Lycianthes biflora (Lour.) Bitter

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

2. Lycianthes biflora (Lour.) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 461. 1919.

Figs 2 B , 3 A , 4 B , 5 B, C , 8

Solanum biflorum Lour. , Fl. Cochinch. 129. 1790. Type. China. Guangdong: Guangzhou [“ Pakwan supra Cantonem ”], Jul 1869, H. F. Hance 2128 (neotype, designated by Hul and Dy Phon 2014, pg. 57: P [P 00058796]; isoneotypes: P [P 00058797]).

Solanum decemdentatum Roxb. ex Wall. , Fl. Ind. [ed. Carey & Wallich] 2: 247. 1824. Type. Singapore, Sep 1822, N. Wallich cat. 2614 [A] (lectotype, designated here: K-W [K 001116583]; probable isolectotypes: BM [BM 000900122], GZU [GZU 000255527]).

Solanum denticulatum Blume , Bijdr. Fl. Ned. Ind. 13: 697. 1826. Type. Indonesia. Java [West Java]: Gagar Buiteng [in montosis Tjerimai et in cacuminae montis Gegar Bentin ex protologue], C. L. Blume s. n. (lectotype, designated here: L [L 0003595, L-bottom-most stem]).

Solanum mollissimum Blume , Bijdr. Fl. Ned. Ind. 13: 697. 1826. Type. No locality cited in protologue [ Indonesia, Jav?], “ in montanis ”, C. L. Blume s. n. (lectotype, designated here: L [L 0003586]).

Solanum decemfidum Roxb. ex Nees , Trans. Linn. Soc. London 17 (1): 43. 1834, nom. illeg. superfl. Type. Based on Solanum decemdentatum Roxb. ex Wall. (cited in synonymy)

Solanum macrodon Wall. ex Nees , Trans. Linn. Soc. London 17 (1): 43. 1834. Type. Bangladesh. [ Chittagong: near Companiganj] [“ Pundua, Wallich 2621, coll. Silva 1466 ”], F. de Silva s. n. [Wallich cat. 2621] (lectotype, designated by Deb 1978, pg. 293 [as “ type collection ”]: CAL [CAL 0000017896]; isolectotypes: BM [BM 001018882], E [E 00273887], GZU [GZU 000255675], K [K 000923271], K-W [K 001116626]).

Solanum zollingeri Dunal , Prodr. [A. P. de Candolle] 13 (1): 176. 1852. Type. Indonesia. Java: [West Java], Tjidurian, Sep 1842, H. Zollinger 723 (lectotype, designated here: G-DC [G 00145615]; isolectotypes: BM [BM 000778207], G [G 00343041, G 00357862], K [K 000759388], P [P 00369008]).

Solanum javanicum Dunal , Prodr. [A. P. de Candolle] 13 (1): 176. 1852. Type. Indonesia. Java: sin. loc., H. Zollinger 1981 (holotype: G-DC [G 00145623]; isotypes: G [G 00301655, G 00343320], W [acc. # 1889-0149948, acc. # 1889-0149949]).

Solanum osbeckii Dunal , Prodr. [A. P. de Candolle] 13 (1): 179. 1852. Type. Indonesia. Java: Prince’s Island, 4–14 Jan 1771, J. Banks & D. Solander s. n. (lectotype, designated here: BM [BM 000778107]).

Solanum osbeckii Dunal var. stauntonii Dunal , Prodr. [A. P. de Candolle] 13 (1): 179. 1852, as “ Stauntoni ”. Type. China. sin. loc., [s. d.], G. Staunton s. n. (lectotype, designated here: W [acc. # 0003086]).

Solanum calleryanum Dunal , Prodr. [A. P. de Candolle] 13 (1): 178. 1852. Type. China. sin. loc. [protologue – “ circa Macao ”], C. Gaudichaud 95 (lectotype, designated here: G-DC [G 00145658]; isolectotype: G [G 00415758]).

