Lycianthes lycioides (L.) Hassl. (as Solanum lycioides L.)

Perennial herbs, shrubs, vines, lianas or trees, sometimes epiphytic. Stems terete or angled, glabrous or pubescent with simple (unbranched), forked, dendritic or stellate trichomes, these usually eglandular, but sometimes glandular. New growth usually with minute papillae, these sometimes glandular. Sympodial units unifoliate or difoliate, if difoliate the leaves geminate and often differing in both size and shape (anisophyllous). Leaves simple, entire, glabrous or pubescent with simple (unbranched), forked, dendritic or stellate trichomes, these usually eglandular, but sometimes glandular; petioles well-developed or not. Inflorescences axillary or adnate to the stems and caulescent (L. kaernbachii of New Guinea only), fasciculate or with a short rhachis; pedicels articulated at the base. Flowers 4–6-merous, usually 5-merous, but some species (e.g., L. banahaensis often 4-merous), cosexual or heterostylous, long- and short-styled flowers borne on the same or different plants (in Australia, New Guinea and the Pacific probably dioecious). Calyx with various numbers (usually multiples of five, but sometimes fewer) appendages protruding from the calyx tube below or just at the truncate rim, or appendages lacking; appendages small bumps to obovoid to linear or linear subulate in shape. Corolla rotate to deeply stellate, white to deep purple or yellow (L. banahaensis), often with the midvein of the lobes darker and the centre paler or yellow-green, interpetalar tissue present or absent, the lobes minute (rotate corollas) or long-triangular, spreading, cupped or reflexed at anthesis. Stamens equal or unequal due to anther and/or filament differences; anthers plumply ellipsoid and obovate to tapering at the tips, usually dehiscing by apical pores opening to longitudinal slits with age or not, these occasionally longitudinally dehiscent. Ovary conical, glabrous; style straight or curved, the stigma minutely capitate, clavate or strongly bifid with diverging lobes. Fruit a berry, globose to ellipsoid to ovoid, green, orange, red or purple, sometimes with stone cells in the mesocarp. Seeds few to many, usually flattened, often triangular in outline, sometimes winged (e.g., L. moszkowskii of New Guinea). Chromosome number: n = 12, 24 (few species have chromosome counts).

Species of Lycianthes are found in the Americas, Asia, Australia, New Guinea and the islands of the Pacific. Species richness is concentrated in Mexico and Central America. No Lycianthes species are native to Africa, Europe or North America north of Mexico.

By far the greatest species diversity in Lycianthes occurs in the Americas (see Dean et al. 2020), but significant diversity occurs in Asia and New Guinea (Knapp 2022; see Fig. 1). The description above attempts to cover variation across the distribution, both in the Americas and outside.

As discussed above and in the Introduction, the species treated here are mostly found throughout Asia, and although they may not be a phylogenetically distinct group, they are geographically logical to treat as a unit.

Based on Solanum banahaense Elmer

Trees or treelets, 3–10 m tall, to 15 cm diameter; stems terete, glabrescent, prominently white-lenticellate, the lenticels corky; new growth glabrous to densely pubescent with transparent, reddish yellow or brownish tan, simple, uniseriate 5–10-celled trichomes 0.2–0.75 mm long, the cells of trichomes small and beadlike (somewhat moniliform); bark of older stems glabrescent, pale greyish green or greyish white, prominently lenticellate. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and occasionally in shape. Leaves simple; blades of major leaves 5.5–30 cm long, 2.5–15 cm wide, elliptic, widest in the middle, discolorous, membranous or chartaceous, occasionally somewhat bullate; adaxial surfaces glabrous or with a few simple uniseriate trichomes like those of the new growth along the midrib, markedly shiny; abaxial surfaces glabrous or with scattered simple uniseriate trichomes on the midrib and major veins, not shiny; principal veins 6–7 pairs, glabrous or sparsely pubescent, drying darker abaxially; base acute and somewhat attenuate onto the petiole, or occasionally truncate (this only rarely and in very large leaves); margins entire; apex acute or acute with an attenuate acumen; petiole 0.6–5 cm long, glabrous or sparsely pubescent with simple uniseriate beadlike trichomes like those of the new growth; blades of minor leaves 3.5–9 cm long, 1.7–6 cm wide, elliptic to broadly elliptic or occasionally almost orbicular; surfaces like those of the major leaves; principal veins of minor leaves 4–5 pairs; base acute to truncate; margins entire; apex acute to somewhat attenuate; petiole of minor leaves 0.4–2 cm long, glabrous or sparsely pubescent. Inflorescences axillary, in fascicles or on a short rhachis 0.1–0.3 cm long, with 2–8(16) flowers, densely pubescent with simple uniseriate trichomes like those of the new growth and young stems; pedicels at anthesis 1.2–2 cm long, 0.5–0.75 mm in diameter at the base, 1.2–1.5 mm in diameter at the apex, spreading but held beneath the leaves, glabrous or more often sparsely pubescent with scattered simple uniseriate trichomes like those of the new growth and leaves, articulated at the base; pedicel scars tightly packed and overlapping on the short rhachis, somewhat corky. Buds ellipsoid, the corolla included in the calyx tube until just before anthesis, the calyx appendages prominent and often somewhat horn-like in bud. Flowers 4- or 5-merous, heterostylous and the plants probably dioecious, “with a peculiar smell” (fide Adduru 109). Calyx tube 2.5–3 mm long, 3–5 mm wide at the mouth, obconical, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the pedicels, with 4–5 appendages arising 0.5–1 mm below the hyaline rim, the appendages 0.5–3.5 mm long, triangular to horn-like, usually perpendicular to the calyx tube, glabrous or with a few trichomes. Corolla 0.8–1 cm in diameter, white, cream (“milky”) or yellow, sometimes with a tinge of violet abaxially, stellate, lobed nearly to the base, abundant interpetalar tissue absent, the lobes 4–5 mm long, 2–4 mm wide, spreading, thick and fleshy, adaxially densely papillate, the midvein somewhat keeled in dry specimens, abaxially glabrous, the tips and margins densely papillate, the tips cucullate. Stamens equal and the same in short- and long-styled flowers; filament tube minute; free portion of the filaments less than 0.2 mm long, glabrous; anthers 2.5–4 mm long, 1.5–2.5 mm wide, plumply ellipsoid and somewhat beaked in some dry material, 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, vestigial in short-styled flowers; style in short-styled flowers absent, in long-styled flowers 2–3 mm long, held almost entirely within the anther cone, glabrous; stigma broadly clavate and irregularly lobed, bright dark green in live plants, the surfaces minutely papillate. Fruit a globose or occasionally somewhat ellipsoid berry, 1–2 cm in diameter, green when immature, orange to bright red when mature, the pericarp glabrous, thin, matte, and opaque; fruiting pedicels 2.5–4 cm long, 1–1.5 mm in diameter at the base, 2–3 mm in diameter at the apex, spreading, somewhat woody, green and prominently lenticellate; fruiting calyx not accrescent or expanding, a flattened disc below the berry. Seeds (5)20–60 per berry, ca. 3 mm long, ca. 2 mm wide, flattened and somewhat reniform, pale yellowish brown, the surfaces pitted, the testal cells sinuate in outline, the testal cell walls elongate but not prominently “hairy”. Stone cells absent. Chromosome number not known.

(Fig. 7). Lycianthes banahaensis occurs in Indonesia and the Philippines; it is found on most islands of the Philippines and in eastern Indonesia from Sulawesi to the Sunda Islands, but not on Java or Borneo or in the Maluku Islands.

Lycianthes banahaensis grows in low to middle elevation evergreen forests or forest edges, sometimes on limestone, from 50 to 1,500 m elevation.

Indonesia. Sulawesi: onbeked (Koorders 18050), paratha kehu (Sangihe language, Talangmin 51), kaometi, makopi (Totemboan language, Koorders 1898, as “S. minahassae”).

(IUCN 2020). EOO (1,652,883 km² - LC); AOO (228 km² - EN). Lycianthes banahaensis is known from more than five localities and is relatively widely distributed in the region. In the Philippines it is known from several protected areas (e.g., Mount Apo, Samar Island and the Northern Sierra Madre). The assessment of Endangered (EN) 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).

Lycianthes banahaensis is a small tree (Fig. 2A) with shiny geminate leaves and prominently lenticellate stems and fruiting pedicels. The leaves are extremely variable in size; this has led to the description of the entity several times from different localities in the Philippines (see synonymy), as has variation in pubescence density on new growth. Lycianthes anisophylla, for example, was distinguished from L. lagunense in being “more glabrous” (Elmer 1915).

