Solanum alpinum Zoll. & Moritzi, Natuur- Geneesk. Arch. Ned.- Indie 2: 571. 1845.

Saerkinen, Tiina, Poczai, Peter, Barboza, Gloria E., Weerden, Gerard M. van der, Baden, Maria & Knapp, Sandra, 2018, A revision of the Old World Black Nightshades (Morelloid clade of Solanum L., Solanaceae), PhytoKeys 106, pp. 1-223 : 1

1. Solanum alpinum Zoll. & Moritzi, Natuur- Geneesk. Arch. Ned.- Indie 2: 571. 1845. Figure 5

Solanum viscidissimum Zoll. & Moritzi, Natuur- Geneesk. Arch. Ned.- Indië 2: 571. 1845.

Solanum viscidissimum Type. Indonesia. Java: Tengger [Tengger Range], above Gebok-Klacca [Klakah], 5500 ft, Oct 1844, H. Zollinger 2514 (lectotype, designated here: G-DC [G00357866]; isolectotypes: BM [BM000778312], G [G00144593, G00343069], P [P00379791, P00379790], W [1889-0052800]).

Solanum dichrophyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 48. 1852, nom. illeg. superfl.

Solanum viscidissimum Type. Based on (renaming of) Solanum alpinum Zoll. & Moritzi

Solanum anacamptocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 59. 1852.

Solanum viscidissimum Type. Indonesia. Java: Waliran [Gunung Welirang], 27 Aug 1844, H. Zollinger 2177 (holotype: G-DC [G00144590]; isotypes: A [A00248838], BM [BM000886120], G [G00343304], MPU [MPU012652], P [P00369059, P00369060 [mixed sheet with right-hand stem belonging to S. nigrum ], P00368907, P00368908]).

Solanum bromoense Kuntze, Revis. Gen. Pl. 453. 1891.

Solanum viscidissimum Type. Indonesia. Java: Mount Tengger, Bromo [Gunung Bromo], 2000 m, 15 Sep 1875, O. Kuntze 5999 (lectotype, designated here: NY [0017228]; isolectotypes: GH [GH00077829], K [K000788117]).

Type.

Indonesia. Java: "in montibus Ardjune" [Gunung Arjuno], 2000-3000m, 14 Sep 1844, H. Zollinger 2255 (lectotype, designated here: P [P00368905] (original label on this sheet with date and locality - "an 2177 var?"); isolectotypes: BM [BM000886119], G [G00144225, G00301669], MPU [MPU014201], P [P00368906]) .

Description.

Annual or short-lived erect to somewhat spreading perennial herbs, height not known, likely subwoody and branching at base. Stems spreading to decumbent, terete, sometimes recorded as greenish-violet ( Afriastini 475), older stems yellowish-brown, with no prickle-like projections, not markedly hollow; new growth densely pubescent with simple, spreading, uniseriate, glandular or eglandular trichomes, the trichomes 6-8(-10)-celled, ca. 0.5 mm long, if glandular then with a terminal gland, sometimes drying with a yellowish-brown tinge. Sympodial units trifoliate to plurifoliate, the leaves not geminate. Leaves simple, (2-)5-12 cm long, (0.5-)1-3 cm wide, lanceolate to ovate, most commonly narrowly elliptic or broadly lanceolate, membranous, concolorous, smell not known; adaxial surface pubescent with simple, uniseriate, glandular or eglandular trichomes evenly and moderately spread along veins and lamina; abaxial surface similar but the pubescence denser along the veins; major veins 7-8 pairs; base cuneate to attenuate; margins entire or shallowly toothed, the teeth, if present, narrow and acute; apex acute to acuminate; petioles ca. 1 cm long, pubescent like the stems and leaves. Inflorescences 2-4 cm long, internodal, unbranched or furcate, with 5-10 flowers mostly clustered at the distal end of the rhachis, pubescent with simple uniseriate trichomes like those of the stems and leaves; peduncle 1.0-3.5 cm long, straight; pedicels 1.0-1.3 cm long, ca. 0.5 mm in diameter at base and apex, filiform, spreading, pubescent with simple uniseriate trichomes like the rest of the inflorescence, articulated at the base; pedicel scars mostly clustered at the distal end of the rhachis and overlapping, sometimes up to 1.5 mm apart in the distal half of the rhachis. Buds narrowly ellipsoid, the corolla soon exserted from the calyx tube. Flowers 5-merous, all perfect. Calyx tube 1-2 mm long, broadly conical, the lobes 0.5-1.0 mm long, ca. 1 mm wide, broadly deltate to triangular, tips acute, densely to moderately pubescent with simple uniseriate 6-8-celled trichomes. Corolla 9-12 mm in diameter, purple or violet, stellate, lobed 2/3 to 3/4 of the way to the base, the lobes 3-4.5 mm long, 1.5-2.0 mm wide, spreading or reflexed, densely papillate along the margins and on the tips. Stamens equal; filament tube <0.2 mm long; free portion of the filaments ca. 1 mm long, densely pubescent adaxially with tangled simple uniseriate trichomes; anthers 2.8-4 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 5-6 mm long, densely pubescent with tangled simple uniseriate trichomes in the basal half, exserted up to 2 mm from the anther cone; stigma minutely bilobed, the surfaces minutely papillate. Fruit a globose berry, 6-9 mm in diameter, black, the pericarp thin and matte; fruiting pedicels 1.2-1.3 cm long, ca. 0.5 mm in diameter at the base and apex, spreading or strongly deflexed, spaced (0-)0.5-1.5 mm apart, not falling with the berry, persistent on older inflorescences; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes ca. 1 mm long, loosely appressed to the berry, not markedly reflexed. Seeds 20-50 per berry, 2.5-3.0 mm long, ca. 1 mm wide, flattened reniform, golden brown to brown, the surfaces minutely pitted, the testal cells small, rectangular to pentagonal in shape. Stone cells (0)2-4 per berry. Chromosome number not known.

