Lycianthes laevis (Dunal) Bitter, 1919

Knapp, Sandra, 2024, A revision of Lycianthes (Solanaceae) in tropical Asia, PhytoKeys 245, pp. 1-106 : 1-106

4. Lycianthes laevis (Dunal) Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 484. 1919.

Figs 3 B , 4 C , 12

Solanum laeve Dunal , Encycl. [J. Lamarck & al.] Suppl. 3: 751. 1814. Type. Indonesia. [Java]: Goudolone, J. B. L. Leschenault de la Tour 683 (lectotype, designated here: P [P 000578637]).

Solanum violaceum Blume , Cat. Gew. Buitenzorg 55. 1823, nom illeg. not Solanum violaceum Ortega (1798) . Type. Indonesia. “ Java ”, C. L. Blume s. n. (no specimens cited; neotype, designated here: L [L 0003600]).

Solanum crassipetalum Wall. , Fl. Ind. (Carey & Wallich ed.) 2: 256. 1824. Type. Nepal. “ Sheopar ”, 1821, R. Blinkworth s. n. [Wallich cat. 2618] (lectotype, designated by Deb 1978, pg. 293 [as “ type ” and with collector R. Brown]: CAL [CAL 0000017942]; isolectotypes: BM [BM 000617159], E [E 00273881, E 00273882], G-DC [G 00131653], K [K 0009234440], K-W [K 001116619, K 001116620]).

Solanum blumei Nees ex Blume , Bijdr. Fl. Ned. Ind. 13: 696. 1826, as “ blumii ”. Type. Based on Solanum violaceum Blume

Solanum pachypetalon Spreng. , Syst. Veg. ed. 16 [Sprengel] 4 (2): 72. 1827, nom. illeg. superfl. Type. Nepal. Based on Solanum crassipetalum Wall. (cited in synonymy).

Solanum membranaceum Wall. ex Nees , Trans. Linn. Soc. London 17 (1): 41. 1834. Type. India. [probably southern India], N. Wallich Cat. 2625 A (lectotype, designated here: K-W [K 001116640]).

Solanum bigeminatum Nees , Trans. Linn. Soc. London 17 (1): 42. 1834. Type. India. “ Travancore ”, Wallich cat. suppl., Oct 1814, Anon. s. n. [Herbarium Rottlerianum] (lectotype, designated by Hepper 1987, pg. 388: K [K 000759406]).

Solanum neesianum Wall. ex Nees , Trans. Linn. Soc. London 17 (1): 42. 1834. Type. India. [Khasi Hills] “ Mt Sylhet ”, N. Wallich [G. Gomez s. n.] Cat. Suppl. 248 (lectotype, designated here: GZU [GZU 000255706].

Solanum zollingeri Dunal var. multiflorum Dunal , Prodr. [A. P. de Candolle] 13 (1): 176. 1852. Type. Indonesia. Java: [no locality in protologue] East Java, in summo montis Taroeb [= Gunung Tarub], prov. Probolingo, 4 Jan 1845 [from P 00369020], H. Zollinger 2597 (holotype: G-DC [G 00145616]; isotypes: BM [BM 001018925], G [G 0343077, G 000357860], P [P 00369013, P 00379534 (as 2597 bis), P [P 00369020, as 2597 Z]).

Solanum subtruncatum Dunal , Prodr. [A. P. de Candolle] 13 (1): 180. 1852. Type. Bangladesh. “ In India orientalis, Silhet, Wall. cat. 2620 ”, F. de Silva s. n. [Wallich cat. 2620] (holotype: G-DC [G 00145656]; isotypes: BM [BM 000778231], CAL [CAL 0000071895], GZU [GZU 000255845], K [K 001116625]).

Solanum kaitisis Dunal , Prodr. [A. P. de Candolle] 13 (1): 157. 1852. Type. India. Tamil Nadu: Nilghiri, Kaithi [“ ad montis Nilgherry circa Kaitis ” – protologue], 1840, G. S. Perrottet 270 [as 230] (holotype: P [P 00058859]).

Solanum gouakai Dunal , Prodr. [A. P. de Candolle] 13 (1): 177. 1852, as “ Gouakai ”. Type. India. [Tamil Nadu]: “ Nillgherry ” [Nilgiri Hills], J. B. L. Leschenault de la Tour 311 (lectotype, designated here: P [P 00057927]).

Solanum gouakai Dunal var. angustifolium Dunal , Prodr. [A. P. de Candolle] 13 (1): 177. 1852. Type. India. [Tamil Nadu]: “ Nillgherry ” [Nilgiri Hills], J. B. L. Leschenault de la Tour 311 (lectotype, designated here: P [P 00057928]).

