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

Knapp, Sandra, 2022, A revision of Lycianthes (Solanaceae) in Australia, New Guinea, and the Pacific, PhytoKeys 209, pp. 1-134 : 1

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

Figs 3C , 3E , 10 , 11

Solanum biflorum Lour., Fl. Cochinch. 129. 1790. Type. China. Guangzhou: "Pakwan supra Cantonem", Jul 1869, H.F. Hance 2128 (neotype, designated by Hul and Dy Phon 2014, pg. 57: P [P00058796]; isoneotypes: P [P00058797]).

Type.

Based on Solanum biflorum Lour.

Description.

Small shrubs or herbs, 0.5-1.5 m tall; stems terete, sparsely to densely pubescent with a mixture of transparent simple and/or forked or dendritic 3-10-celled uniseriate trichomes to 1 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 not in shape. Leaves simple; blades of major leaves 5-16 cm long, 2.5-6.5 cm wide, ovate to elliptic, usually widest in the lower half but occasionally near the middle, somewhat discolorous, membranous; 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; principal veins 4-6 pairs, sparsely to densely pubescent, often drying yellow on both surfaces; base attenuate, markedly decurrent onto the petiole; margins entire, markedly ciliate with transparent and 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 2.5-5 cm long, 1.5-3 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 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 perfect. Calyx tube 2-3 mm long, 2.5-3.5 mm in diameter, conical to openly cup-shaped, sparsely to densely pubescent like the stems and pedicels, with 10(12) linear awl-like appendages 1-5 mm long at anthesis, these variable in length even in a single flower, the appendages emerging at the rim or to 0.5 mm below, pubescent like the rest of the calyx. Corolla 1.4-1.8 cm in diameter, white or lavender with a green central area, often as two green dots at the base of each lobe, stellate, lobed nearly to the base, interpetalar tissue absent, 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 3-3.5 mm long, 1-1.5 mm wide, ellipsoid, the tips slightly pointed, yellow, 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 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 (see Fig. 2C ). 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. Stone cells absent. Chromosome number: n=24 ( Symon 1985; based on Kairo & Symon 10652).

Distribution

(Fig. 12 ). Lycianthes biflora is a widely distributed species in southeast Asia, ranging from India and China through much of Indonesia; the island of New Britain in the Bismarck Archipelago represents the easternmost edge of its range. On the island of New Guinea it is widely distributed in both Papua New Guinea and Indonesia.

Ecology and habitat.

Lycianthes biflora is a plant of disturbed habitats. Often described as a weed, it grows in secondary forests, along stream beds and roads, in burn regrowth and in the vicinity of agricultural fields, from 100 to 1,400 m elevation.

Common names.

None recorded from this region (many vernacular names are known from elsewhere in the species range).

Preliminary conservation assessment

( IUCN 2020). EOO (region treated here only 3,560,380 km2 - LC); AOO (region treated here only 108 km2 - EN). Lycianthes biflora is widely distributed within the region treated here and more broadly; this coupled with its weedy nature suggests a preliminary threat status of Least Concern (LC).

Discussion.

Lycianthes biflora is a widely distributed species throughout tropical Asia, occurring north to Japan and east to New Britain. Bitter (1919) treated it as a species complex ( “Gesamtart” = inclusive species) but nevertheless described many infraspecific variations from across its range based on small differences in pubescence, leaf shape and distribution. Material from New Guinea and the Solomon Islands matches the neotype specimen from China selected by Hul and Dy Phon (2014) and no infraspecific taxa or synonyms have been described based on material from Australia, New Guinea or the Pacific Islands. Therefore, full synonymy for L. biflora has been left to a future monograph treating Lycianthes in Asia (S. Knapp, in prep.). Specimens identified as L. biflora in the file Suppl. material 3 are preliminary and may be changed.

Lycianthes biflora is a small, weak-stemmed shrub, usually growing in open places. Even in New Guinea it is very variable in degree and type of pubescence, with individuals ranging from densely pubescent with mixed branched and simple trichomes to individuals that are almost glabrous. Length of calyx appendages vary between individuals and also within a flower; calyx appendages can be the same or slightly different lengths within the same flower and vary in length between individual plants. The length of pedicels in fruit also varies across the species range, as does number of flowers in a fascicle; despite its name, plants of L. biflora do not always have two flowers per fascicle but can have up to six. In the field, the plants are distinctive, with the flowers hanging below the leaves and the pedicels becoming erect through fruit maturation with ripe fruits held above the leaves, where they are exposed to their dispersers (probably birds, although this has not been verified in the field).

