<figure><img src="image_30.png" /><figcaption>**Figure 22.** Introduced species of *Solanum*. *Solanum chrysotrichum* Schltdl. **A** detail of inflorescence (field photograph, unvouchered, India). *Solanum elaeagnifolium* Cav. **B** habit (*Sampath Kumar et al. 126972*, India) **C** detail view of a flower (*Sampath Kumar et al. 126972*, India) **D** detail view of a fruit (field photograph, unvouchered, India). *Solanum jamaicense* Mill. **E** detail view of a flower (*Stern 265*, Trinidad and Tobago) **F** detail view of fruits (*Stern 265*, Trinidad and Tobago). *Solanum macrocarpon* L. **G** detail view of flower (in cultivation at GAFL Avignon, unvouchered) **H** detail view of fruit (in cultivation at Radboud University, Nijmegen, unvouchered, material now at CGN, Wageningen). Photograph credits: **A** S.Knapp **B–D** X. Aubriot **E, F** S. Stern **G, H** S. Knapp.</figcaption></figure>
The two collections from northern Papua New Guinea cited in the protologue of *S. dunalianum* var. *puberius* were destroyed in Berlin, and we have found no duplicates of these, despite extensive searches. Ledermann's travels are well-documented (Ledermann 1919; van Steenis-Kruseman 1985; Veldkamp et al. 1988), and many of the areas he visited have not been accessed since (Takeuchi and Golman 2002), so we delay neotypifying this name until such collections are available or duplicates of Ledermann's collections are found.

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

**13. Solanum elaeagnifolium Cav., Icon. 3: 22, tab. 243. 1795.**

Fig. 20B–D

No names based on tropical Asian types have been published; for complete synonymy see Knapp et al. 2017.

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

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

**Distribution.** *Solanum elaeagnifolium* has been rather widely collected in India (states of Karnakata, Maharashtra, Rajasthan, Tamil Nadu, and Uttar Pradesh); it is native to North and South America but widely adventive and invasive elsewhere (see Knapp et al. 2017).

**Discussion.** Although we have only seen *S. elaeagnifolium* specimens from India and Pakistan to date, we suspect this invasive species will become more widespread with time. *Solanum elaeagnifolium* is a plant of dry habitats and can become established easily through vegetative reproduction. It is easy to distinguish from native spiny solanums in tropical Asia by its silvery pubescence of lepidote trichomes and relatively narrowly elliptic leaves. It can form large stands in disturbed areas via rhizomes.

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

**14. Solanum forskalii Dunal, Hist. Nat. Solanum 237. 1813.**

Fig. 23

*Solanum villosum* Forssk., Fl. Aegypt.-Arab. 47. 1775, nom. illeg. non *Solanum villosum* Mill. Type. Yemen. Wadi Surdud, Feb 1763, *Herb. P. Forsskål* 414 (holotype: C [C10003107]).

*Solanum macilentum* A.Rich., Tent. Fl. Abyss. 2: 105. 1850. Type. Ethiopia. Choho, R. Quartin-Dillon & A. Petit s.n. (lectotype, designated by Lester 1997, pg. 286: P [P000343696]; isolectotypes: P [P0003436967, P000343698]).
*Solanum albicaule* Kotschy ex Dunal, Prodr. [A. P. de Candolle] 13(1): 204. 1852. Type. Sudan. Kordofan: Nubia, Cordofan Chursi, 29 Dec 1839, *C. Kotschy 309* (lectotype, designated by Lester 1997, pg. 287: G-DC [G00145871]; isolectotypes: B, BM [BM000778322], E [E00193284], ER, GOET, K [K000414013, K000414014, K000414015], LZ, M [M0105605, M0105606], MO [acc. # 3942648], MPU, NY, P [P00344693, P00344694, P00344695], STU [STU000027], TCD [TCD0000843], W [acc. # 1889-0293762, acc. # 0000634], WAG [WAG0003359]).

*Solanum heudelotii* Dunal, Prodr. [A. P. de Candolle] 13(1): 205. 1852. Type. Senegal. near Gabor, 1839, *J. Heudelot 417* (45) (lectotype, designated by Lester 1997, pg. 287: P [P00344004]; isolectotypes: K [K000414051], MPU, P [P00344005, P00344006], UPS).

*Solanum hadaq* Deflers, Bull. Soc. Bot. France 43: 122. 1896. Type. Yemen. Schoukra, Bilad Fodhli, J. Areys, 30 k a' l E.N.E. de Schughra, 150 m, 22 Mar 1890, *M. Deflers 377* (lectotype, designated by Vorontsova and Knapp 2016, pg. 159: P [P00051780]; isolectotypes: K [K000441183], P [P00051781, P00051782]).

*Solanum scindicum* Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 65: 542. 1896. Type. Pakistan. Sind: Scinde, *T. Cooke s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 159: CAL [CAL0000018701]; isolectotype: K [K000441223]).

*Solanum albicaule* Kotschy ex Dunal var. *parvifrons* Bitter, Repert. Spec. Nov. Regni Veg. Beih. 16: 102. 1923. Type. South Sudan. “Nördliches Darfur, El Fascher”, *J.D.C. Pfund 407* (syntypes: B, destroyed, Z, not found; no additional duplicates found).

**Type.** Based on (replacement name for) *Solanum villosum* Forssk.

**Description.** Erect or scandent shrub, 0.5–1 m, prickly. Young stems slender, ascendent to erect, densely stellate-pubescent and prickly, the pubescence of porrect, sessile or occasionally stalked trichomes, the stalks to 0.15 mm long, the rays 6–10, 0.15–0.3(–0.5) mm long, the midpoints same length as the rays or up to 1.5 mm, occasionally rounded, the prickles 3–10 mm long, 1–3 mm wide at the base, straight, occasionally curved, flattened, strongly reflexed, pale yellow to brown, spaced 1–10 mm apart; bark of older stems densely stellate-pubescent, sometimes glabrescent, brown to grey or orange-grey. Sympodial units difoliate, not geminate. Leaves simple, entire to weakly lobed, the blades 1–4(–6) cm long, 0.5–3(–4) cm wide, 1–2 times longer than wide, ovate, membranous to chartaceous, drying concolorous to discolorous, yellow-green to grey-green or brown-green, glabrescent to moderately stellate-pubescent on both surfaces, with porrect, sessile, sometimes stalked trichomes, the stalks to 0.2 mm long, the rays 6–10, 0.2–0.4(–1) mm long, the midpoints ca. same length as the rays, sometimes to 1.5 mm long, with 0(-3) prickles on both surfaces; the primary veins 2–3 pairs, the tertiary venation not visible to the naked eye; base cordate to rounded, the lobes 2(-3) on each side, 0.2–0.4 cm long, extending to 1/4 of the distance to the midvein, broadly deltate, apically rounded to obtuse; apex rounded to obtuse; petiole 0.2–1.6(–2.5) cm long, 1/4–2/3 of the leaf blade length, densely stellate-pubescent, with 0(-5) prickles. Buds ellipsoid, the corolla strongly exserted from the calyx before anthesis. Inflorescences apparently terminal or lateral, 2–6.5 cm long, unbranched or
<figure><img src="image_31.png" /><figcaption>**Figure 23.** *Solanum forsskalii* Dunal. Herbarium specimen collected in India in 1973 (Tiwari 941, CAL0000031126). Photograph credit: © The Director, Botanical Survey of India, Kolkata.</figcaption></figure>
forked, with (1–)2–20 flowers, 1–10 flowers open at any one time, densely stellate-pubescent, with 0(-5) prickles; peduncle 0.1–0.4(-1.5) cm long; pedicels 0.2–1 cm long, erect, articulated at the base, densely stellate-pubescent, with 0(-6) prickles; pedicel scars spaced 0.5–1.5 mm apart. Flowers 5-merous, apparently all perfect. Calyx 2–4.5 mm long, the lobes 0.5–2 mm long, ca. 1 mm wide, deltate, apically acuminate, moderately stellate-pubescent, with 0(-5) prickles. Corolla 1.3–2.4 cm in diameter, mauve to purple, stellate, lobed ca. 4/5 of its length, the lobes 6–11 mm long, 1.5–2 mm wide, deltate, reflexed or spreading, stellate-pubescent abaxially, the trichomes porrect, sessile or stalked, the stalks to 0.15 mm, the rays 4–10, 0.1–0.2 mm long, the midpoints shorter than the rays or to 0.5 mm long. Stamens equal; filament tube ca. 0.1 mm long; free portion of the filaments 0.7–1 mm long; anthers 4.5–7 mm long, connivent to spreading, tapering, yellow, poricidal at the tips, the pores directed distally. Ovary glabrous; style 0.9–1.2 cm long, slender, curved at the apex, glabrous. Fruit a globose berry, 1–10 per infructescence, 0.6–0.9 cm in diameter, the pericarp smooth, orange to red at maturity; fruiting pedicels 0.7–1.6 cm long, 0.3–0.4 mm in diameter at the base, 2–2.5 mm in diameter at the apex, spreading to somewhat deflexed from the weight of the fruit, with 0(-10) prickles; fruiting calyx not accrescent, covering 1/4(-1/2) of the mature fruit, with 0(-5) prickles. Seeds ca. 5–15 per berry, 2.5–4 mm long, 1.8–3 mm wide, flattened-reniform, almost black, the testal cells somewhat sinuate in outline. Chromosome number: n = 12 (Khatoon and Ali 1993), 2n = 24 (Al Wadi 2002).

**Distribution (Fig. 24).** *Solanum forskalii* is found from northeastern Africa and Arabian Peninsula to Pakistan and western India (Rajasthan and Gujarat).

**Ecology and habitat.** *Solanum forskalii* is a desert plant growing on stony ground and rocky slopes, often on granite, elevation not recorded on herbarium sheets we have seen from India (from sea level to 2,000 m elevation fide Vorontsova and Knapp 2016)

**Common names and uses.** India. Rajasthan: nar-kanta [Hindi] (Singh 1991, as *S. albicaule*).

**Conservation status (Knapp 2021b).** *Solanum forskalii* has been formally assessed as least concern (LC; https://www.iucnredlist.org/species/101526207/101526210).

**Discussion.** Although Vorontsova and Knapp (2016) suggested collections of *S. forskalii* from India might be accidental introductions, it is reported as fairly common in the Indian state of Rajasthan (Singh 1991). It is easily recognised by its white-coloured stems, resulting from the dense pubescence of overlapping stellate trichomes with long, delicate rays, and abundant, recurved prickles.

*Solanum forskalii* can be distinguished from *S. cordatum* by its porrect trichomes with 6–10 rays over 0.15 mm long on the young stems (versus trichomes with 12–18 rays under 0.15 mm long on the young stems of *S. cordatum*), 1–20 flowers per inflorescence (versus 1–2 flowers per inflorescence in *S. cordatum*), and anthers 4.5–7 mm long (versus anthers 3–5 mm long in *S. cordatum*). The trichome rays of *S. forskalii* are much thinner and more delicate than those of *S. cordatum*, which appear stout and almost lepidote. There is considerable variability within *S. forskalii* in Africa with respect to inflorescence length, prickle size and trichome midpoint length (Vorontsova and Knapp 2016).

**Specimens examined.** See Suppl. materials 1–3.
<figure><img src="image_32.png" /><figcaption>**Figure 24.** Distribution of *Solanum forsskalii*.</figcaption></figure>

**15. Solanum giganteum Jacq., Collectanea 4: 125. 1791.**

Figs 1A, 25

*Solanum niveum* Vahl, Symb. Bot. 2: 41. 1791. Type. South Africa. “Cape”, sin. loc., *C. Thunberg s. n.* (holotype: UPS-THUNB [microfiche 1036, no. 5209]).

*Solanum farinosum* Wallich in Roxburgh, Fl. Ind. (Carey & Wallich ed.) 2: 255. 1824. Type. India. “Babobad”, *Herb. Heyne s. n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 163: K [K000658975]).

*Solanum farinaceum* Griffith, Itin. Pl. Khasyah Mts. 111. 1848. Type. Bhutan. Dewanagiri, 1837, *W. Griffith 148* (no specimens cited; no original material found at K).

*Solanum giganteum* var. *tenuifolium* Dunal, Prodr. [A. P. de Candolle] 13(1): 259. 1852. Type. India. Tamil Nadu: Madras State, Nilgiri Hills [“Nellighery”], Nedaubetta, 1840, *G. S. Perrottet 520* (lectotype, designated by Vorontsova and Knapp 2016, pg. 163: P [P00049799]; isolectotype: P P00049800]).

*Solanum giganteum* var. *longifolium* Dunal, Prodr. [A. P. de Candolle] 13(1): 259. 1852. Type. India. Sin. loc., *Anonymous s. n.* [*Wallich cat. 2610*] (lectotype, designated by Vorontsova and Knapp 2016, pg. 163: G-DC [G00145950]).

*Solanum bequaertii* De Wild., Repert. Spec. Nov. Regni Veg. 13: 141. 1914. Type. Democratic Republic of the Congo. Katanga: Shaba Prov., Lubumbashi
[Elisabethville], 2 Mar 1912, *J. Bequaert 219* (holotype: BR [BR0000008994356]; isotype: BR [BR0000008993045]).

*Solanum sordidescens* Bitter, Bot. Jahrb. Syst. 57: 260. 1921. Type. Tanzania. Lindi: Kilwa-Kiwindje Distr., Tschumo, Matumbi Mts., 250 m, Jul 1903, *W. Busse* III 3097 (lectotype, designated by Vorontsova and Knapp 2016, pg. 164: EA [EA000001237]); isolectotypes: BM [BM001070317], BR [BR0000006495435]).

*Solanum seretii* De Wild., Miss. Em. Laurent, 1: 439, tab. 122. 1907. Type. Democrat-ic Republic of the Congo. Orientale: Kisangani, Bima to Bambili, 25 Oct 1905, *F. Seret 166* (holotype: BR [BR0000008993465]; isotype: BR [BR0000008993168]).

*Solanum muansense* Dammer, Bot. Jahrb. Syst. 48: 243. 1912. Type. Tanzania. Mwanza: Mwanza, May 1892, *F. L. Stuhlmann 4504* (type: B?, destroyed; no duplicates found).

**Type.** Cultivated in Vienna, original material from South Africa, Cape of Good Hope, *N.J. Jacquin s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 163: W [acc. # 0000608]; isolectotype: W [acc. # 0000609])

**Description.** Erect shrub to small tree, to 4 m tall, armed or occasionally unarmed. Stems erect, terete, prickly, densely stellate-pubescent; prickles to 7 mm long, to 5 mm wide at the base, straight, occasionally curved, deltate, orange-brown to almost white, glabrescent to stellate-pubescent in the lower 3/4; pubescence of stalked multangulate-stellate trichomes, the stalks to 0.7 mm long, the rays 10–25, 0.05–0.2 mm long, the midpoints ca. same length as the rays; new growth densely stellate-pubescent with a pale whitish grey pubescence; bark of older stems glabrescent to densely stellate-pubescent, light brown. Sympodial units plurifoliate, the leaves usually not geminate, if geminate the leaves of a pair differing slightly in size but not in shape, in addition often with small stipule-like leaves along the stem. Leaves simple, entire, the blades 12–40 cm long (if geminate the minor blades usually ca. 3 cm long), (3–)4–13 cm wide, ca. 2.5 times longer than wide, elliptic, chartaceous, strongly discolorous, unarmed; adaxial surface drying green-brown to red-brown, glabrescent; abaxial surface drying light grey-green, moderately to densely stellate-pubescent with multangulate stalked trichomes, the stalks to 0.3 mm long, the rays 8–25, 0.1–0.2 mm long, the midpoints ca. same length as the rays; major veins 8–12 pairs, the finer venation visible on both surfaces; base cuneate; margins entire; apex acute to acuminate; petiole 1.5–5.5 cm long, 1/8–1/4 of the leaf blade length, weakly to densely stellate-pubescent with multangulate trichomes like those of the blades, unarmed. Inflorescences 5–13(–20) cm long, apparently terminal or lateral, several times branched, with 30–150 flowers, 10–30 flowers open at any one time, weakly to densely stellate-pubescent with multangulate trichomes like those of the stems, unarmed; peduncle 20–60 mm long, unarmed; pedicels 0.4–1.5 cm long, 1–1.2 mm in diameter at the base, 2–2.5 mm in diameter at the apex, erect to recurved, unarmed, moderately to densely stellate-pubescent with multangulate trichomes like those of the inflorescence axes, articulated at the base; pedicel scars unevenly spaced 1.5–10 mm apart. Buds ovoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers 4–5-merous, apparently all perfect. Calyx with the tube 2.5–3 mm long, conical, the lobes 0.5–2.5(–4) mm long, 1.3–2.2 mm wide, deltate, apically obtuse to acute, un-
<figure><img src="image_33.png" /><figcaption>**Figure 25.** *Solanum giganteum* Jacq. **A** herbarium specimen collected in 'Brooks' (Sri Lanka) in 1931 (*Simpson* 8890, BM000900133) **B** detail of leaves and infructescence (field photograph, unvouchered, India) **C** detail of an inflorescence (field photograph, unvouchered, India). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B** P. Kumar **C** A. Prashant.</figcaption></figure>

armed, moderately to densely stellate-pubescent abaxially with multangulate trichomes like those of the rest of the plant, these often deciduous. Corolla 0.8–1.5 cm in diameter, usually mauve, sometimes white, stellate, lobed 2/3–3/4 of the way to the base, the lobes 3.5–6 mm long, 1.5–2.5 mm wide, deltate, spreading or not opening fully, moderately stellate-pubescent abaxially, the trichomes porrect, sessile or stalked, the stalks to 0.1 mm, the rays 8–15, 0.05–0.15 mm long, the midpoints ca. same length as the rays. Stamens equal; anthers 2.5–3 mm long, ca. 1 mm wide, connive nt, tapering, yellow, glabrous, poricidal at the tips, the pores not elongating to slits with drying; filament tube ca. 1 mm long, glabrous; free portion of the filaments 0.4–0.8 mm long, glabrous; Ovary conical, glabrous; style 0.45–0.65 cm long, slender, straight or gently curved, glabrous; stigma small capitate. Fruit a globose berry, many per infructescence, 0.6–0.8 cm in diameter,
<figure><img src="image_34.png" /><figcaption>**Figure 26.** Distribution of *S. giganteum*.</figcaption></figure>

the pericarp smooth, evenly green when young, bright red at maturity, glabrous; fruiting pedicels 0.8–1.6 cm long, 0.5–1.2 mm in diameter at the base, 1.2–2 mm in diameter at the apex, woody, erect, unarmed; fruiting calyx lobes not elongating, ca. 1/3 the length of the mature fruit, reflexed, unarmed. Seeds ca. 15–30 per berry, 2.3–2.9 mm long, 1.5–2.5 mm wide, flattened-reniform, dull yellow to orange-brown, the surface minutely pitted, testal cell margins sinuate. Chromosome number: not known.

**Distribution (Fig. 26).** *Solanum giganteum* is a widespread species occurring from Sri Lanka and southern India and throughout eastern and southern Africa (but is absent from western Africa, see Vorontsova and Knapp 2016). It is recently being recorded as adventive in Australia (http://www.flora.sa.gov.au/) where it has not yet naturalised.

**Ecology and habitat.** *Solanum giganteum* is a weedy species growing at the edges of forests and re-growth in clearings, disturbed areas, sometimes in rocky places or open grasslands; from sea level to 2,100 m elevation.

**Common names and uses.** English common names for *S. giganteum* in large parts of India are red bitter berry or healing-leaf tree; other local names in India are kurti [Marathi], paintilkkakkoti, paintilikam, peyccuntai-p-palam [Tamil] (Ved et al. 2016, envis.frlht.org); Kerala: cheruchunda (Ved et al. 2016, envis.frlht.org); Tamil Nadu: sambal kilurai [Tamil] (Matthew 1983); peyccundai, anaiccundai [Tamil] (Nair and Nayar 1987), putharichunda, sambal kilurae [Tamil] (Henry et al. 1987).
*Solanum giganteum* is planted as a shade tree and widely used medicinally in Africa for treatment of sores and ulcers (Hutchings et al. 1996) and in folk medicine in India (Ved et al. 2016).

**Conservation status (IUCN 2019).** *Solanum giganteum* has been formally assessed as LC (Least Concern) due to its wide range and weedy nature and is so listed on the IUCN Red List (Botanic Gardens Conservation International (BGCI) and IUCN SSC Global Tree Specialist Group 2020).

**Discussion.** *Solanum giganteum* is a common small tree in the understory of forests in India and Sri Lanka. Its large discolorous leaves accompanied by small stipule-like leaves along the stems, large bunches of bright red berries, deltate stem prickles and farinaceous white pubescence are distinctive and not shared by any other species in the region. The copious pubescence of multangulate trichomes is brittle and powdery, it easily detaches and rubs off leaves and stems, which is the origin of the specific epithet of the synonym *S. farinosum*. *Solanum vagum* of southern India and Sri Lanka is somewhat similar in overall appearance to *S. giganteum*, but has smaller, more lanceolate leaves, smaller inflorescences and less copious, non-deciduous pubescence (see description of *S. vagum*). The broad stem prickles common in young plants of *S. giganteum* are absent in *S. vagum*. Populations of *S. giganteum* in India are generally found to have smaller leaves and inflorescences than those of Africa (Vorontsova and Knapp 2016).

In analyses of phylogeny using molecular data *S. giganteum* is part of a strongly supported monophyletic group with African species *S. anomalum* Thonn., *S. schimperianum* A.Rich., *S. schleibenii* Werderm., *S. schumannianum* Dammer and *S. somalense* Franch. (Vorontsova et al. 2103). Also part of this group is the Indian species *S. pubescens* (Aubriot et al. 2016a).

*Solanum giganteum* has a long history of cultivation, both in its native range as a shade tree and for fencing and outside of its range in European botanical gardens (see Vorontsova and Knapp 2016). Edmonds (2012) suggests it is native only to the Cape region of South Africa, but we feel this is a biased assessment due to its early introduction from there into Europe.

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

**16. Solanum graciliflorum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 763. 1814.**
Fig. 27

*Solanum athroanthum* Dunal, Prodr. [A. P. de Candolle] 13(1): 208. 1852. Type. Indonesia. Java: [Prov. Banjinwanyne] "in sylvis prope Sukaradja" [Sukaraja], 1846, *H. Zollinger 2907* (holotype: G-DC [G00145833]; isotypes: BM [BM000778325], G [G00301684, G003043306], MPU [MPU012648], P [P00368939, P00368940, P00368941]).

**Type.** Based on an unpublished illustration of Leschenault collection kept in the Node-Véran collection in Montpellier (lectotype, designated by Aubriot et al. 2016b, pg. 100: Sol. Tab. 47 [MPU028534]).
<figure><img src="image_35.png" /><figcaption>**Figure 27.** *Solanum graciliflorum* Dunal Herbarium specimen (epitype) collected in Indonesia (*Horsfield s.n.*, BM000886121). Photograph credit: CC-BY, © copyright The Trustees of the Natural History Museum, London.</figcaption></figure>
**Description.** Scandent shrub to 2 m tall, armed. Stems apparently erect to de-cumbent, prickly and very sparsely stellate-pubescent; prickles to 7 mm long, to 8 mm wide at the base, curved, deltate, laterally flattened, pale yellow, glabrous; trichomes porrect-stellate, sessile to subsessile, the rays (4–)5–8, 0.1–0.25 mm long, the mid-points to 0.15 mm long; new growth sparsely stellate-pubescent; bark of older stems dark brownish grey, glabrescent. Sympodial units difoliate, the leaves geminate, usu-ally similar in size. Leaves simple, more or less deeply lobed, the blades (4.5–)7–11 cm long, (1.5–)3–5 cm wide, ca. 2 times longer than wide, elliptic to ovate, chartaceous, slightly discolorous, usually prickly with 1–10(–12) prickles per leaf side, mostly along the midvein, to 9 mm long, to 2 mm wide at the base, straight or slightly curved at the tip, awl-shaped, conical, pale yellow, glabrous; adaxial and abaxial surfaces sparsely to very sparsely stellate-pubescent with sessile to subsessile porrect-stellate trichomes, the rays 6–8, 0.1–0.25 mm long, the midpoint to 0.25 mm long, usually as long as the rays; major veins 3–4 pairs drying dark; base attenuate to truncate; margins shallowly to deeply lobed, the lobes 1–3 on each side, 0.5–2.5 cm long, broadly deltate, api-cally rounded, the sinuses extending up to 2/3 to the midrib; apex rounded to acute; petiole 0.5–1.8 cm long, 1/10–1/6 of the leaf blade length, unarmed or prickly with 1–2 prickles like those of the blades, sparsely stellate-pubescent with porrect, subsessile trichomes denser at the very base. Inflorescences 2–4 cm long, leaf-opposed or appar-ently lateral and borne between leaf pairs, unbranched to up to 6 times branched, with 15–50+ flowers, many flowers open at any one time, sparsely to very sparsely stellate-pubescent with trichomes like those of the stems, unarmed; peduncle 1–2(–2.5) cm long, with 0–1 prickles like those of the leaves and stems; pedicels 4–7 mm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, erect, unarmed and very sparsely stellate-pubescent with trichomes like those of the stems, articulated at the base; pedicel scars spaced 1–5 mm apart. Buds ellipsoid, strongly exerted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 1–1.5 mm long, campanulate, the lobes 0.25–0.75 mm long, 0.25–0.75 mm wide, deltate, apically acute, unarmed and pubescent abaxially with sessile porrect-stellate trichomes like those of the stems. Corolla 0.5–1 cm in diameter, white to pale lilac, stellate, lobed nearly to the base, the lobes 4–5 mm long, ca. 1 mm wide, narrowly del-tate to linear, reflexed at anthesis, glabrous adaxially, densely stellate-pubescent abaxi-ally with sessile porrect trichomes where exposed in bud. Stamens slightly unequal; anthers unequal, three of the five 4.5–5 mm long and two 3–4 mm long, all 0.5–0.75 mm wide, somewhat connivent, tapering, yellow, glabrous, poricidal at the tips, the pores not elongating to slits with drying; filament tube <0.5 mm long, glabrous; free portion of the filaments almost equal, 0.5–1.25 mm long, glabrous. Ovary conical, minutely glandular-puberulent; style ca. 5.5 mm long, slender, curved at the apex, gla-brous; sigma capitate, the surfaces minutely papillate. Fruit a globose berry, several to many per infructescence, 0.3–0.5 cm in diameter, red when mature, the pericarp shiny, glabrous; fruiting pedicels 0.8–1.2 cm long, ca. 0.5 mm in diameter at the base, taper-ing to a slightly enlarged apex, 0.75–1 mm in diameter at the apex, somewhat woody, spreading, unarmed; fruiting calyx lobes slightly expanding to 1.5 mm long, ca. 1/5
<figure><img src="image_36.png" /><figcaption>**Figure 28.** Distribution of *Solanum graciliflorum*.</figcaption></figure>

the length of the mature fruit, deltate to lanceolate, unarmed. Seeds 6–9 per berry, 3.5–4 mm long, 3–3.5 mm wide, flattened-reniform, orange-brown, the surface minutely pitted, the testal cells pentagonal in outline. Chromosome number: not known.