Solanum gouakai Dunal var. latifolium Dunal , Prodr. [A. P. de Candolle] 13 (1): 177. 1852. Type. India. [Tamil Nadu]: “ Nillgherry ” [Nilgiri Hills], J. B. L. Leschenault de la Tour 311 (lectotype, designated here: P [P 00057929]).

Bassovia wallichii Dunal , Prodr. [A. P. de Candolle] 13 (1): 409. 1852, nom. illeg. superfl. Type. Based on S. crassipetalum Wall. (“ Solanum crassipetalum, Wall. cat. 2618 ”, see discussion).

Solanum blumei Nees var. parvifolium Miq. , Fl. Ned. Ind. 2: 642. 1857, as “ parvifolia ”. Type. Indonesia. Sin. loc. “ S. Eschweilerianum ”, A. Zippelius s. n. (no herbaria cited; lectotype, designated here: L [L 0003598]; isolectotype: L [L 0003599]).

Solanum blumei Nees var. grandifolium Miq. , Fl. Ned. Ind. 2: 642. 1857, as “ grandifolia ”. Type. Indonesia. Sin. loc., T. Horsfield s. n. [1421] (neotype, designated here: BM [BM 001018973]).

Solanum bigeminatum Nees var. zeylanica C. B. Clarke , Fl. Brit. India [J. D. Hooker] 4: 231. 1883. Type. Sri Lanka. Sin. loc., [no date], G. Gardner 628 (lectotype, designated here: K [K 000923382]; isolectotypes: BM [BM 000900109], K [K 000923458], TCD [TCD 51387]).

Solanum blumei Nees var. erythrocarpum Kuntze , Revis. Gen. Pl. 2: 453. 1891. Type. India. Sikkim: sin. loc., “ 5,000 ft ”, 21 Nov 1875, C. E. O. Kuntze 6841 (lectotype, designated here: NY [00172265]).

Solanum blumei Nees var. xanthocarpum Kuntze , Revis. Gen. Pl. 2: 453. 1891. Type. Indonesia. Java: West Java, “ Wilisgebirge ” [Mount Wilis], “ 5,000 ft ”, 29 Aug 1875, C. E. O. Kuntze 5855 (lectotype, designated here: NY [00172280]).

Solanum mindanaense Elmer , Leafl. Philipp. Bot. 8: 2832. 1915. Type. Philippines. Mindanao, prov. Agusan, Cabadbaran (Mt. Urdaneta), Sep 1912, A. D. E. Elmer 13828 (syntype: E [E 00273869]).

Lycianthes pachypetala (Spreng.) Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 475. 1919. Type. Based on Solanum pachypetalon Spreng.

Lycianthes pachypetala (Spreng.) Bitter var. intermedia Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 477. 1919. Type. India. Sikkim: Tashok [as Pashok in protologue] (= Tshoka), 1,594 [1,600 in protologue] m, T. Anderson 1030 (lectotype, designated here: CAL [acc. # 316742]).

Lycianthes pachypetala (Spreng.) Bitter var. grandis Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 478. 1919. Type. India. “ Sikkim ”, Feb 1909, “ Native collector ” s. n. (holotype: WU [WU 0155118]; isotype: W [acc. # 1909-0009669]).

Lycianthes subtruncata (Dunal) Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 478. 1919. Type. Based on Solanum subtruncatum Dunal (as Solanum subtruncatum Wall. )

Lycianthes subtruncata (Dunal) Bitter var. hypolasia Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 480. 1919. Type. India. “ Assam ”, Masters s. n. (“ ex herb hort. Calcutta, hb. Bogor ”; not found at BO, no duplicates found).

Lycianthes bigeminata (Nees) Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 480. 1919. Type. Based on Solanum bigeminatum Nees

Lycianthes bigeminata (Nees) Bitter subsp. nodocalyx Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 481. 1919. Type. India. Karnataka: “ In campis aridis prope Mercara ”, R. F. Hohenacker 803 (lectotype, first stage designated by Hepper 1987, pg. 390 [as “ type ”], second stage designated here: K [K 000759401]; isolectotypes: BM [BM 000900101], FI [ FI 009598 ], M [M 00165986], SLO [n. v.]).

Lycianthes bigeminata (Nees) Bitter subsp. kaitisis (Dunal) Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 481. 1919, as ‘ Kaitisis’. Type. Based on Solanum kaitisis Dunal

Lycianthes bigeminata (Nees) Bitter var. parvifrons Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 482. 1919. Type. India. Tamil Nadu: Nilghiri Hills, Ooty [“ Ootacamund, Nilgiri Hill, prov. Madras. 7,000 ft ”], Jun 1882, D. Brandis 350 (lectotype, designated here: HBG [HBG- 511357]).

