Lycianthes shunningensis C. Y. Wu & S. C. Huang

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

10. Lycianthes shunningensis C. Y. Wu & S. C. Huang , Acta Phytotax. Sin. 16 (2): 77. 1978.

Figs 3 F , 4 F , 5 F , 23

Lycianthes subtruncata (Dunal) Bitter var. remotidens Bitter , Abh. Naturwiss. Verein Bremen 24 [preprint]: 479. 1919. Type. China. Yunnan: “ Szemao ” [Pu’er, Simao District], 27 Aug [no year], A. Henry 12352 A (lectotype, designated here: K [K 001152508]; isolectotypes: B [destroyed], E [E 00806597], MO [MO- 503793, acc. # 52819], US [02840661, acc. # 459052]).

Type.

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 [E 00792527], PE [00633443, 00633444]).

Description.

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.

Distribution

(Fig. 24 ). Lycianthes shunningensis occurs in China, India, Laos, Myanmar ( Burma), Thailand and Vietnam.

Ecology and habitat.

Lycianthes shunningensis grows in wet, evergreen forests, usually in the shaded understory, from 350 to 2,800 m elevation.

Common names.

China. jie chi hong si xian (as L. neesiana ), shun ning hong si xian ( Zhang et al. 1994). Vietnam. cà ng ủ nees (as L. neesiana ) ( Hop 2017).

Preliminary conservation assessment

( IUCN 2020). EOO (1,388,419 km 2 - LC); AOO (72 km 2 - EN). Lycianthes shunningensis is known from ten localities and is somewhat widely distributed in the region. It is known from protected areas (e. g., Doi Inthanon National Park in Thailand, Ta Xua Nature Reserve in Vietnam). The assessment of Vulnerable (EN) based on AOO is likely due to collecting bias. I therefore assign it a preliminary status of Least Concern (LC).

Discussion.

Lycianthes shunningensis is recognised here with some hesitation, and the material included here may in fact belong to two different taxa. The specimens seen and grouped here all have strongly reflexed calyx appendages arising from usually more than 1 mm below the calyx rim (Figs 3 F , 4 F , 5 F ). This morphology is also seen in L. connata J. L. Gentry of Mexico and Guatemala ( Dean et al. 2020: 76–78). The appendages in L. shunningensis vary in length, those on the type specimen (Yu 16455) from Yunnan in China and Averyanov et al. AL 887 from Laos (Figs 3 F , 5 F ) are long and conspicuous, while those on the type of var. remotidens (Henry 12352 A) and other collections are small nubs and quite inconspicuous (Fig. 4 F ); all, however, are strongly reflexed and arise well below the calyx rim. Variation in calyx appendage length is common in other Asian Lycianthes species (e. g. L. laevis , see description and discussion). On all of the specimens I have seen the leaf pubescence of L. shunningensis is also distinctive in being confined or nearly confined to the adaxial surface, most of the other taxa treated here when pubescent have denser pubescence on the abaxial leaf surfaces.

Lycianthes shunningensis occurs together with L. biflora (Yu 16404) at the type locality, and field studies of these populations would be useful in assessing the distinctness and consistency of the reflexed calyx appendage character that distinguishes L. shunningensis . A specimen of L. biflora (Henry 12911, US duplicate) was annotated as L. subtruncata var. remotidens by Bitter in 1925, but it does not have the reflexed calyx appendages and otherwise conforms to my concept of L. biflora here. These Lycianthes species are very difficult to identify!

At least some of the material identified and treated as L. neesiana (here a synonym of L. laevis ) by Zhang et al. (1994) and Hop (2017) corresponds to the type of L. subtruncata var. remotidens with 10 reflexed calyx appendages; the type of S. neesianum (see under L. laevis ) has 5 calyx appendages.

Lycianthes subtruncata var. remotidens was described ( Bitter 1919) based on Meebold 7342 (from “ hb. Berol., Vratisl. ” – both destroyed, and no duplicates found), a collection from an unnamed collector (“ Sammler des Bot. Gart. Calcutta! ”) from Bogor, and Henry 12352 A from “ hb. Berol. ” Nothing corresponding to the cited Bogor specimen has been found in BO. Bitter (1919: 480) distinguished the Henry 12352 A collection as the basic form of the variety (“ als Grundform dieser Varietät anzusehen! ” and I have selected the Kew duplicate of this collection (K 001152508) as the lectotype.

Excluded taxa

Solanum philippinense Merr., Philipp. J. Sci., Bot. 7: 351. 1912. Type. Philippines. Luzon: Cordillera ( CAR) Mount Santo Tomas, 1 Jul 1904, R. S. Williams 1275 (syntypes: K [K 000759468], US [00027740, acc. # 707522]).