**Distribution (Fig. 28).** *Solanum graciliflorum* is known from only a few collections from the islands of Java, Bali, Sulawesi and Ambon (Indonesia).

**Ecology and habitat.** *Solanum graciliflorum* is a plant of tropical rainforest under-
story; elevation not recorded.

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

**Preliminary conservation status (IUCN 2019).** Near Threatened (NT). EOO (131,936 km²); AOO (16 km²). *Solanum graciliflorum* is known only from seven collections, several of which are of uncertain localities. It has not been re-collected since the first half of the 20th century, indicating it is certainly of conservation concern, thus we suggest a threat status of Near Threatened to reflect this. Additional collection of this species and exploration of the type locality are priorities.

**Discussion.** *Solanum graciliflorum* is morphologically similar to the sympatric *S. cyanocarphium* in being a weak, scrambling plant with hooked prickles, but differs from it in its branched inflorescences, smaller flowers and fruits and calyx that is not accrescent in fruit. The inflorescence of *S. cyanocarphium* is unbranched and the calyx strongly accrescent and covers the berry in fruit.

Aubriot et al. (2016b) superfluously lectotypified *S. athroanthum* with the specimen in G-DC (G00145833) but Dunal (1852) clearly cited only this specimen in the protologue, thus making it the holotype.

**Specimens examined.** See Suppl. materials 1–3.
17. *Solanum harmandii* Bonati, Bull. Soc. Bot. Genève, sér. 2, 3: 310. 1914.
Fig. 29

**Type.** CAMBODIA. Sin. loc., *F.J. Harmands.n.* (lectotype, designated here: P [P00055939]).

**Description.** Shrub, size unknown, armed. Stems erect, terete, prickly and densely stellate-pubescent; prickles to 1 cm long, to 0.5 cm wide at the base, straight, del-tate, laterally flattened, orange brownish, sparsely to moderately stellate-pubescent in the lower half; trichomes porrect-stellate, mixture of sessile and stalked, the stalks to 0.1 mm long, the rays 5–8, 0.1–0.4 mm long, the midpoints absent or up to 0.1 mm long, with bulbous bases; new growth densely pubescent, light brownish; bark of older stems greyish, moderately stellate-pubescent. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, shallowly lobed, the blades 6–16.5 cm long, 3–10 cm wide, ca. 1.5 times longer than wide, elliptic to ovate, chartaceous, discolorous, un-armed on both surfaces, or occasionally with a few prickles along the midvein; adaxial surface green, moderately to densely stellate-pubescent, the stellate trichomes porrect, sessile to stalked, the stalks to 0.2 mm long, the rays 5–8, 0.1–0.5 mm long, the midpoints to 0.25 mm long; abaxial surface densely whitish stellate-pubescent with trichomes like those of the adaxial surface; major veins 5–7 pairs drying light-green; base shortly attenuate to truncate; margins shallowly lobed, the lobes 3–4 on each side, 0.5–1.5 cm long, broadly deltate, apically rounded, the sinuses less than halfway to the midrib; apex acute; petiole 1–2.5 cm long, 1/10–1/5 of the leaf blade length, unarmed or prickly with 1–3 prickles, densely stellate-pubescent. Inflorescences 2.5–4 cm long, apparently lateral, forked or 2 times branched, with ca. 8–9 flowers, 2 flowers open at any one time, densely whitish stellate-pubescent, trichomes like those of the stems but with longer stalks, unarmed; peduncle ca. 0.5 cm long, unarmed; pedicels 0.25–0.4 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, erect, unarmed, densely whitish stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars spaced 1.5–3 mm apart. Flowers 5-merous, apparently all perfect. Calyx with the tube ca. 2 mm long, campanulate, the lobes 1–2.5 mm long, 1.5–2 mm wide, deltate with an elongate acute apex, unarmed and densely whitish stellate-pubescent with trichomes like those of the pedicels. Corolla 0.7–1.2 cm in diameter, colour unknown, stellate, lobed 1/2–2/3 of the way to the base, the lobes 5–7 mm long, 2.5–4 mm wide, deltate, spreading at anthesis, glabrous adaxially, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal; anthers ca. 8 mm long, ca. 1 mm wide, glabrous, not tightly connivent, tapering, orange, glabrous, poricidal at the tips, the pores not elongating to slits with drying; filament tube <0.5 mm long, glabrous; free portion of the filaments ca. 0.5 mm long, glabrous. Ovary conical, with simple glandular hairs in the upper 1/4; style ca. 3.5 mm long, slender, curved at the apex, glabrous; stigma capitate, minutely papillate. Fruits and seeds unknown. Chromosome number: not known.

**Distribution (Fig. 30).** *Solanum harmandii* is endemic to Cambodia, but without specific locality.
<figure><img src="image_37.png" /><figcaption>**Figure 29.** *Solanum harmandii* Bonati Herbarium specimen (lectotype) collected in Cambodia (*Harmand s.n.*, P00055939). Photograph credit: CC-BY, Muséum national d'Histoire naturelle, Paris.</figcaption></figure>
<figure><img src="image_38.png" /><figcaption>**Figure 30.** Distribution of *S. harmandii*.</figcaption></figure>

**Ecology and habitat.** *Solanum harmandii* is known only from the type collection, on which no habitat information was recorded.

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

**Preliminary conservation status (IUCN 2019).** Data Deficient (DD). Known only from the type collection, probably collected between 1875 and 1877 (see below). No locality or habitat information was recorded which makes the recollection of this plant a priority.

**Discussion.** *Solanum harmandii* is a stout shrub that is morphologically very different from all the other spiny solanums treated here. The growth form is reminiscent of members of the Brevantherum clade (sensu Gagnon et al. 2022), but it possesses copious prickles and in flower morphology it is a typical spiny solanum with tapering anthers opening by distally directed pores. *Solanum giganteum* of the Indian subcontinent has similar overall morphology, with many-branched inflorescences, but differs from *S. harmandii* in its scurfy white pubescence of multangulate trichomes and entire leaves that are usually almost glabrous adaxially.

Hul and Dy Phon (2014) cited “holo-P” in their treatment of *S. harmandii*, but Bonati (1914) cited no herbarium in any part of the publication of this species. Although he was known to have worked in Paris, this does not constitute effective lectotypification. We here designate the single specimen in P (P00055939) as the lectotype. The type specimen was collected during a French expedition led by Dr. Jules Harmand,
then medical officer in the region previously known as Indochina, between 1875 and
1877 (Arousseau 1922); detailed itineraries are not available for his collecting trips.

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

**18. Solanum hovei Dunal, Prodr. [A. P. de Candolle] 13(1): 311. 1852.**

Figs 2E, 4B, 31

*Solanum jacquemontii* Dunal, Prodr. [A. P. de Candolle] 13(1): 287. 1852. Type. India.
Sin. loc., *V.V. Jacquemont* 563 [593 in protologue] (lectotype, designated here:
MPU [MPU858235]; isolectotype: P [P00049910]).

**Type.** INDIA. Gujarat: Ahmedabad district, Dolca [Dholka] near Sabermatty, *A.P. Hove s.n.* (holotype: BM [BM000900293]).

**Description.** Subshrubs to shrubs, to 1.5 m tall, armed. Stems erect, terete, prickly, sparsely stellate-pubescent; prickles to 7 mm long, to 3.5 mm in diameter at the base, straight to slightly recurved, broad-based, laterally compressed, light orange to brown, glabrous; pubescence of mixed sessile and short-stalked porrect-stellate trichomes, the stalks ca. 0.3 mm long, the rays 4–8, ca. 0.3–0.4 mm long, the midpoint up to 0.3 mm long; new growth sparsely to densely stellate-pubescent; bark of older stems grey to pale greenish brown, glabrescent. Sympodial units difoliate, the leaves usually geminate. Leaves simple, shallowly lobed, the blades 5–21 cm long, 2–9 cm wide, ca. 2–3 times longer than wide, rhombic to subelliptic, chartaceous, discolorous, armed along the central and occasionally the lateral veins, the prickles to 1 cm long, to 2 mm at the base, straight, flattened, often tinged with purple at the base in live plants; adaxial surface dark green, moderately to sparsely stellate-pubescent, the trichomes porrect, thick-stalked, the stalks to 0.2 mm, the rays 3–8, 0.1–0.4 mm long, the midpoint to 1 mm; abaxial surface light green, moderately to densely stellate-pubescent, trichomes a mixture of porrect and multangulate, sessile or short-stalked, the stalks to 0.4 mm, the midpoint and lateral rays relatively equal in length, ca. 0.3–0.4 mm; major veins 4–6 pairs; base cuneate to short attenuate, somewhat oblique; margins entire to shallowly lobed, the lobes 2–3 on each side, 0.1–0.5 cm long, triangular, broadly acute to obtuse, the sinuses less than 1/4 of the way to the midrib; apex narrowly acute; petiole 0.7–2.5 cm, ca. 1/10–1/5 of the leaf blade length, moderately to densely stellate-pubescent, prickly or sometimes unarmed, the prickles similar to those on the blades. Inflorescences 0.4–4 cm long, extra-axillary, unbranched, with ca. 4–10 flowers, 1–3 flowers open at any one time, moderately to densely stellate-pubescent, glabrescent, with a mix of sessile and short-stalked stellate-porrect trichomes like those of the stems, unarmed; peduncle ca. 0–1.3 cm long, unarmed; pedicels 5–9 mm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading at anthesis, unarmed or prickly with a few straight and broad-based prickles, sparsely to moderately pubescent with stellate-porrect trichomes like those of the stems, articulated at the base; pedicel scars spaced 0.5–8 mm apart. Buds ellipsoid to ovoid, exserted from the calyx before anthesis. Flowers 5-merous,
<figure><img src="image_39.png" /><figcaption>**Figure 31.** *Solanum hovei* Dunal **A** herbarium specimen (holotype) collected in India (*Hove s.n.*, BM000900293) **B** habit and leaves (field photograph, unvouchered, India) **C** detail of a fertile branch (field photograph, unvouchered, India) **D** detail view of a flower (field photograph, unvouchered, India) **E** detail view of a fruit (field photograph, unvouchered, India). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B–E** X. Aubriot.</figcaption></figure>
apparently all perfect. Calyx with the tube 0.7–2.5 mm long, conical, the lobes 0.5–1.6 mm long, 1.3–2 mm wide, broadly deltate, unarmed or very sparsely prickly with straight to slightly recurved prickles, moderately to densely stellate-pubescent abaxially with porrect-stellate trichomes like those of the pedicels, sometimes purple-tinged on living plants. Corolla 1.2–1.5 cm in diameter, light blue or purple, rotate-stellate, lobed ca. halfway to the base, interpretalar tissue somewhat present, the lobes 4–5.5 mm long, 3–4 mm wide, ovate, spreading at anthesis, sparsely stellate-pubescent along the midvein and on the lower half of lobe adaxially, the hairs sessile, multangulate, moderately to densely stellate-pubescent abaxially, the hairs subsessile, porrect-stellate and multangulate. Stamens equal; anthers 4–5.5 mm long, 0.6–1.2 mm wide, orange or orangish yellow, connivent, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments 0.3–0.4 mm long, glabrous. Ovary oblong-conical, moderately stellate-pubescent towards the apex; style 6–9 mm long, sparsely to moderately stellate-pubescent on proximal 2/3–3/4, the hairs sessile, porrect-stellate or multangulate; stigma subcapitate, the surfaces minutely papillose. Fruit a subglobose berry, 1–5 per infructescence, 0.6–1 m in diameter, orange when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 1–1.5 cm long, ca. 0.6 mm in diameter at the base, ca. 2 mm in diameter at the apex, strongly deflexed; fruiting calyx not markedly accrescent, the lobes 2–3 mm long, 1–1.5 mm wide, often breaking off in dry material. Seeds 10–20 per berry, 2.8–4.1 mm long, 2.2–3 mm wide, flattened reniform, yellowish brown, the surfaces minutely pitted, the testal cells with slightly sinuate margins. Chromosome number: not known.

**Distribution (Fig. 32).** *Solanum hovei* is endemic to India and occurs primarily along the Western Ghats range in the states of Goa, Gujarat, Karnataka and Maharashtra with most of the collections of this species having been made in the surroundings of Mumbai and Pune.

**Ecology and habitat.** *Solanum hovei* grows in open areas on plateaus and along roadsides and forest outskirts from 150 to 1,200 m elevation.

**Common names and uses.** India. Goa: mothirigani (Naithani et al. 1997).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (173,911 km², LC); AOO (564 km², VU). *Solanum hovei* is a relatively common and widely distributed species in the Western Ghats biodiversity hotspot.

**Discussion.** *Solanum hovei* is a member of a group of species identified in the molecular analyses of Aubriot et al. (2016a) as the ‘S. violaceum group’. It is morphologically similar to the sympatric and widespread *S. violaceum* but differs from it in having leaves with more sparsely pubescent adaxial surfaces and cuneate (rather than acute or truncate) bases. The pedicels of *S. hovei* are strongly deflexed in fruit, while those of *S. violaceum* are spreading. *Solanum multiflorum* of western India also has strongly deflexed pedicels, but they are shorter, and the leaves are densely pubescent with longer trichomes.

Sen Gupta (1964) suggested *S. hovei* was a rare plant in the Western Ghats, but the number of recent collections in local herbaria suggest otherwise. It is probable that *S. hovei* was largely confused with *S. violaceum* in herbaria (usually identified as *S. indicum*).
<figure><img src="image_40.png" /><figcaption>**Figure 32.** Distribution of *S. hovei*.</figcaption></figure>

The protologue of *S. jacquemontii* (Dunal 1852) cites duplicates of "*Jacquemont 593*" in both P and MPU. The specimen we have selected as the lectotype has an annotation with this name in Dunal's hand and is numbered "563", we are treating this as an error to be corrected in the protologue.

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

**19. Solanum insanum L., Mant. 1: 46. 1767.**

Figs 1F, 3E, 4A, 33

*Solanum undatum* Lam., Tabl. Encycl. 2: 22. 1794. Type. Mauritius. Sin. loc., "Insula Franciae", *J. Martin s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 197: P-LA (P00357695, Morton neg. 8392]).

*Solanum trungum* Poir., Encycl. (Lamarck) 4: 308. 1797. Type. Indonesia. Malaku: "Amboina" [Ambon Island] (no specimens cited; lectotype, designated by Merrill 1917, pg. 463, as "type": "Trungum agreste spinosum", Rumphius, Herb. Ambion. 5: 240, t. 86, f.1. 1747).
*Solanum gula* Buch.-Ham., Trans. Linn. Soc. London 14(2): 267. 1824. Type. India.
Karnakata: Kowdhalli “Caudhully [protologue – Mysore]”, Oct 1800, *J. Buchanan-Hamilton s.n.* (lectotype, designated here: BM [BM000900233]).

*Solanum himalense* Dunal, Prodr. [A. P. de Candolle] 13(1): 300. 1852. Type. India.
“Sivala & sub Himala” [protologue: In Himalae et Sivalae montibus altit. 4600 ped., Edgeworth, pl. exs. Ind. n. 116], *M.P. Edgeworth 116* (holotype: MPU [MPU981115]).

*Solanum indicum* Nees var. *pubescens* Dunal, Prodr. [A. P. de Candolle] 13(1): 310. 1852. Type. [India]. Sin. loc., *N. Wallich s.n.* [Wallich Catal. 2626g] (lectotype, designated here: G-DC [G00130396]; isolectotype: K-W [K001116650 pro parte, R-hand two stems]).

*Solanum melanocarpum* Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852, nom. illeg. superfl. Type. Based on *Solanum insanum* L. (cited in synonymy; “Nomen insanum mutavi, quia adeo non insanum videtur ut potius fructus edules credibilius sunt, quemadmodum nomina insanum et melongean, saepissime fuerunt confusa” [I changed the name insanum because it does not seem harmful, the fruits are probably edible, and insanum and melongena were very often getting confused]).

*Solanum melanocarpum* Dunal var. *atropurpureum* Dunal, Prodr. [A. P. de Candolle] 13(1): 356. 1852. Type. Indonesia. Java: sin. loc., *H. Zollinger 698* (lectotype, designated here: G-DC [G00131525]; isolectotypes: G [G00301647], G-DC [G00131526, as *Zollinger 698b*], P [P00379566, P00379567]).

*Solanum undatum* Lam. var. *aurantiacum* Dunal, Prodr. [A. P. de Candolle] 13(1): 359. 1852. Type. Cultivated, sin. loc., *Anonymous s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 198: G-DC [G00131519]).

*Solanum undatum* Lam. var. *violaceum* Dunal, Prodr. [A. P. de Candolle] 13(1): 359. 1852. Type. Mauritius. Sin. loc., *L. Bouton s.n.* (lectotype, designated by Vorontsova and Knapp 2016, pg. 198: G-DC [G00131506]; isolectotype: G [G00131504]).

*Solanum trungum* Poir. var. *divaricatum* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Sri Lanka. “Trinquemalay”, *A.A. Reynaud s.n.* (lectotype, designated here: P [P00049887]).

*Solanum trungum* Poir. var. *rumphii* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852, nom. illeg. Type. Described using same material as *S. trungum* Poir. (should be var. *trungum*)

*Solanum trungum* Poir. var. *sinuatopinnatifidum* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852, nom. illeg. Type. Based on *Solanum trungum* Poir. [sensu Blume 1826] (should be var. *trungum*)

*Solanum trungum* Poir. var. *tongdongense* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Myanmar. “Tong dong”, 7 Jan 1827, *N. Wallich s.n.* [Wallich Catal. Burm. 135] (lectotype, designated here: G-DC [G00131531]; isolectotype: G-DC [G00131371]).

*Solanum album* Lour. var. *gaudichaudii* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Vietnam. Sin. loc., 1839, “Cochinchina”, *C. Gaudichaud s.n.* (holotype: G-DC [G00131563]).
*Solanum cyanocarphium* Blume var. *obtusangulum* Dunal, Prodr. [A. P. de Candolle] 13(1): 362. 1852. Type. Indonesia. Java: Sin. loc., 1837, *J.C. von Hoffmannsegg* 119 (lectotype, designated here: G-DC [G00131561]).

*Solanum cumingii* Dunal, Prodr. [A. P. de Candolle] 13(1): 363. 1852. Type. Philippines. Sin. loc., *H. Cuming* 443 (lectotype, designated here: G [G00076245]; isolectotypes: BM [BM000778205], E [E00190694], G [G00076246, G00076247], K [K000195960, K000195961], L [L0003628], LE, ME [MEL2444029], P [P00578602, P00578613], W [acc. # 0000622, acc. # 1889-0084423]).

*Solanum immane* Hance ex Walp., Ann. Bot. Syst. 3(1): 165. 1852. Type. China. Hongkong: "Hong Kong", [*Herb.] H.F. Hance 860* (no specimens or herbaria cited; lectotype, designated here: BM [BM000942450]).

*Solanum melongena* L. var. *insanum* Prain, Bengal Pl. 746. 1903. Type. Based on *Solanum insanum* L.

*Solanum mirikense* C.R.Mukhop., J. Indian Soc. Bot. 72(1-2): 185. 1993. Type. India. West Bengal: Darjeeling, Mirik, 1700 m, 5 Oct 1985, *C.R. Mukhopadhyay* 428A (holotype: CAL [CAL0000018719]).

*Solanum melongena* L. var. *cumingii* (Dunal) J.Samuels, Proc. XV EUCARPIA Mtg. 258. 2013. Type. Based on *Solanum cumingii* Dunal

*Solanum melongena* L. subsp. *insanum* Banfi, Galasso & Bartolucci, Nat. Hist. Sci. 15(1): 19. 2017. Type. Based on *Solanum insanum* L.

**Type.** INDIA. Gujarat: Surat, *Anonymous s.n.* (lectotype, designated by Hepper and Jaeger 1985, pg. 389: LINN [acc. # 248.29]).

**Description.** Erect shrub, to 1 m tall, armed. Stems erect, terete, prickly, moderately stellate-pubescent to glabrescent; prickles to 8 mm long, to 5 mm wide at the base, straight, flattened, yellow-orange, glabrous; pubescence of stalked porrect-stellate trichomes, the stalks to 0.2 mm long, the rays 6–12, 0.2–0.4 mm long, the midpoints ca. same length as the rays or elongated to 1 mm; new growth densely stellate-pubescent, light brownish in dry material; bark of older stems grey to brown, glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, moderately lobed, the blades 2.5–12 cm long, 1.3–8 cm wide, ca. 1.5 times longer than wide, ovate, chartaceous, concolorous to weakly discolorous, armed with 2–20 prickles on both surfaces, these yellow-tan or purple-tinged; adaxial and abaxial surfaces yellow-green, moderately pubescent, with porrect-stellate trichomes, sessile or stalked, the stalks to 0.2 mm, the rays 5–8, 0.2–0.7(–1) mm long, the midpoints ca. same length as the rays; major veins 3–5 pairs; base truncate, sometimes obtuse; margins lobed, the lobes 2–3 on each side, 0.5–1.2 cm long, broad-deltate, apically rounded, the sinuses less than halfway to the midrib; apex rounded to acute; petiole 0.7–3 cm long, 1/4–1/3 of the leaf blade length, unarmed or prickly with 1–5 prickles, moderately stellate-pubescent to glabrescent, the pubescence with stellate-porrect trichomes like those of the stems. Inflorescences 2.5–3.5 cm long, apparently terminal or lateral, unbranched, with 1–3 flowers, 1 flower open at any one time, moderately to densely stellate-pubescent with porrect trichomes like those of the stems, unarmed; peduncle 0–13 mm long, unarmed
<figure><img src="image_41.png" /><figcaption>**Figure 33.** *Solanum insanum* L. **A** herbarium specimen collected in India in 1994 (*Chorley & Bharatan 3*, BM000013803) **B** habit (*Meeboonya et al. RM 257*) **C** detail view of a hermaphroditic flower (*Meeboonya et al. RM 251*, Thailand) **D** detail view of an inflorescence with an opened staminate flower (*Meeboonya et al. RM 257*, Thailand) **E** detail view of an immature fruit (*Meeboonya et al. RM 277*, Thailand). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B–E** X. Aubriot.</figcaption></figure>
or with a few prickles; pedicels 0.8–2.5 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, erect, unarmed or with a few scattered prickles, densely stellate-pubescent with porrect trichomes like those of the inflorescence axes, articulated at the base; pedicel scars spaced 1–2 mm apart. Buds ovoid, poorly exserted from the calyx before anthesis. Flowers 5(-6)-merous, heterostylous and the plants andromonoecious, with the lowermost flower(s) long-styled and hermaphrodite, the distal flowers short-styled and staminate. Calyx with the tube 4–5 mm long, campanulate, the lobes 4–6 mm long, 2–2.5 mm wide, deltate, apically acute, unarmed or with 1–15 prickles, densely stellate-pubescent with porrect trichomes like those of the pedicels. Corolla 1.8–2.5 cm in diameter, mauve or white, almost rotate with abundant interpetalar tissue, lobed ca. 1/4 of the way to the base, the lobes ca. 7 mm long, ca. 10 mm wide, broadly deltate, spreading at anthesis, glabrous to sparsely stellate-pubescent adaxially with porrect-trichomes mostly on the midvein and the tips, densely stellate-pubescent abaxially with variously stalked porrect trichomes. Stamens equal; anthers 4–5 mm long, ca. 1 mm wide, tapering, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube ca. 1.5 mm long, glabrous; free portion of the filaments 1–2 mm long, glabrous. Ovary conical to ovoid, stellate-pubescent and glandular in the upper 1/4; style 2–3 mm in short-styled flowers, 5–7 mm long in long-styled flowers, moderately stellate-pubescent in the lower 1/2; stigma capitate or occasionally somewhat bilobed, the surfaces minutely papillose. Fruit a globose berry, 1 per infructescence, 1.5–3 cm in diameter, dark green with pale green and cream markings when young, yellow at maturity, the pericarp smooth, glabrous; fruiting pedicels 1.5–2.2 cm long, 1.5–3 mm in diameter at the base, 4–5 mm in diameter at the apex, unarmed or with a few stout prickles, woody, pendulous; fruiting calyx lobes elongating to 9–15 mm long, 1/4–1/3 the length of the mature fruit, reflexed, with 2–30 prickles. Seeds ca. 50–150 per berry, 2.4–2.8 mm long, 1.8–2.2 mm wide, flattened-reniform, orange-brown, the surface minutely pitted, the testal cells with sinuate margins. Chromosome number: n = 12 (Meyer et al. 2012, as *S. undatum*).