Lycianthes bigeminata (Nees) Bitter var. calycodonta Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 482. 1919. Type. India. Karnataka: Kulhutty, “ Kulhutty, Bababood ”, ca. 1,900 m, Oct 1908, A. Meebold 8878 (lectotype, designated here: CAL [acc. # 315662]).

Lycianthes bigeminata (Nees) Bitter forma gouakai (Dunal) Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 483. 1919, as “ Gouakai ”. Type. Based on Solanum gouakai Dunal

Lycianthes laevis (Dunal) Bitter var. brevipedicellata Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 487. 1919. Type. Indonesia. “ Sumatra? Poeding zimbo! hb. Bogor ” (no specimens found at BO).

Lycianthes laevis (Dunal) Bitter var. inaequidens Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 487. 1919. Type. Indonesia. [Java]: “ am Berg Tamp ” [protologue] East Java, in summo montis Taroeb [= Gunung Tarub], prov. Probolingo, 4 Jan 1845 [from P 00369020], H. Zollinger 2597 (lectotype, designated here: P [P 00379534, labelled later as 2597 bis]; isolectotypes: BM [BM 001018925], G [G 0343077, G 000357860], G-DC [G 00145616], P [P 00369013, P 00369020, as 2597 Z] )).

Lycianthes laevis (Dunal) Bitter subsp. crassipetala (Wall.) Deb. , Bot. J. Linn Soc. 76: 293. 1978. Type. Based on Solanum crassipetalum Wall.

Lycianthes laevis (Dunal) Bitter subsp. bigeminata (Nees) Deb , Bot. J. Linn. Soc. 76: 293. 1978. Type. Based on Solanum bigeminatum Nees

Lycianthes laevis (Dunal) Bitter subsp. subtruncata (Wall. ex Dunal) Deb , Bot. J. Linn. Soc. 76: 293. 1978. Type. Based on Solanum subtruncatum Dunal

Lycianthes laevis (Dunal) Bitter subsp. kaitisis (Dunal) Deb , Bot. J. Linn. Soc. 76: 294. 1978. Type. Baes on Solanum kaitisis Dunal

Lycianthes laevis (Dunal) Bitter var. gouakai (Dunal) Deb , Bot. J. Linn. Soc. 76: 294. 1978. Type. Based on Solanum gouakai Dunal

Lycianthes laevis (Dunal) subsp. kaitisis (Dunal) Bole & Ameida , J. Bombay Nat. Hist. Soc. 81 (2): 378. 1984, nom. illeg., not Lycianthes laevis subsp. kaitisis (Dunal) Deb (1978) . Type. Based on Solanum kaitisis Dunal

Lycianthes marlipoensis C. Y. Wu & S. C. Huang , Acta Phytotax. Sin. 16 (2): 78. 1978. Type. China. Yunnan: Malipo, 1,100 –1,400 m, 13 Nov 1937, G. Feng [Feng Guomei or K. M. Feng] 13227 (holotype: KUN [no acc. #]; isotype: PE [00633459]).

Lycianthes subtruncata (Wall.) Bitter var. paucicarpa C. Y. Wu & S. C. Huang , Acta Phytotax. Sin. 16 (2): 79. 1978. Type. China. Yunnan: Longling, 1,600 m, 21 De 1933, H. T Tsai [Cai Xitao] 56685 (holotype: KUN [KUN 1278691, acc. # 182540]; isotypes: A [00077124], LBG [00095865], PE [00031304],

Lycianthes crassipetala (Wall.) R. R. Mill, Edinburgh J. Bot. 57 (3): 465. 2000, as “ crassipetalum ”. Type. Based on Solanum crassipetalum Wall.

Type.

Based on Solanum laeve Dunal

Description.

