.154246

Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber W-H, Li D-Z, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ, Smith GF (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code). *Regnum Vegetabile* 159. Koeltz Botanical Books, Glashütten. https://doi.org/10.12705/Code.2018

Turner IM (1998) Nomenclatural changes for four Malayan species in *Phrynium* (Maranta-ceae), *Solanum* (Solanaceae), *Stachyphrynium* (Marantaceae) and *Boesenbergia* (Zingiber-aceae). *Gardens’ Bulletin* (Singapore) 50: 39–41.

Turner IM (2021) Heyne, Roth, Roemer and Schultes, and the plant names published in *No-vae plantarum* species praesertim Indiae orientalis. *Taxon* 70(2): 365–428. https://doi.org/10.1002/tax.12449

Vallejo-Marin M, Da Silva EM, Sargent RD, Barrett SCH (2010) Trait correlates and function-al significance of heteranthery in flowering plants. *The New Phytologist* 188(2): 418–425. https://doi.org/10.1111/j.1469-8137.2010.03430.x

Van der Knaap E, Tanksley S (2003) The making of a bell-pepper shaped tomato fruit: Identification of loci controlling fruit morphology in Yellow Stuffer tomato. *Theoretical and Applied Genetics* 107(1): 139–147. https://doi.org/10.1007/s00122-003-1224-1

Van Huerck HF (1870) *Observationes botanicae*. Vol. 1. F. Baggerman, Antwerp.

van Steenis-Kruseman MJ (1985) Malyasian plant collectors and collections, being a cyclopedia of botanical exploration in Malaysia. *Martinius Nijhoff* (Kluwer Academic Publishers), Boston, Dordrecht, Lancaster. [originally published 1950 in Djakarta]

Varma SK (1981) *Flora of Bhagalpur. Today’s & Tomorrow’s Printers and Publishers, New Delhi*.

Ved DK, Sureshchandra ST, Brave V, Srinivas V, Sangeetha S, Ravikumar K, Kartikeyan Kulkarni V, Kumar AS, Venugopal SN, Somashekhar BS, Sumanth MV, Begum M, Rani S, Surekha KV, Desale N (2016) FRLHT’s ENVIS Centre on Medicinal Plants. *Bengaluru*. https://frht.envis.org
Veldkamp JF, Vink W, Frodin DG (1988) XI. Ledermann's and some other German localities in Papua New Guinea. Flora Malesiana Bulletin 10: 32-38.

Villeneuve M, Gunawan W, Cornee JJ, Vidal O (2002) Geology of the central Sulawesi belt (eastern Indonesia); constraints for geodynamic models. International Journal of Earth Sciences 91(3): 524-537. https://doi.org/10.1007/s005310100228

Vorontsova MS, Knapp S (2010) Lost Berlin (B) types of *Solanum* (Solanaceae) - found in Göttingen (GOET). Taxon 59(5): 1585-1601. https://doi.org/10.1002/tax.595024

Vorontsova MS, Knapp S (2016) A revision of the "spiny solanums," *Solanum* subgenus *Lepstostemonum* (Solanaceae), in Africa and Madagascar. Systematic Botany Monographs 99: 1-428. https://doi.org/10.1007/BF03027161

Vorontsova MS, Stern S, Bohs L, Knapp S (2013) African spiny *Solanum* (subgenus *Leptostemonum*, Solanaceae): A thorny phylogenetic tangle. Botanical Journal of the Linnean Society 173(2): 176-193. https://doi.org/10.1111/boj.12053

Wagner WL, Herbst DR, Sohmer SH (1999) Manual of the flowering plants of Hawai'i, 2 vols. Bishop Museum Special Publication 13. University of Hawaii and Bishop Museum Press, Honolulu.

Wahlert GA, Chiarini F, Bohs L (2015) A revision of *Solanum* section *Lathyrocarpum* (the Carolinense clade, Solanaceae). Systematic Botany 40(3): 853-887. https://doi.org/10.1600/036364415X689302

Walpers WG (1852) Solanaceae. In: Annales botanicae systematicae 3: 149-179.

Wang J-X, Gao T-G, Knapp S (2008) Ancient Chinese literature reveals pathways of eggplant domestication. Annals of Botany 102(6): 891-897. https://doi.org/10.1093/aob/mcn179

Weese T, Bohs L (2007) A three-gene phylogeny of the genus *Solanum* (Solanaceae). Systematic Botany 33(2): 445-463. https://doi.org/10.1600/036364407781179671

Wei Q, Wang J, Wang W, Hu T, Hu H, Bao C (2020) A high-quality chromosome-level genome assembly reveals genetics for important traits in eggplant. Horticulture Research 7(1): e153. https://doi.org/10.1038/s41438-020-00391-0

Whalen MD (1979) Taxonomy of *Solanum* section *Androceras*. Gentes Herbarium 11(6): 359-426.

Whalen MD (1984) Conspectus of species groups in *Solanum* subgenus *Leptostemonum*. Genes Herbarum 12: 179-282.

Whalen MD, Costich DE (1986) Andromonoecy in *Solanum*. In: D'Arcy WG (Ed.) Solanaceae: biology and systematics. Columbia University Press, New York, 284-302.

Whalen MD, Costich DE, Heiser Jr CB (1981) Taxonomy of *Solanum* section *Lasiocarpa*. Gentes Herbarum 12: 41-129.

Wikramanayake E, Dinerstein E, Loucks CJ, Olson DM, Morrison J, Lamoreux J, McKnight M, Hedao P (2002) Terrestrial ecoregions of the Indo-Pacific: a conservation assessment. Island Press, London, 824 pp.

Wood JRI (1984) Eight new species and taxonomic notes on the flora of Yemen. Kew Bulletin 39(1): 123-139. https://doi.org/10.2307/4107860

Wood JRI (1997) A handbook of the Yemen flora. Royal Botanic Gardens, Kew.

Wu CY, Huang SC (1978) Materiae ad floram Solanorum et Lycianthum Sinensium. Zhiwu Fenlei Xuebao 16: 72-80.

Wu CY, Huang SC (1979) Solanaceae. Flora Yunnanica. Vol. 2. Science Press, Beijing, 541-611.