**Distribution (Fig. 34).** *Solanum insanum* is widely distributed in India and south-eastern Asia (east to the Philippines) and is also found as far west as Madagascar and Mauritius (possibly introduced there from Asia for its medicinal properties).

**Ecology and habitat.** *Solanum insanum* is a plant of degraded scrubland and secondary vegetation and occurs from sea level to ca. 500 m elevation.

**Common names and uses.** Cambodia. trâp rôm nhong (Hul and Dy Phon 2014), trôp (Collard 56), trôp som nhon (Collard 32), China. ye qie (Zhang et al. 1994); Hainan: ngou ke tzi (Lei 392). India. Andhra Pradesh: challa mulaga (Foulkes 83); Chattisgarh: anpa, jangli bhata (Kumar CNH-15565); Rajasthan: bhutkataiya [Hindi] (Singh 1991, as *S. incanum*); Tamil Nadu: thalamoolagah, nullamoolaga (Wight 1573 Ab); Uttar Pradesh: bari bhatkatanja (Bell 863), jangli bengan (Sarin NC-5079). Indonesia. Borneo: terong nasi (Ambriansyah 687); Java: terong-glate (Hoffmansegg 119). Laos. mok kua kim (Spire 914), mok khma lê (Pottier 263). Malaysia/Singapore. têrong (with various qualifiers, Burkill 1935). Malaysia. Sabah: soguntung [Rungus]
<figure><img src="image_42.png" /><figcaption>**Figure 34.** Distribution of *S. insanum*.</figcaption></figure>

(Jones 152). Singapore. terong nyonya (Corner SFN-37775). Sri Lanka. Uva: ela batu [Sinhala] (Hepper & Silva 4719). Vietnam. heng khôm (Spire 1081), cà co [Annamite] (Pierre 706), la plonh [Moi] (Poilane 85), blon blok (Schmid 1608).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (10,914,652 km², LC); AOO (1,052 km², VU). *Solanum insanum* is a widespread weedy species of disturbed areas and agricultural margins. It was not considered of conservation concern in an assessment of eggplant wild relatives (Syfert et al. 2016).

**Discussion.** *Solanum insanum* is the wild progenitor of the cultivated eggplant (aubergine; *S. melongena*) and readily crosses with that species (Davidar et al. 2015; Mutegi et al. 2015), thus complicating its recognition in the past (see references and discussion in Knapp et al. 2013; Ranil et al. 2016). We have chosen to recognise the two as distinct species rather than as infraspecific taxa (e.g., Lester and Hasan 1991; Daunay and Hazra 2012) because cultivated and wild plants are on distinct evolutionary trajectories. Wild plants are subject to natural selection, while cultivated plants are commensal with human beings. We recognise that this can lead to difficulties in identifying individual specimens, particularly those from old collections where details of provenance are not clear.

Distinguishing *S. insanum* from the cultivated *S. melongena* can be difficult in the absence of fruits. Keys in previous publications (Knapp et al. 2013; Ranil et al. 2016) can be used for identification, but in general specimens we assign to *S. insanum* are pricklier, with smaller flowers and fruit, and more ruderal than those we assign to *S. melongena*. Local cultivars of *S. melongena* in tropical Asia can be especially difficult to distinguish from *S. insanum*.

*Solanum incanum* L. is another species with which *S. insanum* is frequently confused, and the name *S. incanum* has frequently been used in the past literature for
all “wild eggplants” (see Knapp et al. 2013 for the history of nomenclature of the wild eggplant relatives both in Asia and Africa). *Solanum incanum* occurs from across northern Africa and the Middle East to Pakistan; we have seen no specimens identifiable as *S. incanum* from the area treated here. The many specimens labelled “Punjab” from early collections in “British India” have all proved to be from Pakistan, not from adjacent India, but *S. incanum* potentially occurs near border areas. *Solanum incanum* is a more densely pubescent plant than is *S. insanum*, with long-stalked trichomes that dry a distinctive yellow colour (see Vorontsova and Knapp 2016 for a description) and plants are generally pricklier, although this character is extremely variable within these taxa; it generally grows in much drier habitats than does *S. insanum*. Crossability of these two taxa is not known, confusion over the identity of accessions used in crossing studies (e.g., Daunay and Hazra 2012) means that this remains to be tested.

Mill (2001: 1050) suggested *S. mirikense*, treated as a synonym of *S. insanum* here, was synonymous with *S. mauritianum* Scop., an introduced member of the Brevantherum clade. He had not seen the type specimen in CAL, which clearly places it as a synonym of *S. insanum*.

The sole element cited in Poiret's (1797) description of *S. trungum* is the treatment and illustration of "Trungum agreste spinosum" in Rumphius (1747), and Merrill (1917) effectively lectotypified the name with this illustration by stating "type". The illustration is of a very spiny plant, which Rumphius (1747) stated was wild and described as "ingrata et spinosa herba" (ungrateful and spiny herb).

The name *S. gula* has been placed in the synonymy of various spiny solanums over the years (R.N. Lester, m.s.). In the protologue Buchanan-Hamilton (1824) states "I found it first in my journey to Mysore, where it is called Gula, and in 1806 I gave specimens, a drawing and a description of it to Sir J.E. Smith". No specimens are extant in the Smith herbarium at the Linnean Society of London, but the drawing held there (see https://linnean-online.org/168572/) is clearly of *S. insanum* and has been annotated as such in James Edward Smith's hand. A specimen at BM (BM000900233) is labelled "Solanum gula B" by Buchanan-Hamilton and also has an annotation of "insanum L" in Smith's hand. This is certainly the specimen mentioned in the protologue and is here designated as the lectotype of *S. gula*.

In the protologue of *S. indicum* var. *pubescens* Dunal (1852) cited two Wallich catalogue numbers, "2629 a, ex parte" and "2626 g", and stated that he had seen specimens in G-DC and "herb. Wall.". The catalogue number 2629 is a mixture of various species (including *S. pubescens*), so we have chosen the G-DC sheet of Wallich cat. 2626g (G00130396) as the lectotype because it is unambiguous, it matches the protologue and it was clearly seen by Dunal. The specimen labelled "Wallich cat. 2626G" at Kew in the Wallich Herbarium (K00116650) is a mixture of two taxa; the left-hand stem corresponds to *S. multiflorum* and the two right hand stems to *S. insanum*, only these two right-hand stems are considered isotype material here.

Two of the four infraspecific taxa described by Dunal (1852) under *S. trungum* were described with no specimens cited, the only references are to previously published works and the descriptions are copied from those. Var. *sinuato-pinnatifidum* cites only "Blum. bijdr. 700, *karondong*, nomen Indicum"; this is a reference to Blume's (1826:
702, an error in citation of page) *S. trungum* Poir., where the name karondong is stated, but no material or localities cited. We consider this homotypic with *S. trungum* and do not recognise Blume's treatment of *S. trungum* as a distinct species. We consider var. *rumphii* to be illegitimate because Dunal (1852) cited the same type as that of *S. trungum*; Dunal appears to be using this as the “typical” variety. Both of the other varieties had specimens cited. Two Wallich catalogue numbers (burm. 135, 2628E) were cited in the protologue of *S. trungum* var. *tongdongense*, both of which were seen in G-DC (“v.s. h. DC”). We have selected one of the duplicates of the best preserved of these, Wallich cat. iter. Burm. 135, at G-DC as the lectotype. The protologue of var. *divaricatum* cites two collections (*Perottet s.n.* from “Pondicherry” [India], *Reynaud s.n.* from “Ceylon”) both seen in Paris. Both collections are *S. insanum*; we have selected the Reynaud specimen from Sri Lanka (P00049887) as the lectotype because it has both flowers and fruits.

Two collections from Java, *Hoffmansegg 119* in G-DC and *Roemer s.n.* in G (”herb. Boiss.”) were cited in the protologue of *S. cyanocarphium* var. *obtusangulum*; we have selected the first of these as the lectotype of the name because the number makes it unambiguous. Although Java collections are attributed to him (as here), Hoffmansegg never visited Java (van Steenis-Kruseman 1985); plants in his herbarium were collected for him there by a friend.

Of the many duplicates of *Cuming 443*, the sole collection cited in the protologue of *S. cumingi*, we have selected the one in G cited by Dunal (1852, as ”herb. Boiss.”) that bears an annotation label in hand as the lectotype (G00076245).

The protologue of *S. immane* cites no specimens or herbaria, but the name (and probably the description) is taken from Henry Fletcher Hance's unpublished manuscript describing Chinese plants (Walpers 1852). Walpers worked in Berlin, and any specimens he saw are likely to have been destroyed. Of the specimens we have seen that could correspond to original material or duplicates of the Hance collection used to describe *S. immane*, only one specimen at BM (*Herb. Hance 860*, BM000942450) matches both the protologue description and the collection locality; we have selected this as the lectotype for the name. Other duplicates of *Hance 860* in P are either from different localities (i.e., P00049897, from “prope Cantonem”) or represent different species (e.g., P00055662 = *S. violaceum*).

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

**20. Solanum involucratum Blume, Bijdr. Fl. Ned. Ind. 13: 701. 1826.**

Figs 2G, 3C, 4C, 35

*Solanum ferox* L. var. * involucratum* (Blume) Miq., Fl. Ned. Ind. 2: 647. 1857. Type.
Based on *Solanum involucratum* Blume.

**Type.** INDONESIA. Java: Sin. loc., *C.L. Blume s.n.* (lectotype, designated here: L [L0003633]).
**Description.** Shrub to 2 m tall, strongly armed. Stems erect, terete, densely prickly and pubescent; prickles to 8 mm long, to 3 mm at the base, straight, awl-shaped to del-tate, pale yellow, glabrous; trichomes porrect-stellate, mixture of subsessile and stalked, the stalks to 0.75 mm long, the rays 4–6, 0.4–1.5 mm long, the midpoints shorter than the rays or up to 2.25 mm long, sometimes purple-tinged on living plants; new growth densely pubescent with mixture of subsessile and long-stalked stellate trichomes; bark of older stems brownish, glabrescent. Sympodial units difoliate, the leaves not gemi-nate. Leaves simple, shallowly to deeply lobed, the blades 12–30 cm long, 9.5–19 cm wide, ca. 1–1.5 times longer than wide, broadly elliptic to ovate, chartaceous, discolor-ous, armed with ca. 20–80 prickles per leaf side, prickles to 20 mm long, to 6 mm wide at the base, straight, awl-shaped, conical, pale yellow, on dried material, sometimes purple-tinged on living plants, glabrous; adaxial surface moderately pubescent, with sessile porrect-stellate trichomes, the rays 3–6, 0.1–0.25 mm long, the midpoints to 1.75 mm; abaxial surface densely stellate-pubescent with trichomes like those of the adaxial surface but stalked, the stalks to 0.4 mm; major veins 5–6 pairs drying light green; base attenuate; margins shallowly to deeply lobed, the lobes 5–6 on each side, 1–1.2 cm long, 3–5 cm wide, broadly deltate, apically acute, the sinuses less than halfway to the midrib; apex acute; petiole 3.5–17.5 cm long, 1/3–4/5 of the leaf blade length, prickly with 5–17 prickles like those of the blade, densely stellate-pubescent with porrect trichomes like those of the stem. Inflorescences apparently lateral, 3–8 cm long, unbranched, with 3–10 flowers, 1 to many flowers open at any one time, densely stellate pubescent with stellate-porrect trichomes like those of the stems but often tinted with purple, unarmed; peduncle 1–4 mm long, unarmed; pedicels 3–10 mm long, 1–2 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading, unarmed or sparsely prickly with a few prickles, densely stellate-pubescent with pur-ple porrect-trichomes like the inflorescence axes, articulated at the base; pedicel scars spaced 0.5–1 mm apart. Buds ovoid, included in the calyx lobes until just before an-thesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 3.5–4 mm long, cup-shaped and slightly inflated at the base and appearing saccate, the lobes 4–6 mm long, 4–4.5 mm wide, deltate to broadly deltate, apically acute, armed with numerous prickles and densely stellate pubescent with purple-tinted porrect-trichomes like those of the pedicels. Corolla 1–1.2 cm in diameter, white, stellate, lobed ca. 1/2–2/3 of the way to the base, interpretalar tissue present, the lobes 4–6 mm long, 2–3 mm wide, deltate, spreading at anthesis, glabrous adaxially, densely stellate-pubescent abaxially on parts exposed in bud with trichomes like those of the calyx but white. Stamens equal; anthers 5–6 mm long, 1.5–2 mm wide, connivent, tapering, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube ca. 0.5 mm long, glabrous; free portion of the filaments ca. 0.25 mm long, glabrous. Ovary conical, densely covered with long simple hairs; style ca. 6 mm long, slender, curved at the apex, glabrous; stigma capitate, the surfaces minutely papillose. Fruit a globose berry, 3–10 per infructescence, 1–2 cm in diameter, orange when mature, the pericarp thick, very densely stellate-pubescent with porrect-trichomes like those of the adaxial surface of the leaves; fruiting pedicels 0.8–2 cm long, ca. 1 mm in diameter at
<figure><img src="image_43.png" /><figcaption>**Figure 35.** *Solanum involucratum* Blume **A** herbarium specimen collected in Thailand in 1930 (Kerr 19844, BM000886046) **B** Habit and leaves (field photograph, unvouchered, Vietnam) **C** detail of the tri-chomes and prickles on the adaxial surface of a blade (field photograph, unvouchered, Vietnam) **D** detail view of an inflorescence and a flower (field photograph, unvouchered, Vietnam) **E** detail view of an accrescent fruiting calyx (field photograph, unvouchered, Vietnam). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B–E** M. Nuraliev.</figcaption></figure>
<figure><img src="image_44.png" /><figcaption>**Figure 36.** Distribution of *Solanum involucratum*.</figcaption></figure>

the base, 2.5–2.8 mm in diameter at the apex, woody, erect to spreading, armed with 3–25 prickles; fruiting calyx lobes expanding to 2 cm long, completely enclosing the fruit at maturity, armed with ca. 10–20 prickles per lobe, the lobes white with purplish venation and trichomes. Seeds >100 per berry, 2–3 mm long, 1.5–2 mm wide, flattened-reniform, dull yellow, the surface minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.

**Distribution (Fig. 36).** *Solanum involucratum* is widely distributed in Indochina and Malay Archipelago (Indonesia, Malaysia, and Christmas Island).

**Ecology and habitat.** *Solanum involucratum* is a plant of tropical evergreen or deciduous rainforest, it grows on limestone or on clay soil, also in secondary forests; from 50 to 1,400 m elevation.

**Common names and uses.** Laos. hma:k sè:n, khüa khünx, hma:k khüa hlè:z, khüa hna:m (Hul and Dy Phon 2014). Thailand. Prachinburi: sa-uk [Thai] (*Kerr 19844*); Prachuap Khiri Khan: ma uk muak [Thai] (*Kerr 10780*); Vietnam. ca duoc nui [Vietnamese] (*Fleury 29910*); cà du’o’c nui, cây cà rai, cà ung, co chu [Munong] (Hul and Dy Phon 2014).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (1,868,808 km², LC); AOO (216 km², EN). *Solanum involucratum* is found over a very broad range in the archipelago, the AOO measurement certainly reflects collecting bias.

**Discussion.** *Solanum involucratum* is a distinctive species, with pubescent berries entirely enclosed within a spiny accrescent calyx. In flower the corolla lobes are shorter than the calyx, so the flowers appear to be entirely within the calyx at anthesis. In flower the calyx tube appears to be somewhat inflated, but it is appressed against the berry in fruit. The bristly, prickly accrescent calyx in fruit is similar to that of
*S. barbisetum*, *S. cyanocarphium* and *S. praetermissum*, but all of those species have glabrous fruits, while those of *S. involucratum* are densely pubescent with stiff, stellate trichomes. The inflorescences of *S. cyanocarphium* are few-flowered like those of *S. involucratum*, but not robust. Inflorescences of *S. barbisetum* and *S. praetermissum* are elongate and multi-flowered.

On the island of Sulawesi *S. involucratum* is sympatric with the very similar *S. sulawesi*. It differs from that species in its accrescent calyx in fruit, larger mature berries and more densely pubescent stems and leaves. Both *S. involucratum* and *S. lasiocarpum* have leaves with petioles that are longer relative to leaf size than *S. sulawesi*, but these three species can be difficult to distinguish.

*Solanum involucratum* was considered by Miquel (1857) to be a variety of *S. lasiocarpum* (as *S. ferox* var. * involucratum*). The two species are not closely related; *S. involucratum* is a member of a clade comprising *S. procumbens*, *S. expedunculatum* Symon and *S. leptacanthum* Merr. & L.M.Perry (the latter two species from New Guinea), itself apparently related, but with poor support, to *S. barbisetum* and *S. praetermissum* (Aubriot et al. 2016a). *Solanum lasiocarpum* on the other hand has been shown in numerous studies to be a member of the otherwise American Lasiocarpa clade (e.g., Bruneau et al. 1995; Bohs 2004; Stern et al. 2011).

Heiser (1996, 2001) followed Miquel (1857) in using the name *S. ferox* L., nom utique rej. (see Knapp 2011) for *S. involucratum*, recognising it at the varietal level, while Whalen et al. (1981) regarded it as ambiguous. Care should be taken with specimens annotated as *S. ferox* in collections, they could be either *S. lasiocarpum* (more common in our experience, see discussion under that species) or *S. involucratum*.

Hul and Dy Phon (2014) incorrectly cited a specimen at L as “holo – L!” for *S. involucratum*. This is not a holotype (see McNeill 2014) and we have selected as lectotype a specimen at L (L0003633) that matches the protologue and appears to have been original material; this is probably the same sheet referred to as holotype by Hul and Dy Phon (2014).

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

**21. *Solanum jamaicense* Mill., Gard. Dict., ed. 8 Solanum 17. 1768.**

Fig. 22E, F

No names based on tropical Asian types have been published; for complete synonymy
see D'Arcy 1974.

**Type.** JAMAICA: “Solanum bacciferum caule et foliis tomentis incanis spiosis, flor luteo, fructu croceo, minore Sl. Cat: 107” [as Herb. Sloane 107 in D’Arcy 1974a, a correctable error], *W. Houstoun s.n.* (lectotype, designated by D’Arcy 1974a, pg. 865, as “type”: BM [BM000815972]).

**Description.** D’Arcy (1974a: 685–686); Nee (1993: 77–78); http://www.solanaceae source.org/solanaceae/solanum-jamaicense.

**Distribution.** *Solanum jamaicense* has been recorded from Indonesia (islands of Borneo, Java, Sulawesi, and Sumatra) and the Philippines in tropical Asia, the sin-
gle collection we have seen from China was cultivated; it is native to the Caribbean, Central and South America, where it occurs in forests and forest margins. It is also known from the state of Florida (United States of America).

**Discussion.** *Solanum jamaicense* is apparently only sporadically adventive in rain-forest areas; most collections are from around logging camps. It is not easily confused with any other from the region. Its sharply hooked prickles, attenuate, decurrent leaves, and tiny flowers and fruits are distinctive.

*Solanum jamaicense* has been identified as a potential problematic weed in the state of Florida (Diaz et al. 2008) where it is locally invasive in the hammock habitat in the central part of the state where it was first recorded in 1930. It has not yet, however, become more widely distributed there.

D'Arcy (1974a) inadvertently lectotypified *S. jamaicense* by citing the specimen collected by William Houstoun at BM (BM000815972) labelled with the polynomial used in the protologue that came from Philip Miller's herbarium. Although Miller (1768) did not cite Houstoun explicitly in the protologue of *S. jamaicense*, he had access to Houstoun's collections before 1768, so this specimen qualifies as original material. He did cite Houstoun in association with this polynomial in earlier editions of the "Gardener's dictionary" (J. Wajar, pers. comm.). An annotation at the base of the twig in Daniel Solander's hand "Solanum Jamiacense Mill dict" indicates the sheet was part of Miller's herbarium used for the dictionary and had been filed as *S. jamaicense* prior to the herbarium coming to BM. We consider the use of "Herb. Sloane" rather than "Sl[oane]. Cat[alogue]:" by D'Arcy (1974a), and subsequently followed by others (Nee 1993), to be a correctable error; therefore, a second step lectotypification is not needed.

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

**22. *Solanum kachinense* X.Aubriot & S.Knapp, sp. nov.**

urn:lsid:ipni.org:names:77298803-1

Fig. 37

**Diagnosis.** Like *Solanum torvum* Sw., but differing in the eglandular pubescence of the inflorescence, more highly branched inflorescences, smaller flowers and almost glabrous mature foliage.

**Type.** MYANMAR. Kachin: Putao District, "North Triangle (Hkinlum)", 29 Jul 1953, *F. Kingdon-Ward 21211* (holotype: BM [BM000900353]; isotype: A).

**Description.** Shrubs of unknown height, armed. Stems erect, terete, drying black, prickly and sparsely stellate-pubescent; prickles broad-based, to 1 cm long, 0.8 cm wide at the base, straight, straw-colored; pubescence of mixed sessile and very short-stalked porrect-stellate trichomes, the stalks to 0.1 mm, the rays 6–8, ca. 0.5 mm long, the midpoints to 0.2 mm long, always shorter than the rays; new growth densely stellate-pubescent, mixed sessile and short-stalked porrect-stellate like those of the stems, the trichomes golden, soon deciduous and the stems glabrate; bark of older stems brown or dark brown. Sympodial units difoliate, the leaves geminate, the leaves of a pair only differing in size. Leaves simple, shallowly lobed, the blades 16–22 cm long, 8–11.5 cm
<figure><img src="image_45.png" /><figcaption>**Figure 37.** *Solanum kachinense* X.Aubriot & S.Knapp – Herbarium specimen (holotype) collected in Myanmar in 1953 (Kingdon-Ward 21211, BM000900353). Photograph credit: CC-BY, © copyright The Trustees of the Natural History Museum, London.</figcaption></figure>
wide, ca. 1.5 times longer than wide, elliptic, widest at the middle or just below, chart-
aceous, discolorous, armed or unarmed; adaxial surface glabrous to sparsely stellate-
pubescent along the veins with a few scattered sessile porrect-stellate trichomes, the
rays 5–6, ca. 0.5 mm long, the midpoints equal to the rays; abaxial surface similarly
but more densely pubescent with mixed sessile and short-stalked porrect-stellate tri-
chomes, the stalks if present less than 0.2 mm long, the trichomes still only along the
veins; major veins 6–8 pairs, drying black, very sparsely stellate-pubescent especially
abaxially, sometimes armed, the prickles if present 1–3 per face, to 1.5 cm long; base
truncate, strongly oblique with the basiscopic side 1–2 cm further along the petiole;
margins shallowly lobed, the lobes broadly deltate with acute tips, the sinuses less than
halfway to the midrib; apex acute; petioles 1.8–3 cm long, ca. 1/10 as long as the leaf
blades, unarmed and glabrous to stellate-pubescent with a few golden porrect-stellate
trichomes like those of the stems, drying blackish brown. Inflorescences 3–6 cm long,
internodal and lateral, forked to 5-times branched, with 50–60 flowers, many flowers
open at any one time, glabrous but sparsely pubescent with porrect-stellate trichomes
at the distal ends; peduncle 1.5–4 cm long, glabrate; pedicels 0.5–0.7 cm long, ca.
1 mm in diameter at the base, ca. 1 mm in diameter at the apex, nodding at anthesis,
sparsely stellate-pubescent with golden porrect-stellate trichomes like the inflorescence
axes, articulated at the base; pedicel closely spaced ca. 1 mm apart or slightly overlap-
ping. Buds elongate and tapering, strongly exserted from the calyx before anthesis.
Flowers 5-merous, apparently all perfect. Calyx with the tube 1.5–2 mm long, conical,
scarious and irregularly ripping into lobes at anthesis, the lobes 2–3.5 mm long (in-
cluding the tip), ca. 2 mm wide, deltate with a subulate tip 1.5–2 mm long, sparsely
stellate-pubescent like the pedicels, the subulate tip glabrous. Corolla 1.5–1.8 cm in
diameter, “purple sometimes white” (fide *Kingdon Ward 21211*), stellate, lobed nearly
to the base or 3/4 of the way to the base, with little interpretalar tissue, the lobes 6.5–
8 mm long, 2.5–4 mm wide, spreading or perhaps somewhat reflexed, glabrous adaxi-
ally or with a few stellate trichomes along the petal midvein, densely stellate-pubescent
abaxially, the trichomes porrect-stellate with 4–8 weak, tangled rays. Stamens equal;
anthers 5–5.5 mm long, ca. 1 mm wide, tapering, yellow, poricidal at the tips, the
pores directed distally, not elongating to slits with drying; filament tube minute, gla-
brous; free portion of the filaments ca. 1 mm long, glabrous. Ovary conical, glabrous;
style 6–8 mm long, glabrous; stigma clavate, the surfaces minutely papillose. Fruit and
seeds not known. Chromosome number: not known.