Shrubs or lax subshrubs, 1–3 m tall; stems terete, glabrous or variously pubescent with whitish grey translucent simple uniseriate 2–8 - celled trichomes 0.5–1.5 mm long, these appressed or spreading, older stems glabrescent; new growth glabrous or sparsely to densely pubescent with translucent, simple, uniseriate 2–8 - celled trichomes 0.5–1.5 mm long; bark of older stems glabrescent, yellow-green to greyish brown or dark brown. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and occasionally in shape, if different in shape slightly more ovate. Leaves simple; blades of major leaves (2 –) 5–22 cm long, (1 –) 2.5–11 cm wide, elliptic to occasionally narrowly elliptic, widest in the middle or just below, concolorous, but occasionally somewhat discolorous in live plants, membranous; adaxial surfaces glabrous or sparsely to moderately evenly pubescent with simple uniseriate trichomes like those of the stems, these denser along the veins, the lamina always visible; abaxial surfaces glabrous to moderately and evenly pubescent with simple uniseriate trichomes like those of the stems, if pubescent the lamina clearly visible; principal veins 5–10 pairs, glabrous or sparsely to moderately pubescent, the tertiary venation drying darker; base attenuate and somewhat decurrent onto the petiole; margins entire or in some populations (see discussion) toothed in the distal half, in pubescent plants ciliate with simple uniseriate trichomes like those of the stems to 1 mm long, if toothed the teeth to ca. 1 cm long, ca. 1.5 cm wide, broadly triangular with acute tips; apex acute to acuminate; petiole 0.9–3.5 cm long, glabrous or pubescent with simple uniseriate trichomes like those of the stems and new growth; blades of minor leaves (1 –) 3–10 cm long, (0.5 –) 1–6 cm wide, elliptic to narrowly elliptic or sometimes almost ovate; surfaces like those of the major leaves, the minor leaves often deciduous; principal veins of minor leaves 3–6 pairs; base attenuate or sometimes only acute; margins entire or very occasionally toothed, usually entire even when the margins of major leaves are toothed; apex acute to acuminate; petiole of minor leaves 0.5–1 cm long, glabrous or sparsely pubescent like the stems. Inflorescences axillary, in fascicles or on a fleshy stub ca. 1 mm long, with (1) 4–10 (16) flowers, glabrous or with a few trichomes at the pedicel bases; pedicels at anthesis 1 – (1.5) 2–3 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading, glabrous to variously pubescent with simple uniseriate trichomes like those of the stems, articulated at the base; pedicel scars tightly packed and overlapping. Buds ellipsoid, strongly tapered and pointed, the corolla never completely included, even in small buds, strongly exserted from the calyx tube before anthesis, the calyx appendages usually more apparent in bud. Flowers 5 - merous, cosexual. Calyx tube 1.5–2.5 mm long, 2.5–3 mm wide at the mouth, obconical or an open cuplike structure, sometimes weakly 5 - ridged, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the pedicels, with 5 appendages arising at or ca. 0.5 mm below the hyaline rim, these more visible in bud, the appendages 0.5–4 mm long, ca, 0.5 mm wide, small nubs to curved and linear, usually erect or spreading and parallel to the calyx tube, glabrous or pubescent with a few simple uniseriate trichomes like those of the stems and pedicels. Corolla 1.3–2.5 (3) cm in diameter, white to variously purple, stellate, lobed 3 / 4 or nearly to the base, interpetalar tissue mostly absent but a thin edge of tissue apparent on lobe margins, the lobes 6–9 mm long, 1.5–3.5 mm wide, spreading, glabrous on both surfaces, the tips and margins densely papillate, the tips somewhat cucullate. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, glabrous; anthers 2.5–3.5 mm long, 1–2 mm wide, ellipsoid and tapering to a beak-like apex, tightly connivent, yellow, glabrous, poricidal at the tips, the pores tear-drop shaped and edged with white in dry material, lengthening to slits with age. Ovary conical, glabrous; style 5.5–10 mm long, exserted from the anther cone, glabrous; stigma prominently capitate, the surfaces minutely papillate. Fruit a globose berry, 0.8–1.3 cm in diameter, green when immature, bright red when mature, the pericarp glabrous, thin, shiny, and transparent at fruit maturity; fruiting pedicels 1.8–3 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading or erect; fruiting calyx not accrescent or expanding, but remaining a plate-like structure, often reflexed below berry in dry specimens, the appendages spreading. Seeds 16–60 per berry, 2–2.5 mm long, 2–2.5 mm wide, flattened and triangular or somewhat tear-drop shaped, pale straw-colored or yellow, the surfaces deeply pitted, the testal cells rectangular to pentagonal, the lateral walls very thick, prominent “ hairy ” appendages absent. Stone cells absent. Chromosome number not known.

Distribution

(Fig. 13 ). Lycianthes laevis is widely distributed across tropical and subtropical Asia. It occurs in Bangladesh, Bhutan, Brunei Darussalam, China, India, Indonesia, Japan, Laos, Malaysia, Myanmar ( Burma), Nepal, Philippines, Sri Lanka, Thailand and Vietnam.

Ecology and habitat.

Lycianthes laevis grows in a wide variety of forest types from montane forest to evergreen and semi-evergreen forest to dry hillsides. It is often found in shrubby undergrowth or in light gaps and along paths, from (90 –) 500 to 2,100 m elevation. Most collections are from between 900 and 2,000 m elevation.