= Tubocapsicum anomalum (Franch. & Sav.) Makino

Note. Merrill (1912: 352) suggested this species was “ allied to Solanum blumei Nees ” (= L. laevis ). He cited several collections in this description and indicated that Williams 1275 was the type, but without citing a herbarium. All the collections that Merrill (1912) cited that I have seen are referrable to Tubocapsicum anomalum (Elmer 6561, Merrill 8003, 8164; I have not seen a duplicate of Ramos 5406).

Names (designations) not validly published

Lycianthes biflora (Lour.) Bitter var. glabra (Koidz. ex Hatus.) Hatus., Fl. Ryukus , ed. 2, 193. 1994, as “ biflorum var. glabrum ”. Not validly published, no complete citation of basionym place of publication (Art. 41.5, Turland et al. 2018). Based on Solanum biflorum Lour. var. glabrum Hatus. = L. biflora (Lour.) Bitter

Solanum crassipetalum Wall., A numerical list of dried specimens of plants in the East India Company’s Museum 80. 1828–1849, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). “ Wallich Cat. ” 2618. = L. laevis (Dunal) Bitter

Solanum decemdentatum Wall., A numerical list of dried specimens of plants in the East India Company’s Museum 80. 1828–1849, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). “ Wallich Cat. ” 2614. = L. biflora (Lour.) Bitter

Solanum floccosum Zipp. ex Span., Linnaea 15 (4) 337. 1841, nomen nudum. No description or diagnosis (Art 38.1, Turland et al. 2018). = L. bimensis (Miq.) Bitter.

Solanum lysimachioides Wall., A numerical list of dried specimens of plants in the East India Company’s Museum 80. 1828–1849, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). “ Wallich Cat. ” 2609. = L. lysimachioides (Wall.) Bitter

Solanum macrodon Wall., A numerical list of dried specimens of plants in the East India Company’s Museum 80. 1828–1849, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). “ Wallich Cat. ” 2621. = L. biflora (Lour.) Bitter

Solanum membranaceum Wall., A numerical list of dried specimens of plants in the East India Company’s Museum 81. 1828–1849, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). “ Wallich Cat. ” 2625. = L. laevis (Dunal) Bitter

Solanum minahassae Koord., Meded. Lands. Plantentuin 19: 547. 1989, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). Based on specimen labelled “ Solanum arboreum Teysm. et Binn. msc. in H. H. B. ” at Bogor (Koorders 18030 β, 18049 β, 18152 β all annotated as “ Solanum arboreum Teysm. et Binn. ” at BO). = L. banahaensis (Elmer) Bitter

Solanum subtruncatum Wall., A numerical list of dried specimens of plants in the East India Company’s Museum 80. 1828–1849, nomen nudum. No description or diagnosis (Art. 38.1, Turland et al. 2018). “ Wallich Cat. ” 2620. = L. laevis (Dunal) Bitter

Solanum urbanum Morong var. typicum Chodat, Bull. Soc. Bot. Genève, ser. 2, 8: 151. 1916, not validly published. Use of prohibited epithet (Art. 24.3, Turland et al. 2018). = L. rantonnetii (Carrière) Bitter

Figures

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

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

Figure 5. Fruits of Lycianthes in Asia A L. banahaensis showing the opaque pericarp, lenticellate pedicel and small nub-like calyx appendages (Rule s. n., Philippines; DOL 134667) B L. biflora showing sequence of fruiting ripening and elongate calyx appendages (Nuraliev 182 a, Vietnam) C L. biflora plant with short calyx appendages and little pubescence (Pelser & Barcelona s. n., Philippines; DOL 44710) D L. parasitica showing the small inflorescence axes and berries changing from white to orange with ripening (Tandang s. n., Philippines; DOL 76584) E L. parasitica cross-section of berry showing the two seeds (Pelser & Barcelona s. n., Philippines; DOL 84746) F L. shunningensis showing unripe berry and strongly reflexed calyx appendages (Averyanov et al. 887, Laos). Photograph credits: A Greg Rule B Maxim Nuraliev C, E Pieter B. Pelser & Julie F. Barcelona D Danilo Tandang F Leonid Averyanov. DOL numbers are identifiers from PhytoImages (https://phytoimages.siu.edu). Details of collecting localities can be found in Suppl. material 2.

Figure 23. Lycianthes shunningensis C. Y. Wu & S. C. Huang herbarium specimen. China, Yunnan, Yu 16455 (E 00792527; isotype of L. shunningensis). Courtesy of Royal Botanic Garden, Edinburgh, reproduced with permission.

Figure 24. Distribution of Lycianthes shunningensis.