**Etymology.** *Solanum kachinense* is named in honour of the province of Myanmar where it occurs, and for the Kachin peoples of the region – who generously assisted the Kingdon-Wards while they were collecting in the region.

**Distribution (Fig. 38).** *Solanum kachinense* is known only from the type collection, made in northern Myanmar in the drainage of the Mali Kha, one of the eastern tributaries of the Irrawaddy River whose waters are fed by the Himalayan glaciers of Tibet.

**Ecology and habitat.** The type specimen indicates *S. kachinense* is a “village weed of very sour soil” indicating it grows, as do many solanums, in disturbed or open areas; the village of Hkinlum is at approximately 1,200 m elevation in a region where tropical
<figure><img src="image_46.png" /><figcaption>**Figure 38.** Distribution of *S. kachinense*.</figcaption></figure>

and temperate elements of the flora mix; Kingdon-Ward (1956) characterised the vegetation of Hkinlum as “moist warm temperate evergreen forest. Not subtropical…”

**Common names and uses.** Myanmar. “fruits bitter, not eaten” (*Kingdon-Ward 21211*).

**Preliminary conservation status (IUCN 2019).** Data Deficient (DD). Known only from a single collection, data on the distribution and status of *S. kachinense* is insufficiently known to speculate about its conservation status.

**Discussion.** Despite being currently known to us from a single collection, we describe *S. kachinense* here because it is so distinct, and that future botanists will be encouraged to find more collections because the entity is known to exist. Very often botanists pass by spiny solanums as uninteresting, in part due to the ubiquity of many introduced species such as *S. torvum*.

Morphologically, *S. kachinense* is similar to members of the Torva clade (sensu Stern et al. 2011; Aubriot et al. 2016a) with difoliate sympodia and lateral or leaf-opposed inflorescences with many flowers and fruits. It differs from all other Torva clade species in the region (*S. chrysotrichum*, *S. comitis*, *S. poka*, *S. peikuoense*, *S. pseudosaponaceum*, *S. torvoideum* and *S. torvum*) in its glabrate leaves and stems with long (to 1 cm) very broad-based prickles, especially on stems It has larger flowers than the almost sympatric *S. pseudosaponaceum* (1–1.8 cm in diameter versus 1–1.5 mm). The glandular simple
trichomes present on the inflorescence axes of *S. torvum* (see description in Vorontsova and Knapp 2016 and on Solanaceae Source) are absent in *S. kachinense*.

The glabrate upper leaf surfaces are similar to those of *S. giganteum* but the copious scurfy white pubescence of inflorescences, leaf undersides and stems on that species is distinctive; even in more glabrate individuals of *S. giganteum* the pubescence is of multangulate trichomes, not of porrect-stellate trichomes like those of *S. kachinense*.

*Solanum kachinense* was collected on Frank and Jean Kingdon-Ward's last trip to the drainages of the upper Irrawaddy in Myanmar to collect in the high mountains bordering China. Hkinlum was their base for their forays into the higher mountains collecting rhododendrons, lilies and alpines for the horticultural trade in England. This plant was not mentioned in Kingdon-Ward (1956); it was collected in the hot, wet season between the Kingdon-Ward's forays into the mountains, spring for flowers and autumn for seeds when they were perhaps a bit fed up with being stuck in a hot, wet village.

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

## 23. *Solanum lasiocarpum* Dunal, Hist. Nat. Solanum 222. 1813.

Figs 1C, 2A, 39

*Solanum indicum* L., Sp. Pl. 187. 1753. nom. utique rej. Type. Sri Lanka. Sin. loc., "Habitat in India utraque", *Anonymousus s.n.* (lectotype, designated by Hepper 1978, pg. 555: Herb. Hermann 3: 16, No. 94 [BM000594658]).

*Solanum hirsutum* Roxb. ex Wall., Fl. Ind. (Carey & Wallich ed.) 2: 253. 1824, nom. illeg., not *Solanum hirsutum* Dunal, 1813. Type. probably based on *Solanum lasiocarpum* Dunal.

*Solanum quadriloculare* Spreng., Syst. Veg., ed. 16 [Sprengel] 4(2, Cur. Post.): 72. 1827. Type. based on *Solanum Solanum hirsutum* Roxb. ex Wall.

*Solanum zeylanicum* Blanco, Fl. Filip. [F.M. Blanco] 136. 1837, nom. illeg., non *Solanum zeylanicum* Scop., 1786. Type. Philippines (no specimens or illustrations cited; no specimens extant). Philippines. Luzon, Benguet province, May 1914, E.D. Merrill [*Species Blancoanae*] 465 (neotype, designated here: A [00230113]; isoneotypes: CAL [acc. # 316470], K [K000195921], P [P00369171]).

*Solanum lasiocarpum* Dunal var. *velutinum* Dunal, Prodr. [A. P. de Candolle] 13(1): 253. 1852. Type. Philippines. Sin. loc., 1841, *H. Cuming* 690 (lectotype, designated here: P [P00379526]; isolectotypes: BM [BM000886159], K [K000195954, K000195955], G [G00301654], P [P00379525]).

*Solanum ferox* L. var. *lasiocarpum* (Dunal) Miq., Fl. Ned. Ind. 2: 647. 1857. Type. Based on *Solanum lasiocarpum* Dunal.

*Solanum lasiocarpum* Dunal var. *domesticum* Heiser, Solanaceae: Biol. & Syst. (ed. D'Arcy) 413. 1986. Type. Cultivated in the Indiana University greenhouses from seeds of "fruits purchased in market in Bangkok, Thailand, Sept. 1, 1980" [collection date mistakenly as "September 21, 1980" in the protologue], 30 Jun 1981, C.B. Heiser 8008 (holotype: IND [IND-1000065]).
**Type.** INDIA. Sin. loc. (lectotype, designated by Whalen et al. 1981, pg. 100: [illustration] Rheede von tot Draakestein, Hort. Malab. 2: tab. 35, "Ana-Schunda". 1680).

**Description.** Shrub to 2 m tall, often heavily armed. Stems erect to spreading, terete, unarmed or sparsely to moderately prickly and stellate-pubescent; prickles to 0.5–1 cm long, broad-based, deltate to narrowly deltate, laterally compressed, very variable in size on a single stem; trichomes porrect-stellate, mixture of sessile and variably stalked, the stalks multiseriate, to 2.5 mm long, the rays 5–8, 0.4–1.5 mm long, the midpoints variable in length, shorter or longer than the rays; new growth densely pubescent; bark of older stems brownish, moderately to sparsely prickly and stellate-pubescent. Sympodial units difoliate, the leaves geminate, the leaves of a pair equal in size and shape or the minor leaf slightly smaller. Leaves simple, shallowly lobed, the blades 20–35 cm long, 15–30 cm wide, ca. 1–1.5 times longer than wide, broadly ovate to ovate, chartaceous, discolorous, armed with prickles similar to those of the stem, these scattered along the midrib and principal lateral veins, often dark-purplished tinged; adaxial surface felty, densely stellate pubescent with porrect sessile trichomes, the rays 2–5, 0.1–0.5 mm long, the midpoints to 2.5 mm long; abaxial surface felty, densely stellate pubescent and farinaceous in appearance, with porrect trichomes like those of the adaxial surface but with longer stalks, the stalks to 2 mm long; principal veins 5–7 pairs, often drying yellowish abaxially; base truncate or obtuse; margins shallowly to moderately lobed, the lobes 3–6 on each side, occasionally somewhat dentate, 1.6–4 cm long, 3–5 cm wide, deltate to broadly deltate, acute or rounded at the tips, the sinuses less than halfway to the midrib; apex acute; petioles 3–14 cm, 1/5–1/2 the length of the blades, densely stellate-pubescent, unarmed or with a few to many prickles like those of the stems, the pubescence with a mixture of stipitate and sessile porrect-trichomes like those of the stem. Inflorescences 0.4–0.9 cm, extra-axillary, often very close to a leaf pair, unbranched, with 6–16 flowers, only one or two open at a time, densely stellate-pubescent, frequently prickly, the prickles like those of the stems; peduncle 0.1–0.4 cm, unarmed or with a few prickles; pedicels 4–9 mm long, 1–1.5 mm in diameter at the base, 1.5–2 mm in diameter at the apex, spreading or somewhat reflexed at anthesis, densely stellate-pubescent like the inflorescence axes, unarmed or with a few prickles, articulated at the base; pedicel scars 0.25–1.25 mm apart. Buds ovoid, the corolla enclosed within the calyx lobes and tube until just before anthesis. Flowers 5-merous, heterostylous and the plants andromonoecious, with the lowermost flower long-styled and hermaphrodite, the distal flowers short-styled and staminate. Calyx with the tube 2.5–4.5 mm long, broadly campanulate, the lobes 3–5 mm long, 2.5–5 mm wide, deltate to broadly deltate, apically acute, unarmed and densely stellate-pubescent with porrect-stellate trichomes like those of the pedicels but the midpoints usually somewhat longer. Corolla 2.5–3.5 cm in diameter, white, stellate, lobed 1/2–3/4 of the way to the base, interpretalar tissue absent, the lobes 6–9 long, 3–6 mm wide, broadly ovate, spreading at anthesis, the tips somewhat reflexed, glabrous adaxially, densely stellate-pubescent abaxially with porrect-stellate trichomes where exposed in bud. Stamens equal or slightly unequal; filament tube minute; free portion of the filaments 0.1–0.2 mm long, glabrous; anthers 6–8.5 mm long, 1.5–2.2 mm wide, strongly
<figure><img src="image_47.png" /><figcaption>**Figure 39.** *Solanum lasiocarpum* Dunal **A** herbarium specimen collected in Thailand in 1910 (Kerr 1236, BM000886387) **B** habit (Meeboonya et al. RM 287, Thailand) **C** detail view of a calyx and a floral bud (Meeboonya et al. RM 272, Thailand) **D** detail view of an inflorescence with immature fruits (Wang et al. 2063, China) **E** detail view of a mature fruit (Wang et al. 2063, China). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B, C** X. Aubriot **D, E** S. Knapp.</figcaption></figure>
tapering, connivant but the tips somewhat spreading, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying. Ovary conical, pubescent, the hairs appearing simple but with poorly developed and very short rays at the base; style of long-styled flowers 5–10 mm long, glabrous, 1–2 mm long in short-styled flowers; stigma capitate, the surface minutely papillose. Fruits a globose berry, 1–5 per inflorescence, 2.5–3.5 cm in diameter, orange when ripe, the pericarp thin, densely stellate pubescent with sessile porrect-trichomes, the rays 5–15, 0.1–0.4 mm long, the midpoints always longer than the rays, 1.5–4 mm long; fruiting pedicels 1–1.5 cm long, ca. 2.5 mm in diameter at the base, ca. 4 mm in diameter at the apex, woody, spreading or deflexed from weight of the berry; fruiting calyx not markedly accrescent, the lobes only slightly lengthening in fruit and often breaking off, usually unarmed. Seeds 100–200+, 2.2–3.5 mm long, 1.75–2.5 mm wide, flattened reniform, tan, the surfaces minutely pitted, the testal cells with sinuate margins. Chromosome number: n = 12 (Whalen et al. 1981; Madhavaian 196814, as *Solanum ferox* L.).

**Distribution (Fig. 40).** *Solanum lasiocarpum* is a commonly cultivated plant and occurs from tropical India east through Indochina, extreme southern China, Malaysia and Indonesia to the Philippines and the island of New Guinea.

**Ecology and habitat.** *Solanum lasiocarpum* grows in forest openings, disturbed sites and second growth thickets, from sea level to 1,000 m elevation. Widely cultivated in the region for its fruit.

**Common names and uses.** Unless otherwise indicated, common names are from Whalen et al. (1981: 103); languages when recorded are in square brackets. Brunei Darussalam: tarang asai, tarung bawi (*Bernstein 133*); China: mo ke shue (*Tsang 16102*); India: ram-begun [Bengali]; Kerala vellathu-vazhuthana (Mohanan and Henry 1994, as *S. ferox*). Indonesia. Maluku: tomate hutan (*Nooteboom 5837*), trong baguri (*Robinson 286*); Sumatra: lattoeeng (many collections), terong hutan (*Lörzing 15014*), sontoman (*si Toroes 2848*); Laos: chou pout din (*Vidal 5438*), gim ông (*Vidal 5438*); Malaysia (general): tar-ong-asam [Malay]; Sabah: tarong sulok [Kadayan] (*Abdul 2854*), terong pipit or terong pasai (*Cuadra A-2160*), tokung [Sungei] (*Cuadra A-1169*); Malaysia/Singapore: tërong asam (Burkill 1935); Philippines: basula [Ibang], dabatung [Sulu], tagtum [Panay-Bisaya], tarong-ayam [Bikol], tarong-tarong [Samar-Bisaya, Tagalog], talong gubat [Tagalog] (Blanco 1837: 136); Sri Lanka: mala-batu [Sinhalese]; Taiwan: da hugno teg io (*Henry 358*); Timor Leste: kaubasu (*Wiriadianata 428*); Thailand: ma ûk muak (Burkill 1935), ma uek (Heiser 1986), makhua-puu, ma-puu, mauek, yang-khui-dee, bakuek (*Widmer 106*); Vietnam: cây ca [Annamite] (*Poilane s.n.*), xo plo xoc [Moi] (*Poilane 184*). More orthographic and local variants of these names occur throughout the region (see Suppl. material 1 and dataset on the NHM Data Portal, https://doi.org/10.5519/0rqfzvgd).

*Solanum lasiocarpum* is widely cultivated across its range for its juicy fruits that are used as a base for juices and for cooking, especially in curries (Heiser 1986). The seeds are burned as used for relief of toothache and in Bangladesh *S. lasiocarpum* is used as a remedy for coughs, asthma, fever, vomiting, and gonorrhoea (Lim 2012).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (6,950,615 km², LC); AOO (1,032 km², VU). As a cultivated plant, the true wild
<figure><img src="image_48.png" /><figcaption>**Figure 40.** Distribution of *S. lasiocarpum*.</figcaption></figure>

distribution of *S. lasiocarpum* is difficult to assess. As a weedy ruderal species, however, we suggest it is not of immediate conservation concern.

**Discussion.** *Solanum lasiocarpum*, together with *S. repandum* G.Forst. of the Pacific islands (see Whalen et al. 1981), is one of two non-American members of the otherwise Neotropical Lasiocarpa clade (Stern et al. 2011) and shares with those species large repand leaves and pubescent berries. Heiser (1987) postulated that *S. lasiocarpum* was an introduction of the American species *S. candidum* Lindl. by Spanish mariners. Whalen et al. (1981) suggested that *S. lasiocarpum* was closely related to and were not “at ease retaining them” as distinct, but did pending further study; they suggested that the distribution of *S. lasiocarpum* was a puzzling historical problem and was likely not due to recent human introduction. The taxa are indeed morphologically very similar with felty, repand leaves and pubescent fruits, but recent molecular analyses have shown that *S. lasiocarpum* is in fact more closely related to the Pacific species *S. repandum* G.Forst. (Bruneau et al. 1995; Bohs 2004; Aubriot et al. 2016a) and that the morphological similarities are more likely due to convergence.

Cultivated forms of *S. lasiocarpum* in general lack or have very few prickles, but this character is highly variable. The fruits of *S. lasiocarpum* are bright orange and have juicy sweet-sour flesh, they are not as large as the fruits of the more commonly cultivated South American species of the Lasiocarpa clade *S. quitoense* Lam. and *S. sessiflorum* Dunal, nor of those of the Pacific *S. repandum* (see Whalen et al. 1981). *Solanum lasiocarpum* is not as strongly andromonoecious as are species such as *S. insanum* or *S. melongena*, and often several of the lowermost flowers are hermaphroditic and develop berries. In other species of the Lasiocarpa clade the expression of andromonoecy is quite labile (Miller and Diggle 2003; Diggle and Miller 2004), we suspect this is the case for *S. lasiocarpum* as well.
In many herbaria specimens of *S. lasiocarpum* and of *S. involucratum* are annotated as *Solanum ferox* L., nom utique rej. This confusion is the result of the typification of another suppressed name, *S. indicum* L. (discussed in detail in Knapp 2011). Miquel (1857) recognised *S. involucratum* and *S. lasiocarpum* as varieties of the same taxon (as *S. ferox*). Heiser (1996, 2001) used the name *S. ferox* for *S. involucratum*, while Whalen et al. (1981) regarded it as ambiguous. Thus, care should be taken with specimens annotated as *S. ferox* in collections, they could be either *S. lasiocarpum* (more common in our experience) or *S. involucratum*. The two species are not closely related; *S. involucratum* is a member of a clade comprised of *S. procumbens*, *S. expedunculatum* Symon and *S. leptacanthum* Merr. & L.M.Perry (the latter two species from New Guinea), itself apparently related to *S. barbisetum* and *S. praetermissum* (Aubriot et al. 2016a). *Solanum lasiocarpum* on the other hand has been shown in numerous studies to be a member of the otherwise American Lasiocarpa clade (e.g., Bruneau et al. 1995; Bohs 2004; Stern et al. 2011).

Whalen et al. (1981) did not treat several of the taxa we recognise as synonyms of *S. lasiocarpum* here, necessitating typification of these names. No herbarium or specimens were cited in Blanco’s (1837) “Flora de Filipinas”, and no herbarium is known to exist (Merrill 1918a, 1923). Merrill (1918a) provided a series of “illustrative specimens” in order to help fix applications of the Blanco’s names, mostly from his series “Merrill: Species Blancoanae”. He felt that just because a work was difficult, it should not be ignored, stating “We can no longer look on the work or this or that author, no matter how incomplete or imperfect, as unworthy of consideration, nor can we accept Hooker’s dictum, regarding species proposed by such authors as Blanco, that it was undesirable to devote time to their identification” (Merrill 1918a: 6). We have selected the duplicate of Merrill’s “illustrative specimen” of *S. zeylanicum*, *Merrill Species Blancoanae* 465 at the Arnold Arboretum (A barcode 00230113) as the neotype for this name.

In describing *S. lasiocarpum* var. *velutinum* Dunal (1852) cited two collections, one from Macau (*Callery s.n.* in P) and the other (*Cuming 690*) from the Philippines. We have selected the best-preserved duplicate of the most widely distributed of these (*Cuming 690*, P00379526) as the lectotype.

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

**24. *Solanum macrocarpon* L., Mant. Pl. Altera 205. 1771.**

Fig. 22G, H

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

*Solanum dimorphum* Matsum., Bot. Mag. (Tokyo) 15: 56. 1901. Type. Cultivated in Tokyo, Japan “cult. Hort. Bot. Tokyo” (possibly from material collected in Taiwan but origin uncertain) (no specimens cited; no original material found).

**Type.** Cultivated in Uppsala, Sweden, *Hort. Uppsala s.n.* (lectotype, designated by Hepper and Jaeger 1985, pg. 391: LINN [acc. # 248.11]).
**Distribution.** *Solanum macrocarpon* is widely distributed across Africa and has been introduced in cultivation in other parts of the world possibly in association with the trans-Atlantic slave trade (e.g., Brazil, the Caribbean, Central America). In tropical Asia *S. macrocarpon* has been collected in Sri Lanka, Singapore, and China, but may be more widely cultivated (see below). *Solanum macrocarpon* is not known outside cultivation.

**Common names.** Gboma eggplant is the English common name in widest use for this species; for common names in Africa see Vorontsova and Knapp (2016).

**Description.** Vorontsova and Knapp (2016: 225–229); http://www.solanaceae-source.org/solanaceae/solanum-macrocarpon.

**Discussion.** *Solanum macrocarpon* is a cultivated plant, used for its leaves and fruits in Africa (Vorontsova and Knapp 2016); its wild progenitor is *S. dasyphyllum* Schumach. & Thonn. of sub-Saharan Africa. It is easily distinguished from other cultivated eggplants in tropical Asia by its large, glabrous or very sparsely pubescent leaves with attenuate bases decurrent onto the stem, its large flowers with copious interpretalar tissue, and its foliaceous calyx lobes.

We have seen only a few specimens of *S. macrocarpon* L, although Burkill (1935) reports its use in the Malay Peninsula as “spreading throughout the Dutch Indies” and used as a pot-herb (leaves eaten). Accessions of *S. macrocarpon* are held in the collections of the World Vegetable Center (WorldVeg or AVDRC) in the Philippines (Taher et al. 2017) but it is not widely cultivated nor has it escaped or naturalised.

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

**25. Solanum mammosum L., Sp. Pl. 1: 187. 1753.**
Fig. 41A, B

No names based on tropical Asian types have been published; for complete synonymy see Nee 1976; Vorontsova and Knapp 2016).

**Type.** **BARBADOS.** “Solanum barbadense spinosum, foliis villosis, fructu aureo rotundiore, Pyri parvi inversiforma et magnitude” (lectotype, designated by Knapp and Jarvis 1990, pg. 343: [Illustration] Plukenet, Phytographia, pars tertia, tab. 226, fig. 1. 1692). Typotype material: Sloane vol. 98: 59 (BM-SL).

**Distribution.** *Solanum mammosum* has been recorded from China, Myanmar, the Philippines, Thailand, and Vietnam, but is almost certainly more widespread in the region; it is native to southern South America and is widely cultivated for ornament.

**Common names.** China. ru qie (Zhang et al. 1994). Malaysia/Singapore. těrong bělanda, těrong asam (Burkill 1935). Vietnam. cà du, blanh (Hul and Dy Phon 2014)

**Description.** Vorontsova and Knapp (2016: 235–239); http://www.solanaceae-source.org/solanaceae/solanum-mammosum-1.

**Discussion.** *Solanum mammosum* is widely cultivated for its unusual teat-shaped fruit that is often used in flower arrangements. It is a member of the Acanthophora clade along with *S. aculeatissimum*, *S. capsicoides*, and *S. viarum*. *Solanum mammosum*
<figure><img src="image_49.png" /></figure>

A

<figure><img src="image_50.png" /></figure>

B

<figure><img src="image_51.png" /></figure>

C

<figure><img src="image_52.png" /></figure>

D

<figure><img src="image_53.png" /></figure>

E

<figure><img src="image_54.png" /></figure>

<figure><img src="image_55.png" /></figure>

G

<figure><img src="image_56.png" /></figure>

H

**Figure 41.** Introduced species of *Solanum*. *Solanum mammosum* L. **A** detail view of a flower (Stevens 35067, Nicaragua) **B** detail view of a fruit (Stevens 35067, Nicaragua). *Solanum robustum* H.L.Wendl. **C** habit (*Sampath Kumar et al. 126920*, India) **D** detail view of a short-styled flower (*Sampath Kumar et al. 126920*, India) **E** detail view of a fruit (*Sampath Kumar et al. 126920*, India). *Solanum sisymbriifolium* Lam. **F** habit (*Sampath Kumar et al. 126922*, India) **G** detail view of a flower (*Sampath Kumar et al. 126922*, India) **H** detail view of a fruit (*Sampath Kumar et al. 126922*, India). Photograph credits: **A, B** O.M. Montiel **C-H** X. Aubriot.
differs from those species in its larger, purple flowers, densely pilose leaves and stems, and large fruits. *Solanum capsicoides* has similarly large berries, but these are darker orange and the seeds are strongly winged; seeds of *S. mammosum* lack wings. Hul and Dy Phon (2014) record it as naturalised in Vietnam, where the fruits are apparently eaten by villagers once cooked, which is a bit surprising, given that they are used as insecticides in other parts of the world (Nee 1979).

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

**26. Solanum melongena L., Sp. Pl. 1: 186. 1753.**

Fig. 42

*Solanum mexianum* Hill, Veg. Syst. 9: 39, pl. 39, fig. 1. 1765. Type. Cultivated "The Rugged Nightshade" (no specimens cited; lectotype, designated here: [illustration] Hill, The Vegetable System 9: Tab 39, f. 1. 1765).

*Melongena ovata* Mill., Gard. Dict., ed. 8, no. 1. 1768. Type. "Melongena (Ovata) caule inermi herbaceo, foliis oblong-ovatis tomentosis integris, fructu ovato [Melongena fructu oblongo violaceo Tourn. Inst. 151]" (no specimens cited; possibly described from live plants; neotype, designated here: BM [BM000942564]).

*Melongena teres* Mill., Gard. Dict., ed. 8, no. 2. 1768, as "*Tereta*". Type. "Melongena (Tereta) caule inermi herbaceo, foliis oblong-ovatis tomentose, fructu terete [Melongena fructu tereti violaceo Tourn. Inst. 151]" (no specimens cited, no material found; probably described from live plants).

*Melongena incurva* Mill., Gard. Dict., ed. 8, no. 3. 1768. Type. "Melongena (Incurva) caule inermi herbaceo, foliis oblongis sinuatis tomentosis, fructu incurvo [Melongena fructu incurvo Tourn. Inst. 152]" (no specimens cited, no material found; probably described from live plants).