Solanum denticulatum Blume var. lanceolatum Miq. , Fl. Ned. Ind. 2: 644. 1857. Type. Indonesia. Java: [protologue – “ bij Lamadjang en Tenga ”], H. Zollinger s. n. (no herbarium cited; lectotype, designated here: P [P 00369015]; isolectotype: P [P 00369014]).

Solanum biflorum Lour. var. mollissimum (Blume) Kuntze , Revis. Gen. Pl. 2: 453. 1891. Type. Based on Solanum mollissimum Blume

Solanum biflorum Lour. forma pilosa Kuntze , Revis. Gen. Pl. 2: 453. 1891, as “ Solanum biflorum var. mollisimum forma pilosa ” Type. Indonesia. Java: East Java, “ Bromo 4,000 ” [= Mount Bromo], ca. 1220 m, O. Kuntze 6007 (lectotype, designated here: NY [00172277]).

Lycianthes biflora (Lour.) Bitter var. sparsiloba Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 464. 1919. Type. Indonesia. Java: “ G. Tjibodas, Tjampen ” [Cibodas], 200 m, C. A. Backer 4658 (holotype: BO [acc. # BO- 1911410]).

Lycianthes biflora (Lour.) Bitter var. mollissima (Blume) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 465. 1919. Type. Based on Solanum mollissimum Blume

Lycianthes biflora (Lour.) Bitter var. grandifolia Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 466. 1919. Type. Myanmar ( Burma). “ Papun bei Moulmein ”, A. Meebold 17068 (lectotype, designated here: CAL [acc. # 315674]; isolectotype: WRSL [destroyed]).

Lycianthes biflora (Lour.) Bitter var. subtusochracea Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 466. 1919. Type. China. Yunnan: S of Red River, A. Henry 13652 (holotype: B [destroyed]; lectotype, designated here: K [K 000759409]).

Lycianthes biflora (Lour.) Bitter subsp. hupehensis Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 466. 1919. Type. China. Hubei: sin. loc., “ Faber in Henry’s Coll. from Centr. China ” A. Henry [Faber] 4304 (lectotype, designated here: W [acc. # 1886-0001758]; isolectotypes: B [destroyed], BM [BM 001018849], CAL [acc. # 316326], E [E 00426461], GH, K [K 000759407], P [P 00058798]).

Lycianthes biflora (Lour.) Bitter var. velutinella Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 467. 1919. Type. Indonesia. Sulawesi: [Sulawesi Utara] “ prov. Minahassa, Urwald bei Biwak Penamarangan bei Kajoevatoe [protologue] ”, S. H. Koorders 18041 β (no herbarium cited; lectotype, designated here: BO [acc. # BO- 4324391]; isolectotype: L [L. 2859422]).

Lycianthes biflora (Lour.) Bitter subsp. elongatidens Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 468. 1919. Type. Indonesia. Sulawesi: [Sulawesi Utara] “ Prov. Minahassa, Tomahon [Tomohon], 800 m ”, S. H. Koorders 18038 β (lectotype, designated here: BO [acc. # BO- 1990064]; isolectotype: L [L. 2859397]).

Lycianthes macrodon (Wall. ex Nees) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 468. 1919. Type. Based on Solanum macrodon Wall. ex Nees

Lycianthes macrodon (Wall. ex Nees) Bitter var. mollitersetosa Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 470. 1919. Type. India. Sikkim: sin. loc., T. Anderson 303 (lectotype, designated here: CAL [acc. # 315720]; isolectotype: B [destroyed]).

Lycianthes macrodon (Wall. ex Nees) Bitter var. sikkimensis Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 470. 1919. Type. India. Sikkim: Toong, ca. 1,600 m, A. Meebold 15768 (holotype: WRSL [destroyed], no duplicates found).

Lycianthes macrodon (Wall. ex Nees) Bitter var. manipurensis Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 470. 1919. Type. India. Manipur: Ukrul Nagab, ca. 1,900 m, Dec 1907, A. Meebold 6906 (holotype: WRSL [destroyed]; lectotype, designated here: CAL [no acc. #]).

