5.

Figs 2 C, 14

Solanum lysimachioides Wall., Fl. Ind. (Carey & Wallich ed.) 2: 257. 1824. Type. Nepal. [Bagmati prov.]: “ Chondaghery ” [Chandragiri], Feb 1821, N. Wallich Cat. 2609 (lectotype, designated by Deb 1978, pg. 293 [as “ type ”]: CAL [CAL 0000071943, acc. # 315728]; isolectotypes: BM [BM 001018867], K [K 000923249], K-W [K 001116562, K 001116563]).

Solanum caulorhizum Dunal, Prodr. [A. P. de Candolle] 13 (1): 181. 1852. Type. Indonesia. Java: “ prov. Badong ”, 1847, H. Zollinger 705 (holotype: G-DC [G 00145653]; isotypes: BM [BM 000778224], G [G 00301677], MPU [MPU 012644], P [P 00369006]).

Solanum nematosepalum Miq., Fl. Ned. Ind. 2: 643. 1857. Type. “ Solanum ciliatum Blume in herb. Reg. L. B. ” (no specimens cited or located); Indonesia. [Java]: sin. loc., C. L. Blume s. n. (neotype, designated here: L [L. 2881683]).

Solanum macrodon Wall. ex Nees var. lysimachioides (Wall.) C. B. Clarke, Fl. Brit. India [J. D. Hooker] 4: 232. 1883. Type. Based on Solanum lysimachioides Wall.

Lycianthes lysimachioides (Wall.) Bitter var. caulorhiza (Dunal) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 493. 1919. Type. Based on Solanum caulorhizum Dunal

Lycianthes lysimachioides (Wall.) Bitter var. sinensis Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 493. 1919. Type. China. Hubei: sin. loc., 1885, A. Henry 6080 (lectotype, designated here: BM [BM 001018842]; isolectotypes CAL [acc. # 316339, 316340], E [E 00426492], GH [no barcode], US [02840640, acc. # 801545]).

Lycianthes lysimachioides (Wall.) Bitter var. formosana Bitter, Repert. Spec. Nov. Regni Veg. 18: 320. 1922. Type. Taiwan. (“ Formosa ”) Arisan, 2,500 m, 1914, U. Faurie 640 (holotype: G [G 00415805]; isotypes: BM [BM 001018839], GH [no barcode]).

Solanum debilissimum Merr., Philipp. J. Sci. 23: 265. 1923. Type. China. Hainan: “ Ng Chi Leng [Five-Finger Mountain] ”, 11 May 1922, F. A. McClure 9532 (no herbarium cited; lectotype, designated here: E [E 00426499]; isolectotypes: A [0077823], BISH [BISH 1005077, acc. # 182918], BM [BM 001018850], K [K 000759408], MO [MO- 503598, acc. # 915771]).

Numaeacampa kerrii Gagnep., Bull. Soc. Bot. France 95 (1): 33. 1948. Type. Laos. Vienchan: Pu Tat (Phu That), “ Wiengchan ” [Vienchan], 21 Apr 1932, A. F. G. Kerr 21186 (holotype: P [P 00236810]; isotypes: BM [BM 001018901], K [K 000923331, K 000923332]).

Lycianthes solitaria C. Y. Wu & A. M. Lu, Acta Phytotax. Sin. 16 (2): 76. 1978, nom. illeg., not Lycianthes solitaria Standl. (1927). Type. China. Tibet: Zayu, 1,700 m, 14 Jul 1973, Qinghai-Tibet group 73-672 (holotype: KUN [acc. # 484269]; isotype: KUN [acc. # 484270]).

Lycianthes lysimachioides (Wall.) Bitter var. purpuriflora C. Y. Wu, Acta Phytotax. Sin. 16 (2): 79. 1978. Type. China. Sichuan: Emei, 1,200 m, 26 May 1935, Du Dahua 418 (lectotype: PE [00031385]; isolectotypes: IBK [IBK 00381736], PE [00013186, 0013187, 0031388]).

Lycianthes lysimachioides (Wall.) Bitter var. cordifolia C. Y. Wu & S. C. Huang, Acta Phytotax. Sin. 16 (2): 79. 1978. Type. China. Sichuan: Emei, 1952, J. Xiong [Xiong Jihua], X. Zhang [Zhang Xiushi] & X. Jiang [Jiang Xinglin] 32197 (holotype: PE [00031390, acc. # 267671]).

Lycianthes lysimachioides (Wall.) Bitter var. rotundifolia C. Y. Wu & S. C. Huang, Acta Phytotax. Sin. 16 (2): 79. 1978. Type. China. Shaanxi: Xianyang, 1,000 m, 23 Sep 1933, T. P. Wang [Wang Zuobin] 857 (lectotype, designated here: PE [00031306]; isolectotype: PE [00031307]).

Lycianthes tibetica Li Bing Zhang & Yi F. Duan, Phytotaxa 170 (4): 280. 2014. Type. Based on (replacement name for) Lycianthes solitaria C. Y. Wu & A. M. Lu

Type.

Based on Solanum lysimachioides Wall.

Description.

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.

Distribution

(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.

Ecology and habitat.

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.

Common names.

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 ng ủ d ạng trân châu (Hop 2017).

Preliminary conservation assessment

(IUCN 2020). EOO (14,512,768 km 2 - LC); AOO (372 km 2 - 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).

Discussion.

Lycianthes lysimachioides is distinctive amongst Asian species of Lycianthes in being a creeping herb of forest understories and shady places (Fig. 2 C); 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 K 0007507178 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. 2 C), 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 (P 00369007) 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 (BM 001018842) 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 (E 00426499) 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.