*Melongena spinosa* Mill., Gard. Dict., ed. 8, no. 4. 1768. Type. "Melongena (Spinosa) spinosa, foliis sinuatis-lacinitis, fructu tereti, caule herbaceo [Solanum pomiferum fructu spinoso J.B. 3. 619]" (no specimens cited, no material found; lectotype, designated here: [illustration] "Solanum pomiferum fructu spinoso" Bauhin, Historia plantarum 3: 619. 1651).

*Solanum zeylanicum* Scop., Delic. Fl. Faun. Insubr. 1: 1. 1786. Type. Cultivated in Italy [Pavia] from seeds sent by D. Marsilius (no specimens cited, described from live plants; lectotype, designated here: [illustration] Scopoli, Deliciae Flora et Fauna Insubricae 1: tab. 1. 1786).

*Solanum album* Lour., Fl. Cochinch. 129. 1790. Type. Indonesia. Malaku Islands: Amboina [Ambon Island] (no specimens cited; based on illustration; lectotype, designated here [cited as holotype by Hul and Dy Phon 2014, pg. 30]: [illustration] "Trongum agreste spinosum" in Rumphius, Herb. Amboin. 5: 241, tab. 86, fig. 1. 1747).

*Solanum oviferum* Nocca, Ann. Bot. (Usteri) 6: 61. 1793., nom. illeg. superfl. Type. Based on *Solanum melongena* L. (cited in synonymy)
*Solanum oviferum* Salisb., Prodr. Stirp. Chap. Allerton 134. 1796, nom. illeg. superfl. Type. Based on *Solanum melongena* L. (cited in synonymy)

*Solanum esculentum* Dunal, Hist. Nat. Solanum 208, tab. 3. 1813, nom. illeg. superfl. Type. Based on *Solanum melongena* L. (cited in synonymy with the comment [Dunal 1852] "nomen Melongena no accepi, quia sub hoc nomine, species plures confusae suerunt").

*Solanum ovigerum* Dunal, Hist. Nat. Solanum 210. 1813, nom. illeg. superfl. Type. Based on *Solanum melongena* L. (cited in synonymy)

*Solanum pressum* Dunal, Hist. Nat. Solanum 217. 1813. Type. Based on "Trongum prâ rubrum" of Rumphius (no specimens cited; lectotype, designated here: [illustration] Rumphius, Herbarium Amboinense 5: tab. 86, f. 2. 1747).

*Solanum melanoxylon* Link, Enum. Hort. Berol. Alt. 1: 188. 1821. Type. Cultivated in Berlin, Germany (no specimens cited; no original material found, likely destroyed).

*Solanum longum* Roxb. ex Wall., Fl. Ind. (Carey & Wallich ed.) 2: 248. 1824. Type. India. West Bengal: Cultivated "HBC" [Hort. Bot. Calcutta], 1821, *Without collector s.n.* [Wallich cat. 2628D] (lectotype, designated here: K-W [K001116666]).

*Solanum ovigerum* Dunal var. *insanum* Blume, Bijdr. Fl. Ned. Ind. 13: 698. 1826. Type. Indonesia. "in hortis frequenter cultum/Terong Pangang" (no specimens cited; no original material found).

*Solanum pseudo-undatum* Blume, Bijdr. Fl. Ned. Ind. 13: 699. 1826. Type. Indonesia. Java. "in locis cultis/nomen Terong Rangu" [ex protologue] Sin. loc., *C.L. Blume s.n.* (lectotype, designated here: L [L0003642]).

*Solanum pseudo-undatum* Blume var. *albiflorum* Blume, Bijdr. Fl. Ned. Ind. 13: 699. 1826. Type. Indonesia. "in hortis colitur/Terong lelles" [ex protologue] (no specimens cited; no original material found).

*Solanum pseudo-undatum* Blume var. *atropurpurascens* Blume, Bijdr. Fl. Ned. Ind. 13: 699. 1826. Type. Indonesia. "cum praecidentibus/Terong Kupa" [ex protologue] (no specimens cited; no original material found).

*Solanum pseudo-undatum* Blume var. *leucocarpon* Blume, Bijdr. Fl. Ned. Ind. 13: 700. 1826. Type. Indonesia. "montis Salak" [ex protologue] (no specimens cited; no original material found).

*Solanum edule* Schumach. & Thonn., Beskr. Guin. Pl. 125 [145]. 1827. Type. "Guinea". Sin. loc., cultivated, *P. Thonning 141* (lectotype, designated here: C [C10004584]).

*Solanum melongena* L. var. *esculentum* (Dunal) Walp., Repert. Bot. Syst. (Walpers) 3: 81. 1844. Type. Based on *Solanum esculentum* Dunal.

*Solanum aethiopicum* L. var. *violaceum* Dunal, Prodr. [A. P. de Candolle] 13(1): 351. 1852. Type. Based on *Solanum aethiopicum* sensu Lour., non L.

*Solanum album* Lour. var. *richardii* Dunal, Prodr. [A. P. de Candolle] 13(1): 362. 1852. Type. Réunion. "Bisom. croix dans les lieux habités", 1837, *A. Richard 186* (holotype: P [00352546]).

*Solanum album* Lour. var. *rumphii* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852, nom. illeg. Type. Based on *Solanum album* Lour. (as "Trongum agreste").
*Solanum edule* Thonn. var. *multifidum* Dunal, Prodr. [A. P. de Candolle] 13(1): 357. 1852. Type. Cultivated "in hort. Tonelle Audiberti cultum" [Audibert nursery in Tonelle, near Tarascon, France], sin. dat., *Without collector s.n.* (holotype: AV [n.v.]).

*Solanum serpentinum* Desf. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852. Type. Cultivated in Montpellier, France, 17–26 Aug 1838, R.L. *Desfontaines s.n.* (holotype: MPU [MPU854428]).

*Solanum esculentum* Dunal var. *aculeatum* Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852. Type. Indonesia. Java: Sin. loc., sin. dat., *H. Zollinger 702* (holotype: G-DC [G00131553]; isotypes: BM [BM000778111], K [K000788269]).

*Solanum esculentum* Dunal var. *inerme* Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852. Type. Brazil. Bahia: sin. loc., sin. dat., *J.S. Blanchet 368* (lectotype, designated here: G [G00357998]).

*Solanum esculentum* Dunal var. *subinerme* Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852. Type. Brazil. Bahia: sin. loc., 1831, *J.S. Blanchet 236* (lectotype, designated here: G-DC [G00131527]).

*Solanum lagenarium* Dunal, Prodr. [A. P. de Candolle] 13(1): 368. 1852. Type. Cultivated "hort. Tonelle cultum" [Audibert nursery in Tonelle, near Tarascon, France], 1824, *Anon. s.n.* (holotype: AV [n.v.]).

*Solanum ovigerum* Dunal [unranked] *sinuatorepandum* Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852. Type. Cuba. La Habana: "La Havane" [Havana], 1826, *de la Ossa s.n.* (lectotype, designated here: G-DC [G00131521]).

*Solanum ovigerum* Dunal [unranked] *subrepandum* Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852. Type. Cultivated in Geneva, Switzerland "in tepidario DC.", sin. dat., *Anon. s.n.* (lectotype, designated here: G-DC [G00131523]).

*Solanum ovigerum* Dunal var. *violaceum* Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852, as *Solanum ovigerum* [unranked] *subrepandum* var. *violaceum*. Type. No specimens or locality cited; reference to Tournefort ("Tourn. inst. 152").

*Solanum ovigerum* Dunal var. *oblongo-cylindricum* Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852, as *Solanum ovigerum* [unranked] *subrepandum* var. *oblongo-cylindricum*. Type. Based on *Solanum ovigerum* var. *insanum* Blume.

*Solanum trongum* Poir. var. *sinuatopinnatifidum* Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Based on *Solanum trongum* Poir. sensu Blume [Java. "in agestribus/karnadong", *C. Blume s.n.* (no specimens cited, based on *S. trongum* Poir. sensu Blume)].

*Solanum requienii* Dunal, Prodr. [A. P. de Candolle] 13(1): 363. 1852., as 'Requieni'. Type. Cultivated in France "hort. Tonelle ex Audibert fratr.", 1822, *Anon. s.n.* (holotype: AV [n.v.]).

*Solanum sativum* Dunal, Prodr. [A. P. de Candolle] 13(1): 360. 1852, nom. illeg. superfl. Type. Based on *Solanum pseudo-undatum* Blume (cited in synonymy "No-men pseudo-undatum mutavi, quonian semigraecum, semilatinem")

*Solanum sativum* Dunal var. *albiflorum* (Blume) Dunal, Prodr. [A. P. de Candolle] 13(1): 360. 1852. Type. Based on *Solanum pseudo-undatum* Blume var. *albiflorum* Blume.
*Solanum sativum* Dunal var. *atropurpurascens* (Blume) Dunal, Prodr. [A. P. de Candolle] 13(1): 360. 1852. Type. Based on *Solanum pseudo-undatum* Blume var. *atropurpurascens* Blume.

*Solanum sativum* Dunal var. *leucocarpon* (Blume) Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Based on *Solanum pseudo-undatum* Blume var. *leucocarpon* Blume.

*Solanum melongena* L. var. *giganteum* Alef., Landw. Fl. 136. 1866, as "gigantea". Type. Cultivated (no specimens cited; no original material found).

*Solanum melongena* L. var. *leucoum* Alef., Landw. Fl. 136. 1866. Type. Cultivated (no specimens cited; no original material found).

*Solanum melongena* L. var. *stenoides* Alef., Landw. Fl. 136. 1866. Type. Cultivated (no specimens cited; no original material found).

*Solanum melongena* L. var. *stenoleucum* Alef., Landw. Fl. 136. 1866, as "stenoleuca". Type. Cultivated (no specimens cited; no original material found).

*Solanum melongenum* St.-Lag., Ann. Soc. Bot. Lyon 7: 135. 1880, nom. illeg. superfl. Type. Based on *Solanum melongena* L. (cited in synonymy).

*Melongena esculenta* (Dunal) Grecescu, Consp. Fl. Romaniei 423. 1898. Type. Based on *Solanum esculentum* Dunal, nom. illeg.

*Solanum melongena* L. var. *inerme* (Dunal) Hiern, Cat. Afr. Pl. (Hiern) 1(3): 748. 1898. Type. Based on *Solanum esculentum* var. *inerme* Dunal

*Solanum melongena* L. var. *viride* Dikii, Trudy Prikl. Bot. 88: 104. 1984, as "viridis". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from Asia Minor. 4 Jul 1971, L. Studentsova s.n. [VIR cat. # 249] (holotype: WIR [n.v.]).

*Solanum melongena* L. subsp. *agreste* Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "agrestis". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India, 7 Aug 1975, *N. Frantskevich s.n.* [VIR cat. # 713] (holotype: WIR [n.v.]).

*Solanum melongena* L. var. *angustum* Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "angusta". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India "c.v. R.R.I.", 7 Aug 1975, *N. Frantskevich s.n.* [VIR cat. # 2333] (holotype: WIR [n.v.]).

*Solanum melongena* L. var. *cylindricum* (Filov) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "cylindrica". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India, 7 Aug 1975, *N. Frantskevich s.n.* [VIR cat. # 116] (holotype: WIR [n.v.]).

*Solanum melongena* L. var. *racemiflorum* Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "racemiflora". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India "c.v. Purple Cluster", 27 Sep 1958, *V. Barkovskaya s.n.* [VIR cat. # 2515] (holotype: WIR [n.v.]).

*Solanum melongena* L. var. *racemosum* (Filov) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as "racemosa". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India, 4 Jul 1971, *L. Studentsova s.n.* [VIR cat. # 709] (holotype: WIR [n.v.]).
*Solanum melongena* L. var. *globosi* Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 106. 1984.
Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from Vietnam, 14 Jul 1971, *L. Studentsova s.n.* [VIR cat. # 2582] (holotype: WIR [n.v.]).

**Type.** Cultivated in Uppsala, Sweden, *Anonymousous s.n.* (lectotype, designated by Schönbeck-Temesy 1972, pg. 70: LINN [acc. # 248.28]).

**Description.** Erect annual herb, 0.2–0.5 m, unarmed or more rarely prickly. Stems erect, terete, unarmed or occasionally prickly, moderately stellate-pubescent to glabrescent; prickles to 3 mm long, less than 0.5 mm wide at the base, straight, acicular, yellow-orange to dark brown, glabrous; pubescence a mixture of minute simple hairs and porrect-stellate trichomes, the stellate trichomes sessile to short-stalked, the stalks to 0.2 mm long, the rays 8–15, 0.3–0.7 mm long, the midpoints ca. same length as the rays or elongated to 1 mm; new growth moderately to densely stellate-pubescent, whitish green; bark of older stems glabrescent, green-brown to dark brown. Sympodial units difoliate, not geminate. Leaves simple, moderately lobed, the blades 10–23 cm long, 9–15 cm wide, 1.5–2 times longer than wide, ovate, chartaceous, drying discolorous, unarmed; adaxial surface yellow-green to red-brown, sparsely to moderately stellate-pubescent, occasionally glabrescent; abaxial surface glabrescent to moderately stellate-pubescent usually more pubescent than the adaxial surface, the trichomes porrect, sessile to short-stalked, the stalks to 0.2 mm long, the rays 5–8, 0.3–1 mm long, the midpoints ca. same length as the rays; major veins 4–7 pairs; base cordate to obtuse; margins lobed, the lobes 2(-3) on each side, 0.5–2 cm long, deltate, apically rounded, the sinuses extending 1/4–1/3 to the midrib; apex acute; petiole 1–10 cm long, 1/4–1/3(-2/3) of the leaf blade length, moderately stellate-pubescent to glabrescent, unarmed or prickly with 1–2 prickles. Inflorescences 6–15 cm long, apparently terminal or lateral, unbranched, with 1–8 flowers, 1–3 flowers open at any one time, moderately stellate-pubescent to glabrescent, with a mix of sessile and short-stalked stellate-porrect trichomes like those of the stems, unarmed; peduncle 0–80 mm long, unarmed; pedicels 0.8–3.5 cm long, 1–1.8 mm in diameter at the base, 1–2 mm in diameter at the apex, erect to pendent, unarmed or more rarely prickly with up to 5 prickles, moderately stellate-pubescent to glabrescent with porrect-stellate trichomes like those of the stems, articulated at the base; pedicel scars spaced 3–5 mm apart. Buds ovoid, the corolla ca. halfway exerted from the calyx before anthesis. Flowers 4–8-merous, heterostylous and the plants andromonoecious, with the lowermost flower long-styled and hermaphrodite, the distal flowers short-styled, staminate and usually somewhat smaller. Calyx with the tube 3–8 mm long in long-styled flowers, urceolate to deeply cup-shaped, 2–3 mm long in short-styled flowers, cup-shaped to elongate cup-shaped, the lobes in long-styled flowers 5–17 mm long 3–5 mm wide, in short-styled flowers 3.5–5 mm long, 2–2.5 mm wide, deltate to narrowly deltate, apically acute to long-acuminate, unarmed or more rarely prickly with up to 30 prickles, moderately stellate-pubescent, with porrect-stellate trichomes like those of the pedicels. Corolla 2.5–5 cm in diameter in long-styled flowers, 2.4–4 cm in diameter in short-styled flowers, white
<figure><img src="image_57.png" /><figcaption>**Figure 42.** *Solanum melongena* L. **A** herbarium specimen collected in India (*Macé s.n.*, P00058105) **B** habit (field photograph, unvouchered, Thailand) **C** detail view of a hermaphroditic flower (*Meeboonya et al. RM 294*, Thailand) **D** detail view of an oblong and white fruit (*Meeboonya et al. RM 250*, Thailand) **E** detail view of a round green fruit marbled with white (*Meeboonya et al. RM 248*, Thailand). Photograph credits: **A** CC-BY, Muséum national d'Histoire naturelle, Paris **B–E** X. Aubriot.</figcaption></figure>

to mauve or purple, stellate, lobed 1/4–1/2 of the way to the base, the lobes 10–15 mm long, 8–13 mm wide in long-styled flowers, 5–8 mm long and 8–13 mm wide in short-styled flowers, broad-deltate, spreading at anthesis, not opening fully in long-styled
flowers, sparsely stellate-pubescent abaxially, the trichomes porrect, sessile or stalked, the stalks to 0.2 mm, the rays 4–8, 0.2–0.7 mm long, the midpoints ca. same length as the rays. Stamens equal; anthers 5.5–7.5 mm long in long-styled flowers, 5.5–6 mm long in short-styled flowers, ca. 2 mm wide, connivent, tapering, yellow, glabrous, poricidal at the tips, the pores not elongating to slits with drying; filament tube 2–3 mm long, glabrous; free portion of the filaments 1.2–3 mm long, glabrous. Ovary conical, stellate-pubescent in upper part near the style base; style in long-styled flowers ca. 9 mm long, broad and straight, moderately stellate-pubescent in the lower half to 3/4, in short-styled flowers 2–3 mm long, pubescent at the base; stigma globose capitate to somewhat bilobed, bright green in live plants, the surface minutely papillate. Fruit a globose to ovoid, ellipsoid, or oblong to variously curved berry (many cultivars with a wide variety of fruit shapes), 1 (rarely > 1) per infructescence, 3–20 cm long, 3–7 cm wide, green, sometimes mottled or striped, white, pink, mauve, purple, or black when young, usually white or maroon at maturity, the pericarp smooth, glabrous; fruiting pedicels 2.5–8 cm long, 2–4 mm in diameter at the base, 5–10 mm in diameter at the apex, unarmed or sparsely prickly, woody, pendent and deflexed from the weight of the berry; fruiting calyx lobes elongating to 12–50 mm long, 1/4–1/3 the length of the mature fruit, often cup-shaped around the fruit in some cultivars, unarmed or sparsely prickly. Seeds usually aborted in cultivars. Chromosome number: n = 12 (Doganlar et al. 2002; Barchi et al. 2019; Chapman 2019; Gramazio et al. 2019; Wei et al. 2020).

**Distribution.** *Solanum melongena* is cultivated worldwide in tropical and subtropical areas (in the temperate zone under glass); the greatest diversity of landraces and cultivars is found in Asia (India, China and southeast Asia), with secondary centres in the Middle East and around the Mediterranean. The origin of *Solanum melongena* is in Asia, but the exact place of domestication in not clear (see Meyer et al. 2012; references in Knapp et al. 2013). We have not provided a distribution map as herbarium specimens do not accurately reflect the range of the species in cultivation.

**Ecology and habitat.** *Solanum melongena* is only known from cultivation (see below).

**Common names and uses.** China. qie (Zhang et al. 1994), qie zi (Lester and Hawkes 2001). India. brinjal [Hindi] (Lester and Hawkes 2001); Bihar: baigan, bhanta (Varma 1981); Karnataka: advibadani, doorla (Singh 1988); Tamil Nadu: kathiri (Tamil, Matthew 1983); kaththiri ([Tamil] Henry et al. 1987). Japan. nasu (Lester and Hawkes 2001). Indonesia. Java: enchung, térong (Burkill 1935); Sumatra: tiung, chung (Burkill 1935). Malaysia/Singapore. térong (with many qualifiers indicating varieties, Burkill 1935). *Solanum melongena* is a worldwide food crop; after tomato (S. lycopersicum L.) it is the most widely cultivated fruit crop in the Solanaceae (Daunay and Janick 2007; Knapp et al. 2019b). Recent work to improve eggplants in the face of climate change has seen increased interest in wild relatives and crosses with wild species across the spiny solanums (e.g., García-Fortea et al. 2019). See section on Uses (pp. 18–20) for further details. Discussion of the many aspects of eggplant biology can be found in Chapman (2019).

**Preliminary conservation status (IUCN 2019).** *Solanum melongena* is known primarily from cultivation, and as such we have not assigned it a threat status using the IUCN criteria.
**Discussion.** *Solanum melongena* is known to cross easily with its wild progenitor *S. insanum* (Davidar et al. 2015; Mutegi et al. 2015) and distinguishing the two taxa can be difficult especially in older specimens or in feral populations. We have chosen to recognise the two as distinct species rather than as infraspecific taxa (e.g., Lester and Hasan 1991; Daunay and Hazra 2012) because cultivated and wild plants are on distinct evolutionary trajectories. Wild plants are subject to natural selection, while cultivated plants are commensal with human beings. We recognise that this can lead to difficulties in identifying individual specimens, particularly those from old collections where details of provenance are not clear.

Distinguishing the cultivated *S. melongena* from wild *S. insanum* can be particu-
larly difficult in the absence of fruits. Keys in previous publications (Knapp et al. 2013;
Ranil et al. 2016) can be used for identification, but in general specimens we assign to
*S. insanum* are pricklier, with smaller flowers and fruit, and more ruderal than those we
assign to *S. melongena*. Flowers of *S. melongena* are often fasciated (with more than 5
parts), but usually are not in local cultivars. Local cultivars of *S. melongena* in tropical
Asia can be especially difficult to distinguish from *S. insanum*.

The only original material we have found for Sir John Hill's *Solanum mexianum*
is his illustration (Hill 1765) that clearly represents *S. melongena*; we lectotypify the
name with this illustration.

Miller (1768) recognised the pre-Linnaean genus *Melongena*, in which he segre-
gated those species treated as *Solanum* by Linnaeus (1753) as *S. melongena*. His species
were distinguished based mainly on fruit size and shape; he considered colour variants
to all belong to his *Melongena ovata*. He did not cite any specimens in his treatment,
he cited instead pre-Linnaean polynomials mostly from Tournefort (1700), none of
which are illustrated in that work. Original material is thus apparently lacking for
these names. From Miller's (1768) discussion of *Melongena* he had obviously culti-
vated all of the taxa he recognised at the Chelsea Physic Garden. A single specimen of
*S. melongena* from the Miller herbarium has been found in BM, labelled on the verso
side with "Hort. Chelsea, Miller" and "Melongena ovata" in Solander's handwriting,
indicating it was so filed in Miller's herbarium (J. Wajer, person. comm.). We designate
this here as the neotype for *Melongena ovata*, as it is not clear it is original material; all
of these species of *Melongena* are certainly described from living material, as were most
of the plants in the "Gardener's dictionary" (Miller 1768). We have not designated
neotypes for *Melongena teres* and *M. incurva*; they should perhaps be sought in the
Tournefort herbarium in Paris. *Melongena spinosa* was described (Miller 1768) with
reference to Bauhin's (1651) "Historia plantarum" ("Solanum pomiferum fructu spi-
noso J.B. 3. 619"). In that work there is an illustration labelled "Solanum pomiferum
fructu spinoso" that is clearly *S. melongena*, with two fruits, one an ovoid shape with
a prickly calyx and the other a different elongate shape with a non-prickly calyx. We
designate this illustration as the lectotype for *M. spinosa*, it is the only original material
we have found.

In describing *S. zeylanicum* Scopoli (1786) provided an extremely detailed descrip-
tion of the plant and an excellent illustration both clearly referrable to *S. melongena*,
although to a prickly relatively small-fruited variety. Scopoli (1786) stated that the fruit pulp was white (“albida”) and the illustration shows large, pendent fruits.

In describing *S. album*, Loureiro (1790) referred only to “Trongum agreste. Album. Rumph. Amb. l. 8. c. 48. P. 241” and refers to its use “Virtus radices Ondonotologica. Bacca edulis”. In Rumphius’s “Herbarium Amboinense” (Rumphius 1747) “Trongum agreste album/Trongum pra” comprises two white-flowered taxa, one with green fruits marbled with white and the other with red fruits, but the use in dentistry cited by Loureiro (1790) is part of the description of the previous taxon, “Trongum agreste/Trongum udan” (Rumphius 1747: 240). Hul and Dy Phon (2014) cited as “holotype” of *S. album* the illustration of “Trongum agreste spinosum” (Rumphius 1747: tab. 86, f. 1), a taxon described as part of “Trongum agreste”, not “Trongum agreste album”. No botanical materials have survived from Loureiro’s stay in Indochina (Merrill 1935), so a holotype is out of the question; we have designated as the neotype the illustration cited by Loureiro (1790) that was clearly available to him at the time he described *S. album* (see Turland et al. 2018, Art. 8.1). In his interpretation of the “Herbarium Amboinense”, Merrill (1917) cited a specimen that “almost certainly represents “*Solanum agreste* album” Rumph.” (*Robinson* 286, US acc. # 654604). This is a specimen of *S. lasiocarpum* Dunal and does not correspond with the illustrations in Rumphius (1747).

Dunal (1813) based his *S. pressum* entirely on Rumphius’ (1747) “Trongum prà rubrum”, citing no other material and taking his description entirely from that of Rumphius. He took his epithet from the suggestion that “Trongum agreste album” was translatable to Latin as “Trongum pressum” (Rumphius 1747). We have selected the one element cited by Dunal (1813) as the lectotype of *S. pressum*; the illustration shows an unarmed plant with solitary flowers and berries. Rumphius’ (1747) use of the word “rubrum” may indicate the berries were red, thus perhaps making this name a synonym of *S. aethiopicum* L., but this is not mentioned in the description.

In the description of *S. longum* Roxburgh (1824) states that he distinguishes this from *S. melongena* by its elongate fruit form, in all other respects said to be exactly like the brinjal eggplant. Roxburgh (1814) first used the name *Solanum longum* in his “Hor tus Bengalensis”, which is a list of the plants growing in the Calcutta Botanical Garden; we have not found any unambiguous original material, nor any specimens with fruits as those in the description, the description may have come from live plants. We have selected as a neotype for *S. longum* specimen in the Wallich herbarium at Kew of a plant cultivated in the Calcutta Botanical Garden in 1821 (Wallich cat. 2628D) that has an immature fruit that looks like it might be of the elongate form (K001116666).