Lycianthes denticulata (Blume) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 473. 1919. Type. Based on Solanum denticulatum Blume

Lycianthes biflora (Lour.) Bitter subsp. yunnanensis Bitter , Repert. Spec. Nov. Regni Veg. 18: 319. 1922. Type. China. Yunnan: Mengtze, 1,570 m, A. Henry 9160 (holotype: B [destroyed]; lectotype, designated here: K [K 000922382]; isolectotypes: A [00619929], E [E 00426448], US [02840689, acc. # 456785]).

Lycianthes macrodon (Wall. ex Nees) Bitter var. longifrons Bitter , Repert. Spec. Nov. Regni Veg. 18: 319. 1922. Type. India. Meghalaya: Birch Hill near Darjeeling, 2,200 m, 7 Nov 1896, H. Hallier s. n. (holotype: M [M- 0166005]; isotype: CAL [no acc. #]).

Solanum boninensis Nakai ex Tuyama , Bot. Mag. (Tokyo) 50: 132, f. 27. 1936. Type. Japan. Ryuku Islands: Ins. Titizima [protologue, on label as “ Bonin: Chichigima ”], 8 Jul 1920, T. Nakai s. n. (holotype: TI [TI 00043095]).

Solanum biflorum Lour. var. glabrum Koidz. ex Hatus. , J. Geobot. 17: 49. 1969. Type. Japan. Ryuku Islands: along the Kuira River, Isl. Iriomote, 6 Aug 1968, T. Narita s. n. (no herbarium cited in protologue; lectotype, designated here: KAG [acc. # 163736]; isolectotype: US [02840676, no accession number]).

Solanum biflorum Lour. var. kotoense Y. C. Liu & C. H. Ou , Quart. J. Chin. Forest. 7 (4): 151. 1974, as ‘ kotoensis ’. Type. Taiwan. “ Botel Tobago, low altitudes ” [= Orchid Island], 7 Aug 1932, T. Sato s. n. (holotype: TAI [n. v.]).

Lycianthes biflora (Lour.) Bitter subsp. macrodon (Nees) Deb , Bot. J. Linn. Soc. 76: 293. 1978. Type. Based on Solanum macrodon Nees

Lycianthes hupehensis (Bitter) C. Y. Wu & S. C. Huang , Acta Phytotax. Sin. 16 (2): 77. 1978. Type. Based on Lycianthes biflora Lour. subsp. hupehensis Bitter

Lycianthes yunnanensis (Bitter) C. Y. Wu & S. C. Huang , Acta Phytotax. Sin. 16 (2): 77. 1978. Type. Based on Lycianthes biflora Lour. subsp. yunnanensis Bitter

Lycianthes laevis (Dunal) Bitter var. kotoensis (Y. C. Liu & C. H. Ou) T. Yamaz. , Fl. Japan 193. 1993. Type. Based on Solanum biflorum Lour. var. kotoense Y. C. Liu & C. H. Ou

Solanum chingchunense S. S. Ying , New Taxa New Names 6 (2): 311. 2023, as “ chingchunensis ”. Type. Taiwan. Hsinchu County: Wufeng Township, Chingchun, 550 m, 16 May 2023, S. S. Ying s. n. (holotype: NTUF [acc. # 112-073]).

Type.

Based on Solanum biflorum Lour.

Description.

