Lycianthes schizocalyx (Merr.) Bitter

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

9. Lycianthes schizocalyx (Merr.) Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 461. 1919.

Figs 3 E , 21

Solanum biflorum Lour. var. corynephorum Kuntze , Revis. Gen. Pl. 2: 453. 1891. Type. Indonesia. Java: “ Java: Tjibodas, 4,600 f ”, 25 May 1875, O. Kuntze 4574 (lectotype, designated here: NY [00172276]; isolectotype: NY [00172275 )).

Solanum schizocalyx Merr. , Philipp. J. Sci. 5 (C): 383. 1910. Type. Philippines. Luzon [Cordillera CAR]: Mount Data, District of Lepanto, Nov 1905, E. D. Merrill 4548 (no herbaria cited; lectotype, designated here: US [00027886, acc. # 71033]; isolectotypes: K [K 000759387], L [L. 2859466], NY [00138723], P [P 00368998]).

Lycianthes brachyanthera Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 473. 1919. Type. Indonesia. Sulawesi: [Sulawesi Utara], Lokon [Gunung Lokon, near Tomohon], s. d., P. B. Sarasin & F. K. Sarasin [VI 44 a] 396 (holotype: B [destroyed]; no duplicates found, in synonymy ex descr.).

Lycianthes minutipila Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 473. 1919. Type. Indonesia. Sumatra: [Aceh], Boernipaja [protologue “ Gajoe Loeas in Alas Landen ”], 16 Feb 1904, G. C. E. van Daalen & R. M. Pringgo Atmodjo 21 [cited only as van Daalen by Bitter] (lectotype, designated here: BO acc. # BO- 1324388]; isolectotype: BO [acc. # BO- 1414432]).

Lycianthes denticulata (Blume) Bitter var. liophylla Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 474. 1919. Type. Myanmar ( Burma). “ Tenasserim, Molyet ” [= Tanintharyi Region], 1,750 m, G. Gallatly 189 (holotype: B [destroyed]; lectotype, designated here: CAL [acc. # 315676]; isolectotype: CAL [no acc. #]).

Lycianthes laevis (Dunal) Bitter var. glabratula Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 488. 1919. Type. Philippines. Luzon [Cordillera CAR]: Pauai, prov. of Benguet, 2,100 m, Jun 1910, R. C. McGregor 8393 (lectotype, designated here: US [02840845, acc. # 628912]; isolectotype: B, [destroyed]).

Lycianthes laevis (Dunal) Bitter subsp. luzonensis Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 489. 1919. Type. Philippines. Luzon [Cordillera CAR]: Mount Santo Tomas [“ Mount Tonglon, prov. Benguet ”], Aug 1906, H. M. Curran 5029 (lectotype, designated here: US [00027887, acc. # 708756]; isolectotypes: B [destroyed], CAL [no acc. #]).

Lycianthes laevis (Dunal) Bitter var. majuscula Bitter , Abh. Naturwiss. Vereins Bremen 24 [preprint]: 488. 1919. Type. Indonesia. Java: “ West-Java. Berg Malabar ”, E. M. Wichura 2168 (holotype: B [destroyed]; no duplicates found, in synonymy ex descr.).

Lycianthes baviensis V. V. Hop , J. Biol. ( Vietnam) 26 (4 A): 44. 2004. Type. Vietnam. Ha Noi: Hà Tây, Mt. Ba Vi, 800–1,200 m, 24 Apr 1976, Nguyen Van Phu HPP 136 (lectotype, designated here: HN [sheet with label saying “ Typus ” in V. V. Hop hand ”]; isotypes: HN [two sheets, not with annotation as “ typus ”]).

Type.

Based on Solanum schizocalyx Merr.

Description.

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.

Distribution

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

Ecology and habitat.

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.

Common names.

Vietnam. cà ng ủ ba vì ( Hop 2017).

Preliminary conservation assessment

( IUCN 2020). EOO (7,731,851 km 2 - LC); AOO (368 km 2 - 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).

Discussion.

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. 3 E ), 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.

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 21. Lycianthes schizocalyx (Merr.) Bitter herbarium specimen. Indonesia. Sumatra, Robinson & Kloss 116 (BM 001018978). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

Figure 22. Distribution of Lycianthes schizocalyx.