Lycianthes banahaensis (Elmer) Bitter

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

1. Lycianthes banahaensis (Elmer) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 509. 1919.

Figs 2 A , 4 A , 5 A , 6

Solanum banahaense Elmer , Leaflets Philipp. Bot. 1: 341. 1908. Type. Philippines. Luzon [Calabarzon]: Tayabas, Lucban, May 1906, A. D. E. Elmer 7492 (no herbaria cited; lectotype, designated here: A [00077843]; isolectotypes: CAL [CAL 0000018718], E [E 00273870], G [G 00343322, G 00415767], K [K 000759394], L [L 0003585], LAE [acc. # 229579], LE, US [00027470, acc. # 629692], W [acc. # 1910-0010370]).

Solanum lagunense Elmer , Leaflets Philipp. Bot. 1: 341. 1908. Type. Philippines. Luzon [Calabarzon]: Los Baños, prov. of Laguna, Apr 1906, A. D. E. Elmer 9425 (no herbaria cited; lectotype, designated here: K [K 000759392]; isolectotype: E [E 00426930], K [K 000759393], LAE [acc. # 229581]).

Solanum manucaling Elmer , Leaflets Philipp. Bot. 2: 732. 1910. Type. Philippines. Mindanao [Davao]: Todaya (Mount Apo), Davao del Sur, May 1909, A. D. E. Elmer 10489 (no herbaria cited; lectotype, designated here: BISH [BISH 1005081, acc. # 581198]; isolectotypes: A [00077848], BM [BM 000846477], CAL [acc. # 316482], E [E 00273862], G [G 00415768], GH [00077849], HBG [HBG- 511353], K [K 000759389], L [L 0003601], LE [LE 00016976, LE 00016977], MO [MO- 716043, acc. # 3717134], NY [00172286], US [00027669, acc. # 873056], W [acc. # 1912-0001339]).

Solanum anisophyllum Elmer , Leaflets Philipp. Bot. 8: 2830. 1915. Type. Philippines. Mindanao [Caraga]: Cabadbaran, Mount Urdaneta [= Mount Hilong-Hilong or Masay], Agusan del Norte, Sep 1912, A. D. E. Elmer 13887 (no herbaria cited; lectotype, designated here: A [00077840]; isolectotypes: BM [BM 0001014585], CAL [acc. # 316453], E [E 00196406], G [G 00415766], GH [00077841], K [K 000759392], L [L 0003597], LAE [acc. # 229574], LE [LE 00016833], MICH [1109888], P [P 00368935], U [U 0113983], US [00027453, acc. # 894985], W [acc. # 1915-0012862]).

Lycianthes aceratia Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 508. 1919. Type. Indonesia. East Nusa Tenggara: Sumba [“ Soemba, Parimbang ” in protologue], J. E. Teijsmann 8918 (holotype: BO [acc. # BO- 1323491]; isotype: L [L. 2859654]).

Lycianthes banahaensis (Elmer) Bitter subsp. manucaling (Elmer) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 509. 1919. Type. Based on Solanum manucaling Elmer

Lycianthes lagunensis (Elmer) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 511. 1919. Type. Based on Solanum lagunense Elmer

Lycianthes anisophylla (Elmer) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 512. 1919. Type. Based on Solanum anisophyllum Elmer

Lycianthes anisophylla (Elmer) Bitter var. masbateensis Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 513. 1919. Type. Philippines. Luzon [Bicol]: Island of Masbate, Aug 1903, E. D. Merrill 3046 (holotype: B [destroyed]; lectotype, here designated: US [0027867, acc. # 438017]; isolectotypes: BM [BM 001019006], K [K 000759391]).

Type.

Based on Solanum banahaense Elmer

Description.

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.

Distribution

(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.

Ecology and habitat.

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

Common names.

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

Preliminary conservation assessment

( IUCN 2020). EOO (1,652,883 km 2 - LC); AOO (228 km 2 - 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).

Discussion.

Lycianthes banahaensis is a small tree (Fig. 2 A ) 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. 4 A ); 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. 4 A ), 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 (K 000759392) 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 (BISH 1005081, 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.

Figures

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

Figure 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 6. Lycianthes banahaensis (Elmer) Bitter herbarium specimen. Philippines. Mindanao, Elmer 13782 (E 00593145, staminate specimen). Courtesy of Royal Botanic Garden, Edinburgh, reproduced with permission.

Figure 7. Distribution of Lycianthes banahaensis.