Small shrubs or herbs, 0.5–1.5 m tall, sometimes described as a vine or scrambler; stems terete, sparsely to densely pubescent with a mixture of transparent simple and / or forked or dendritic 3–10 - celled uniseriate trichomes to 2 mm long, the dendritic trichomes antler-like or merely forked; new growth sparsely to densely pubescent with simple and dendritic trichomes like those of the stems, in plants with sparse pubescence the trichomes mostly confined to the leaf veins; bark of older stems pale brown, somewhat glabrescent. Sympodial units difoliate, the leaves geminate, the leaves of pair usually differing in size but usually not in shape. Leaves simple; blades of major leaves 3–17 cm long, 2–8 cm wide, ovate to elliptic or occasionally narrowly elliptic (e. g., Chai 337360) or broadly ovate (e. g., Bodinier 799), usually widest in the lower half but occasionally near the middle, somewhat discolorous or occasionally strikingly so (plants previously identified as var. subtusochracea ), membranous, the leaves usually larger on lower branches; adaxial surfaces almost glabrous to evenly and moderately pubescent with transparent mixed simple and dendritic trichomes like those of the stems, these much denser along the veins; abaxial surfaces sparsely to moderately pubescent with the same trichomes as those of the adaxial surfaces, but the pubescence denser, markedly so on more distal leaves; principal veins 4–6 pairs, sparsely to densely pubescent, often drying yellow on both surfaces; base attenuate or occasionally abruptly attenuate, markedly decurrent onto the petiole; margins entire, markedly ciliate with transparent mixed simple and / or dendritic trichomes like those of the leaf surfaces; apex abruptly acuminate or acuminate; petiole 0.5–2.5 cm long, winged from the decurrent leaf bases, sparsely to densely pubescent like the stems and leaves; blades of minor leaves 1–5 cm long, 0.9–3 cm wide, shape, texture and pubescence like that of the majors; base attenuate onto the petiole; margins entire, ciliate; apex abruptly acuminate or acuminate; petiole 0.4–1 (2.5) cm long, pubescent like the stems and leaves. Inflorescences axillary fascicles of (1) 2–6 flowers, usually only one open at a time, sparsely to densely pubescent with mixed simple and dendritic trichomes like the stems; pedicels at anthesis 0.9–1.2 cm long, ca. 0.75 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, nodding and the flowers borne below the leaves, sparsely to densely pubescent with transparent mixed simple and / or dendritic uniseriate like those of the stems and leaves, articulated at the base; pedicel scars tightly packed in the leaf axils. Buds elliptic, the corolla strongly exserted from the calyx tube before anthesis, the calyx appendages clasping the buds. Flowers 5 - merous, apparently all cosexual. Calyx tube 2–3 mm long, 2.5–3.5 mm in diameter, obconical to openly cup-shaped, sparsely to densely pubescent like the stems and pedicels, with 10 (12) linear awl-like appendages 1–9.5 mm long at anthesis, variable in length between plants and populations, the appendages emerging at the rim, pubescent like the rest of the calyx. Corolla 1.4–1.8 cm in diameter, white or lavender or purple, with a green central area, sometimes appearing as two green dots at the base of each lobe, stellate, lobed nearly to the base, interpetalar tissue absent or a thin edge on the lobes, the lobes 4–6 mm long, ca. 3 mm wide, spreading or slightly reflexed, membranous, adaxially glabrous, densely puberulent / papillate in the distal half abaxially, the tips and margins densely papillate. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, glabrous; anthers 2.5–4.5 mm long, 1–1.5 mm wide, ellipsoid, the tips slightly pointed to a sharp beak to 0.5 mm long (the tips often drying paler than the rest of the anther), yellow, glabrous or variously pubescent with simple uniseriate trichomes to 0.2 mm long, these along the thecae edges or over the entire anther (e. g., Cuong 417), poricidal at the tips, the pores tear-drop shaped, distally directed, lengthening to slits with age. Ovary conical, glabrous; style 4.5–6 mm long, straight, glabrous; stigma small-capitate, the surfaces minutely papillate. Fruit a globose berry, 1–1.5 cm in diameter, bright red when ripe, changing from green to orange to red through development, the pericarp glabrous, thin, shiny and transparent; fruiting pedicels 1–1.8 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, green, not markedly woody, erect with the fruits borne above the leaves; fruiting calyx a flat plate beneath the fruit, the calyx appendages elongating to ca. 2 times their length in flower, spreading and forming a star under the berry. Seeds 100 + per berry, 1.5–2 mm long, 1–1.5 mm wide, flattened and prismatically irregularly tear-drop shaped, straw-yellow, the surfaces deeply pitted, the testal cells sinuate in outline, prominent “ hairy ” testal cell walls absent. Stone cells absent. Chromosome number: n = 24 ( Symon 1985; based on Kairo & Symon 10652 from Papua New Guinea).