Plants appear to have either staminate flowers with vestigial ovaries or have berries, suggesting L. banahaensis is likely dioecious like many Lycianthes species from New Guinea (Knapp 2022). Flowers on fruiting specimens have short styles that are contained almost completely within the anther tube (see Fig. 4A); this means that recognising short- or long-styled flowers can be difficult, especially on dry specimens. Field verification of the reproductive biology of L. banahaensis is a priority (as is the case for most of these species outside the Americas).

Flowers of L. banahaensis are often tetramerous, and this character appears to be geographically fixed in populations, with many specimens from the Philippines mostly being 4-merous (Fig. 4A), while 5-merous collections are found in both the Philippines and Indonesia. The truly yellow flowers of many populations of L. banahaensis are unusual in Lycianthes, but cream or white flowers have been recorded on labels, so this appears not to be a constant character.

Lycianthes banahaensis is most similar to L. bimensis of the Sunda Islands; both species are trees with fleshy, heterostylous flowers, dense fine pubescence on the new growth and orange or bright red mature berries. The prominently lenticellate stems and pale bark of L. banahaensis are diagnostic; L. bimensis has smooth stems and dark brown bark. Lycianthes banahaensis has smaller flowers (corolla 0.8–1 cm in diameter versus 1.4–1.6 cm in diameter in L. bimensis) with the style of pistillate flowers held within the anther cone and strongly lobed (versus exserted and long-clavate in L. bimensis). Both species occur on the island of Flores (West Nusa Tenggara, Indonesia).

Adolph Elmer lived and collected in the Philippines from 1904 until his death in a Japanese internment camp in 1942. For the names coined by him (S. banahaensis, S. lagunense, S. anisophyllum), all specimens upon which they were based that were in his private collection at the Philippine National Herbarium (then the Philippine Bureau of Agriculture under United States jurisdiction) were 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). Duplicates of these are widely distributed, Elmer sold large numbers of specimens to Harvard, Geneva and Kew. For Solanum banahaense I have chosen the duplicate held in the herbarium of the Arnold Arboretum at Harvard (A, barcode 00077843) as the lectotype; it is the best preserved of the single collection (Elmer 7492) cited in the protologue. Similarly, I have selected the more complete duplicate of Elmer 9425 held at Kew (K000759392) as the lectotype for S. lagunense; other duplicates I have seen lack flowers or fruits. Elmer (1915) cited two collections in the protologue of S. manucaling (Elmer 10489, Elmer 11693 – in flower and fruit respectively). I have selected the Bishop Museum duplicate of Elmer 10489 (BISH1005081, acc. # 581198) as the lectotype, as it is one of the few duplicates of either of these collections that still has numerous flowers. For S. anisophyllum Elmer (1915) cited a single collection (Elmer 13887) now represented in many herbaria; most of the duplicates are sterile or only have flowers in packets. I have selected the Arnold Arboretum duplicate (A, barcode 00077840) as the lectotype as it has flowers attached to the stems.

Bitter (1919) cited a single collection in the Bogor herbarium (“Teysmann n. 8918! hb. Bogor”) in the protologue of L. aceratia. The Bogor specimen (acc.# BO-1323491) bears an annotation slip in Bitter’s distinctive hand, suggesting he had this material in his possession at some point. Bitter never travelled to Indonesia (Weber 1929) but must have had material on loan from Bogor itself; he never cites the Leiden collections in his Lycianthes monograph. This indicates he only saw the Bogor sheets of collections that are duplicated in Leiden, as when he examined sheets of the same collection from various herbaria his practice was to cite them all. I have therefore treated the Bogor sheet of Teijsmann 8918 as the holotype of L. aceratia.

In the protologue of L. anisophylla var. masbateensis Bitter (1919) cited a duplicate of Merrill 3046 at Berlin, now destroyed. I have selected the US duplicate (barcode 0027867, acc. # 438017) as the lectotype; Merrill worked for the United States government as the Director of the Philippine Bureau of Agriculture during the first quarter of the 20^th Century, leaving in 1923 to return to the United States (Schultes 1957). He regularly sent duplicates to the National Herbarium and the specimen there is the best preserved of those I have seen.

Based on Solanum biflorum Lour.

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).

(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.

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.

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 ngủ cuống to (as L. macrodon), cà ngủ (Hop 2017).

(IUCN 2020). EOO (26,990,176 km² - LC); AOO (1,684 km² - 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).

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. 3A, 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. 3A), 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. 5B, 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. 5F). 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, K001116583) 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 (G00145615), 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 (BM000778107), 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 (G00145658) 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 (P00369015) 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 (K000759409) 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 18041β. 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 18038β from “herb. Bogor.) and Sarasin 365 from Berlin. I have chosen the Bogor specimen of Koorders 18038β (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 (K000922382). 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).

Based on Solanum bimense Miq.

Trees or treelets, 3–15 m tall, to 20 cm diameter; stems terete, glabrescent, sparsely pubescent with weak-walled transparent simple uniseriate 2–10-celled trichomes when young; new growth densely floccose pubescent with transparent, golden, simple, uniseriate 2–10-celled trichomes to 0.7 mm long, the trichomes moniliform and tangled; bark of older stems glabrescent, brown or greyish brown, smooth. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and occasionally in shape. Leaves simple; blades of major leaves 9–24 cm long, 4.5–14 cm wide, elliptic to elliptic-ovate, widest in the middle or just below, somewhat discolorous, membranous or chartaceous; adaxial surfaces glabrous to sparsely pubescent with simple uniseriate trichomes like those of the stems, these denser along the veins; abaxial surfaces glabrous to sparsely pubescent with simple uniseriate trichomes like those of the stems, if the lamina glabrous the trichomes confined to the principal veins and midrib; principal veins 10–14 pairs, glabrous or pubescent, the tertiary venation prominent, drying dark abaxially; base truncate and usually somewhat oblique; margins entire; apex acute to acuminate; petiole 2–7 cm long, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the new growth; blades of minor leaves 3.5–9 cm long, 1.9–8 cm wide, ovate to orbicular; surfaces like those of the major leaves; principal veins of minor leaves 6–9 pairs; base truncate; margins entire; apex acute; petiole of minor leaves 0.7–2.5 cm long, glabrous or sparsely pubescent. Inflorescences axillary, in fascicles or on a short rhachis 0.2–0.5 cm long, with 8–16 flowers, glabrous or densely floccose pubescent with simple uniseriate trichomes like those of the new growth and young stems; pedicels at anthesis 1.4–1.5 cm long, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading, glabrous to densely floccose pubescent with simple uniseriate trichomes like those of the new growth and leaves, articulated at the base; pedicel scars tightly packed and overlapping on the short rhachis, somewhat corky. Buds long-ellipsoid, the corolla included in the calyx tube when young, ca. halfway exserted before anthesis, the calyx appendages more prominent in bud. Flowers 5-merous, heterostylous and the plants probably dioecious. Calyx tube 3–3.5 mm long, 3–4 mm wide at the mouth, obconical, glabrous to densely pubescent with simple uniseriate trichomes like those of the pedicels, with 5 appendages arising 0.1–0.5 mm below the hyaline rim, the appendages ca. 0.5 mm long, triangular to subulate, usually perpendicular to the calyx tube, glabrous or with a few trichomes. Corolla 1.4–1.6 cm in diameter, deep to pale violet, stellate, lobed nearly to the base, abundant interpetalar tissue absent, the lobes 7–8 mm long, 2–2.5 mm wide, spreading, thick and fleshy, glabrous on both surfaces, but with a few papillae on the slightly keeled midvein adaxially, the tips and margins densely papillate, the tips cucullate. Stamens equal and the same in short- and long-styled flowers; filament tube minute; free portion of the filaments ca. 0.5 mm long, glabrous; anthers 3.5–4 mm long, 1.2–1.5 mm wide, ellipsoid, 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, vestigial in short-styled flowers; style in short-styled flowers ca. 2.5 mm long, in long-styled flowers 8–10 mm long, exserted from the anther cone, glabrous; stigma ellipsoid to long-clavate, the tip sometimes 2-lobed, the surfaces minutely papillate. Fruit a globose berry, 0.8–1.2 cm in diameter, green when immature, bright red when mature, the pericarp glabrous, thin, matte, and opaque; fruiting pedicels 2–2.5 cm long, 1–1.5 mm in diameter at the base, 2.5–3 mm in diameter at the apex, spreading, somewhat woody and verrucose; fruiting calyx not accrescent or expanding, a flattened disc below the berry. Seeds 60–80+ per berry, 2.5–3 mm long, ca. 2.5 mm wide, rounded and not flattened, pale straw-colored, the surfaces deeply pitted, the testal cells pentagonal in outline and with prominent “hairy” lateral walls. Stone cells absent. Chromosome number not known.

