Lycianthes oliveriana (Lauterb. & K.Schum.) Bitter, Abh. Naturwiss. Vereins Bremen 24 [preprint]: 504. 1919, as "oliveriana"

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

12. Lycianthes oliveriana (Lauterb. & K.Schum.) Bitter, Abh. Naturwiss. Vereins Bremen 24 [preprint]: 504. 1919, as "oliveriana"

Figs 2 , 37 , 38.

Solanum oliverianum Lauterb. & K.Schum., Fl. Schutzgeb. Südsee [Schumann & Lauterbach] 535. 1900 [ “1901”], as “Oliverianum”. Type. Papua New Guinea. Sanduan/East Sepik: "Kaiser Wilhelmsland, Augustafluss", Sep 1887, M. Hollrung 776 (lectotype, designated by Symon 1985, pg. 56: K [K000759399]; isotypes: HBG [HBG511470], L [L0003651], LE [LE00016994], MEL [MEL104160], P [P00379610]).

Solanum memecylonoides Bitter & Schltr., Bot. Jahrb. Syst. 55: 93. 1917. Type. Papua New Guinea. Sanduan: "Kaiser Wilhelmsland, Torricelli-Geb[irges]", 800 m, 18 Sep 1909, F.R.R. Schlechter 20256 (holotype: B [destroyed]; lectotype, designated here: P [P00379576]; isolectotype: BR [BR0000005528844]).

Solanum memecylonoides Bitter & Schltr. var. finisterrae Bitter, Bot. Jahrb. Syst. 55: 94. 1917. Type. Papua New Guinea. Madang: "Kaiser Wilhelmsland, Finisterre-Gebirge",1,000 m, 3 Jul 1908, F.R.R. Schlechter 17961 (holotype: B [destroyed]; lectotype, designated here: P [P00379575]; isolectotype: UC [cited by Symon 1985, not seen nor on UC/JEPS database]).

Solanum balanidium Bitter, Bot. Jahrb. Syst. 55: 95. 1917. Type. Papua New Guinea. East Sepik: “Hunsteinspitz” [Mount Hunstein], 1300 m, Feb-Mar 1913, C.L. Ledermann 11332 (holotype: B [destroyed]). Papua New Guinea. East Sepik: Hunstein range, (Mt. Samsai) at site "Camp 3"on slopes above main streamcourse, 450 m, 17 Jul 1990, W.N. Takeuchi 6156 (neotype, designated here: LAE [acc. # 293351]; isoneotypes: A [00619947, 00619957], BISH [acc. # 618017], K [K001153745, K000922457, K000922458], L [L.2881432, L.2882048], MO [acc. # 4235181], NSW [NSW825821], NY [01404956, 02286515], US [01253664, acc. # 3723521]).

Solanum ledermannii Bitter, Bot. Jahrb. Syst. 55: 107. 1917, as “Ledermannii”. Type. Papua New Guinea. East Sepik: “Etappenberg” [between Kamelrücken and Bambooberg 142°29E, 4°38S, fide Veldkamp et al. 1988], 850 m, Oct 1912, C.L. Ledermann 9214 (holotype: B [destroyed]). Papua New Guinea. East Sepik: Amboin, Angoram subdistrict, 90 m, 29 Jul 1967, A.N. Millar & A.W. Dockrill NGF-35176 (neotype, designated here: LAE [acc. # 89947]; isoneotypes: BRI [n.v.], L [L.2881436]).

Lycianthes balanidium (Bitter) Bitter, Abh. Naturwiss. Vereins Bremen 24 [preprint]: 504. 1919. Type. Based on Solanum balanidium Bitter.

Lycianthes ledermannii (Bitter) Bitter, Abh. Naturwiss. Vereins Bremen 24 [preprint]: 504. 1919, as “Ledermannii”. Type. Based on Solanum ledermannii Bitter.

Lycianthes memecylonoides (Bitter & Schltr.) Bitter, Abh. Naturwiss. Vereins Bremen 24 [preprint]: 504. 1919. Type. Based on Solanum memecylonoides Bitter & Schltr.

Type.

Based on Solanum oliverianum Lauterb. & K.Schum.

Description.