On New Guinea, Lycianthes biflora could be most easily confused with L. bitteriana , a similarly shrubby species of open areas. Lycianthes biflora can be distinguished from L. bitteriana in its red versus blackish purple berries, its branched trichomes that are loose and shaped like deer anthers versus congested branched trichomes that look like Christmas trees. Plants of L. biflora are generally smaller and less robust than those of L. bitteriana .

Like many other species of Solanaceae that are widespread, small differences in morphology can look very different when looked at in isolation but become difficult to disentangle when morphology is examined across a wide geographical range (e.g., see Knapp 2013; Särkinen et al. 2018; Knapp et al. 2019). Future studies on the genetics and sequence variability of the considerable morphological variation of L. biflora across its range are needed.

Specimens examined.

Australia. Christmas Island: middle of Island , Lombok utan, 1898, Andrews s.n. (BM, K); Aug 1908, Andrews 181 (BM); Christmas Island, (So of Java), 20 Nov 1888, Lister s.n. (K); Aug 1980, Powell 164 (K); Phosphate Hill, Oct 1904, Ridley 34 (K) .

Indonesia. Maluku: Buru, NW Buru, S. of Bara, Waeduna River , 350-400 m, 16 Nov 1984, van Balgooy 4916 (A); Seram, Manusela National Park , along a trail between Hatumete (sea level) and Hoale Pass (1,770 m) southern slope of Murkele Ridge, Kecamatan District , Tehoru; C. Seram, 550-1,200 m, 20 Feb 1985, Kato et al. C-7273 (A). Maluku Utara: Bacan, Babang, Kec. Labuha, 100 m, 26 Aug 1986, Ramlanto 869 (K, L); North Maluku, Gunung Sibela, N Moluccas, Bacan Island, Gunung Sibela near Waiaua , 1,000 m, 23 Oct 1974, de Vogel 3565 (L, LAE, MO); Gunung Sibela, N Moluccas, Bacan Island, Gunung Sibela near Waiaua , 250 m, 28 Oct 1974, de Vogel 3718 (L, MO). Papua: NE Kepal Burung, Kabupaten Manokwari, Kecamantan Manokwari, mountains S of the Arfak Plains , steep ridges between Arfak Plains and Gunung Itsiwei, 550 m, 29 Apr 1994, Sands et al. 6431 (K). Papua Barat (West Papua): Andjai [Andaj], Kebar Valley , 600 m, 7 Sep 1959, Moll BW-9529 (L); North East Kepala Burung , Kabupatem Manokwari, Arfak Mountains , Mupi Dessa, trail from Mupi village to G[unung] Humibou, near Sungai Mupi between confluence of Kali Ngwes and Sungai Mupi and site of Kamnpong Mubri Lama, 875 m, 14 Apr 1995, Sands & Maturbongs 6791 (A, K); Wondiwoi Mountains , Wandammen Peninsula, 300 m, 24 Feb 1962, Schram BW-10645 (L); Wondiwoi Mountains , Wandammen Peninsula, 350 m, 28 Feb 1962, Schram BW-10744 (L, WAG) .