Distribution

(Fig. 9 ). Lycianthes biflora is widely distributed and very common; it occurs in Australia [ Christmas Island only], Bangladesh, Bhutan, Brunei Darussalam, Cambodia, China, India, Indonesia, Japan, Laos, Malaysia, Myanmar ( Burma), Nepal, Papua New Guinea, Philippines, Singapore, Taiwan, Thailand, Timor Leste and Vietnam.

Ecology and habitat.

Lycianthes biflora grows in a wide variety of mostly disturbed habitats, in evergreen broadleaf or semi-evergreen broadleaf forests or in dry areas along roads and paths, from sea level to 2,300 m elevation. Most collections in the southern part of its range are from below 2,000 m elevation, but in the Himalaya it can grow at higher elevations.

Common names.

China. da chi hong si xian (as L. macrodon ), dian hong si xian (as. L. yunnanensis ), e hong si xian (as. L. hupehensis ), hong si xian ( Zhang et al. 1994); Guangdong: chicken eye clear, kai ngan ts’ing (Lingnan University Herbarium 12648); Hainan: shan nau kwo (McClure 9600). India. malum pavadam (Wight 1569); Sikkim: kolimbe (Lepcha language, Srivastava 10301). Indonesia. Sulawesi: lewa-lewsa-koelo, kamoeti (Totemboan language, Koorders 1898, as S. denticulatum ), kalekamates (Tonsawant language, Koorders 1898, as. S. denticulatum ), tahakkok, tohokok (Tombulu language, Koorders 18308), tampai (Johansson 143), wewelesan, wewelesan-in-taloen (Tombolu language, Koorders 1898, as S. denticulatum ); Sumatra: akikaoe poga (si Boeea 10780), so haboe haboe (si Boeea 10242), si marpoga poga (si Boeea 19071), poga oetan (si Boeea 10418), poga (si Boeea 6619), poga poga (si Boeea 8469, 8765, 9034). Japan. Ryuku Islands: mejiro-hondzuki (Amano 6961). Malaysia. Sabah: lapak puru (Dusun language, Bakia 612), tensisiah (Dusun language, Giking 236), tutan (Amin 93928, Bundu Tutan language, Aban Gibot 66986), tutan geuton (Dusun language, Surunda 100), tutan puru (Dusun language, Soibeh 789); Sarawak: kerid pait (Kelabit language, Christensen 71), mata antu sebayan (Iban language, Ashton 19162), munting eja (Yii 70379). Nepal. banbi (Mananduar 6805). Philippines. Mindanao: beganbagan (Manobo language, Elmer 13881). Thailand. Central: ma wenj pa (Put Phraisurind 156). Vietnam. cà ng ủ cu ống to (as L. macrodon ), cà ng ủ ( Hop 2017).

Preliminary conservation assessment

( IUCN 2020). EOO (26,990,176 km 2 - LC); AOO (1,684 km 2 - VU). Lycianthes biflora 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., Halimun National Park on Java, Lore Lindu National Park on Sulawesi, Indonesia; Iriomote National Park on Okinawa, Japan). 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 biflora is a name that has been widely used for shrubby Lycianthes in tropical Asia. As discussed in the Introduction, Bitter (1919) characterised it as an “ Overall species ” (“ Geasamart ”) and split it into a multitude of infraspecific taxa, all of which are here recognised in synonymy. Most of these were distinguished based on geography or on pubescence variation, but examination of specimens from across the range show no real boundaries or diagnostic differences, something Bitter himself recognised was likely to occur with more material ( Bitter 1919: 461).