(Fig. 11). Lycianthes bimensis is confined to the lesser Sunda Islands of Flores, Sumbawa and Timor in the Indonesian provinces of East and West Nusa Tenggara.

Lycianthes bimensis is a forest treelet in moist to wet tropical forest on mountainsides, growing from 400 to 1,000 m elevation.

None recorded.

(IUCN 2020). EOO (38,498 km² - NT); AOO (24 km² - EN). Lycianthes bimensis is a species known from only five localities and has a narrow distribution in the region. It occurs within the boundaries of Mount Tambora National Park on Sumbawa, but collections from there date to the 19^th century. The assessment of Vulnerable (EN) based on AOO is likely due to collecting bias, but also due to the island nature of the distribution. But based on the number of localities and threats to the habitat I consider L. bimensis to be of some conservation concern and therefore suggest a preliminary status of VU (B2a, b[iii]).

Lycianthes bimensis is a distinctive plant with long-petiolate leaves with prominent and crowded principal veins (Fig. 10). The flowers are heterostylous, and I have not seen berries or developing berries on specimens with short-styled flowers, suggesting L. bimensis is dioecious. The calyx of L. bimensis has 5 small appendages borne somewhat below a hyaline rim; it can be distinguished from other species with 5 calyx appendages such as L. laevis in its tree habit, fleshy corolla lobes (versus membranous lobes) and heterostylous flowers. The calyx tube in L. bimensis is usually larger (3–3.5 mm versus 1.5–2.5 mm) and more robust than that of L. laevis.

Lycianthes bimensis is most similar to L. banahaensis, both species are trees with fleshy, heterostylous flowers, dense fine pubescence on the new growth, 5 calyx appendages and orange or bright red mature berries. Lycianthes bimensis has smooth stems and dark brown bark versus the diagnostic prominently lenticellate stems and pale bark of L. banahaensis. Leaf petioles in L. bimensis are generally longer relative to leaf size than in L. banahaensis, and the flowers of L. bimensis are larger (corolla 1.4–1.6 cm in diameter versus 0.8–1 cm in diameter in L. banahaensis). The style of long-styled flowers of L. bimensis is exserted from the anther cone and long-clavate whereas that of L. banahaensis has the style of long-styled flowers contained within the anther cone and strongly lobed. Both species occur on the island of Flores (West Nusa Tenggara, Indonesia).

The type collection of L. bimensis is from an expedition to Mount Tambora on the island of Sumbawa and was made by the prolific Swiss collector Heinrich Zollinger, who studied in Geneva with the de Candolles and whose collections are widely distributed. The massive eruption of Tambora in 1815 is the largest historic volcanic eruption (Oppenheimer 2003), expelling more volume of rock and ash and with more fatalities than that of the more well-known Krakatoa. In his time collecting in Java and the surrounding area in the 1840s, Zollinger wished to study Tambora after the eruption to assess its effects on the local ecosystem and to document plant recovery (Zollinger 1855). In 1847 he and his companions were the first scientists to ascend the volcano after the eruption. The summit was still wreathed in smoke but he records vegetation recovery at various levels, and also reflects that he occasionally fell through a crust to land in a warm sulphurous powder. The type collection of S. bimense is not on Tambora itself, but on an adjacent peak (Doro Omomboha, on Zollinger’s map labelled as Buha) nearer to the town of Bima.

I have lectotypified S. bimense with the sheet of Zollinger 3458 in the Bogor herbarium (BO acc. # BO-1323395) as no herbarium was mentioned in the protologue (Miquel 1864) and the specimen is housed in Indonesia, where L. bimensis is endemic. Miquel never travelled to Indonesia (Stafleu 1966); it is likely that he either saw material in Leiden, where he was director of the Rijksherbarium, or sets of duplicates readied for distribution.

The designation “Solanum floccosum Zipp.” was published in a list of names with no description or diagnosis (Spanoghe 1841) rendering it a nomen nudum.

Based on Solanum laeve Dunal

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.

(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.

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.

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 (Blume1823, 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 ngủ cặp đôi (as L. bigeminata), cà ngủ nhẵn (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.

(IUCN 2020). EOO (15,152,928 km² - LC); AOO (800 km² - 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).

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 3B, 4C). 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, P000578637). 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 (2625B from “herb. Wight”) was excluded and referred to S. laeve. Nees van Esenbeck (1834) indicated that only the specimens labelled 2625A and 2625C were part of his S. membranaceum. In the Wallich herbarium at Kew there are three specimens labelled 2625 (K001116640, K001116641, K0011166420) none of which bears the annotation corresponding to the catalogue entry. One of these is unambiguously labelled 2625A (K001116640) 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 (GZU000255706); 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 (GZU000255706) 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 (P00248294) 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 (P00058859) 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 (P0057928) is annotated “α angustifolium” by Dunal, that chosen for var. latifolium (P00057929) is annotated “β latifolium”) by Dunal and that for the species is only annotated with the specific epithet (P00057927). 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 (BM001018973) 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 (K000923382) 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 (E00273869); 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 (K00759401) 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 (P00379534) 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 P000369020 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.

Based on Solanum lysimachioides Wall.

Prostrate or creeping herbs, the stems to 1.5 m long, occasionally described as subshrubs to 1 m; stems terete, collapsing as if hollow in most specimens, rooting at the nodes or near the nodes along the stem, densely pubescent to glabrescent, if pubescent the trichomes whitish transparent, simple, uniseriate 5–8-celled, to 2 mm long; new growth glabrous to densely pubescent with whitish transparent, simple uniseriate 5–8-celled trichomes to 2 mm long; surfaces (bark) of older stems glabrescent to variously pubescent, the trichomes sparser on older stems, pale greyish brown or greenish brown. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and occasionally in shape, when minor leaves are similar in size the leaves appear opposite. Leaves simple; blades of major leaves 1.8–14 cm long, 1.4–8.7 cm wide, elliptic to ovate (occasionally narrowly elliptic), widest in the middle or in the lower half, concolorous but occasionally purple or purple-veined abaxially (e.g., Sino-American Guizhou Botanical Expedition 1240), thin and membranous; adaxial surfaces glabrous to evenly and sparsely to moderately pubescent with simple uniseriate trichomes like those of the stems, these denser along the veins, the lamina always visible; abaxial surfaces glabrous to evenly and moderately pubescent with simple uniseriate trichomes like those of the stems, the lamina clearly visible, the trichomes denser on the principal veins and midrib; principal veins 3–5 pairs, glabrous or variously pubescent, the venation not markedly coloured; base attenuate to truncate to cordate; margins entire, in pubescent plants ciliate with simple uniseriate trichomes to 1 mm long; apex acuminate; petiole 0.9–6.3 cm long, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the stems and new growth; blades of minor leaves 0.8–9 cm long, 0.6–5.5 cm wide, elliptic to ovate, sometimes almost orbicular; surfaces like those of the major leaves; principal veins of minor leaves 3–4 pairs; base attenuate to truncate or cordate; margins entire; apex acute to acuminate; petiole of minor leaves 0.4–2.1 cm long, glabrous or sparsely pubescent like the stems. Inflorescences axillary, in fascicles, with 1(2) flowers, glabrous; pedicels at anthesis 0.6–2.3 cm long, ca. 0.5 mm in diameter at the base, 1–1.5 mm in diameter at the apex, spreading and sometimes apparently bent (geniculate) in the upper third, glabrous to pubescent with simple uniseriate trichomes like those of the stems, articulated at the base; pedicel scars tightly packed in the leaf axils. Buds strongly tapered and pointed, the corolla strongly exserted from the calyx tube before anthesis, the calyx appendages enclosing the bud only partially. Flowers 5-merous, cosexual. Calyx tube 1.9–2.5 mm long, 2.5–3.5 mm wide at the mouth, obconical or an open cuplike structure, 10-ridged, glabrous or pubescent with simple uniseriate trichomes like those of the pedicels, the trichomes often denser on the ridges, with 10 appendages arising ca. 0.5 mm below the hyaline rim, the appendages 3–6 mm long, 0.5–0.7 mm wide, subulate, parallel to the calyx tube, glabrous or pubescent with simple uniseriate trichomes like those of the stems and pedicels. Corolla 1.8–3 cm in diameter, white, “pink” or pale violet (lilac), with green dots at the base of the lobes, stellate, lobed nearly to the base, interpetalar tissue absent but a thin edge of tissue apparent on lobe margins, the lobes 8–15 mm long, 2.5–3.5 mm wide, spreading, glabrous on both surfaces, the tips and margins sparsely and minutely papillate. Stamens equal; filament tube minute; free portion of the filaments 0.75–1 mm long, glabrous; anthers 2–4 mm long, 0.75–1.5 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 4–9 mm long, exserted from the anther cone, glabrous; stigma prominently capitate, the surfaces minutely papillate. Fruit a globose berry, 0.6–0.8 cm in diameter, green when immature, bright red when mature, the pericarp glabrous, thin, shiny, and transparent; fruiting pedicels 1–3 cm long, 0.5–1 mm in diameter at the base, 1.5–2 mm in diameter at the apex, spreading; fruiting calyx not accrescent or expanding, but remaining a cup covering ca. 1/4 of the berry, the appendages spreading like a star beneath the berry. Seeds 10–40 per berry, 2–2.5 mm long, 1.2–2 mm wide, flattened-reniform, pale straw-coloured or yellowish tan, the surfaces pitted, the testal cells sinuate in outline in the centre, rectangular on the margins, prominent “hairy” appendages absent. Stone cells absent. Chromosome number not known.