Common names.

China. ma po hong six ian (as L. marlipoensis ), que chi hong si xian ( Zhang et al. 1994). India. Mizaram: vainan (Mrs. Parry 405); Karnataka: kandukachi (Raghavan 67850); “ Sikkim ”: kanom bi bur, knombi (both Anon. s. n.); Tamil Nadu: gouakai (Leschenault de la Tour 311). Indonesia. bu-lum ( Blume 1823, as S. violaceum ); Sumatra: ikkaoe-paet (si Boeea 10641), kajoe paga (si Boeea 7082), poga poga batoe (si Boeea 10563). Malaysia. bulung (Sundanese language, Burkill 1935, as S. blumei ), tĕrong chator (language uncertain, Burkill 1935, as S. blumei ); Selangor: bayma hutan (Temuan language, Millard 1792). Vietnam. cà ng ủ c ặp đôi (as L. bigeminata ), cà ng ủ nh ẵn ( Hop 2017). The leaves were eaten and cooked locally in northern India (Anon. s. n.). Burkill (1935) records in Java the use of leaves as a vegetable and the sweet, edible nature of the fruits; he also recorded the use of the seeds in cure of toothache but was unsure if it is this species being used for this purpose.

Preliminary conservation assessment

( IUCN 2020). EOO (15,152,928 km 2 - LC); AOO (800 km 2 - VU). Lycianthes laevis is a weedy species known from more than five localities and is very widely distributed in the region. Throughout the range it is known from protected areas (e. g., Gunung Gede Pangrango on Java, Kerinci Seblat on Sumatra, Indonesia; Western Ghats in Tamil Nadu, India). The assessment of Vulnerable (VU) based on AOO is likely due to collecting bias, but also due to the island nature of the distribution. I therefore assign it a preliminary status of Least Concern (LC).

Discussion.

Lycianthes laevis is a widespread species with considerable regional variation. It can be recognised by its calyx with 5 appendages, these are usually quite short, but in southern India and Indonesia can be filiform and up to several mm long (Figs 3 B , 4 C ). The calyx appendages always appear longer in bud. A collection from Mount Makiling in the Philippines (Sulit 8499) has a single flower with 7 appendages, but this is the only example of L. laevis I have seen with more than 5 calyx appendages. Lycianthes laevis can be distinguished from other species in the area with 5 calyx appendages (e. g., L. banahaensis , L. bimensis ) by its shrubby versus tree habit and in its cosexual flowers with membranous (versus thick and somewhat fleshy) corolla lobes. Like those of L. schizocalyx , the pedicels in fruit can be elongate, in both species reaching to 3 cm long. The berries of L. laevis are marginally smaller (0.8–1.3 cm in diameter versus 1–1.3 cm in diameter) but the number of calyx appendages (10 in L. schizocalyx versus 5 in L. laevis ) is distinct.

Like L. biflora , pubescence density in L. laevis is extremely variable. Many specimens are nearly glabrous (including the type of S. laeve ) whereas others (e. g., Brandis 350, the type of var. parvifrons ) are densely and uniformly pubescent with simple uniseriate trichomes. I have never seen branched trichomes on plants of L. laevis , unlike in L. biflora , where they are common. Similarly, pubescent anthers have not been seen in L. laevis .

Flowers from some (but not all) populations in the north of the range in Nepal and northwestern India tend to be fleshier and somewhat larger than those from southern India and Sri Lanka and have been called L. crassipetalum (e. g., Mill 2001) When material is examined from across the area, however, this variation appears to be continuous or merely populational. This continuity has been noted by others, on a specimen of L. laevis in CAL (acc. # 315562, Hooker & Thomson s. n. from “ Khasia ”) an annotation by J. S. Gamble reads “ intermediate between S. subtruncatum and S. crassipetalum ” suggesting he too thought the variation in this species was continuous and these taxa were not distinct. Collections from Mount Salak in Java ( Indonesia) also have large flowers (e. g., Kuntze 4575, Kurz 462); these may be virally infected but further field investigation is needed.