Lycianthes biflora is very similar to L. schizocalyx , which also has 10 linear to subulate calyx appendages (Fig. 3 A, E ). The appendages in L. biflora are all the same length in a single flower and arise from the edge of the calyx tube (Fig. 3 A ), leaving no hyaline rim above their emergence. The appendages of L. schizocalyx , in contrast are usually of variable length in a flower and at least some of them arise from below the edge of the tube, leaving a hyaline rim. The calyx appendages in L. biflora vary in length in individual plants and populations across the range (see Fig. 5 B, C ); plants from northern India with longer calyx appendages have been called L. macrodon . The flowering pedicels of L. biflora are 0.9–1.2 cm long, whereas those of L. schizocalyx are (1) 2–3 cm long; individual specimens can sometimes be difficult to distinguish using this single character. It may also be that the species hybridise where they co-occur (i. e., places like Mount Kinabalu in Sabah, Malaysia). Lycianthes shunningensis is also a shrubby species with 10 calyx appendages, but these arise from well below the edge of the calyx tube and are distinctly turned downwards, especially in fruit (Fig. 5 F ). Lycianthes biflora collections made from young plants can be difficult to distinguish from L. lysimachioides , but that species is a creeping or scandent herb, has a single flower per fascicle (only rarely more) and larger flowers (corolla 1.8–3 cm versus 1.4–1.8 cm in diameter in L. biflora ).

Pubescence in L. biflora is remarkably variable; individual plants range from almost completely glabrous or with a few scattered simple uniseriate trichomes to densely pubescent. Densely pubescent individuals are found throughout the species range and have been called var. mollissima . Trichomes are either simple and uniseriate, often as many as 10 to 15 cells long, or variously branched. Branched trichomes in L. biflora usually bear only a few 1–2 - celled branches but can be antler-like with many branches or merely forked (with a single branch). Density of pubescence is not a predictor of whether or not trichomes will be branched, and often a given plant will only have a few branched or forked trichomes with very short branches amidst denser simple uniseriate pubescence. Notations of glandular trichomes on some herbarium specimens are mistaken identification of fungal sporing bodies; I have seen no glandular trichomes in the specimens I have examined.

The anthers of some specimens of L. biflora are pubescent, with simple uniseriate trichomes along the dehiscence zone (e. g., Forrest 24674 from Yunnan; Chun 41918 from Hong Kong in China) or over the entire anther surface (e. g., Cuong 417 from Vietnam). These hairs are reminiscent to those found in members of the Tomato clade of Solanum ( Peralta et al. 2008) , that hold the anthers together in a connivent structure that aids pollination. This would be an interesting species in which to investigate the genetic basis of the anther cone as has been done in Solanum ( Glover et al. 2004) .

In India, very pubescent individuals of L. laevis have often been annotated as L. biflora , but that species differs from L. biflora in its 5 calyx appendages that are either long and filiform or short and stubby. The name L. denticulatum , the type of which falls within my concept of L. biflora , was often applied to these southern Indian populations from the Courtallum area (Tamil Nadu).

The name Solanum decemdentatum first occurred as a nomen nudum in Roxburgh’s Hortus Bengalensis ( Roxburgh 1814) but was validated by Nathaniel Wallich in the edition of Flora Indica edited by him and William Carey ( Carey and Wallich 1824). Here the name is attributed to Roxburgh, with citation of a plant seen by Wallich in Singapore and a reference to its similarity to S. rumphii Dunal (= S. americanum Mill. , see Särkinen et al. 2018). A specimen in the Wallich herbarium at Kew (Wallich cat. 2614, K 001116583) is labelled with both Singapore and a date preceding the description, matching the entry in Wallich’s distributed list of specimens (https://wallich.rbge.org.uk/index.php?section=entries&id=2614). I have selected this specimen as the lectotype of S. decemdentatum .

Blume (1826) recognised two varieties of his S. denticulatum as var. α and var. β, from different localities on Java. A sheet at Leiden (L 0003595) has three labels and three fragments of stem mounted together labelled “ Solanum denticulatum ”. I have selected the lower L-most twig that seems to correspond to the adjacent label with the annotation “ 450 / var. α / Gagar Bentang ” as that bearing the most information linking it to the protologue. The other twigs mounted on the same sheet are likely to be syntypes; they are all labelled “ Solanum denticulatum ” in a variety of different hands. An additional sheet (L 0003596) labelled “ var. α ” but does not have the same locality; it should also be considered a syntype. The description of S. mollissimum similarly does not cite a herbarium or collector ( Blume 1826) nor is a locality named in the protologue. I have selected the sheet in Leiden from the Blume herbarium (L 0003586) labelled “ Solanum mollissimum Bl. ” in Blume’s hand as the lectotype.