(Fig. 15). Lycianthes lysimachioides occurs in China, India, Indonesia, Japan, Laos, Myanmar (Burma), Nepal, Taiwan, Thailand and Vietnam. I have not seen specimens from Cambodia, but it is likely to occur there.

Lycianthes lysimachioides is usually found growing in deep shade in damp places along streams or in forest understory, often as dense patches, from 600 to 2,400 m elevation.

China. dan guo hong si xian (as L. solitaria), dan hua hong si xian (Zhang et al. 1994). Indonesia. Sulawesi: kamoenti (Koorders 1898, as S. lysimachioides). Vietnam. cà ngủ dạng trân châu (Hop 2017).

(IUCN 2020). EOO (14,512,768 km² - LC); AOO (372 km² - EN). Lycianthes lysimachioides is a species known from more than ten localities and is quite widely distributed in the region. Throughout the range it is known from protected areas (e.g., Upper Shillong Protected Forest in India; Emeishan in Sichuan, China). The assessment of Endangered (EN) based on AOO is likely due to collecting bias, but also due to the island nature of the region. I therefore assign it a preliminary status of Least Concern (LC).

Lycianthes lysimachioides is distinctive amongst Asian species of Lycianthes in being a creeping herb of forest understories and shady places (Fig. 2C); it only occasionally attains shrubby proportions, but if so then it is less than a metre tall. The specific epithet reflects its striking similarity with species of Lysimachia L. (Primulaceae). The geminate leaves of L. lysimachioides can be mistaken for the opposite leaves of Lysimachia and coupled with the creeping habit, leads to confusion in herbarium specimens. I have seen several specimens of Lysimachia evalvis Wall. identified as Lycianthes lysimachioides; the plants are very similar at first glance (see image of K0007507178 at https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:701116-1). Like most of the species treated here, it is extremely variable in pubescence density, with some individuals densely pubescent and others nearly glabrous. Trichomes of L. lysimachioides are simple and uniseriate, and usually composed of many small cells, I have never seen branched trichomes in this species. Leaf shape in L. lysimachioides is quite variable, some specimens have almost orbicular, cordate leaves (Fig. 2C), whereas others are almost narrowly elliptic with acute or attenuate bases. Most of the infraspecific taxa placed in synonymy here have been based on pubescence or leaf shape differences. There appears to be no particular geographical or habitat correlation with leaf shape or pubescence density.

Lycianthes lysimachioides is sometimes mistaken for L. biflora but is a creeping or scandent herb rooting at the nodes rather than an erect woody shrub, 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). Identification of sterile or poor specimens of L. lysimachioides can be helped by looking for roots at the nodes and stems that collapse in dried specimens. The single collection I have seen from peninsular Malaysia (Henderson SF-22378 from Pahang) is somewhat woody but has clear roots emerging at the nodes and along the stems. Plants identified and depicted as L. lysimachioides from Manipur, northeastern India by Eshuo (2023) are part of the concept of L. biflora as treated here.

Lycianthes schizocalyx has similar calyx appendages in which at least some arise below the calyx rim leaving a hyaline edge; it can be distinguished from L. lysimachioides by its shrubby habit, calyx appendages of variable length in a single flower and larger berries (1–1.3 cm versus 0.6–0.8 cm in diameter.

Dunal (1852) described Solanum caulorhizum based on the collection Zollinger 705 in the de Candolle herbarium (G-DC); a duplicate in the Paris herbarium (P00369007) is a specimen of L. biflora and is excluded from the isotypes here (this sheet has been designated Zollinger 705 [a] in the material examined).

Solanum nematosepalum was described only referring to a specimen “Solanum ciliatum Blume in herb. reg. L.B.”(Miquel 1864) from Java. Searches in both Bogor and Leiden have revealed no specimens so labelled, so I here designate a neotype from the Blume herbarium (L.2881683) collected in Java that corresponds to the protologue, represents this species and is labelled in what looks like Miquel’s handwriting.

In his description of L. lysimachioides var. sinensis Bitter (1919) cited a number of collections (Henry 7207, 3063, 6080; Wilson 1374, 2158; Rosthorn s.n.) all from “herb. Berol.” From these large-leaved specimens I have selected Henry 6080 (BM001018842) as the lectotype, as this is the best-preserved specimen, and duplicates of this collection are widely distributed.

Merrill (1923) cited no herbaria in his description of S. debilissimum, and only a single collection, McClure 9532. Material Merrill examined may have been destroyed in Manila (see under L. banahaensis) but duplicates of this material are widely distributed. I have selected the sheet in Edinburgh (E00426499) as the lectotype for this species, as this specimen has flowers and a developing fruit, whereas others have only flowers or are sterile.

The varieties of L. lysimachioides described by Wu and Huang (1978) are generally referred to only using the word “type” and cite only the herbarium where the type is housed (i.e., var. purpurifolia, var. rotundifolia); where multiple sheets are housed in the cited herbaria (i.e., PE) I have selected that annotated as “holotype” as the lectotype. Cases where only one specimen is housed in the cited herbarium are clear.

Based on Solanum oliverianum Lauterb. & K.Schum.

Woody climbers or lianas, sometimes described as shrubs, to 3+ m tall (often described on labels “beautiful” e.g., van Royen & Sleumer 7716 from New Guinea); stems terete, glabrous; new growth glabrous or minutely papillate with tiny 1–2-celled weak simple uniseriate trichomes less than 0.2 mm long, these soon deciduous; bark of older stems whitish grey, peeling and flaking, somewhat rugose and thick. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size but not in shape. Leaves simple; blades of major leaves (6.5)9–25 cm long, (2.8)3–10 cm wide (perhaps larger but not collected), elliptic, slightly discolorous, thick and coriaceous or chartaceous; adaxial surfaces glabrous, somewhat shiny; abaxial surfaces glabrous; principal veins 6–8 pairs, the midrib slightly keeled above, sometimes drying yellowish tan; base acute, often somewhat oblique; margins entire, revolute; apex acute or acuminate with an elongate drip-tip; petioles 1–2.5 cm long, glabrous; blades of minor leaves 4–9 cm long, 2.5–5 cm wide, shape, texture and pubescence like that of the major leaves; base acute; margins entire, revolute; apex acute or acuminate, occasionally rounded; petioles 0.6–1 cm long, glabrous. Inflorescences dense axillary fascicles, occasionally woody and enlarged with what appear to be short, fat branches, these to 0.3 cm long, with 10–20-flowers, several open at the same time, glabrous; pedicels at anthesis 1–1.4 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading, glabrous, articulated at the base; pedicel scars tightly packed on the woody fascicle base. Buds plumply ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis. Flowers 5-merous (4-merous in Takeuchi 23389), heterostylous, specimens with either all short-styled flowers or long-styled flowers and fruit, the plants probably dioecious. Calyx tube 2.5–3 mm long, 3–3.5 mm in diameter, deeply cup-shaped, usually described as purple or purplish blue, thick and fleshy, densely verrucose/tuberculate, without appendages, the rim somewhat hyaline ca. 0.5 mm wide, sparsely papillate. Corolla 0.8–1.1 cm in diameter, white or purple, stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 2–5 mm long, 1–2 mm wide, spreading or reflexed, thick and fleshy (live plants), appearing woody in dry material, adaxially glabrous to densely papillate with a few weak trichomes distally, abaxially densely papillate somewhat verrucose, the tips and margins densely papillate, the midvein raised especially adaxially, the tips cucullate. Stamens equal; filament tube minute; free portion of the filaments 1–1.5 mm long, glabrous; anthers 2–2.5 mm long, ca. 1 mm wide, plumply ellipsoid or slightly obovoid, creamy white, yellow or purple, glabrous, poricidal at the tips, the pores round, distally directed, not elongating with age. Ovary conical, glabrous, vestigial in short-styled flowers; styles less than 0.2 mm long and vestigial in short-styled flowers, 5–6 mm long in long-styled flowers, straight, glabrous; stigma slightly bilobed, the surfaces minutely papillate. Fruit a globose berry, 0.7–1 cm in diameter, green and becoming bluish black when ripe, the pericarp glabrous, thick and appearing woody in dry material, matte, opaque; fruiting pedicels 1.1–1.5 cm long, 1–1.5 mm in diameter at the base, 1.5–2 mm in diameter at the apex, spreading or erect (?), woody, corky and markedly verrucose/tuberculate; fruiting calyx a cup surrounding ca. the lower half of the berry (making the fruit look like an acorn), woody (fleshy in live plants) and verrucose/tuberculate both adaxially and abaxially, green flushed with purple (fide Polak 864). Seeds 20–40 per berry, 3–3.5 mm long, 2–2.5 mm wide, flattened reniform or slightly tear-drop shape, reddish brown, the surfaces at the margins deeply pitted with pentagonal testal cells, the seed centre only shallowly pitted and the cells not clear, prominent “hairy” testal cell walls absent. Stone cells absent. Chromosome number not known.