Distribution

(Figure 6 ). Endemic to Indonesia and is found along the mountainous volcanic spine of Java, Bali and Lombok (Lesser Sunda Islands).

Ecology.

Grows in montane habitats, in grassy areas along forest margins or in clearings; between 1,700 and 3,100 m elevation.

Common names.

Indonesia: ranti hutan.

Uses. No specific uses recorded, but labels ( Afriastini 475) indicate young berries are "edible but rather bitter".

Preliminary conservation status

( IUCN 2016). Solanum alpinum , although found over a relatively large geographical area, is never common and is known from a small number of collections. With a relatively small AOO of 64 km2 (EN) and EOO of 18,806 km2 (VU) and, in view of the scattered populations at isolated high elevations, we assign the species a preliminary conservation status of VU (Vulnerable; Table 7 ). Forests on these mountains are under threat from land conversion and fire associated with human use ( van Steenis 2006).

Discussion.

Solanum alpinum is a plant of high elevations with both glandular and eglandular morphs. Glandular forms were named as S. viscidissimum , while in the same publication eglandular forms were named S. alpinum . As the two names have not been synonymised before (neither name was included in Backer and Bakhuisen van der Brink 1965 or in van Steenis 2006), we have chosen to use S. alpinum over S. viscidissimum because the name is more generic and appropriate for the distribution of the species. It can be distinguished from S. americanum and S. nigrum , both of which also occur in Indonesia, by its larger usually violet flowers on long pedunculate inflorescences. Anthers of S. alpinum are 2.8-3.8(-4) mm long, while those of S. americanum are minute (ca. 1.5 mm) and those of S. nigrum ca. 2 mm long. The leaves are usually narrower and longer (more lanceolate) than either of those two species, but two Horsfield collections with smaller, rounder leaves are included here based on anther length and flower size, so variability could be greater than we have seen to date.

The plant illustrated as "Solanum nigrum" in van Steenis (2006: plate 51) is likely to be S. alpinum ; the excellent illustration has the narrow leaves, long pedicels and large flowers characteristic of that species. The dismissal of it as a probable introduction has been the fate of all of the more narrowly distributed members of this group. The distribution of S. alpinum along the volcanic chain of Java above 2,000 m elevation is typical of other members of this rich flora that are found in areas created and influenced by fire resulting from volcanism and subsequent open habitat and grassland creation (as described in van Steenis 2006). Solanum alpinum is likely to also occur in the "tjemara forests" (sensu van Steenis 2006) dominated by Casuarina junghuhniana Miq. ( Casuarinaceae ). Further fieldwork and more accurately geolocated collections are needed to ascertain the distribution and habitat preferences of S. alpinum .

A single collection of Zollinger was cited for each of S. alpinum ("Herb. N. 2255") and S. viscidissimum ("Herb. N. 2514"), but no herbarium was cited; we have lectotypified S. alpinum with the P (P00368905) duplicate of Zollinger 2255, because it has a label with the protologue locality. We have selected the G-DC (G00357866) sheet of Zollinger 2514 as the lectotype for S. viscidissimum as the best preserved of the duplicates we have seen.

Dunal (1852) illegitimately coined the name S. dichrophyllum for S. alpinum because he felt the Zollinger’s and Moritzi’s name was inappropriate as the plant did not grow in the Alps ("Nomen Solanum alpinum mutavi quia in montibus certe alpinus non habitat" - I have changed the name because this certainly does not inhabit the Alps). He ( Dunal 1852) used another sheet collected by Zollinger (Zollinger 2177) as the basis for his S. anacamptocarpum .