Unravelling the identities of the various taxa referrable to *S. melongena* described by Blume (1826) is complicated by the lack of specimens of these cultivated plants. We here typify only those names for which we have found unambiguous original material; from Blume’s descriptions the identity of the plants in question is clear. In describing *S. ovigerum* var. *insanum*, he is apparently coining the varietal epithet as new, not citing the Linnaean name *S. insanum*. The citation of this as occurring in cultivation and hav- ing oblong, cylindrical fruit makes it a clear synonym of *S. melongena*, but no original
material has been found. *Solanum pseudo-undatum* was also described from cultivated plants; here we designate a sheet in L collected by Blume and labelled as “Solanum pseudoundatum” as the lectotype (L 0003642). The three varieties of *S. pseudo-undatum*, var. *albiflorum*, var. *atropurpurascens* and var. *leucocarpon*, are distinguished purely on flower or fruit colour and common name (see above); we have found no original material for any of these.

Schumacher and Thonning (Schumacher 1827) described *S. edule* from plants cultivated in west Africa. Of the two collections labelled as *S. edule* in Thonning’s herbarium at Copenhagen, the sheet collected by Thonning himself (C10004584) is here selected as the lectotype, the other sheet, collected by P.E. Isert (C10004585) is less well preserved. Vorontsova and Knapp (2016) incorrectly cited the Thonning collection as “holotype”.

The protologue of *S. esculentum* var. *inerme* (Dunal 1852) cites two Brazilian collections, one of which (*Salzmann 392* in G-DC, G00131550) was questioned as belonging to *S. melongena* by Dunal himself. We have therefore selected the other collection, *Blanchet 368* (G00357998) that bears a label in Dunal’s hand as the lectotype. Var. *subinerme* similarly was based on two collections; we have selected *Blanchet 236* (G00131527) as the lectotype because it has a label in Dunal’s hand saying “esculentum β 1842” and the other syntype, *Sieber 308* from Trinidad (represented by two sheets G00357997, G00131564) is not annotated by Dunal.

The infraspecific names Dunal (1852) coined under his *S. ovigerum* are presented in a manner different to most of the other infraspecific names in this work. The main infraspecific divisions, *subrepandum* and *sinuatorepandum*, are indicated with capital letters and are thus unranked; greek letters in this work indicate varietal rank. The varieties listed under [unranked] *subrepandum* are a mixture of polynomials and references to pre-Linnaean works, we do not typify these here. Dunal (1852) cited several collections in the protologue of [unranked] *sinuatorepandum*, *Wallich cat. 2628d, de la Ossa s.n.* and *Gaudichaud 475*. We have selected *de la Ossa s.n.* (G00131521) in G-DC as the lectotype, the Wallich specimen in G-DC is of poor quality and the Gaudichaud collection in Paris is a plant of *S. aethiopicum*. The protologue of [unranked] *subrepandum* mentions only “v.s. in h. DC”, making the identity of original material difficult. We have selected a specimen in G-DC that has a label in Dunal’s hand stating “A. folia subrepanda” as the lectotype for this name (G00131523).

The varietal names coined for the various fruit-shape and colour forms of *S. melongena* in Friedrich Alefeld’s “Landwirthschaftliche flora” (Alefeld 1866), a compendium of names for all plants cultivated in Germany, are most likely to have been based on living plants, the descriptions are minimal and no localities are cited. We have not typified these; they are more properly considered as cultivar names. In his work on the cultivated flora of the Soviet Union Filov (1958) coined many names, but many of these had neither description nor diagnosis and so are not validly published. The Russian taxonomist S.P. Dikii (1984) apparently thought Alefeld’s names were not validly published and re-published them all at the same rank (see Names not validly published).

**Specimens examined.** See Suppl. materials 1–3.
27. *Solanum miyakojimense* T.Yamaz. & Takushi, J. Jap. Bot. 66: 46. 1991.
Figs 2B, 43

**Type.** JAPAN. Ryuku Islands: Cultivated in Nago, Okinawa Island [originally collected in Miyako Island [Miyakojima], Agarihetona], 20 Dec 1989, *A. Takushi* [6400] s.n. (holotype: TI [TI00085080]; isotypes: TI [TI00085081, TI00085082]).

**Description.** Woody herb or subshrub, the branches to 1 m long, spreading and prostrate, armed or unarmed. Stems terete, unarmed or sparsely prickly and moderately to densely eglandular pubescent, the prickles, if present, 0.1–0.35 cm long, straight or very slightly curved, somewhat broad-based, the pubescence of sessile or short-stalked porrect-stellate trichomes, the stalks if present ca. 0.1 mm, the rays 5–8, 0.2–0.4 mm long, the midpoints absent or ca. 0.1 mm long, occasionally to 0.5 mm long; new growth densely (moderately in cultivation) pubescent with sessile or short-stalked porrect-stellate trichomes like those of the stems; bark of older stems pale tan-brown. Sympodial units difoliate, the leaves geminate or not geminate. Leaves simple or very shallowly lobed, the blades 1.2–4 cm long, 0.9–3 cm wide, 1.1–1.3 times longer than wide, broadly ovate to nearly orbicular, widest in the lower half, somewhat thick and fleshy, concolorous or slightly discolorous, unarmed or sparsely armed with a few straight prickles to 0.2 cm long; adaxial surfaces densely pubescent with the lamina still somewhat visible, the pubescence of sessile and short-stalked porrect-stellate trichomes, the stalks 0.1–0.3(0.5) cm long, the rays 5–8, ca. 0.5 mm long, the midpoints absent or 0.3–0.5 mm long, the prickles few or absent; abaxial surfaces densely pubescent with the lamina completely obscured, the pubescence of sessile and short-stalked porrect-stellate trichomes, the stalks usually longer than those of the adaxial surface trichomes, 0.2–0.5 mm long, the rays 6–10, 0.5–0.6 mm long, the midpoints ca. 0.5 mm long; principal veins 2–3 pairs, only visible on the adaxial surfaces; base acute; margins entire or very shallowly lobed, the lobes rounded at the tips, the sinuses reaching less than 1/3 of the way to the midrib, rounded; apex obtuse and rounded; petioles 0.3–0.9 cm long, densely stellate-pubescent like the stems and leaves. Inflorescences internodal, 0.1–0.3(–1.5) cm long, occasionally longer but always with a flower emerging from the very base, unbranched and usually looking like a tiny spur on the stem, the 1–2(4) flowers often seeming to emerge directly from stem, 1–2 flowers open at any one time,; densely stellate-pubescent with sessile and short-stalked porrect-stellate trichomes, the stalks if present ca. 0.1 mm, the rays 5–8, 0.2–0.4 mm long, the midpoints absent or ca. 0.1 mm long, occasionally to 0.5 mm long; peduncle absent to 1.5 mm long; pedicels ca. 0.9 cm long, ca. 0.8 mm in diameter at the base, ca. 1 mm in diameter at the apex, abruptly narrowing to calyx base, densely to moderately pubescent with sessile and short-stalked porrect-stellate trichomes like the stems, articulated at the base; pedicel scars tightly packed at tip of minute inflorescence, with the lowermost one near the stem. Buds ellipsoid, the corolla ca. halfway included in the calyx tube until just before anthesis. Flowers 5-merous, apparently all perfect. Calyx tube 2.5–3.5 mm long, cup-shaped, the lobes mere enations on the rim to 1 mm long, broadly deltate, tearing during
<figure><img src="image_58.png" /><figcaption>**Figure 43.** *Solanum miyakojimense* T.Yamaz. & Takushi A herbarium specimen (holotype) collected in Japan in 1989 (*Takushi* [6400] s.n., TI00085080) B habit (Hsu *16234*, Taiwan) C detail view of the prickles and the pubescence of a stem (Hsu *16234*, Taiwan) D detail view of a flower (Hsu *16234*, Taiwan) E detail view of a fruit with seeds inside (Hsu *16234*, Taiwan). Photograph credits: A TI B–E M.-I. Weng.</figcaption></figure>

development, densely to moderately pubescent with sessile and short-stalked porrect-stellate trichomes, the stalks if present ca. 0.1 mm, the rays 5–8, 0.2–0.4 mm long, the midpoints absent or ca. 0.1 mm long, occasionally to 0.5 mm long, the sinuses scarious, the tips rounded. Corolla 1–1.2 cm in diameter, white, stellate, lobed ca.
3/4 of the way to the base, the lobes 4.5–5 mm long, ca. 1.5 mm wide, spreading at anthesis, glabrous adaxially, densely pubescent where exposed in bud with tangled sessile and short-stalked porrect-stellate trichomes, the stalks ca. 0.1 mm long, the rays 6–8, ca. 0.2 mm long, the midpoints absent. Stamens equal; filament tube minute; free portion of the filaments 0.25–0.3 mm long, glabrous; anthers 4–4.5 mm long, ca. 1.5 mm wide, strongly tapering, yellow, poricidal at the tips, the pores directed distally. Ovary conical, glabrous; style 5.5–6 mm long, slightly curved, white, sparsely pubescent in the lower half with weak sessile porrect-stellate trichomes; stigma minutely bilobed, the surfaces minutely pubescent. Fruit a globose to somewhat elongate berry, 1–1.5 cm in diameter, orange (with “green longitudinal bands” fide *Takushi s.n.* ) when mature, the pericarp glabrous, shiny, thin and somewhat translucent; fruiting pedicels 1.2–1.5 cm long, ca. 1.5 mm in diameter at the base, ca. 2.5 mm in diameter at the apex, thickened and woody, curved downwards, glabrescent to sparsely stellate-pubescent; fruiting calyx slightly accrescent, woody, the tube 3–3.5 mm long, the lobes 2–2.5 mm long, splitting irregularly with incrassate margins, appressed to the base of the fruit. Seeds 20–60 per berry, 2–2.5 mm long, ca. 2 mm wide, flattened reniform, straw-yellow, the surfaces minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.

**Distribution (Fig. 44).** Solanum miyakojimense occurs from the southernmost Ryuku Islands of Miyako and Irabu (Okinawa prefecture) to the island of Ponso no Tao (Lan Yu or Orchid Island) off the southeastern coast of Taiwan to the islands of the Batanes group in the Philippines.

**Ecology and habitat.** Solanum miyakojimense occurs in rocky coastal areas, often on coral bluffs (Hsu et al. 2007) near sea level; often occurring in sea spray.

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

<figure><img src="image_59.png" /><figcaption>**Figure 44.** Distribution of S. miyakojimense.</figcaption></figure>
**Preliminary conservation status (IUCN 2019).** Endangered (EN [B1a,biii,iv]). EOO (641 km², VU); AOO (40 km², EN). *Solanum miyakojimense* is known from only a few wild collections scattered along the chain of islands across a large geographical distance, and although this makes the extent of occurrence larger than an endangered status might warrant, we consider the threats to habitats and the widely distant populations warrant this preliminary assessment.

**Discussion.** In describing *S. miyakojimense* Yamazaki and Takushi (1991) suggested it was endemic to the Ryuku Islands; we have since found it to be more widely distributed, but still only along the island arc that runs south from Miyakojima Island to the Batanes Islands in the Philippines. *Solanum miyakojimense* is superficially similar to both *S. procumbens* and *S. violaceum*. It differs from *S. procumbens*, which occurs from China east to Thailand and the islands of Timor and Flores (Indonesia), in its seaside habitat; it can also be distinguished in its straight or only slightly curved stem prickles (versus strongly hooked), tiny inflorescences with only a few flowers with no or a very tiny peduncle (versus a longer thin peduncle), smaller flowers with broadly triangular corolla lobes (versus flowers with long-triangular corolla lobes), shorter pedicels in fruit and seeds with sinuate (versus pentagonal) cell walls. Berries of *S. procumbens* are bright red and globose at maturity, while those of *S. miyakojimense* are orange or orange-red and usually somewhat ellipsoid (Hsu et al. 2007).

The inflorescences of *S. miyakojimense* are shorter and fewer-flowered than those of the widespread *S. violaceum*, and the fruiting pedicels are conspicuously curved (not markedly spreading and straight as in *S. violaceum*) at fruit maturity. In *S. miyakojimense* the flowers are usually borne at the very base of the rhachis where it joins with the stem such that there is no clear peduncle; *S. violaceum* has a clearly pedunculate inflorescence. The leaves of *S. violaceum* are more markedly discolorous than those of *S. miyakojimense*. *Solanum violaceum* is a relatively robust shrub that can reach 2 m tall, while *S. miyakojimense* is a prostrate plant. *Solanum violaceum* is found in the northwestern part of the main island of Taiwan but has not been collected on Lan Yu (Orchid) island off the southeast coast. The only collections of *S. violaceum* from the Philippines are from cultivated plants.

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

**28. Solanum multiflorum Roth, Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 669. 1819.**

Figs 3B, 45

*Solanum agreste* Roth in Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 669. 1819. Type. India. “H. in India orientali. B. Heyne” 29 Jun 1816, B. Heyne s.n. (neotype, designated by Turner 2021, pg. 417: K-W [K00116650, shoot mounted in lower left only]).

*Solanum multiflorum* Roth, Nov. Sp. Pl. 129. 1821, nom. illeg. non *Solanum multiflorum* Roth in Roem. & Schultes, 1819. Type. Based on same material as *S. multiflorum* Roth ex Roem. & Schult.
*Solanum agreste* Roth, Nov. Pl. Sp. 130. 1821, nom. illeg. non *S. agreste* Roth in Roem. & Schult., 1819. Type. Based on same material and homotypic with *S. agreste* Roth in Roem. & Schult. (isonym).

*Solanum himalense* Dunal var. *soumbe* Dunal, Prodr. [A. P. de Candolle] 13(1): 300. 1852. Type. India. Tamil Nadu: “Nilgiris, Mts. de Nellygerry” [Nilgiri mountains], *J.B.L. Leschenault de la Tour 163* (lectotype, designated here: P [P00055586]; isolectotypes: P [P00055587, P00055588]).

*Solanum erosum* Van Heurck & Müll.Arg., Observ. Bot. (Van Heurck) 83. 1870. Type. India. Tamil Nadu: Nilgiris, “in montibus Nilagiri”, *R.F. Hohenacker 1074* (lectotype, designated here: BR [AWH10071571]; isolectotypes: BM [BM000778310, BM000778311], E [E00196408], G [G00442714], K [K000014858], L [L, L], LE [LE00016194], P [P00055723, P00055724, P00055725, P00055726], W [acc. # 0003339]).

*Solanum indicum* L. var. *multiflorum* (Roth) C.B.Clarke, Fl. Brit. India [J. D. Hooker] 4: 235. 1883. Type. Based on *Solanum multiflorum* Roth

*Solanum indicum* L. subsp. *erosum* (Van Heurck & Müll.Arg.) Bitter, Repert. Spec. Nov. Regni Veg. Beih. 16: 11. 1923. Type. Based on *Solanum erosum* Van Heurck & Müll.Arg.

*Solanum anguivi* Lam. var. *multiflorum* (Roth) Vajr., Fl. Tamil Nadu Ind., Ser I: Analysis 2: 115. 1987, as “*multiflora*”. Type. Based on *Solanum multiflorum* Roth

*Solanum violaceum* Ortega subsp. *multiflorum* (C.B.Clarke) K.M.Matthew, Kew Bull. 46: 545. 1991. Type. Based on *Solanum multiflorum* Roth

**Type.** INDIA. [“H. in India orientali. B. Heyne” protologue] “Peninsula Ind. orientalis” [1836], *herb. R. Wight prop. 1575* (neotype, designated by Turner 2021, pg. 418: K-W [K000759413]; probable isoneotypes: CAL, E [E00757306, E00617617, E00617618, E00617619, E00617621], NY [NY00825634], P [P00049971, P00055603, P03933610]).

**Description.** Erect shrub, 2–3 m, armed, much branched. Stems erect, robust, terete, densely stellate-pubescent and prickly, the trichomes porrect-stellate, sessile or short-stalked, the stalks to 0.8 mm long, the rays 6–8, 0.1–0.5 mm long, the midpoints to 2 mm long, the prickles to 10 mm long, to 6 mm wide, straight to slightly curved, flattened, orange-brown, glabrous or bearing a few stellate trichomes; new growth densely stellate-pubescent with trichomes like those of the stems; bark of older stems somewhat glabrescent, grey-brown. Sympodial units difoliate, the leaves geminate. Leaves simple, more or less deeply lobed, the blades 5–20 cm long, 3.5–16 cm wide, ca. 1.5 times longer than wide, ovate, chartaceous, discolorous, unarmed or prickly with up to 20 straight prickles to 10 mm long on primary and secondary veins on both sides of the leaf; adaxial surface densely to sparsely stellate-pubescent, with sessile to stalked porrect-stellate trichomes, the stalks to 0.8 mm, the rays 6–8, 0.2–0.8 mm, the midpoints 0.7–2 mm long; abaxial surface densely stellate-pubescent with sessile and short-stalked trichomes like those of the adaxial surface; major veins 3–5 pairs; base cuneate to truncate, often oblique; margins shallowly to deeply lobed, the lobes 3(4)
<figure><img src="image_60.png" /><figcaption>**Figure 45.** *Solanum multiflorum* Roth **A** herbarium specimen collected in India in 1835 (*Wight 1575*, E00757306) **B** habit (*Sampath Kumar et al. 126950*, India) **C** detail view of a fruit with seeds (*Sampath Kumar et al. 126958*, India) **D** detail view of a calyx and floral buds (*Sampath Kumar et al. 126937*, India) **E** detail view of an infructescence (*Sampath Kumar et al. 126950*, India). Photograph credits: **A** Royal Botanic Garden Edinburgh **B–E** X. Aubriot.</figcaption></figure>

on each side, 1.5–5 cm long, broad-deltate to obovate, often with well-developed secondary lobes, apically acute to rounded, the sinuses extending 1/3–4/5 to the midrib; apex acute; petioles 1–4 cm, 1/6–1/3 of the leaf length, unarmed or with a few straight
prickles, densely stellate-pubescent with porrect sessile and short-stalked trichomes like those of the stems. Inflorescences 3–7 cm long, apparently lateral, unbranched or forked, more rarely several times branched, with 10–20 flowers, 3–6 flowers open at any one time, densely stellate-pubescent with stellate-porrect trichomes like those of the stems, unarmed or with a few straight prickles; peduncle 1–5 mm long, usually unarmed; pedicels 0.5–1.5 cm long, stout, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, strongly recurved at anthesis, unarmed, densely stellate-pubescent like the inflorescence axes, articulated at the base; pedicel scars spaced 1–2 mm apart. Buds globose to ovoid, strongly exserted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 2–6 mm long, obconical to cup-shaped, the lobes 2–6 mm long, 1.5–2 mm wide, long-deltate, apically long-acuminate, unarmed or with a few straight prickles, densely stellate-pubescent with porrect-stellate trichomes like those of the pedicels. Corolla 1.3–1.5 cm in diameter, violet to purple, stellate, lobed ca. 2/3 of the way to the base, interpretalar tissue somewhat present, the lobes 4.5–6 mm long, 2.5–3 mm, deltate to ovate, spreading at anthesis, glabrous adaxially but with a few minute stellate trichomes at the tips and on the midribs, densely stellate-pubescent abaxially, the trichomes porrect, a mixture of sessile and stalked, the stalks up to 0.1 mm, the rays 6–8, 0.1–0.3 mm, the mid-points to 0.8 mm long, longer towards lobe apices. Stamens equal; anthers 3.5–4 mm, ca. 1.5 mm wide, tapering, drying brown to red-brown, sometimes with pronounced papillae on the dorsal surface, poricidal at the tips, the pores not elongating to slits with drying; filament tube minute; free portion of the filaments 0.5–1 mm, glabrous. Ovary conical, densely stellate-pubescent in the upper 3/4–1/2; style 5.5–7 mm long, stellate-pubescent for almost all of its length; stigma clavate, the surfaces minutely papillose. Fruit a globose berry, many per infructescence, 0.6–0.9 cm in diameter, orange-red to dark brown when ripe, the pericarp thin, smooth and shiny, with stellate trichomes sparsely covering the fruit pericarp persisting into maturity; fruiting pedicels 1–1.3 cm long, 0.7–1.5 mm wide at the base, 1.7–2.5 mm in diameter at the apex, woody, thick, strongly recurved downwards, unarmed or prickly with up to 5 filiform straight prickles; fruiting calyx moderately accrescent, elongating to (3.5–)5–11 mm long, covering up to 1/3 of the mature berry, usually not reflexed, unarmed or prickly with up to 10 straight prickles. Seeds ca. 10–15 per berry, 2.5–3.7 mm long, 2.1–3 mm wide, flattened-reniform, orange-brown, the surface minutely pitted, the testal cells pentagonal to slightly sinuate in outline. Chromosome number: n = 12, 2n = 24 (Madhavadian 1968; Kirti and Rao 1983; both as *Solanum indicum* var. *multiflorum* Clarke).

**Distribution** (Fig. 46). *Solanum multiflorum* is endemic to the western part of Tamil Nadu Province in southern India, occurring in the Nilgiri, Coimbatore, Didingul and Madurai districts.

**Ecology and habitat.** *Solanum multiflorum* is a plant of deciduous or evergreen broad-leaf forests and grows in forest edges and roadsides; from 1,700 to 2,300 m elevation.

**Common names and uses.** India. Karnataka: kahi sunde (*Hosagoudar* 96871); Tamil Nadu: soumbé (*Leschenault de la Tour* 163). Some of the names attributed to *S. indicum* in floras may refer to this species.
<figure><img src="image_61.png" /><figcaption>**Figure 46.** Distribution of *Solanum multiflorum*.</figcaption></figure>

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (669,122 km², LC); AOO (324 km², EN). *Solanum multiflorum* is a species of disturbed habitats in southern India, with many recent collections. It is not of particular conservation concern.

**Discussion.** *Solanum multiflorum* was treated as a variety of “*Solanum indicum*” (now a suppressed name, see Suppressed Names) in various Indian floras, in part due to issues with the application of Roth’s name. It is morphologically similar and related to *S. violaceum* (the name now used for the suppressed *S. indicum*), both taxa are members of *S. violaceum* group of Aubriot et al. (2016a), along with *S. hovei* and *S. deflexicarpum*. *Solanum multiflorum* is sympatric with *S. violaceum* and can be distinguished from it in its usually branched inflorescences with congested pedicels, long, soft pubescence and its short fruiting pedicels that are strongly deflexed rather than widely spreading. The leaves of *S. multiflorum* are usually more deeply lobed than those of *S. violaceum*, but *S. violaceum* is extremely variable and this does not always hold. *Solanum multiflorum* shares the strongly deflexed fruiting pedicels with *S. deflexicarpum* of China but differs from that species in its more copious pubescence, more deeply lobed leaves and usually purple (rather than white) flowers.

The species names proposed by Albrecht Wilhelm Roth were published in 1819 by Roemer and Schultes (1819), two years before they were published by Roth himself
(Roth 1821). Descriptions in Roemer and Schultes (1819) are abbreviated from Roth's
longer ones published in 1821; no new material was apparently seen (see Turner 2021).
Material at Edinburgh from Robert Wight's personal herbarium (here as probable iso-
neotypes) is in better condition than the sheet (K000759413) in the Wallich hebarium
designated as the neotype. The numbering of Wight's herbarium was done after the
fact specimens bearing the same number may or may not be duplicates of the neotype.

*Solanum himalense* var. *soumbe* was described based on the collection *Leschenault de la Tour 163* seen by Dunal (1852) in Paris. From the three specimens with this number at P we have selected the specimen (P00055586) with an annotation in Dunal's hand as the lectotype.

The collection *Hohenacker 1074* was cited from two herbaria ("hb. Van Heurck et hb. DC") in the protologue of *S. erosum* (Van Heurck 1870). 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 the duplicate of *Hohenacker 1074* held in BR (AWH10071571) as the lectotype for *S. erosum*; this collection is widely distributed.