Dunal (1852) cited Zollinger 723 from “ v. s. in h. Boiss et DC ” in the protologue of S. zollingeri . I have selected the better of these two specimens, that in G-DC (G 00145615), as the lectotype for the species. The original collecting locality for this gathering, as is often the case for Zollonger’s collections, is on the specimen held in Paris. In describing S. osbeckii Dunal (1852) cited only “ J. B. exs. in h. Banks ”; the sheet at BM (BM 000778107), collected by Joseph Banks and Daniel Solander is annotated in Dunal’s hand and is selected as the lectotype. In the protologue of S. osbeckii var. stauntonii he cites a collection of “ sir G. Staunton in h. Lambert ”. The sheet in Vienna W (acc. # 0003086) is the only one I have seen that is unambiguously from Aylmer Bourke Lambert’s herbarium that was widely distributed by sale after his death ( Miller 1970). Dunal (1852) described S. calleryanum from China, citing collections made by Joseph Callery housed in Paris and Gaudichaud 95 from the De Candolle herbarium. Several un-numbered Callery collections are housed in Paris, all with different dates of collection. I have selected the sheet of Gaudichaud 95 in G-DC (G 00145658) as the lectotype, as it is well-preserved and unambiguously the specimen seen by Dunal.

Solanum denticulatum var. lanceolatum was described by Miquel (1864) with the citation of an un-numbered collection by H. Zollinger from “ Lamadjang bei Tenga ”. The only specimens I have seen with the correct locality and collector are in Paris and appear to be duplicates; one of these (P 00369015) has an annotation in Miquel’s hand stating “ Solanum denticulatum Bl. var. lanceolt ”. I select this sheet as the lectotype for the variety.

Kuntze’s herbarium is currently held at NY ( Zanoni 1980); I have selected the sheet at NY (barcode 00172277) as the lectotype of S. biflorum forma pilosa Kuntze , as it corresponds to the protologue and is annotated by Kuntze.

In his global monograph of Lycianthes Georg Bitter (1919) described many infraspecific taxa under both L. biflora and L. macrodon , here considered conspecific. Some of these cited a single specimen in a single herbarium (e. g., L. biflora var. sparsiloba ) and I have considered these holotypes. For those with single specimens cited from Berlin (where the Solanaceae were largely destroyed during World War II, see Vorontsova and Knapp 2010) or with multiple syntype collections cited, I have generally lectotypified these with specimens held in the countries where the plants are native, or from where they were described. For L. biflora var. grandifolia Bitter (1919) cited two syntypes, Meebold 17068 and Meebold 17069, both from WRSL; these specimens were destroyed in World War II and are no longer extant. A duplicate of Meebold 17068 in CAL is designated the lectotype of this variety. Bitter (1919) cited a single specimen of Henry 13652 from “ herb. Berol. ” (= Berlin) in the protologue of L. biflora var. subtusochracea ; I have selected a sheet of this collection at Kew (K 000759409) as the lectotype for var. subtusochracea . For L. biflora subsp. hupehensis Bitter (1919) cited two specimens of Henry 4304 (as “ Faber in Henry’s Coll. from Centr, China ”), from “ herb. Berol., Vindob. ”. I have designated the syntype cited by Bitter from Vienna (W acc. # 1886-0001758) as the lectotype for this Chinese subspecies. No herbarium was mentioned for the only collection cited in the description of L. biflora var. velutinella, Koorders 18041 β . I have chosen the specimen in Bogor as the lectotype for var. velutinella (BO acc. # BO- 4324391) as it is housed in Indonesia, where the type was collected. Bitter (1919) cited two collections in the protologue of L. biflora subsp. elongatidens, Koorders 18038 β from “ herb. Bogor.) and Sarasin 365 from Berlin. I have chosen the Bogor specimen of Koorders 18038 β (acc. # BO- 1990064) as the lectotype. Several collections were cited in the protologue of L. macrodon var. mollitersetosa, Anderson 303 , 1025 and 1026 all from Berlin and a collection from an un-named collector (“ Sammler 12011 ”) from Bogor ( Bitter 1919). The collection in Bogor (acc. # BO- 1593104) has a label where the collector name is torn off the corner, but the handwriting and label style is unambiguously that of C. B. Clarke. Since this variety was described from Indian collections, I have selected the duplicate of Anderson 303 held in Calcutta (CAL acc. # 315720) as the lectotype. Lycianthes macrodon var. sikkimensis was described ( Bitter 1919) from Meebold 15768 and cited as seen in WRSL. These collections are now destroyed, and I have found no duplicates. I have not designated a neotype in the hopes that a duplicate will be found. Lycianthes macrodon var. manipurensis was similarly described from a sheet in WRSL, Meebold 6906, but a duplicate of this gathering is held at CAL (without an accession number or barcode) and is here designated the lectotype.