(Fig. 17). In Asia Lycianthes oliveriana occurs only in Indonesia on the Maluku Islands (Seram [Maluku] and Halmahera [Maluku Utara]) but is widespread on the island of New Guinea in both Papua New Guinea and Indonesia (Knapp 2022).

Lycianthes oliveriana grows in lowland to montane and premontane rainforests, from almost sea level to 2,300 m elevation. On New Guinea this wide elevational range is accompanied by much variation in leaf size and shape (Knapp 2022).

None recorded.

(IUCN 2020). EOO (1,006,290 km² - LC); AOO (156 km² - EN). Lycianthes oliveriana is known from more than 10 localities at a wide variety of elevations; the assessment here is based on its entire distribution. It occurs within protected areas in both Indonesia and Papua New Guinea (see Knapp 2022). This suggests a preliminary threat status of either Least Concern (LC) or Near Threatened (NT).

Lycianthes oliveriana is a distinctive species with many-flowered axillary inflorescences, relatively small (ca. 1 cm in diameter) flowers with valvate aestivation and thick fleshy corollas, plump, ellipsoid to slight obellipsoid anthers and somewhat warty calyces with no appendages (Fig. 3D). The berries appear to have a slightly woody pericarp; this woodiness coupled with the somewhat accrescent calyx tube gives the fruits of L. oliveriana the look of tiny acorns. Lycianthes oliveriana appears to be dioecious, with long- and short-styled flowers on different plants. This needs confirmation in the field, and L. oliveriana would be an ideal subject for a reproductive biology study since it is relatively common and widely distributed.

Lycianthes oliveriana could potentially be confused with L. parasitica, a species with a wider distribution in tropical Asia outside of New Guinea. Berries of L. oliveriana are somewhat woody on dry specimens and have many seeds, while those of L. parasitica are juicy with a translucent pericarp and only have two (to very occasionally four) seeds. Flowers of L. parasitica are cosexual while those of L. oliveriana are either long- or short-styled. On New Guinea L. oliveriana has a wide variety of leaf shapes (Knapp 2022). The Malaku specimens are of plants with broadly elliptic leaves (Fig. 16).

Based on Solanum parasiticum Blume

Epiphytic shrubs, 1–2 m tall, to 20 m high in trees; stems terete, glabrescent, occasionally (in eastern Sabah, see discussion) sparsely pubescent with weak-walled transparent simple uniseriate 2–10-celled trichomes when young, often markedly zig-zag; new growth glabrous or occasionally pubescent with transparent, simple uniseriate 8–10-celled trichomes to 0.2–1 mm long; bark of older stems glabrous or glabrescent, papery-white, markedly exfoliating. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and occasionally in shape, the minor leaves often apparently absent. Leaves simple; blades of major leaves 7–21 cm long, 2–8.4 cm wide, narrowly elliptic to elliptic, widest in the middle, concolorous and shiny, membranous or somewhat fleshy; adaxial surfaces glabrous or occasionally sparsely pubescent with simple uniseriate trichomes like those of the stems, these denser along the veins; abaxial surfaces glabrous to sparsely pubescent with simple uniseriate trichomes like those of the stems, if the lamina glabrous the trichomes confined to the principal veins and midrib; principal veins 5–6 pairs, glabrous or pubescent, the tertiary venation not visible; base acute to attenuate; margins entire; apex acute to acuminate; petiole 0.7–2 cm long, fleshy, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the new growth; blades of minor leaves 1–3 cm long, 1–2.5 cm wide, elliptic to orbicular; surfaces like those of the major leaves, the minor leaves often deciduous; principal veins of minor leaves 3–4 pairs; base acute to rounded-truncate; margins entire; apex obtuse or somewhat rounded; petiole of minor leaves 0.5–0.7 cm long, glabrous or sparsely pubescent. Inflorescences axillary, in fascicles or on a short rhachis to 0.5 cm long, with 3–8 (16) flowers, glabrous; pedicels at anthesis 1–1.5 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading, glabrous and shiny or occasionally pubescent with simple uniseriate trichomes like those of the stems, articulated at the base; pedicel scars tightly packed and overlapping on the short rhachis, somewhat corky. Buds ellipsoid, the corolla completely included in the calyx tube when young, ca. halfway exserted before anthesis, the calyx appendages only apparent in bud. Flowers 5-merous, cosexual. Calyx tube 1.5–2.5 mm long, 2.5–3.5 mm wide at the mouth, obconical, glabrous or occasionally pubescent with simple uniseriate trichomes like those of the pedicels, with 5 small appendages arising ca. 0.5 mm below the hyaline rim and only visible in bud, the appendages ca. 0.2 mm long, triangular nubs, usually perpendicular to the calyx tube, glabrous or with a few trichomes. Corolla 0.9–1 cm in diameter, white or pale violet (lilac), stellate, lobed nearly to the base, interpetalar tissue absent but a thin edge of tissue apparent on lobe margins, the lobes 4–4.5 mm long, 1.2–2 mm wide, spreading, glabrous on both surfaces, the tips and margins densely papillate, the tips cucullate. Stamens equal; filament tube minute; free portion of the filaments ca. 1 mm long, glabrous; anthers 2–2.3 mm long, 1–1.5 mm wide, ellipsoid and somewhat tapering, 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 4.5–5 mm long, exserted from the anther cone, glabrous; stigma prominently capitate, the surfaces minutely papillate. Fruit a globose berry, 0.5–0.8 cm in diameter, green when immature, becoming white, then orange or red when mature, the pericarp glabrous, thin, shiny, and transparent; fruiting pedicels 1.5–2.2 cm long, 0.5–1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading; fruiting calyx not accrescent or expanding, but remaining a cup beneath the berry, 2–2.5 mm long, 3–4 mm wide at the mouth. Seeds 2(-4) per berry, 3.5–6 mm long, ca. 2.5–4.5 mm wide, ovoid-reniform, pale straw-colored, the surfaces deeply pitted, the testal cells sinuate in outline and with prominent “hairy” appendages on the thickened lateral walls. Stone cells absent. Chromosome number not known.

(Fig. 19). Lycianthes parasitica occurs in Brunei Darussalam, Indonesia, Malaysia, the Philippines and Thailand (a single sterile specimen only, Put Phraisurind 122).

Lycianthes parasitica grows as an epiphytic shrub in wet forests, usually at low elevations along rivers and streams and in swamp forests, from 100 to 1,000 m elevation.

Brunei Darussalam: usak oncom payo (Dusun language, Bernstein 322). Indonesia. Java: terong-an (Blume 1823); Sumatra: sarindon (si Boeea 6246, 9081). Malaysia. Sabah: akar (Bakar 25023), akar tumpeng (Sungei Kinabatangan language, Enggoh 7345). Philippines. Mindanao: lawmoos (Bagóbo language, Elmer 1910), sisumbidan (Manobo language, Elmer 13572).