C.E.O. Kuntze named S. bromoense for Gunung Bromo in the Tenggar range, citing only "Java. Bromo 2000m" ( Kuntze 1891). We have selected the sheet in NY (NY0017228) bearing the same locaity as the lectotype; Kuntze’s personal herbarium is held in NY.

Specimens examined.

Indonesia. Bali: bei der Quelle Jaritie auf Weg zum Gunung Ajaung, 2 Jun 1912, Arens 19 (L); Kleine Soenda Eilanden, Bali, Z. helling Agoeng, 6 Apr 1936, van Steenis 7839 (K) ;. Java: Central Java, Blumbang, Mt. Lawu, Central Java, 26 Nov 1982, Afriastini 475 (A); West Java, Mt.Malabar , Anderson 369 (CAL); East Java, Ardjoeno, tjemarabosch boven Lalidjiwo, 17 Oct 1915, Arens s.n. (L); East Java, 12 Oct 1915, Arens 48 (L); East Java, Pasoeroean, G [unung] Tengge, boven Tosari, 4 Jun 1913, Backer 8380 (L); East Java, Te Pasoeroean, Ngadisari, Jan 1925, Backer 36563 (A); East Java, Pasoeroean, Tengge, boven Tosari, Backer 36564 (L); Central Java, Soerkarta, Top van de Lawoe [Mount Lawu], 16 Jul 1936, Brinkman 754 (NY); Sitiebondo, G [unung] Raneg [Raoeng] via Brembeinri, 15 May 1932, Clason-Laarman EHH - 157 (L); East Java, SE Java, 1880, Forbes 1019 (BM); Central Java, Central Java, Slamet Mountain , between base camp Beraba-top of summit, 17 Mar 2004, Hoover et al. Deden-113 (A); Central Java, Mt. Prahu , Horsfield s.n. (BM); Central Java, Surakarta, Horsfield s.n. (BM); Central Java, Mt. Prahu , Horsfield s.n. (BM); Central Java, Blambangan & Mt. Prahu , Horsfield s.n. (BM); East Java, Pasoeroean G [unung] Smeroe, Jesweit s.n. (L); Central Java, Tenggu, Penandjaan, 31 Jul 1932, Kleinhoonte 371 (L); Central Java, Besoeki, G [unung] Merapi, 18 Jul 1916, Koorders & Koorders-Schumacher 43218 B (L); East Java, Res. Besoeki Kawah Idjen, 1 Nov 1916, Koorders & Koorders-Schumacher 43214B (L); East Java, Besoeki, Jang Plateau, 12 Aug 1916, Koorders & Koorders-Schumacher 43463 (L); East Java, Tdjan plateau, Mt. Raung , Nov 1938, Kostermans s.n. (L); Central Java, G[unung] Muria, Tjollo, N of Kudus, 25 Nov 1951, Kostermans 6278 (L); West Java, Malawar, Dec 1875, Kuntze 5410 (DUKE, NY); West Java, Gunong Gede, 6 Mar 1979, Murata et al. 2021 (L); East Java, Pasoeroean, 4 Jun 1935, Neth Ind Forest Service 7038 (L); East Java, Pasoeroean, 4 Jun 1935, Neth Ind Forest Service 7056 (A); East Java, Tosari 49, Proefstation voor de Javasuikerindustrie s.n. (L); East Java, Tosari 52, Proefstation voor de Javasuikerindustrie s.n. (L); Central Java, Central Java Merbaboe, Jul 1922, Roorda van Eysinga s.n. (MO); East Java, Tosari, Pasoeroean, Teysmann s.n. (CAL, K); East Java, Tengergebergte tusschen Ngadisari, Zandzee en Tosari , Went s.n. (L); Central Java, Meapi, Catu septr, Without collector s.n. (U); [iter Java secundum], Zollinger 1790 (LE, P) . West Nusa Tenggara: Klein Soenda Eilanden, Plawangan, Segara anak, 16 Jun 1936, de Voogd 2654 (GH, NY); Lombok, Rindjani Vulkangebirge N Seite, 9 May 1909, Elbert 1327 (L) .

Figures

Figure 5. Solanum alpinum Zoll. & Moritzi A Flowering habit B Detail of abaxial leaf surface C Detail of adaxial leaf surface D Leaf with an inflorescence E Bud F Dissected flower G Fruiting habit H Fruit (A-C, E-F Zollinger 2177; D Zollinger 2574; G-H Zollinger 2255). Drawing by R. Wise.

Figure 6. Distribution of Solanum alpinum Zoll. & Moritzi.