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

**29. *Solanum nienkui* Merr. & Chun, *Sunnyatsenia* 2: 318. 1935.**
Fig. 47

**Type.** CHINA. Hainan: Sam Ah, 1932, *N.K. Chun & C.L. Tso 43319* (holotype: NY [00172269]; isotype: A [00077825]).

**Description.** Erect shrub, to 4 m, unarmed or less often prickly. Stems erect, terete, unarmed or armed with small prickles, sparsely to densely stellate-pubescent, glabrescent; prickles to 3 mm long, to 3 mm wide at the base, curved, deltate, laterally flattened, pale yellow, glabrous; pubescence of mixed sessile and short-stalked porrect-stellate trichomes, the stalks to 0.25 mm long, the rays 4–8, 0.1–0.4 mm long, the midpoints up to 0.8 mm long; new growth densely stellate-pubescent, light brownish to brown in dry material; bark of older stems dark brownish, glabrous. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, not lobed or less often shallowly lobed, the blades 3.5–10.5 cm long, 2.5–4 cm wide, ca. 1.5–2.5 times longer than wide, elliptic to broadly ovate, chartaceous, discolorous, unarmed or sometimes armed with small prickles, the prickles 1–9(–17) per leaf side, mostly inserted on the midvein, like those of the stems; adaxial surface moderately to densely stellate-pubescent, the stellate trichomes porrect, sessile or stalked, the stalks to 0.25 mm long, the rays 3–8, 0.1–0.5 mm long, the midpoints to 0.5 mm long, usually shorter than the rays; abaxial surface densely stellate-pubescent with trichomes like those of the adaxial surface; major veins 4–7 pairs, drying light green; base attenuate to short-attenuate; margins entire, sinuate, sometimes shallowly lobed when young, the lobes 2–3 on each side, 0.3–1 cm long, deltate, apically acute, the sinuses less than halfway to the midrib; apex acute; petioles 1.5–2.5 cm long, 1/7–1/3 of the leaf blade length, unarmed or prickly with 1–7 prickles like those of the stems, densely stellate-pubescent with sessile porrect
<figure><img src="image_62.png" /><figcaption>**Figure 47.** *Solanum nienkui* Merr. & Chun **A** herbarium specimen collected in Vietnam in 1932 (*Poilane 20877*, P00054081) **B** inflorescence and young stem (*Wang et al. 2073*, China) **C** detail view of a flower (*Wang et al. 2073*, China). Photograph credits: **A** CC-BY, Muséum national d'Histoire naturelle, Paris **B**, **C** S. Hul.</figcaption></figure>

trichomes like those of the stems. Inflorescences 2.5–7.5 cm long, internodal and lateral or more or less leaf-opposed, unbranched, with ca. 5–13 flowers, 1–3 flowers open at any one time, densely stellate-pubescent, with porrect stellate-trichomes like those of the stems, unarmed; peduncle 0.5–3 cm long, unarmed or armed with 1–3 prickles; pedicels 2.5–10 mm long, ca. 0.5 mm in diameter at the base, 0.75–1 mm in diameter at the apex, recurved, unarmed, densely stellate-pubescent with porrect stellate-trichomes like those of the axes, articulated at the base; pedicel scars spaced 0.1–0.6 mm apart. Buds ellipsoid, tapering, more or less strongly exerted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 1.2–1.5 mm long, campanulate, the lobes 0.5–2.5 mm long, 0.5–1 mm wide, deltate, acute at the apex, unarmed and densely stellate-pubescent with porrect stellate-trichomes like those of
the pedicels. Corolla 0.8–1.3 cm in diameter, white to purplish blue, stellate, lobed ca. 4/5 of the way to the base, the lobes 5.5–8 mm long, 1–2.5 mm wide, deltate, spreading at anthesis, densely stellate-pubescent abaxially on parts exposed in bud. Stamens slightly unequal; anthers unequal, two to three of the five 4.5–5.5 mm long and two to three 3.5–4 mm long, all ca. 0.75 mm wide, tapering, connivent, yellow, glabrous, poricidal at the tips, the pores not lengthening to slits with age; filament tube <0.5 mm long, glabrous; free portions of the filaments all equal, ca. 0.1 mm long, glabrous. Ovary conical, minutely glandular-puberulent; style 5.5–6 mm long, slender, curved at the apex, glabrous; stigma capitate, minutely papillate. Fruit a globose berry, 1–4 per infructescence, 0.5–1 cm in diameter, the pericarp smooth, red when mature, glabrous; fruiting pedicels 0.7–1.2 cm long, ca. 0.5 mm in diameter at the base, ca. 2 mm in diameter at the apex, woody, spreading, unarmed; fruiting calyx lobes expanding to 5 mm long, 2/5–3/5 the length of the mature fruit, broadly to narrowly deltate, spreading to perhaps somewhat reflexed, unarmed, ending with a long acumen. Seeds 10–32 per berry, 2.5–3 mm long, ca. 2 mm wide, flattened-reniform, dull yellow, the surface minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.

**Distribution (Fig. 48).** *Solanum nienkui* is found on the island of Hainan (South China) and in southern Vietnam.

<figure><img src="image_63.png" /><figcaption>**Figure 48.** Distribution of *S. nienkui*.</figcaption></figure>
**Ecology and habitat.** *Solanum nienkui* is found in dry, deciduous forests, growing in more or less open areas on clay or sand, from 100 to 1,000 m elevation.

**Common names and uses.** China. shu ci qie (Zhang et al. 1994); Vietnam. Gia Lai: trong nörvah [Jarai language] (*Dournes s.n.*); Ninh Thuân: bô kö chinh ao [Mnong language] (*Poilane 8933, 20877*), dan a xâm [Mnong language] (*Poilane 9887*), dan chinh ao [Mnong language] (*Poilane 9082*).

**Preliminary conservation status (IUCN 2019).** Near Threatened (LC). EOO (64,999 km², LC); AOO (72 km², EN). *Solanum nienkui* is a plant of dry forests, often considered less valuable than wetter habitats and thus more often subject to anthropogenic change. Given the relatively small areas in which the species occurs and the potential habitat alteration, we have assigned a preliminary status indicating some threat.

**Discussion.** *Solanum nienkui* is a weak shrub with deeply stellate flowers and a slightly zygomorphic androecium. It is morphologically similar to *S. robinsonii* and *S. putii*, and in the molecular analyses of Aubriot et al. (2016a) resolved as sister to *S. putii* in a group with *S. camranhense* that itself was sister to *S. cyanocarphium*. It differs from *S. robinsonii* (also found in Vietnam) in its wider, less discolorous leaves, deeply stellate, smaller corolla, and somewhat zygomorphic androecium. *Solanum camranhense* has similar shaped leaves to those of *S. nienkui* but is a low creeping plant rather than an upright, spindly shrub.

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

## 30. *Solanum peikuoense* S.S.Ying in X.Aubriot & S.Knapp, sp. nov.

urn:lsid:ipni.org:names:77298804-1

Fig. 49

**Diagnosis.** Like *S. pseudosaponaceum*, but with the upper surfaces of leaves glabrate and shiny, inflorescences forked but not many times branched, larger flowers corolla (2–2.4 cm diam. versus 1–1.5 cm diam.) and generally fewer seeds per berry (18–40 versus 25–>100).

**Type.** TAIWAN. Southern Taiwan: Pingtung County, Wutai Hsiang, 13 Oct 1993, Y-R. Lin et al. 134 (holotype: HAST [29822]; isotype: BM [BM000846380], E [E00320675], MO [MO-3105792, acc. # 04669018]).

**Description.** Erect shrub to small tree, to 4 m, unarmed, less often armed with a few prickles. Stems erect, terete, unarmed or with a few scattered prickles, sometimes dark purple (*Ku 1752*), sparsely stellate-pubescent; prickles, if present, 0.1–0.3 cm long, straight, deltate, orange-yellow, glabrous; pubescence of mixed sessile and variably stalked porrect-stellate trichomes, the stalks to 0.1 mm long, the rays 6–8, 0.1–0.5 mm long, the midpoints absent or up to 0.1 mm long; new growth densely stellate-pubescent, the trichomes like those of the stems; bark of older stems glabrescent, reddish brown and shiny. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, very shallowly lobed, the blades 11–18 cm long, 5.2–10 cm wide, ca. 1.5–2
<figure><img src="image_64.png" /><figcaption>**Figure 49.** *Solanum peikuoense* S.S.Ying **A** herbarium specimen collected in Taiwan in 1993 (*Lin et al. 134*, BM000846380) **B** old stem and prickles (field photograph, unvouchered, Taiwan) **C** detail view of leaf base and petiole (field photograph, unvouchered, Taiwan) **D** detail view of inflorescence and open flower (field photograph, unvouchered, Taiwan) **E** detail view of infructescence (field photograph, unvouchered, Taiwan). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B–E** M.-I. Weng.</figcaption></figure>
times longer than wide, elliptic to somewhat ovate, widest at the middle, chartaceous, strongly discolorous, unarmed or more rarely armed along the principal veins with tiny awl-shaped prickles like those of the stems; adaxial surface dark green, almost completely glabrous to sparsely stellate-pubescent along the midrib and veins but with a few sessile porrect-stellate trichomes on the midrib and along the principal veins, the rays 6–8, 0.3–0.5 mm long, the midpoints ca. 0.4 mm long, approximately equal to the rays; abaxial surface similarly but more densely pubescent with sessile porrect-stellate trichomes along the veins; major veins 5–7 pairs sparsely pubescent especially abaxially, drying pale yellow abaxially; base acute to attenuate, usually somewhat oblique; margins shallowly lobed, the lobes 5–7 on each side, 0.5–1 cm long, deltate and apically acute, the sinuses extending up to ca. 1/4 of the distance to the midvein; apex acute to acuminate; petiole 1.5–3.5 cm long, 1/9–1/5 of the leaf blade length, unarmed or with a few awl-shaped prickles, and very sparsely pubescent with porrect-stellate trichomes like those of the stems. Inflorescences 1.5–3.5 cm long, internodal and lateral, forked, with 10–16 flowers, some of these borne below the branching point, only a few flowers open at any one time, sparsely pubescent with porrect-stellate trichomes like those of the stems; peduncle ca. 1.3 cm long; pedicels 1–1.2 cm long, ca. 1 mm in diameter at the base, ca. 1.2 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 irregularly spaced (1)4–5 mm apart. Buds pointed-ovoid, the corolla exserted from the calyx lobe acumens before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube ca. 2 mm long, cup-shaped, tearing to form irregular lobes, the lobes ca. 1 mm long with an elongate acumen to 3 mm long, ca. 1 mm wide, sparsely stellate-pubescent to glabrescent. Corolla 2–2.4 cm in diameter, white or purple (*Ku 1752*), stellate, lobed ca. 2/3 of the way to the base, the lobes 8–10 mm long, 3–4 mm wide, spreading at anthesis, glabrous adaxially, glabrous or sparsely stellate-pubescent along the petal midribs abaxially. Stamens equal; anthers 5–7 mm long, ca. 1.5 mm wide, tapering, yellow, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous and papery; free portion of the filaments ca. 0.5 mm long, glabrous. Ovary conical, glabrous; style ca. 10 mm long, straight, glabrous; stigma minutely capitate, the surfaces minutely papillose. Fruit a globose berry, 2–6 per infructescence, 1–1.3 cm in diameter, the pericarp smooth, red or black (*Lin et al. 134*), glabrous; fruiting pedicels 1–2.3 cm long, 1–1.2 mm in diameter at the base, ca. 2 mm in diameter at the apex, woody, erect or spreading, unarmed; fruiting calyx lobes not markedly expanding, 1/3 the length of the mature fruit, spreading, broadly deltate with a filiform acumen to 3 mm long, this usually breaking off. Seeds 18–40 per berry, 3–3.5 mm long, 2–2.7 mm wide, flattened-reniform, dull yellow, the surfaces minutely pitted, the testal cells with sinuate margins. Chromosome number: not known.

**Distribution (Fig. 50).** *Solanum peikuoense* is endemic to the island of Taiwan, most collections are from the southern part of the island.

**Ecology and habitat.** *Solanum peikuoense* grows in broadleaf, coniferous or mixed broadleaf-coniferous forests, often along roadsides or in open areas, from 1,200 to 2,000 m elevation.
<figure><img src="image_65.png" /><figcaption>**Figure 50.** Distribution of *S. peikuoense*.</figcaption></figure>

**Common names and uses.** None recorded

**Preliminary conservation assessment (IUCN 2019).** Vulnerable (VU [B1,2a,bi,ii,]). EOO (10,013 km², VU); AOO (44 km², EN). *Solanum peikuoense* is a single island endemic with a small extent of occurrence. It is found in at least one protected area (Dawushan National Park), so is likely to be protected in the future.

**Discussion.** *Solanum peikuoense* is a distinctive plant, and unlikely to be confused with any other species in the region. It is a member of the Torva clade (see Aubriot et al. 2016a) and is morphologically similar to other southeast Asian members of the group such as *S. poka*, *S. pseudosaponaceum* and *S. torvoideum* (and the introduced *S. torvum*). Its shiny, almost glabrous mature leaves and stems distinguish it from any of these species, all of which are densely pubescent. It can be distinguished from the introduced *S. torvum* in the shiny leaves with acute to acuminate (rather than acute to truncate) apices, and its red or black berries (versus dirty green-yellow in *S. torvum*); *S. peikuoense* also lacks the characteristic simple glandular trichomes found in the inflorescence axes of *S. torvum*. Some collections (e.g., *Ku 1752*) mention dark purple stems and purple flowers; other gatherings record the flowers as white.

Although a single gathering (*Ying s.n.*) in a single herbarium (NTUF) was cited in Ying (1997:14), the word type or holotype, or its equivalent was not used. This is contrary to Art. 40.6 of the “Code” (Turland et al 2018) and renders “*Solanum peikuoensis*
S.S.Ying” not validly published. We validate this name here and cite as type material a gathering for the same area that is widely distributed and has both flowers and fruits.

**Paratypes.** [also see Suppl. materials 1–3] **TAIWAN. Chiayi Hsien:** Chiayi, Arisan, Kagi, common around Funkiko, 25 Jan 1918, *Wilson 9657* (A). **Hualien Hsien:** Tung-nengkao, Panpien, 20 Apr 1927, *Saito 6965* (A). **Kaohsiung Hsien:** Teng-chu, 22 Jan 1972, *Hsu 13689* (A); Tengchih, 17 Nov 1992, *Huang et al. 15850* (A); Taoyuan Hsiang, along Southern Cross-Island Hwy, between Lanya and Likuan, 27 Jun 2000, *Lin 405* (CAS, HAST); Taoyuan Hsiang, near road mileage sign 13 km on Meilan Forest Road, 10 May 1994, *Liu et al. 427* (HAST); Taoyuan Hsiang, Shihshan Forest Road 9.7 km, 21 May 1997, *Peng 16833* (HAST); Taoyuan Hsiang, Tengchih, 28 Jun 2000, *Wang 4317* (HAST); Kaohsiung Hsien, Taoyuan Hsiang, Tengchih, 29 Jun 2000, *Wang 4376* (HAST, PE); Taoyuan Hsiang, Tengchih, 24 Jul 2004, *Wang et al. 11628* (HAST). **Nantou Hsien:** Hsinyi Hsiang, Shenmu village, 31st forest compartment, 19 Apr 1984, *Peng 6576* (HAST); Jenai Hsiang, 9-10K, 22 Apr 2000, *Wang 5514* (HAST). **Pingtung Hsien:** Wutain Hsiang, Hsiaogueihu forest road, 3 May 2003, *Ku 1752* (HAST). **Taichung Hsien:** Hoping Hsiang, Chia-yang to Tsui-luan, 7 May 1997, *Lin et al. 938* (HAST); Hoping Hsiang, Tahsuehshan Forest Road, 30 Sep 2006, *Lu 12541* (HAST); Tainchung Hsien, Hoping Hsiang, Tahhuehshan 200 forest road 30K, 28 Feb 2001, *Wang & Li 4720* (HAST, PE); Hoping Hsiang, on Hsuehshan Forest Road, at road mileage sign 28.2 km, 5 May 1999, *Wu 1273* (A, HAST); Hoping Hsiang, Pei-ko-ta-shan, 1 Aug 1996, *Ying s.n.* (HAST). **Taitung Hsien:** Haituan Hsiang, Hungshih Forest Road, near road marker 9.5 km, 1 Mar 2000, *Liu 1369* (HAST); Peinan Hsiang, LiChia Forest Road, between road marker 19 km and 22 km, 25 Jul 2008, *Lu 16536* (HAST); Yenping Hsiang, Yenping Forest Road, 24 Dec 2000, *Yang s.n.* (HAST); Chin-shui-ying, 18 Sep 1998, *Yang 23750* (HAST).

**31. Solanum poka** Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 768. 1814.
Fig. 51

*Solanum torvum* Sw. var. *scabrescens* Miq., Fl. Ned. Ind. 2: 648. 1857. Type. Indonesia. Sumatra: Sin. loc., *F.W. Junghuhn s.n.* (lectotype, designated here: L [L.0403917]; isolectotype: U [U0113979]).

*Solanum torvum* Sw. var. *polyacanthum* Miq., Fl. Ned. Ind. 2: 648. 1857, as "*polyacantha*". Type. Indonesia. Java: sin. loc., *T. Horsfield s.n.* (lectotype, designated here: BM [BM000886307]).

**Type.** Based on an unpublished illustration of a Jean-Baptiste Leschenault de la Tour collection from Java kept in the Node-Véran collection in Montpellier (lectotype, designated by Aubriot et al. 2016b, pg. 105: [illustration] Sol. Tab. 55 [MPU028527]).

**Description.** Shrubs to 3 m, armed. Stems erect, terete, usually densely prickly distally, moderately pubescent, glabrescent; prickles to 3.5 mm long, to 2.5 mm wide at the base, straight, awl-shaped to deltate, conical, pale yellow, glabrescent; pubescence of mixed sessile and variously stalked porrect-stellate trichomes, the stalks to
<figure><img src="image_66.png" /><figcaption>**Figure 51.** *Solanum poka* Dunal – Herbarium specimen (epitype) collected in Indonesia (*Horsfield s.n.*, BM000886306). Photograph credit: CC-BY, © copyright The Trustees of the Natural History Museum, London.</figcaption></figure>
0.2 mm long, the rays (4–)5–8, 0.1–0.25 mm long, the midpoints reduced to mere bumps; new growth moderately stellate-pubescent, black to dark brownish in dry material; bark of older stems brownish grey, sparsely stellate-pubescent. Sympodial units difoliate, the leaves geminate. Leaves simple, entire to deeply lobed, the blades 11–24 cm long, 4–13 cm wide, ca. 1.5–3 times longer than wide, elliptic to broadly ovate, chartaceous, slightly discolorous, unarmed or prickly with 1–6 prickles per leaf side, the prickles to 6 mm long, to 1.5 mm wide at the base, straight or slightly curved at the tip, awl-shaped, conical, pale yellow, glabrous; adaxial surface moderately stellate-pubescent with porrect, sessile and less often variously stalked trichomes, the stalks to 0.1 mm long, the rays 4–8, 0.1–0.4 mm long, the midpoints to 0.25 mm long; abaxial surface moderately stellate-pubescent with trichomes like those of the adaxial surface, but more often stalked; major veins 6–8 pairs drying yellow; base short-attenuate to truncate; margins entire or shallowly to deeply lobed, the lobes 1–5 on each side, 0.5–5 cm long, deltate, rounded to apically acute, the sinuses extending up to 2/3 of the distance to the midvein; apex acute; petiole 1.5–4 cm long, 1/10–1/5 of the leaf blade length, densely stellate-pubescent with porrect, sessile trichomes like those of the blades, unarmed or prickly with 1–5 prickles like those of the stems. Inflorescences 2–5 cm long, apparently lateral or leaf opposed, unbranched to up to 2 times branched, with ca. 5–20 flowers, 2–6 flowers open at any one time, moderately to densely stellate-pubescent with porrect-stellate trichomes like those of the stems, unarmed; peduncle 0.5–1.5 cm long, unarmed or with very few prickles; pedicels 0.5–1.2 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, erect, unarmed, densely stellate-pubescent with porrect-stellate trichomes like those of the inflorescence axes, articulated at the base; pedicel scars spaced 2–4 mm apart. Flowers 5-merous, apparently all perfect. Calyx with the tube 1–3 mm long, conical, the lobes 3–5 mm long, ca. 1.5 mm wide, the lower part deltate and abruptly constricting to an elongate acumen, the acumen 3/4 of the total lobe length, the abaxial surface more or less strongly keeled along the midvein, unarmed, densely stellate-pubescent on the midvein with porrect-stellate trichomes like those of the pedicels. Corolla 1–2 cm in diameter, white, stellate, lobed ca. 1/2–2/3 of the way to the base, the lobes 5–8 mm long, 2–3.5 mm wide, deltate, spreading at anthesis, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal; anthers 5–6.5 mm long, ca. 0.75 mm wide, tapering, yellow, connivent, glabrous, poricidal at the tips, the pores not lengthening to slits with age; filament tube <0.5 mm long, glabrous; free portion of the filaments 0.75–1.5 mm long, glabrous. Ovary conical, minutely glandular-puberulent; style 0.6–1 cm long, slender, curved at the apex, with a few scattered hairs at the tip; stigma capitate, minutely papillate, stellate-pubescent. Fruit a globose berry, 8–18 per infructescence, 0.8–1.5 cm in diameter, the pericarp smooth, bluish green when young, turning to dark greyish yellow, glabrous; fruiting pedicels 1.2–2.5 cm long, ca. 1–1.5 mm in diameter at the base, 2–3 mm in diameter at the apex, woody, erect, unarmed; fruiting calyx lobes not expanding. Seeds 100–200 per berry, 1.75–2 mm long, 1.5–1.75 mm wide, flattened reniform, pale yellowish, the surface minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.
<figure><img src="image_67.png" /><figcaption>**Figure 52.** Distribution of *Solanum poka*.</figcaption></figure>

**Distribution (Fig. 52).** Solanum poka is widely distributed in the Malay Archipelago, from western Sumatra to the Maluku Islands and across Sulawesi, northwards to the Talaud islands.

**Ecology and habitat.** Solanum poka is a plant of evergreen broadleaf forests, growing in open woodland, forest edges, degraded vegetation, usually on limestone or volcanic rocks; from sea level to 1,600 m elevation.

**Common names and uses.** Indonesia. Java: pooka (Horsfield 786), daun poka (Dunal 1814); North Sulawesi: poki poki (De La Savinierre 343bis), sangkirumanuwawi (Lam 2772).

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (1,202,780 km², LC); AOO (68 km², EN). Although the EOO measurement indicates a status of least concern, the few collections coupled with the profound transformation in lowland Indonesian habitats where *S. poka* is found (Margono et al. 2014) suggest that the species is a priority for recollection and reassessment.

**Discussion.** Solanum poka is a member of the Torva clade (Aubriot et al. 2016a) and is similar to other members of the group occurring in southeast Asia such as *S. pseudosaponaceum* and *S. torvoideum*. Morphologically, *S. poka* most closely resembles *S. pseudosaponaceum*, a widespread species ranging from Taiwan and southern China to Indonesia but differs in having denser indumentum on the adaxial leaf surface, more numerous straight prickles on the upper stems, fewer, larger flowers with elongate strongly keeled calyx lobes, and much larger fruits. Flowers of *S. pseudosaponaceum* are lilac or purplish-white while those of *S. poka* are always described on labels as white. Solanum poka can be distinguished from the introduced *S. torvum* in its strongly keeled calyx lobes with elongate acumens (rather than deltate
lobes), red (rather than green) berries, and in the lack of simple glandular trichomes
in the inflorescence.

Aubriot et al. (2016b) recognised *S. torvum* var. *scabrescens* as a synonym in their
resurrection of the name *S. poka* but erroneously referred to the Leiden duplicate
(L.0403917) of the *Junghuhn s.n.* collection cited in the protologue as “holotype”;
Miquel (1857) cited no herbaria in the protologue. They did not include Miquel’s var.
*polyacanthum* in the synonymy of *S. poka*, but the minimal description with reference
to pricklier stems and leaves approaching *S. giganteum*, however, suggest this varietal
name refers to the taxon we recognise as *S. poka*. The protologue states no specific lo-
cality, merely “β. Java (Horsfield)”. We have therefore selected the specimen of *S. poka*
from Java collected by Thomas Horsfield that best matches the protologue at BM
(BM000886307) as the lectotype for *S. torvum* var. *polycanthum*.

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

**32. Solanum praetermissum Kerr ex Barnett, Kew Bull. 16: 485. 1963.**
Figs 4G, 53

*Solanum barbisetum* Nees var. *griffithii* Prain, J. Asiat. Soc. Beng. 65(2): 541. 1896.
Type. India. [Arunachal Pradesh?]: “Upper Assam”, 1841, *F. Jenkins* 253 (lectotype,
designated by Hul and Dy Phon 2014, pg. 36: K [K000449043]; isolectotype: K
[K000449042]).

*Solanum griffithii* (Prain) C.Y.Wu & S.C.Huang, Acta Phytotax. Sin. 16(2): 75. 1978,
non *S. griffithii* (C.B.Clarke) Kuntze, 1891. Type. Later homonym, based on
*Solanum barbisetum* Nees var. *griffithii* Prain.

*Solanum neogriffithii* V.V.Hop, J. Biol. Vietnam 27(3): 13. 2005. Type. Based on
*Solanum barbisetum* Nees var. *griffithii* Prain.

*Solanum membranisepalum* Li Bing Zhang & Ngan T.Lu, Phytotaxa 186(4): 239.
2014. Type. Based on *Solanum barbisetum* Nees var. *griffithii* Prain.

*Solanum brevipedunculatum* Li Bing Zhang & Yi F.Duan, Phytotaxa 170(4): 280.
2014, non *S. brevipedunculatum* Rusby, 1907. Type. Based on *Solanum barbisetum*
Nees var. *griffithii* Prain.

**Type.** THAILAND. Tak: Raheng [Rahaeng], *E. Smith s.n.* (holotype: K [K000614048]; isotypes: BK [BK229395], BM [BM000886099]).