Leaves in some populations of L. laevis are shallowly lobed in the distal third with irregular, broad teeth arising from the vein endings (see Fig. 12 ); these have been called S. gouakai in southern India. Some local floras in southern India ( Henry et al. 1987) recognise these leaf variants at the subspecific and varietal level in L. laevis using Dunal’s (1852) epithets, whereas others recognise a single taxon ( L. bigeminata, Nair and Nayar 1987 ; L. laevis, Mohanan and Henry 1994 ). These leaf types occur most frequently in southern India (e. g., Bourne 2088) and Sri Lanka (Walker 65), but I have also seen collections with lobed leaves from western Myanmar (e. g., Fujikawa et al. 094293). Toothed and entire leaves occur on the same plant as is common in the Morelloid clade of Solanum ( Knapp et al. 2023) , in many species of spiny solanums ( McClelland et al. 2020; Aubriot and Knapp 2022) and in Lycianthes series Tricolores Bitter of Mexico ( Dean et al. 2017). Sometimes juvenile leaves in Solanum can be more lobed or toothed than leaves on mature branches ( Roe 1966); this may be the case here, but it is not clear. Field surveys of these populations in Tamil Nadu would be very informative.

Lycianthes laevis has a wide distribution, overlapping with most of the other Lycianthes species in Asia. It is more westerly than L. biflora or L. schizocalyx but does co-occur with both those taxa. Hybridisation is often cited as the reason for wide ranges of intraspecific variation, but the calyx appendage character is constant across the range of L. laevis ; populations in sympatry should be studied in the field. One such, where L. laevis and L. schizocalyx co-occur and have been mixed in collections is on Mount Santo Tomas in the Philippines (e. g., Williams 1334 and 1334 [A]). At first glance these plants look very similar, but closer examination show that not only do the calyx appendages differ (5 versus 10) but the pubescence also differs. The element of this gathering that represents L. schizocalyx was labelled with the same number (as a presumed duplicate) in pencil, possibly long after collection (see discussion of L. schizocalyx ).

Solanum laeve was described by Dunal (1814) based only on a collection seen by him in Paris from Java labelled “ Bondolene ” which Dunal took to be the local name. A specimen collected by J. B. Leschenault de la Tour in Java labelled “ Goudolene ” (misread by Dunal as Bondolene) does not bear a label in Dunal’s hand but corresponds to the protologue in being in fruit and is here selected as the lectotype (Leschenault de la Tour 683, P 000578637). In the Prodromus ( Dunal 1852), the collector of the type is mentioned.

Blume (1823) published the homonym S. violaceum Blume (not S. violaceum Ortega , an Asian spiny solanum; Aubriot and Knapp 2022) without citing a locality or herbarium. A sheet in Leiden collected by Blume in Java has an annotation “ Solanum Blumii Nees ” and a much later annotation as S. violaceum . I select this sheet (L 0003600) here as a neotype because there is no evidence that it was used by Blume in the description of S. violaceum , which could have been described from living material. Blume and Nees van Esenbeck were correspondents ( van Steenis 1989) and the replacement name for the illegitimate S. violaceum Blume was coined by Nees van Esenbeck in Blume’s honour and published with a description by Blume ( Blume 1826) as “ Solanum blumii Nees ab Esenb. ”. The annotation of “ Solanum Blumii Nees ” on the lectotype may be in Nees van Esenbeck’s hand.

Wallich’s designation “ Solanum membranaceum ” comprised three elements (Wallich 1828–1849: 80) was validated by Nees van Esenbeck (1834) where one of these (2625 B from “ herb. Wight ”) was excluded and referred to S. laeve . Nees van Esenbeck (1834) indicated that only the specimens labelled 2625 A and 2625 C were part of his S. membranaceum . In the Wallich herbarium at Kew there are three specimens labelled 2625 (K 001116640, K 001116641, K 0011166420) none of which bears the annotation corresponding to the catalogue entry. One of these is unambiguously labelled 2625 A (K 001116640) and has an annotation in Nees van Esenbeck’s hand and it also has the catalogue entry affixed to the sheet; I have selected this sheet as the lectotype of S. membranaceum . All three of these specimens are conspecific.

Nees van Esenbeck (1834) attributed the name S. neesianum to Wallich, but there is no mention of that name in Wallich’s catalogue (M. Watson, pers. comm. 7 Feb 2024). The specimen cited is “ Wall. Catal. Suppl. n. 248 / Crescit in montibus Silhet Guil. Gomez ” suggesting the collector was William Gomez who had collected in the area and whose collections are listed on page 248 of the catalogue (https://wallich.rbge.org.uk/index.php?section=pages&id=1279). A specimen in Nees van Esenbeck’s personal herbarium now held in GZU is clearly original material (GZU 000255706); it is labelled with the name, has an annotation “ 248 ” and a locality “ Mt Sylhet FD ”. There is clearly a mistake in the published protologue as to collector; “ FD ” indicates the specimen was collected by Francis de Silva, one of Wallich’s collectors in the Silhet region. In the absence of any more specific material, I have designated this sheet collected by Francis de Silva in GZU (GZU 000255706) as the lectotype. I suspect this specimen belongs to the set “ Wallich Cat. 2620 ”, collected at the same place by the same person (“ Mont. Sillet, FD ”) as the type of S. subtruncatum .