Bitter later ( Bitter 1922) described L. biflora subsp. yunnanensis citing only Henry 9160 from Berlin; I have selected the most complete of the several duplicates of this collection I have seen (with both flowers and fruits) as the lectotype (K 000922382). In the same publication Bitter (1922) described L. macrodon var. longifrons citing a Hallier collection in Munich that had on it two twigs. He differentiated the two as different varieties by stating (transl. from the original German by S. Knapp) “ Next to the twig described above is a second fruiting twig, apparently collected there by Hallier, which has much richer and stronger hairs on the internodes, the fruit stalks and calyxes, and on both leaf surfaces; (Leaves considerably wider; lam. maj. 12: 6.2 cm lam. min. 8: 5 cm); this second specimen belongs to the var. mollitersetosa Bitter ) ” ( Bitter 1922: 320). These two are likely to be duplicates.

The protologue of S. biflorum var. glabrum ( Hatushima 1969) has no herbarium cited, but Hatushima worked at Kagushima University and a specimen held there (KAG acc. # 163736) is a match for material held in US (barcode 02840676) that has attached a letter from S. Hatushima to E. H. Walker mentioning that this new variety would be described soon. I have selected the KAG specimen as the lectotype of this variety. Hatushima (1969) placed S. schizocalyx in synonymy with his new variety, based on the lack of pubescence mentioned in the protologue of S. schizocalyx ( Merrill and Merritt 1910) .

Figures

Figure 2. Habit and leaves of Lycianthes in Asia A L. banahaensis (Rule s. n., Philippines; DOL 134625) B L. biflora (Pelser & Barcelona s. n., Philippines; DOL 44707) C L. lysimachioides (Nuraliev 1638, Vietnam) D L. parasitica (Saavedra s. n., Philippines; DOL 145620) E L. shunningensis (Nuraliev 2810, Vietnam). Photograph credits: A Greg Rule B Pieter B. Pelser & Julie F Barcelona C, E Maxim Nuraliev D) Aljohn J. Saavedra. DOL numbers are identifiers from PhytoImages (https://phytoimages.siu.edu). Details of collecting localities can be found in Suppl. material 2.

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 5. Fruits of Lycianthes in Asia A L. banahaensis showing the opaque pericarp, lenticellate pedicel and small nub-like calyx appendages (Rule s. n., Philippines; DOL 134667) B L. biflora showing sequence of fruiting ripening and elongate calyx appendages (Nuraliev 182 a, Vietnam) C L. biflora plant with short calyx appendages and little pubescence (Pelser & Barcelona s. n., Philippines; DOL 44710) D L. parasitica showing the small inflorescence axes and berries changing from white to orange with ripening (Tandang s. n., Philippines; DOL 76584) E L. parasitica cross-section of berry showing the two seeds (Pelser & Barcelona s. n., Philippines; DOL 84746) F L. shunningensis showing unripe berry and strongly reflexed calyx appendages (Averyanov et al. 887, Laos). Photograph credits: A Greg Rule B Maxim Nuraliev C, E Pieter B. Pelser & Julie F. Barcelona D Danilo Tandang 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 8. Lycianthes biflora (Lour.) Bitter herbarium specimen. China. Yunnan, Forrest 8650 (BM 001018846). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

Figure 9. Distribution of Lycianthes biflora.