(IUCN 2020). EOO (26,990,176 km² - LC); AOO (1,684 km² - VU). Lycianthes parasitica is a species known from more than five localities and is relatively widely distributed in the region, though as an epiphyte is rarely collected. Throughout the range it is known from protected areas (e.g., Mount Kinabalu and the Sepilok Forest Reserve, Sabah, Malaysia; Sohoton National Park on Samar Island in the Philippines). 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).

Lycianthes parasitica is distinctive in its epiphytic habit, usually strongly zig-zagging stems with markedly exfoliating white bark and berries with few (usually only two), large seeds (Figs 2D, 5E). The epiphytic habit has evolved independently several times in the Solanaceae (Orejuela 2021). The leaves are shiny on the adaxial surfaces and quite fleshy, in dried specimens the lower orders of venation are barely visible. In flower it is possible that L. parasitica could be confused with L. laevis, which similarly has 5 calyx appendages, but that species is a shrub, not an epiphyte, has larger flowers and berries with more than 20 rather than only 2 or 3 seeds. The calyx appendages of L. parasitica are usually not prominent in flower (Figs 3C, 4D). The berries of L. parasitica change from green to white, then orange or bright red during maturation (Fig. 5D), whereas those of L. laevis turn only from green to orange or red (Fig. 2B).

The minor leaves of the geminate pair are often deciduous in older stems of L. parasitica; this was the principal feature used by Merrill (1912) in distinguishing his new species S. epiphyticum.

In the Sandakan area of Sabah (Malaysian Borneo) several collections of L. parasitica (e.g., Evangelista 883, Pereira & Postar SAN-151226 and Sinclair 9347) are distinctive in being densely pubescent with simple uniseriate trichomes on all parts. These collections correspond to the glabrous individuals of L. parasitica in flower and leaf size and in numbers of seeds in berries, so I have retained them here, recognising that pubescence in Lycianthes in Asia is extremely variable. Further investigation of these populations would be of interest.

Solanum parasiticum was described in a footnote to a species list of plants found near the botanic gardens in Bogor (Blume 1823) without citation of specimens, nor were specimens cited in his later treatment (Blume 1826). No specimens collected by Blume are extant in the Bogor herbarium (A. Kartonegoro, pers. comm.). When he left Indonesia for the Netherlands Blume took with him several cases of specimens he and others had collected (van Steenis 1989). I have selected a neotype from among the specimens at Leiden (Blume s.n., L 0003603) corresponding to the description in the protologue.

Like other species described by Elmer (1910), no herbarium was cited in the protologue of S. angatii. I have selected the best preserved of the duplicates of the single collection cited (Elmer 10762) as the lectotype. This species was named in honour of Elmer’s Bagóbo guide Angat from the village of Todaya, for whom he had great praise (Elmer 1910).

Merrill (1912) described S. epiphyticum from several collections, citing one of them (“Cuming 873”) as “type”, but with no herbarium mentioned. There are no collections of Cuming 873 that are Solanaceae, but Cuming 837 is from the type locality, and I am considering Merrill’s (1912) citation of “Cuming 837” as an error to be corrected. Since the herbarium in Manila was burnt during the Second World War and all specimens held there were destroyed (see discussion under L. banahaensis), I have selected the sheet at BM (BM001018999) as the lectotype; it has well-preserved material with both flowers and fruits.

Bitter (1919) synonymised S. angatii and S. epiphyticum under his concept of L. parastica, and described two additional infraspecific taxa, vars. campylorhachis and praelongipedicellata. He cited a single specimen from Bogor (“Koorders 40378β ! (hb. Bogor)”) for var. campylorhachis; I have selected the best preserved and most complete of the three duplicates of this collection (BO acc. # BO-1414436) as the lectotype. A collection made by the cousins Karl F. and Paul B. Sarasin (“it. celeb. Sarasinorum n. 375!”) from Berlin was cited by Bitter (1919) for var. praelongipedicellatum; Berlin specimens were destroyed and most of this material is lost (van Steenis Kruseman 1985). I have seen no duplicates of any Sarasin collections. A neotype should be sought from the area where the type was gathered (https://openlibrary.org/books/OL16289390M/Reisen_in_Celebes#overview), I have seen no specimens of L. parasitica from Sulawesi.

Bitter (1922) later described L. parasitica var. plurifolia, citing a single specimen in “herb. Delessert”, held in G; in the absence of any evidence that he could have seen others of this collection (Wenzel 500), I suggest the specimen at G (G00415812) is the holotype.

Based on Solanum rantonnetii Carrière

Shrubs 0.5–3 m tall, with multiple stems from the base, these arching and sometimes scandent and sprawling; stems 3–4-angled, the angles yellowish green in live plants and paler than the rest of the stem, sparsely to moderately pubescent with spreading transparent simple uniseriate 1–4-celled trichomes to 0.5 mm long, these occasionally forked or dendritic, glabrescent with age; new growth moderately pubescent with transparent simple uniseriate or occasionally dendritic trichomes like those of the stems; bark of older stems pale greyish brown, prominently angled. Sympodial units unifoliate or more usually difoliate, the leaves usually geminate, if paired the leaves similar in shape and size. Leaves simple; blades of major leaves (1)4–15.5 cm long, (0.5)3.5–7.5 cm wide, ovate, rhombic-elliptic, elliptic or occasionally almost lanceolate, broadest in the upper half or rarely at the middle, membranous, concolorous; adaxial surfaces sparsely and evenly pubescent with 1–3-celled simple uniseriate trichomes, these denser along the midrib; abaxial surfaces sparsely to moderately and evenly pubescent with 1–3-celled simple uniseriate trichomes, these denser along the midrib; principal veins 3–7 pairs, more pubescent than the lamina. drying yellowish green abaxially; base attenuate onto the petiole; margins entire or somewhat undulate; apex acute to acuminate; petiole 0.5–2.4(4) cm long, winged from the attenuate leaf base, pubescent with simple uniseriate (or occasionally dendritic) trichomes like those of the stems and leaves; blades of minor leaves similar in size and shape to those of the major leaves, or slightly smaller; petioles 0.5–3 cm long, winged. Inflorescences axillary fascicles with (1)2–7 flowers, pubescent with transparent trichomes like those of the new growth and stems; pedicels 1.2–1.7 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading at anthesis, sparsely to moderately and evenly pubescent with transparent simple (occasionally dendritic) uniseriate 1–3-celled trichomes like those of the stems, articulated at the base; pedicels scars tightly packed in the leaf axils. Buds ellipsoid to fusiform with pointed tips, the corolla more than halfway exserted from the calyx tube before anthesis. Flowers 5-merous, all apparently cosexual. Calyx with the tube 1.5–4 mm long, 2.5–4.5 mm wide, openly cup-shaped, with (5)10 linear subulate appendages of variable length 0.25–5.2 mm long, arising ca. 0.25–1 mm from the hyaline rim, usually alternating long and short, sparsely to moderately pubescent with simple trichomes like those of the pedicels. Corolla 1.2–2 cm in diameter, violet with the midveins dark purple and the centre yellow, rotate, lobed less than 1/10 of the way to the base, interpetalar tissue abundantly present, the lobes ca. 1 mm long, ca. 1 mm wide and mere acumens from the rotate corolla, glabrous on both surfaces except for the densely papillate, cucullate tips (acumens). Stamens unequal; filament tube minute; free portion of the filaments of two lengths, three long filaments 2–3 mm long, two short filaments 0.8–1.5 mm long, glabrous or adaxially pubescent with tangled weak-walled uniseriate simple trichomes; anthers ellipsoid and slightly curved, orange-yellow, glabrous, poricidal at the tips, the pores round, distally directed, not elongating to slits with age. Ovary conical, glabrous; style 3.5–5.5 mm long, slightly curved in the same direction as the anthers, glabrous; stigma slightly clavate and bilobed, the surface minutely papillate. Fruit a compressed-ellipsoid or compressed globose berry, 2–3 cm long, 1.3–1.5 cm in diameter (usually absent or smaller and seedless in cultivated plants), yellow or yellowish orange when mature, the pericarp glabrous, thin, shiny and translucent; fruiting pedicels 2.5–4 cm long, ca. 1.5 mm in diameter at the base, ca. 3 mm in diameter at the apex, somewhat woody, spreading or hanging from the weight of the berries; fruiting calyx a plate with the appendages somewhat longer than in flower, spreading and often broken off, stiff and woody. Seeds 20–100 per berry (many fewer in cultivated plants), 2–3.5 mm long, 1.5–3.5 mm wide, rounded and compressed, reddish tan, the surfaces minutely pitted, the testal cells with sinuate margins, “hairy” extensions of lateral testal cell walls absent. Stone cells more than 20 per berry, ca. 0.5–1.5 mm in diameter. Chromosome number: 2n = 24 (Gerasimenko and Reznikova 1968 [cited in D’Arcy 1974] as Solanum rantonnetii; Acosta et al. 2005, as L. rantonnei, voucher Moscone 4260 [CORD]).