Dunal (1852) validated Wallich’s designation “ Solanum subtruncatum ” citing the specimen “ Wallich cat. 2620 ” from G-DC; Deb’s apparent lectotypification (by the citation of the collection Wallich cat. 2620 in CAL) of S. subtruncatum was not necessary. In describing S. kaitisis Dunal (1852) cited a collection Perrottet 230 from Paris; the sheet labelled Perrottet 230 (P 00248294) corresponds to Uvaria narum (Dunal) Blume ( Annonaceae ). Dunal also worked with Annonaceae ( Dunal 1817) . I am treating the collection number 230 in the protologue as an error to be corrected to Perrottet 270, a sheet (P 00058859) that is annotated by Dunal as Solanum kaitisis . Dunal (1852) described S. gouakai and its two varieties ( var. angustifolium and var. latifolium ) from a single collection made by J. B. Leschenault de la Tour in southern India (Leschenault de la Tour 311), that he saw in Paris. I have used the three sheets of this collection In P as lectotypes for the species and each of the two varieties; that chosen as the lectotype for var. angustifolium (P 0057928) is annotated “ α angustifolium ” by Dunal, that chosen for var. latifolium (P 00057929) is annotated “ β latifolium ”) by Dunal and that for the species is only annotated with the specific epithet (P 00057927). The three sheets vary somewhat in leaf width but clearly are from the same gathering.

Bassovia wallichii is an illegitimate superfluous name for S. crassipetalum Wall. in that Dunal (1852) based the name on “ Solanum crassipetalum Wallich cat. 2618 in G-DC ” ( Turland et al. 2018: Art. 52.2 (e )).

Of the two varieties of S. blumei described by Miquel (1864) I have found original material for only S. blumei var. parvifolium . Two sheets of the specimen “ S. Eschweilerianum Zipp. herb. ” mentioned in the protologue are held in Leiden, one (L 0003598) is annotated by Miquel and is a better specimen and I have designated this the lectotype. The other is an isolectotype (L 0003599). I have found no unambiguous original material for var. grandifolium . Among material cited was something collected by “ Horsf. ” a reference to Thomas Horsfield who collected in Java ( van Steenis-Kruseman 1985); a specimen of Horsfield’s corresponding to the protologue (BM 001018973) is here selected as the neotype for S. blumei var. grandifolium .

Clarke (1883) cited collections from “ Gardner, Wight, Thwaites ” in the protologue of S. bigeminatum var. zeylanica but with no mention of collection numbers or herbaria. I have selected the sheet of Gardner 628 from the Hooker herbarium at Kew (K 000923382) as the lectotype for var. zeylanica , it is annotated as var. zeylanica in Clarke’s hand and has flowers and fruits. Other duplicates at Kew are less complete. Hepper (1987) mentioned a Gardner collection in his description of var. zeylanica , but without citing a herbarium or number.

Kuntze (1891) described two varieties for S. blumei based on fruit colour from his own collections. In the very short protologue for var. erythrocarpum he states the distribution as Java and Sikkim; I have selected a specimen from Kuntze’s herbarium ( NY barcode 00172265) now at NY annotated by Kuntze from Sikkim as lectotype for this variety. For var. xanthocarpum he states the distribution as “ Wilis ” (= Wilisgebirge, or Mount Wilis in West Java); here a single specimen in NY from Kuntze’s herbarium (barcode 00172280) from the correct locality and annotated by Kuntze is selected as the lectotype for the variety.

Elmer (1915) cited a single collection (Elmer 13828) in the protologue of S. mindanaense but no herbarium; specimens upon which Elmer’s names were based were in his private collection at the Philippine National Herbarium (then the Philippine Bureau of Agriculture under United States jurisdiction) that was destroyed by fire during the Japanese occupation of the Philippines in the Second World War just a day before the liberation of Manila ( Schultes 1957). The only duplicate I have seen is a sterile specimen at Edinburgh (E 00273869); intensive searches at other herbaria where Elmer sent collections have not yet been successful. Elmer suggested his plant differed from S. crassipetalum (= L. laevis ) in “ a number of minor characters ” ( Elmer 1915: 2833). I place S. mindanaense in synonymy here based on the description but will continue to search for a fertile duplicate with which to lectotypify this name.