**Description.** Herbs to small shrubs, to 0.5 m tall, armed. Stems erect, terete, moderately to sparsely prickly and densely stellate-pubescent; prickles to 4 mm long, to 2 mm at the base, straight, narrowly deltate, pale yellow, glabrous; trichomes por-rect-stellate, mixture of sessile and stalked, the stalks to 0.25 mm long, the rays 4–8, 0.1–0.4 mm long, the midpoints up to 0.1 mm long; new growth densely stellate-pubescent, whitish green in dry material; bark of older stems light brownish, mod-erately to densely stellate pubescent. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing somewhat in size but not in shape. Leaves simple, entire to shallowly lobed, the blades 9–17 cm long, 6.5–13 cm wide, ca. 1–1.5 times longer
<figure><img src="image_68.png" /><figcaption>**Figure 53.** *Solanum praetermissimum* Kerr ex Barnett **A** herbarium specimen collected in Laos in 1974 (*Vidal* 5855, P00074767) **B** habit (field photograph, unvouchered, Vietnam) **C** detail view of a fruit with the calyx removed (field photograph, unvouchered, Vietnam) **D** detail view a flower (field photograph, unvouchered, Vietnam) **E** detail view an infructescence (field photograph, unvouchered, Vietnam). Photograph credits: **A** CC-BY, Muséum national d'Histoire naturelle, Paris **B–E** M. Ruraliev.</figcaption></figure>

than wide, ovate to broadly ovate, chartaceous, discolorous, moderately prickly with 4–16(–24) prickles per leaf side, usually longer than those of the stems, up to 7 mm long, often purplish black in living plants; adaxial surface dark green, moderately to
densely stellate-pubescent, the stellate trichomes porrect, sessile or stalked, the stalks to 0.1 mm long, the rays 4–7, 0.1–0.4 mm long, the midpoints up to 0.5 mm long; abaxial surface light green, densely stellate-pubescent with trichomes like those of the adaxial surface but more long-stalked; major veins 4–5 pairs drying whitish brown; base attenuate to truncate; margins sinuate to shallowly lobed, the lobes 3–4 on each side, 0.5–2 cm long, broadly deltate to obovate, apically rounded to acute, the sinuses extending up to 1/3 of the distance to the midvein; apex acute; petiole 2.5–7 cm long, 1/4–1/3 of the leaf blade length, densely stellate-pubescent with porrect sessile trichomes like those of the stems, unarmed or with 1–8 prickles like those of the blades. Inflorescences 1.5–3 cm long, apparently lateral, unbranched, with ca. 5–11 flowers, 1–2 flowers open at any one time; axes densely stellate-pubescent with trichomes like those of the stems, but sometimes purplish tinged in living plants, unarmed or with prickles like those of the stems but smaller, often purplish black in live plants; peduncle 0.2–1 cm long, with 0–4 prickles; pedicels 0.4–1 cm long, ca. 0.75 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, erect to recurved, unarmed or with 1–2 small prickles, densely stellate-pubescent with trichomes like those of the inflorescence axes, articulated at the base; pedicel scars spaced 1–3 mm apart. Buds oval to ellipsoid, more or less exerted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 2–3 mm long, conical, the lobes 2–3.5 mm long, 0.75–1.5 mm wide, narrowly deltate, apically acute, unarmed and densely stellate-pubescent abaxially with trichomes like those of the pedicels. Corolla 0.7–1.5 cm in diameter, white, sometimes pale lilac, stellate, lobed ca. 2/3 of the way to the base, the lobes 4–5.5 mm long, 1.5–3 mm wide, narrowly deltate to deltate, spreading at anthesis, glabrous adaxially, densely stellate pubescent abaxially on parts exposed in bud. Stamens equal; anthers 4–5 mm long, ca. 1 mm wide, not markedly connivent, tapering, yellow, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube <1 mm long; free portion of the filaments 0.5–1 mm long, glabrous. Ovary conical, minutely glandular-puberulent at the top; style ca. 6 mm long, slender, slightly recurved at the apex, glabrous; stigma capitate, minutely papillate. Fruit a globose berry, 4–8 per infructescence, 0.8–1.2 cm in diameter, whitish cream at maturity, the pericarp smooth, glabrous; fruiting pedicels 0.9–1.2 cm long, ca. 1 mm in diameter at the base, 2–3.5 mm in diameter at the apex, woody, erect to recurved, unarmed or with 1–10 prickles; fruiting calyx accrescent, completely covering the fruit at maturity, the tube enclosing the ripe berry, prickly and stellate-pubescent, the lobes expanding to 1 cm long, with up to 20 prickles per lobe, these often purplish black in living plants. Seeds 20–50 per berry, 2–3 mm long, 1.5–2 mm wide, flattened-reniform, light brownish, the surface minutely pitted, the testal cells somewhat sinuate in outline. Chromosome number: not known.

**Distribution (Fig. 54).** *Solanum praetermissum* occurs from northeastern India to South China and Vietnam.

**Ecology and habitat.** *Solanum praetermissum* is a plant of evergreen broadleaf forests or semi-deciduous forests, growing in forest understory or in clearings of secondary woodlands, or on rocky banks; from 300 to 400 m elevation.

**Common names and uses.** Vietnam. co chu (Muong, Hul and Dy Phon 2014).
<figure><img src="image_69.png" /><figcaption>**Figure 54.** Distribution of *S. praetermissum*.</figcaption></figure>

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (1,191,003 km², LC); AOO (68 km², EN). *Solanum praetermissum* is widely distributed, but where we have seen it in the field, is in small populations, perhaps indicating further assessment is necessary.

**Discussion.** *Solanum praetermissum* is most similar to *S. barbisetum*, with which it is sympatric and has previously been confused. It was first described as a variety of the latter (as var. *griffithii*) and once it was recognised as distinct several attempts were made to assign a name at the specific level (see synonymy); none of these authors apparently realised the name *S. praetermissum* was already published and thus had priority. Both species are small shrubs and have small, whitish cream berries enclosed in prickly accrescent calyces. *Solanum praetermissum* can be distinguished from *S. barbisetum* in its pubescence of largely sessile stellate trichomes (versus long-stalked stellate trichomes), lack of bristles on the inflorescence (versus densely bristly inflorescences and calyces, especially in fruit), generally shorter inflorescence axes (1.5–3 cm versus 3–10 cm long, although some young inflorescences can be shorter), and smaller flowers (0.7–1.5 cm versus 1.8–2.2 cm in diameter).

Mill (2001) treated *S. praetermissum* for Bhutan under the synonym *S. griffithii*, with the suggestion that they were doubtfully distinct. He cited no specimens for Bhutan for either species.
Aubriot et al. (2016a) resolved *S. praetermissum* as sister to the southern Indian endemic *S. wightii*, but with very low support; in the same study, *S. barbisetum* was resolved as sister to the 'S. expedunculatum and relatives' lineage albeit with poor support. *Solanum wightii* also has a strongly accrescent calyx but is otherwise morphologically very different from *S. praetermissum* (see description below).

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

## 33. *Solanum procumbens* Lour., Fl. Cochinch. 132. 1790.

Fig. 55

*Solanum hainanense* Hance, J. Bot. 6: 331. 1868. Type. China. Hainan: "At vias prope Kieng chau fú, metropolis ins: Hainan", Nov 1866, Sampson & H.F. Hance s.n. [13816] (lectotype, designated here: BM [BM000942492]; isolectotype: K [K000759385]).

*Solanum scopulorum* Kerr ex Barnett, Kew Bull. 16: 486. 1963. Type. Thailand. Prachuap Khiri Khan: Hua Hin, 11 Nov 1928, A.F.G. Kerr 16210 (holotype: K [K000922039]; isotypes BK [BK257532], BM [BM000886111]).

**Type.** VIETNAM. Thua Thiên-Huê: Huê and vicinity, Jan 1923, *R. W. Squires 27* (neo-type, designated by Hul and Dy Phon 2014, pg. 20: P [P00054100]; isoneotypes: BM [BM000886103], E [E00224906], K [K000195689]).

**Description.** Scandent or creeping herbs or shrubs, to 2(–3) m tall, armed. Stems prostate or erect, terete, prickly and pubescent; prickles up to 1 cm long, to 1 cm at the base, more or less regularly spaced, strongly hooked but sometimes straight, flattened and deltate, pale yellow, glabrous; pubescence of sessile porrect-stellate trichomes, the rays 6–8, 0.1–0.3 mm long, the midpoints absent or <0.1 mm long; new growth sparsely to densely stellate-pubescent, green to light brownish in dry material; bark of older stems brownish grey, glabrous. Sympodial units apparently plurifoliate, the leaves not geminate. Leaves simple, entire to moderately lobed, the blades 2–6.5 cm long, 1–3.5 cm wide, ca. 1.5–2.5 times longer than wide, elliptic to ovate, sometimes obovate, chartaceous, discolorous, unarmed or with up to 5 prickles per face, these slightly smaller and thinner than the ones on the stems; adaxial surface green to dark green, evenly and sparsely pubescent with erect sessile porrect-stellate trichomes, the rays 6–8, 0.1–0.3 mm long, the midpoints to 0.1 mm; abaxial surface densely stellate pubescent with trichomes like those of the adaxial surface but with stalks up to 0.1 mm long and longer rays; major veins ca. 3 pairs drying light brownish; base attenuate; margins entire, shallowly to moderately lobed, the lobes 1–3 on each side, 0.4–1 cm long, oblong to deltate, apically rounded, the sinuses extending up to 1/3–1/2 of the distance to the midvein; apex acute to rounded; petioles 0.5–1 cm long, 1/10–1/4 of the leaf blade length, unarmed or sparsely prickly with a few curved prickles, densely stellate-pubescent, the pubescence of sessile porrect-stellate trichomes like those of the blades. Inflorescences 1.5–4 cm long, lateral or occasionally leaf-opposed, unbranched, with 4–8 flowers, 1–3 flowers open at any
<figure><img src="image_70.png" /><figcaption>**Figure 55.** *Solanum procumbens* Lour. **A** herbarium specimen collected in China in 1935 (Lau 5138, E00320676) **B** habit (Wang et al. 2075, China) **C** inflorescence (Wang et al. 2075, China) **D** detailed view of a flower and prickles (Wang et al. 2075, China) **E** infructescence (Wang et al. 2075, China). Photograph credits: **A** Royal Botanic Garden Edinburgh **B–E** S. Knapp.</figcaption></figure>

one time, unarmed or sparsely prickly, densely pubescent with sessile stellate-porrect trichomes like those of the stems; peduncle 0.1–1.4 cm long; pedicels 0.5–2 cm long, ca. 0.5 mm in diameter at the base and 0.5–0.75 mm in diameter at top, erect or somewhat recurved at anthesis, densely stellate-pubescent like the inflorescence axes,
articulated at the base; pedicel scars irregularly spaced 0.1–2 mm apart. Buds ovoid, exserted from the calyx before anthesis. Flowers 4(–5)-merous, apparently all perfect. Calyx with the tube 1–1.5 mm long, campanulate, the lobes 1–1.5 mm long, ca. 1 mm wide, deltate to broadly deltate, apically acute to acuminate, unarmed, densely stellate-pubescent with porrect-stellate trichomes like those of the pedicels. Corolla 1–1.5 cm in diameter, white or pale purple to deep blue, stellate, lobed ca. 3/4 of the way to the base, the lobes 5–7 mm long, 1–3 mm wide, long-triangular, spreading or reflexed at anthesis, mostly glabrous adaxially but with a few stellate trichomes on the middle vein, densely stellate pubescent abaxially on parts exposed in bud. Stamens equal or slightly unequal with 2 slightly longer than the rest; anthers 5–5.5 mm long, 1–1.5 mm wide, not connivent to somewhat spreading, all tapering, dull yellow, glabrous, poricidal at the tips, the pores not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments ca. 0.5 mm long, glabrous. Ovary conical to globular, minutely glandular-puberulent at the top; style 6–8 mm long, with few stellate trichomes scattered at the base; stigma clavate, the surfaces minutely papillose. Fruit a globose berry, several per infructescence, 0.6–1 cm in diameter, red at maturity, pericarp thin, glabrous; fruiting pedicels 1.3–1.8 cm long, ca. 1 mm in diameter at the base, 1.5–1.8 mm in diameter at the apex, erect or slightly recurved, somewhat woody, unarmed; fruiting calyx lobes ca. 2 mm long, not markedly accrescent, but covering 1/4 of the berry and not reflexed, unarmed. Seeds 5–25 per berry, 2–3 mm long, 2–3 mm wide, flattened reniform to rounded, brownish orange, the surfaces minutely pitted, the testal cells pentagonal in outline. Chromosome number: not known.

**Distribution (Fig. 56).** *Solanum procumbens* is widely distributed from China and Indochina to Indonesia (Flores and Timor) and Timor Leste.

**Ecology and habitat.** *Solanum procumbens* is a plant of open places in many forest types, often scrambling over other vegetation and on rocks, from sea level to 650(1,500) m elevation.

**Common names and uses.** China. hai nan qie (Zhang et al. 1994). Vietnam. cà gai, (cây) cur'o'ng, trông cà dap (Hul and Dy Phon 2014)

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (1,575,888 km², LC); AOO (164 km², EN). *Solanum procumbens* is widely distributed and occurs in many habitat types, including extremely anthropogenically disturbed ones.

**Discussion.** *Solanum procumbens* is similar to *S. trilobatum* and *S. camranhense* in its habit as a small scrambling shrub, but in molecular analyses of Aubriot et al. (2016a) resolves as closely related to *S. involucratum*, a much more robust and morphologically very different plant. *Solanum procumbens* can be distinguished from *S. camranhense* in its usually 4-parted corollas, slender fruiting pedicels, and more lanceolate leaves with less truncate bases.

No specimens were cited in the protologue (Hance 1868), so the lectotype we have selected for *S. hainanense* (BM00942492) is the better preserved of the two duplicates we have found and bears the Hance exsiccata number 13816.

**Specimens examined.** See Suppl. materials 1–3.
<figure><img src="image_71.png" /><figcaption>**Figure 56.** Distribution of *Solanum procumbens*.</figcaption></figure>

34. *Solanum pseudosaponaceum* Blume, Bijdr. Fl. Ned. Ind. 13: 702. 1826.
Figs 3F, 57

*Solanum inaequilaterale* Merr., Philipp. J. Sci. 1, Suppl. 236. 1906. Type. Philippines.
CAR: Luzon, Benguet Province, Suyse to Panai, Oct 1905, *E.D. Merrill* 4807
(lectotype, designated here: US [00027616, acc. # 710478]; isolectotypes: K
[K000195913], NY [00172282]).

*Solanum torvum* Sw. var. *lasiostylum* Y.C.Liu & C.H.Ou, Quart. J. Chinese Forest. 7(4): 151. 1974. Type. Taiwan. Sin. loc., "In southern parts at low altitudes", 1 Apr 1971, *Y.C. Liu & C.H. Ou* 540 (holotype: TCF).

*Solanum lasiostylum* (Y.C.Liu & C.H.Ou) Tawada, J. Phytogeogr. Taxon. 27(1): 36.
1979. Type. Based on *Solanum torvum* Sw. var. *lasiostylum* Y.C.Liu & C.H.Ou.

**Type.** INDONESIA. Java: Sin. loc., *C.L. Blume s.n.* (lectotype, designated here: GZU [GZU000255455]; isolectotypes: L [L0003669, L0003670]).

**Description.** Erect shrub to small tree to 8 m tall, armed. Stems erect, terete, prickly, usually densely stellate-pubescent; prickles to 5 mm long, to 5 mm wide at the base, straight to slightly curved, deltate, laterally flattened, pale yellow, glabrescent; pubescence of mixed sessile and variously stalked porrect-stellate trichomes, the stalks to 0.25 mm long, the rays 5–8, 0.25–0.5 mm long, the midpoints absent or up to 0.25 mm long; new growth densely stellate-pubescent with a whitish pubescence, bark black; bark of older stems brownish grey to black, glabrescent. Sympodial units difoliate to plurifoliate, the leaves not geminate. Leaves simple, entire to deeply lobed, the blades 8–17 cm long, 3.5–10.5 cm wide, ca. 1.5–3 times longer than wide, elliptic
<figure><img src="image_72.png" /><figcaption>**Figure 57.** *Solanum pseudosaponaceum* Blume **A** herbarium specimen collected in Taiwan in 1864 (*Old-ham* 342, BM000846375) **B** fertile stem (field photograph, unvouchered, Philippines) **C** open fruit with seeds inside (field photograph, unvouchered, Taiwan) **D** inflorescence (field photograph, unvouchered, Philippines) **E** infructescence (field photograph, unvouchered, Philippines). Photograph credits: **A** CC-BY, © copyright The Trustees of the Natural History Museum, London **B**, **D**, **E** D. Tandang **C** M.-I. Weng.</figcaption></figure>

to ovate, chartaceous, strongly discolorous, unarmed or prickly with 1–6 prickles per leaf side, to 6 mm long, to 1.5 mm wide at the base, straight, awl-shaped, conical, pale yellow, glabrous; adaxial surface dark green, usually sparsely stellate-pubescent to glabrescent, sometimes moderately stellate-pubescent (e.g., *Elmer* 12879), the stellate
trichomes porrect, sessile or stalked, the stalks to 0.25 mm long, the rays 4–9, 0.25–0.75 mm long, the midpoints to 0.5 mm long; abaxial surface light yellowish green, densely stellate-pubescent with trichomes like those of the adaxial surface, but more often stalked; major veins 5–7 pairs drying yellow; base short-attenuate to truncate; margins entire or shallowly to deeply lobed, the lobes 1–5 on each side, 0.5–4.5 cm long, deltate, rounded to apically acute, the sinuses extending up to 2/3 of the distance to the midvein; apex acute; petiole 1.5–4 cm long, 1/5–1/3 of the leaf blade length, moderately to densely stellate-pubescent with porrect, sessile trichomes like those of the blades, unarmed or prickly with 1–3 prickles like those of the stems. Inflorescences 3–5.5 cm long, apparently lateral, forked to 3 times branched, with ca. 10–40 flowers, 1–8 flowers open at any one time, densely stellate-pubescent with porrect trichomes like those of the stems, unarmed; peduncle 0.5–1.5 cm long, unarmed; pedicels 0.7–0.8 cm long, ca. 0.75 mm in diameter at the base, ca. 0.75 mm in diameter at the apex, erect, unarmed, densely stellate-pubescent with porrect trichomes like those of the inflorescence axes, articulated at the base; pedicel scars spaced 0.5–2 mm apart. Buds ovoid to ellipsoid, strongly exerted from the calyx before anthesis. Flowers 5-merous, apparently all perfect. Calyx with the tube 0.5–0.75 mm long, campanulate, the lobes 1–1.5 mm long, 0.5–0.75 mm wide, deltate, apically acute to acuminate, unarmed and densely stellate-pubescent abaxially with trichomes like those of the pedicels. Corolla 1–1.5 cm in diameter, purple, stellate, lobed ca. 1/2–3/4 of the way to the base, the lobes 4–7 mm long, 2–4 mm wide, deltate, spreading at anthesis, adaxially glabrous but with a few stellate trichomes at the lobe tips, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal; anthers ca. 5 mm long, ca. 1 mm wide, tapering, yellow, connivent, glabrous, poricidal at the tips, the pores not lengthening to slits with age; filament tube ca. 1 mm long, glabrous; free portion of the filaments 0.5 mm long, glabrous. Ovary globose, with minute glandular hairs and few stellate hairs in the lower half; style 7–8 mm long, slender, curved at the apex, densely stellate pubescent in the lower half; stigma capitate, minutely papillate. Fruit a globose berry, several to many per infructescence, 0.7–1 cm in diameter, the pericarp smooth, thin, red when mature, glabrous; fruiting pedicels 1–1.5 cm long, 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, woody, erect, unarmed; fruiting calyx lobes expanding to 2.5 mm long, 1/3 the length of the mature fruit, broadly deltate, reflexed, unarmed. Seeds 25–more than 100 per berry, 2–3 mm long, 1.5–2 mm wide, flattened-reniform, dull yellow, the surface minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.

**Distribution (Fig. 58).** *Solanum pseudosaponaceum* is widely distributed from South China (Fujian, Guangdong, Hainan provinces), Taiwan and Ryukyu Islands of southern Japan to the Philippines, Indonesia (Sulawesi, Java) and Indochina (Cambodia, Laos).

**Ecology and habitat.** *Solanum pseudosaponaceum* is found in pristine or secondary rainforests as well as in open wasteland, usually growing on sand or limestone; from 20 to 2,450 m elevation.

**Common names and uses.** China. shan qie (Zhang et al. 1994); Guangxi: tain tin ke tsz shue (*Tsang 23902*); Hainan: dithoulat (Lois, *Lau 157*), shan ke (*Tsang 874*). Indonesia. Java: tokokka, magay (Blume 1826). Philippines: qumit (Ifugao,
<figure><img src="image_73.png" /><figcaption>**Figure 58.** Distribution of *S. pseudosaponaceum*.</figcaption></figure>

*Conklin 217*). Taiwan: mao zhu wan tao hua ('twigs are fluffy, many flowered, pink like peach flowers')

**Preliminary conservation status (IUCN 2019).** Least Concern (LC). EOO (2,387,194 km², LC); AOO (160 km², EN). *Solanum pseudosaponaceum* is widely distributed across tropical Asia, and where it occurs apparently grows in large populations at forest edges and other disturbed habitats.

**Discussion.** This species was treated as *S. macaoense* Dunal in the “Flora of China” (Zhang et al. 1994), however, the type specimen of *S. macaoense* (*Callery* 28/29, P00055495) is clearly conspecific with *S. torvum*, making *S. pseudosaponaceum* the oldest and correct name for this taxon. *Solanum pseudosaponaceum* is one of the few Torva clade species occurring outside of the Americas (Aubriot et al. 2016a). It shares with the American members of the clade a shrubby habit, usually branched inflorescences, and hermaphroditic flowers. In common with *S. poka*, *S. peikuoense* and *S. torvoideum*, it has red berries; members of this clade native to the Americas have green to yellowish green mature fruits (e.g., *S. chrysotrichum*, *S. torvum*). For characters useful in distinguishing *S. pseudosaponaceum* from other members of the Torvum clade see descriptions of *S. poka* and *S. torvum*.

The specimen (Martin 1383) treated as *S. asperolanatum* Ruiz & Pav. in Hul and Dy Phon (2014) is a juvenile plant of *S. pseudosaponaceum*; in the absence of any inflorescences, it is difficult to place, but the pubescence and prickles accord better with it being identified as *S. pseudosaponaceum* than *S. chrysotrichum* (another determination on the sheet at P). *Solanum pseudosaponaceum* is only known from a few collections in Indochina. *Solanum asperolanatum* is an Andean endemic and does not occur in tropical Asia.
The disjunct distribution of the Torva clade is most likely due to long-distance dispersal (Dupin et al. 2016; Echeverría-Londoño et al. 2020), but from a single event, all species included in phylogenetic analyses (Aubriot et al. 2016a) are each other's closest relatives.

The specimen in Nees van Esenbeck's herbarium (Stafleu and Cowan 1981) at GZU (GZU0005524455) is annotated "Blume leg." and has an original label with the common name "Tokkoka" cited in the protologue. We have selected this as the lectotype for *S. pseudosaponaceum* because it is clearly from Blume's herbarium and has additional information from the protologue on it. The two collections in L here recognised as isolectotypes bear minimal labelling, and none of them has any locality or common name information; one of them (L0003669) has been previously annotated as "lectotype" by an unknown user or by L herbarium staff.

In the protologue of *Solanum inaequilaterale* (Merrill 1906) two collections were cited, *Merrill 4807* and *Elmer 6204*, the latter with the comment "appears to be a form of this species". We have selected the best preserved of the duplicates we have seen of *Merrill 4807* (US [00027616, acc. # 710478]) as the lectotype for this name, as Merrill unequivocally accepted this collection as his new species.

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

**35. *Solanum pubescens* Willd., Phytographia 1: 5. 1794.**

Figs 3D, 4D, 59

*Solanum calycinum* Nees, Trans. Linn. Soc. London 17(1): 60. 1834. Type. India. Sin. loc., "Herb. Madr. 237", *Anonymous s.n.* [Wallich Catal. Suppl. n. 237] (lectotype, designated here: GZU [GZU000255511])

*Solanum neesianum* D.Dietr., Syn. Pl. (D. Dietrich) 1: 697. 1839, nom. illeg., non *Solanum neesianum* Wall. ex Nees, 1834. Type. Based on *Solanum calycinum* Nees

*Solanum esenbeckii* Steud., Nomencl. Bot. ed. 2, 2: 602. 1841, nom. illeg. superfl. Type. Based on *Solanum calycinum* Nees

*Solanum conanthum* Dunal, Prodr. [A. P. de Candolle] 13(1): 127. 1852, nom. illeg. superfl. Type. Based on *Solanum calycinum* Nees

**Type.** Cultivated. "Habitat in Indiae hortis, arboretis solo argilloso rarius", *Anonymous s.n.* (lectotype, designated by Hepper 1987, pg. 369, as "type": B [B-W04313-010]).

**Description.** Erect shrubs to 5 m tall, unarmed. Stems erect, terete, stellate-pubescent and sticky glandular; pubescence of very short-stalked multangulate trichomes mixed with sessile porrect-stellate trichomes, the multangulate trichomes with more than 10 rays, the rays 0.4–0.5 mm long, the porrect-stellate trichomes with 6–8 rays, 0.4–0.5 mm long, the midpoints to 1 mm long, all trichomes usually glandular tipped and the plants sticky; new growth densely glandular-pubescent, the trichomes tangled, soon deciduous and the stems glabrate; bark of older stems greyish white. Sympodial units plurifoliate, the leaves not geminate. Leaves simple,