Lycianthes rantonnetii is widely cultivated in the tropics and subtropics (and even into the temperate zone as a short-lived perennial) worldwide. In this region I have only seen specimens from India and Pakistan. It is native to southern South America (Argentina, Bolivia, southern Brazil and Paraguay).

In its native range L. rantonnetii is a plant of semi-moist, seasonal forests and open areas; from (sea level) 100 to 2,000 m elevation.

In its native range in Argentina L. rantonnetii is called meloncillo del aire (Barboza 2013).

(IUCN 2020). Not applicable to this species for this region.

Lycianthes rantonnetii is native to South America (Barboza 2013) but widely cultivated in subtropical and temperate areas worldwide. It is the only species occurring in the region treated here that has stone cells in the berries, but in cultivation it rarely sets fruit. It can easily be distinguished from all native species by its rotate corollas with copious interpetalar tissue (Fig. 4E), orange-yellow anthers that are slightly curved and angled, and somewhat striped stems. Lycianthes rantonnetii has variable length calyx appendages like L. schizocalyx, but the two species are not easy to confuse; L. schizocalyx has small bright red berries without stone cells whereas L. rantonnetii rarely sets fruit in cultivation, but, when it does, the berries are dirty yellow and have copious stone cells. In Asia I have only seen specimens of L. rantonnetii from India and Pakistan, but I would expect it to be in cultivation anywhere in subtropical areas of Asia.

The specific epithet is often seen spelled “rantonnei” but is correctable to “rantonnetii” following Art. 60.9 of the ICN (Turland et al. 2018: Ex. 31), which stipulates that epithets honouring persons where there is an intentional latinisation of the name that involves the omission of a terminal vowel or consonant are not permitted; the epithet in this case honours the French horticulturalist Barthélémy Victor Rantonnet so is correctable to “rantonnetii” even though Carrière (1859) originally spelled it “rantonnei” (using the latinisation Rantonneus).

Based on Solanum schizocalyx Merr.

Shrubs, 0.75–2 m tall; stems terete, glabrous or variously pubescent with golden or whitish grey translucent simple uniseriate trichomes to 1.5 mm long, these appressed or spreading and tangled, older stems glabrescent; new growth glabrous to sparsely or densely pubescent with translucent, simple uniseriate 2–10-celled trichomes to 1.5 mm long; bark of older stems glabrescent, greyish brown or dark brown. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and occasionally in shape, the minor leaves if different in shape usually more ovate. Leaves simple; blades of major leaves 8–20 cm long, 2.5–6.5 cm wide, elliptic to narrowly elliptic, widest in the middle or just below, concolorous but occasionally somewhat discolorous, membranous; adaxial surfaces glabrous to evenly and sparsely to moderately pubescent with simple uniseriate trichomes like those of the stems, these denser along the veins, the lamina always visible; abaxial surfaces glabrous to evenly and moderately pubescent with simple uniseriate trichomes like those of the stems, the lamina clearly visible, the trichomes denser on the principal veins and midrib; principal veins 7–8 pairs, usually sparsely to moderately pubescent, occasionally glabrous. the venation not markedly prominent; base attenuate; margins entire, in pubescent plants ciliate with simple uniseriate trichomes to 1 mm long; apex acute to acuminate; petiole 1–3.5 cm long, glabrous or pubescent with simple uniseriate trichomes like those of the stems and new growth; blades of minor leaves 3–5 cm long, 1–2 cm wide, elliptic to narrowly elliptic or ovate, sometimes almost orbicular; surfaces like those of the major leaves; principal veins of minor leaves 5–6 pairs; base attenuate; margins entire; apex acute to acuminate; petiole of minor leaves 0.5–1.2 cm long, glabrous or sparsely pubescent like the stems. Inflorescences axillary, in fascicles, with 1–6 flowers, glabrous or with a few trichomes at the pedicel bases; pedicels at anthesis (1)2–3 cm long, ca. 1 mm in diameter at the base, 1.5–2.5 mm in diameter at the apex, deflexed (perhaps spreading?), glabrous to pubescent with simple uniseriate trichomes like those of the stems, articulated at the base; pedicel scars tightly packed in the leaf axils. Buds strongly tapered and pointed, the corolla strongly exserted from the calyx tube before anthesis, the calyx appendages enclosing the bud only partially. Flowers 5-merous, cosexual. Calyx tube 4–5 mm long, 4–5.5 mm wide at the mouth, an open cuplike structure, strongly 7–10-ridged, sometimes coloured purple or dull violet (fide Jacobs 7367), glabrous or pubescent with simple uniseriate trichomes like those of the pedicels, the trichomes often denser on the ridges, with 7–10 appendages arising at and ca. 0.5 mm below the hyaline rim, the appendages 1.5–5 mm long, ca, 0.5 mm wide, differing in length in the same flower, subulate, parallel to the calyx tube, glabrous or pubescent with simple uniseriate trichomes like those of the stems and pedicels. Corolla 2–3 cm in diameter, white, violet or purple, occasionally with a darker purple centre (fide Jacobs 7367), 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 9–13 mm long, 3–5.5 mm wide, spreading, glabrous on both surfaces, the tips cucullate and densely papillate. Stamens equal; filament tube minute; free portion of the filaments 1–1.5 mm long, glabrous; anthers 4–5 mm long, ca. 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 9–11 mm long, exserted from the anther cone, glabrous; stigma prominently capitate, the surfaces minutely papillate. Fruit a globose berry, 1–1.3 cm in diameter, green when immature, purple or bright red when mature, the pericarp glabrous, thin, shiny, transparent at fruit maturity; fruiting pedicels 2–3 cm long, ca.1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading; fruiting calyx not accrescent or expanding, but remaining a cup covering ca. 1/4 of the berry, the appendages spreading like a star beneath the berry. Seeds 40–60 per berry, 3.5–4 mm long, ca. 2.5 mm wide, flattened-reniform, pale straw-coloured or golden tan, the surfaces pitted, the testal cells sinuate in outline in the centre, rectangular on the margins, prominent “hairy” appendages absent. Stone cells absent. Chromosome number not known.

(Fig. 22). Lycianthes schizocalyx occurs in India, Indonesia, Malaysia, Myanmar (Burma), Philippines, Thailand and Vietnam.

Lycianthes schizocalyx is found in montane forests, evergreen forests and disturbed areas in these forests, usually growing in the understory in wet areas, from (100–)1,000 to 2,500 m elevation. Most collections are from above 1,000 m elevation.

Vietnam. cà ngủ ba vì (Hop 2017).

(IUCN 2020). EOO (7,731,851 km² - LC); AOO (368 km² - EN). Lycianthes schizocalyx is known from more than ten localities and is quite widely distributed in the region. Throughout the range it is known from protected areas (e.g., Mount Data National Park in the Philippines, Mount Kinabalu in Sabah, Malaysia). The assessment of Endangered (EN) 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).

Lycianthes schizocalyx is one of several Asian species of Lycianthes with more than 5 (usually 10) calyx appendages. It is most similar to L. biflora in its shrubby habit, the usually sublate or linear calyx appendages, flowers held below the leaves maturing to erect fruits, and in its elliptic, ovate or narrowly elliptic leaves. Merrill (Merrill and Merritt 1910) suggested L. schizocalyx was “manifestly allied” to L. biflora and distinguished it by “being nearly glabrous, with comparatively large flowers and its calyx splitting down one side.” Pubescence varies throughout the range of L. schizocalyx, as it does in most of these species, and the splitting of the calyx tube occurs often in L. schizocalyx but is not unique to the species.

Lycianthes schizocalyx can be distinguished from L. biflora – sometimes with difficulty – by its usually fewer-flowered fascicles, calyx appendages that differ in length within a flower and some of which arise from below the calyx tube rim (Fig. 3E), the usually strongly ribbed calyx tube (although this can be the case in L. biflora as well) and flowering pedicel length [(1)2–3 cm long in L. schizocalyx versus 0.9–1.2 cm long in L. biflora]. The flowering pedicel length is the most consistent character separating the two taxa, along with the insertion of the calyx appendages, but that can sometimes be difficult to see in poorly prepared material. In general, L. schizocalyx is found at higher elevations than L. biflora, but this is not consistent across the range of L. biflora. These two species may hybridise, but this has not been verified.