Papua New Guinea. Bismarck Archipel., 1889, Warburg 21250 (BM). Central: Boridi, 914 m, 16 Nov 1935, Carr 14991 (BM, K, L, NY); [Merska Hills] Sogeri Region., 762 m, 10 Apr 1886, Forbes, H.O. 882 (BM, CAL, MEL, P); subdistrict Port Moresby, on ridge below Boridi village, 920 m, 1 Oct 1973, Foreman & Vinas LAE-60242 (A, LAE); NE of Manumu Village, subdistrict Port Moresby, 450 m, 16 Sep 1973, Isles & Vinas NGF-33899 (L). East New Britain: near Mapping site at edge of Mengen Massif, subdistrict Pomio, 885 m, 9 Jun 1973, Stevens & Lelean LAE-58668 (E, K, LAE); Gazelle Peninsula, Baining Mountains, bulldozer track into the Wild Dog Prospect at Mt Sinvit, 950 m, 10 Feb 2004, Takeuchi et al. 16902 (A, K, L). Eastern Highlands: Kassam Pass, Kainantu subdistrict, 1,280 m, 9 Jan 1988, Henty et al. NGF-29203 (K, LAE); Crater M[ountain] Wildlife Management Area, Kusare, near the El Niño burn area, 1400 m, 28 Jul 1998, Takeuchi 12688 (A, K, LAE). Madang: "Kaiser Wilhelmsland, waldranderam oberen Djamu" [northern New Guinea], 700 m, 9 Feb 1908, Schlechter 17305 (P). Milne Bay: Biniguni Camp, Gwariu River, 200 m, 12 Aug 1953, Brass 23978 (A, LAE). Morobe: 1955 Planting area, Bulolo. Morobe District, T.N.G., 1,067 m, 9 Jun 1955, Floyd 7459 (BM, K, LAE, US), 15 Jun 1955, Floyd 7509 (BM, K, LAE); Mount Missan, C.N.G.T. Logging areas, Stoney Creek, on foot slopes of Mount Missan (near Bulolo), Wau subdist., 914-1,219 m, 1 Jun 1977, Kairo & Symon 10652 (K, LAE); Bulolo, Middle Logging Area, subdistrict Wau, 853 m, 10 Aug 1966, Kairo & Streimann NGF-27869 (K, LAE); Oomsis Creek, Markham valley, 500 m, 3 Feb 1960, Millar NGF-11795 (K, LAE); Oomsis Creek, Markham Valley, 152 m, 3 Feb 1960, Millar NGF-11794 (A); Finschaffen, 300 m, 5 Jul 1978, Rau 380 (A, L); Hump L.A. 5 mi SE Bulolo, Wau subdistrict, 1,067 m, 15 Mar 1971, Streimann & Kairo NGF-25853 (A, K, LAE); Bulolo, 914 m, Jan 1957, Wells NGF-7569 (A, K, LAE). New Britain Island: New Britain, New Pommeron. Bei Mussawa., Nov 1901, Schlechter 13748 (BM, K, P); bei Mussawa, Nov 1901, Schlechter 13749 a, (K). Oro: Isuarava [Isurava], 5 Mar 1936, Carr 15965 (BM, L).

Figures

Figure 2. Representative morphology of Lycianthes A Lycianthes oliveriana with major and minor leaves differing only in size (Damas et al. SAJ- 1050, Papua New Guinea) B Lycianthes cladotrichota with major and minor leaves differing markedly in size and shape (James et al. SAJ- 1385, Papua New Guinea) C Lycianthes biflora in fruit with linear, awn-like calyx appendages (Knapp 10106, Yunnan, China) D Lycianthes oliveriana with urceolate calyces with no appendages (Damas et al. SAJ- 1050, Papua New Guinea) E Lycianthes biflora with stellate corollas and membranous lobes (Knapp 10106, Yunnan, China) F Lycianthes cladotrichota with stellate corollas and thick, fleshy lobes (short-styled plant, James et al. SAJ- 1385, Papua New Guinea). Photograph credits: A, B, D, F Shelley James; C, E Sandra Knapp.

Figure 3. Trichome diversity in New Guinea Lycianthes A simple (unbranched) stiff, antrorse trichomes (Streimann & Katik NGF- 34091) B simple (unbranched) " crisped " trichomes (Kalkmann BW- 3479) C dendritic (antler-like) trichomes (Streimann & Katik NGF- 34091) D dendritic (Christmas-tree like) trichomes (Wells NGF- 7565) E simple (unbranched) stiff, antrorse trichomes with enlarged bases (Millar NGF- 40737). Drawing by Lucy Smith.

Figure 10. Lycianthes biflora (Lour.) Bitter. Reproduced from Wight (1850: tab. 1397, as S. denticulatum Blume). Courtesy of the NHM Library and Archives reproduced with permission.

Figure 11. Lycianthes biflora herbarium specimen. Papua New Guinea. Morobe: Floyd NGF- 7509 (BM 001018933). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

Figure 12. Distribution of Lycianthes biflora (only for region treated in this monograph).