Bitter (1919) described three infraspecific taxa for L. bigeminata from India; subsp. nodocalyx , var. parvifrons and var. calycodonta . Hepper (1987) lectotypified L. bigeminata subsp. nodocalyx with the collection Hohenacker 803 but in citing both K and SLO necessitated a second-step lectotypification here. I have selected the specimen at Kew (K 00759401) as the second-step lectotype. For L. bigeminata var. parvifrons Bitter (1919) cited many different collections (Wight 2021, 2025, 1569; Hooker & Thomson s. n; Brandis 350, s. n.). The collections of Robert Wight are widely distributed, but whether they are real duplicates is not clear ( Noltie 2005). I have therefore selected Brandis 350 in Hamburg (HBG- 511357), cited as from “ herb. Hamb. ” and annotated by Bitter, as the lectotype for var. parvifrons . Many collections were also cited in the protologue of var. calcodonta mostly from “ herb. Berol. ” (Hohenacker 1415; Thomson s. n.; Meebold 8878, 9456, 11696, 12346; Saulière 88), plus a sheet of Wight 2012 from Copenhagen. I have selected a duplicate of Meebold 8878 held in Calcutta (CAL acc. # 315662) as the lectotype for var. calycodonta ; it is a well-preserved specimen and is held in an Indian herbarium.

As part of his concept of L. laevis Bitter (1919) published several infraspecific taxa. For L. laevis var brevipedicellata he cited a specimen from Bogor “ Sumatra Poeding zimbo! hb. Bogor ” of which no material has been found despite an extensive search in BO and in Leiden. I have not neotypified this name here, as material may be found in the future. The protologue of L. laevis subsp. inaequidens includes citations of duplicates of Zollinger 2597 from Bogor and Paris (although he cites the Berlin specimen of Zollinger 2597 under L. laevis ) together with a number of collections seen from “ hb. Bog. ” Searches at BO have turned up only Koorders 19891 and 31886 of all the collections cited; both are scrappy specimens with little reproductive material. The sheet at Paris labelled Zollinger 2597 bis with an undated annotation slip by Bitter (P 00379534) is therefore selected as the lectotype for var. inaequidens . All the sheets of Zollinger 2597 I have seen appear to be from the same gathering (although only P 000369020 from the Drake herbarium in P, labelled “ 2597 Z ” in an unknown hand, has the original locality label) and I consider S. zollingeri var. multiflorum and L. laevis subsp. inaequidens to be homotypic.

Figures

Figure 3. Calyx appendage variation of Lycianthes in Asia A L. biflora with 10 equal calyx appendages (Nuraliev s. n, Vietnam [no voucher]) B L. laevis with 5 calyx appendages (Mustaqim s. n., Indonesia; DOL 207886) C L. parasitica with small, nub-like calyx appendages (Pelser & Barcelona s. n., Philippines; DOL 84744) D L. oliveriana with thick, warty calyces with no appendages (Damas et al. SAJ- 1050, Papua New Guinea) E L. schizocalyx with long flower pedicels and 10 unequal calyx appendages (Heyer s. n., Philippines; DOL 151906) F L. shunningensis with 10 strongly reflexed calyx appendages (Averyanov 887, Laos). Photograph credits: A Maxim Nuraliev B Wendy Mustaqim C Pieter B. Pelser & Julie F. Barcelona D Shelley James E Philippe Heyer F Leonid Averyanov. DOL numbers are identifiers from PhytoImages (https://phytoimages.siu.edu). Details of collecting localities can be found in Suppl. material 2.

Figure 4. Flowers of Lycianthes in Asia A L. banahaensis with tetramerous yellow flowers, note the stigma sitting almost within the anther tube (Rule s. n., Philippines; DOL 134710) B L. biflora with membranous corolla lobes and a minutely capitate stigma (Knapp et al. 10106, China) C L. laevis flower with green blotches at the base of the corolla lobes (Mustaqim s. n., Indonesia; DOL 207828) D L. parasitica flower with spreading corolla lobes and capitate stigma (Tandang s. n., Philippines; DOL 76576597) E L. rantonnetii rotate corolla with orange, slightly curved anthers (Stevenson s. n., cultivated in Mexico; DOL 183568) F L. shunningensis with very short reflexed calyx appendages and cucullate corolla lobe tips (Nuraliev et al. 2810, Vietnam). Photograph credits: A Greg Rule B Sandra Knapp C Wendy Mustaqim D Danilo Tandang; E Dennis Stevenson F Maxim Nuraliev. DOL numbers are identifiers from PhytoImages (https://phytoimages.siu.edu). Details of collecting localities can be found in Suppl. material 2.

Figure 12. Lycianthes laevis (Dunal) Bitter herbarium specimen. India. Tamil Nadu, Beddome 5486 (BM 00900116). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

Figure 13. Distribution of Lycianthes laevis.