Lycianthes lysimachioides is sometimes confused with L. schizocalyx. The two species have similar calyx appendages; these reliably emerge below the rim in L. lysimachioides and often emerge below the rim in L. schizocalyx. Lycianthes schizocalyx can be distinguished from L. lysimachioides by its shrubby habit (versus prostrate herb rooting at the nodes), calyx appendages of variable length in a single flower (versus of equal length), and larger berries (1–1.3 cm versus 0.6–0.8 cm in diameter). In addition, L. lysimachioides usually has one-flowered inflorescences and leaves of a geminate pair often the same size and shape (appearing opposite). Lycianthes shunningensis also has 10 calyx appendages, but these are strongly reflexed; those of L. schizocalyx are erect or spreading.

Despite the difference in calyx appendage number L. schizocalyx (10 calyx appendages) can be very difficult to tell from L. laevis (5 calyx appendages) where they grow in sympatry. These two species appear to co-occur at the same locality in various parts of the species’ ranges (e.g., Williams 1334 and 1334 [A] from Philippines; Kuntze 4574, 4575 from Java). A specimen at NY purported to belong to the gathering Williams 1334 (here distinguished as Williams 1334 [A], NY barcode 01404894) represents a plant of L. schizocalyx; this specimen is annotated as a duplicate of 1334 (“Dup. 1334”) in an unknown hand and lacks specific locality data. The other two specimens numbered Williams 1334 are clearly L. laevis (see Suppl. material 2). The specimen of L. schizocalyx (NY barcode 01404894) may have been numbered a part of this gathering (Williams 1334) not by R.S. Williams, but by a herbarium curator at a later date. Numbers of calyx teeth are consistent but may be artificial, and population studies in areas where these plants co-occur are needed to really untangle this complex situation. The Vietnamese collection (Croat & Dzu 77996) cited by Hop (2017) as L. neesiana (here a synonym of L. laevis) is here identified as belonging to L. schizocalyx.

In describing S. biflorum var. corynephorum, Kuntze (1891) stated only “Java: Tjibodas”; as lectotype for this name I have selected the better of the two specimens of Kuntze 4574 with flowers and fruit in Kuntze’s herbarium held in NY (barcode 00172276) that is labelled with that locality and annotated as “var. corynephorum m” by Kuntze, along with an additional slip with details corresponding to the protologue.

Merrill (Merrill and Merritt 1910) cited several collections in the protologue of S. schizocalyx, indicating that Merrill 4548 was the type, but did not cite a herbarium. I have selected the US duplicate of Merrill 4548 (barcode 00027886, acc. # 71033) as the lectotype, as it is well-preserved, and Merrill was employed by the US government to work in the Philippines; the collections held in the Philippine National Herbarium were destroyed in World War II (Schultes 1957).

Bitter (1919) included the type of S. schizocalyx (Merrill 4548) in the syntypes cited for L. laevis var. glabratula, at the same time recognising L. schizocalyx as distinct, along with McGregor 8393, both from Berlin (“hb. Berol.”). To avoid creating homotypic synonyms, I have designated McGregor 8393 as the lectotype of var. glabratula, using the duplicate of this collection in the US National Herbarium (US 02840845, acc. # 628912). In his protologue for var. luzonensis Bitter (1919) cited Curran 5029 (a paratype of S. schizocalyx) and Elmer 6565, both from Berlin, calling Curran 5029 the “Hauptform” (main form). The US duplicate of Curran 5029 (strictly Curran in Herb. Bur. Sci 5029) is selected therefore as the lectotype of var. luzonensis (US 00027887, acc. # 708756). I have been unable to locate duplicates of the single collection cited (Bitter 1919) for L. laevis var. majuscula (Wichura 2168); it is here placed in synonymy based on the description.

Lycianthes baviensis was described (Hop 2004) citing as “Typus” the collection HPP 136 held in the Hanoi herbarium (HN). Three specimens of the gathering HPP 136 are held in HN, of these the one with the annotation as “type” and best corresponding to the line drawing in the protologue is selected as the lectotype.

China. Yunnan: “Shunning, Hila” [Fenqing District, Shun Ning Lu], 2,180 m, 24 Jun 1938, T.T. Yu [Yu Dejun] 16455 (holotype: KUN [acc. # 230263]; isotypes: A [00077123], E [E00792527], PE [00633443, 00633444]).

Shrubs or herbs, straggly or lax, sometimes described as climbing, ca. 1 m tall; stems terete, glabrous or sparsely pubescent with yellowish cream translucent simple uniseriate 2–8-celled trichomes to 1 mm long, soon glabrescent; new growth moderately to densely pubescent with translucent, simple uniseriate 2–8-celled trichomes to 1 mm long; bark of older stems glabrescent, yellow-green to greyish green. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in only size. Leaves simple; blades of major leaves 8–16 cm long, 2–6.5 cm wide, narrowly elliptic to elliptic, widest in the middle, concolorous, membranous; adaxial surfaces sparsely and evenly pubescent with simple uniseriate trichomes on the veins and lamina, the trichomes 2–3-celled, simple, uniseriate to 0.5 mm long, these often denser along the veins, the lamina always visible and the trichomes appearing depressed in dry specimens; abaxial surfaces glabrous or occasionally very sparsely pubescent with simple uniseriate trichomes like those of the adaxial surfaces, the midrib and principal veins with sparse pubescence; principal veins 6–8 pairs, sparsely pubescent or almost glabrous, the tertiary venation drying darker abaxially; base attenuate and somewhat decurrent onto the petiole; margins entire, ciliate with simple uniseriate trichomes to 0.5 mm long; apex acuminate; petiole 1.3–2.9 cm long, sparsely pubescent with simple uniseriate trichomes like those of the stems and new growth; blades of minor leaves 4–7.5 cm long, 1.5–3.5 cm wide, narrowly elliptic to elliptic; surfaces like those of the major leaves, the minor leaves often deciduous; principal veins of minor leaves 4–6 pairs; base attenuate; margins entire, usually somewhat ciliate; apex acuminate; petiole of minor leaves 0.6–0.8 cm long, sparsely pubescent like that of the major leaves. Inflorescences axillary, in fascicles, with 2–5(8) flowers, glabrous or with a few trichomes at the pedicel bases; pedicels at anthesis 0.8–1.1 cm long, 0.5–0.75 mm in diameter at the base, 1–1.5 mm in diameter at the apex, spreading, very sparsely 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 slightly exserted, strongly exserted from the calyx tube before anthesis, the calyx appendages usually more apparent in bud. Flowers 5-merous, cosexual. Calyx tube 2–3 mm long, 2.5–4.5 mm wide at the mouth, an open cuplike structure, not ridged, glabrous or very sparsely pubescent with simple uniseriate trichomes like those of the pedicels, with 10 appendages arising ca. 1 mm below the hyaline rim, these more visible in bud (especially when the appendages are short), the appendages 0.2–4 mm long, ca. 0.5 mm wide, small nubs to curved and linear, strongly deflexed and parallel to the calyx tube, glabrous or very sparsely pubescent with a few simple uniseriate trichomes like those of the stems and pedicels. Corolla 1.1–1.3 cm in diameter, white to variously purple, stellate, lobed 3/4 or nearly to the base, interpetalar tissue absent but a thin edge of tissue apparent on lobe margins, the lobes 5–6 mm long, 2–2.5 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 0.75–1 mm long, glabrous; anthers 2–2.5 mm long, ca. 1.25 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 5–6 mm long, exserted from the anther cone, glabrous; stigma prominently capitate, the surfaces minutely papillate. Fruit a globose berry, 0.8–1 cm in diameter, green when immature, bright red when mature, the pericarp glabrous, thin, shiny, and transparent at fruit maturity; fruiting pedicels 1.5–2 cm long, ca.1 mm in diameter at the base, ca. 2.5 mm in diameter at the apex, spreading or erect; fruiting calyx not accrescent or expanding, but remaining a plate-like structure, the appendages reflexed below berry in dry specimens, the appendages strongly reflexed and somewhat spreading. Seeds 20–40 per berry, 2–2.5 mm long, 1.5–2 mm wide, flattened-reniform or somewhat tear-drop shaped, pale straw-colored or yellow, the surfaces pitted only on the margins, the marginal testal cells rectangular to pentagonal, the lateral walls with “hairy” appendages ca. 0.2 mm long, the central testal cells sinuate. Stone cells absent. Chromosome number not known.