Woody climbers or lianas, sometimes described as shrubs, to 3+ m tall (often described on labels “beautiful” e.g., van Royen & Sleumer 7716); stems terete, glabrous; new growth glabrous or minutely papillate with tiny 1-2-celled weak simple uniseriate trichomes less than 0.2 mm long, these soon deciduous; bark of older stems whitish grey, peeling and flaking, somewhat rugose and thick. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size but not in shape. Leaves simple; blades of major leaves (6.5)9-25 cm long, (2.8)3-10 cm wide (perhaps larger but not collected), elliptic, slightly discolorous, thick and coriaceous or chartaceous; adaxial surfaces glabrous, somewhat shiny; abaxial surfaces glabrous; principal veins 6-8 pairs, the midrib slightly keeled above, sometimes drying yellowish tan; base acute, often somewhat oblique; margins entire, revolute; apex acute or acuminate with an elongate drip-tip; petioles 1-2.5 cm long, glabrous; blades of minor leaves 4-9 cm long, 2.5-5 cm wide, shape, texture and pubescence like that of the major leaves; base acute; margins entire, revolute; apex acute or acuminate, occasionally rounded; petioles 0.6-1 cm long, glabrous. Inflorescences dense axillary fascicles, occasionally woody and enlarged with what appear to be tiny axes to 0.3 cm long, with 10-20-flowers, several open at the same time, glabrous; pedicels at anthesis 1-1.4 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading, glabrous, articulated at the base; pedicel scars tightly packed on the woody fascicle base. Buds plumply ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis. Flowers 5-merous (4-merous in Takeuchi 23389), heterostylous and unisexual, specimens with either all short-styled flowers or long-styled flowers and fruit, the plants probably dioecious. Calyx tube 2.5-3 mm long, 3-3.5 mm in diameter, deeply cup-shaped, usually described as purple or purplish blue, thick and fleshy, densely verrucose/tuberculate, without appendages, the rim somewhat hyaline ca. 0.5 mm wide, sparsely papillate. Corolla 0.8-1.1 cm in diameter, white or purple, stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 2-5 mm long, 1-2 mm wide, spreading or reflexed, thick and fleshy (live plants), appearing woody in dry material, adaxially glabrous to densely papillate with a few weak trichomes distally, abaxially densely papillate somewhat verrucose, the tips and margins densely papillate, the midvein raised especially adaxially, the tips cucullate. Stamens equal; filament tube minute; free portion of the filaments 1-1.5 mm long, glabrous; anthers 2-2.5 mm long, ca. 1 mm wide, plumply ellipsoid or slightly obovoid, creamy white, yellow or purple, poricidal at the tips, the pores round, distally directed, not elongating with age. Ovary conical, glabrous, vestigial in short-styled flowers; styles less than 0.2 mm long and vestigial in short-styled flowers, 5-6 mm long in long-styled flowers, straight, glabrous; stigma slightly bilobed, the surfaces minutely papillate. Fruit a globose berry, 0.7-1 cm in diameter, green and becoming bluish black when ripe, the pericarp glabrous, thick and appearing woody in dry material, matte, opaque; fruiting pedicels 1.1-1.5 cm long, 1-1.5 mm in diameter at the base, 1.5-2 mm in diameter at the apex, spreading or erect (?), woody, corky and markedly verrucose/tuberculate; fruiting calyx a cup surrounding ca. the lower half of the berry (making the fruit look like an acorn), woody (fleshy in live plants) and verrucose/tuberculate both adaxially and abaxially, green flushed with purple (fide Polak 864). Seeds 20-40 per berry, 3-3.5 mm long, 2-2.5 mm wide, flattened reniform or slightly tear-drop shape, reddish brown, the surfaces at the margins deeply pitted with pentagonal testal cells, the seed centre only shallowly pitted and the cells not clear. Stone cells absent. Chromosome number not known.

Distribution

(Fig. 39 ). Lycianthes oliveriana is widespread on the island of New Guinea in both Papua New Guinea (Central, East Sepik, Mandang, Morobe, Oro, Southern Highlands, Western) and Indonesia (Papua, Papua Barat); it has also been collected on the nearby Maluku Islands (Seram [Maluku] and Halmahera [Maluku Utara]).

Ecology and habitat.

Lycianthes oliveriana grows in lowland to montane and premontane rainforests, from almost sea level to 2,300 m elevation. This wide elevational range is accompanied by much variation in leaf size and shape.

Common names.

None recorded.

Preliminary conservation assessment

( IUCN 2020). EOO (1,006,290 km2 - LC); AOO (156 km2 - EN). Lycianthes oliveriana is known from more than 10 localities at a wide variety of elevations. It occurs within protected areas in both Indonesia and Papua New Guinea. This suggests a preliminary threat status of either Least Concern (LC) or Near Threatened (NT).

Discussion.

Lycianthes oliveriana is, apart from the Pacific L. vitiensis , the most commonly collected and widely distributed of the taxa treated in this monograph. It is a large canopy liana that is often described as “beautiful” due to its many flowered inflorescences and large, shiny leaves. Leaf size and shape vary considerably in L. oliveriana , plants with smaller, narrower leaves were described as several different taxa ( S. memecylonoides , S. memecylonoides var. finisterrae and S. balanidium ) by Bitter (1917). Leaf size varies continuously between the extremes, and I cannot discern any environmental factors determining leaf shape and narrowness, although field study or more in-depth environmental analysis might reveal this. All these leaf size variants have the same many-flowered axillary inflorescences, relatively small ca. 1 cm in diameter) flowers with valvate aestivation, thick fleshy corollas, plump, ellipsoid to slightly obovoid anthers and somewhat warty calyces with no appendages.

Lycianthes oliveriana is similar to L. kaernbachii but differs from it in its glabrous (versus softly pubescent) leaves and strictly axillary (versus cauliflorous) inflorescences. Both species have fleshy calyces without appendages that are often somewhat urceolate and small stellate flowers with fleshy corolla lobes. In low elevation forests L. oliveriana broadly co-occurs with L. impar that has similar glabrous leaves and difoliate geminate sympodial units. It differs from L. impar in having no inflorescence axis, although the woody fascicle can be rather large and lump-like with what are tiny axes, the inflorescence in L. impar has a distinct axis with paired pedicel scars along it. Fruits of L. impar are imperfectly known, but they appear to be soft and fleshy, while those of L. oliveriana have woodier pericarp; this woodiness coupled with the somewhat accrescent calyx tube gives the fruits of L. oliveriana the look of tiny acorns (Fig. 38 ).

Like other species of New Guinea Lycianthes , L. oliveriana appears to be dioecious, with long- and short-styled flowers on different plants. This needs confirmation in the field, and L. oliveriana would be an ideal subject for a reproductive biology study since it is relatively common and widely distributed.

In his discussion of Solanum balanidium , Bitter (1917: 96) suggested that it might be better placed as part of his S. memecylonoides , but lack of material held him back. Like S. memecylonoides , the collection on which S. balanidium is based (Ledermann 11332) had narrow leaves and short-styled flowers with no fruit. Almost all of Ledermann’s collections, all of which were distributed from Berlin, have been lost (see Veldkamp et al. 1988; Takeuchi and Golman 2002). I have found no duplicates of Ledermann’s collection from Mount Hunstein, so have selected another gathering (Takeuchi 6196) of a short-styled plant from the same mountain range with many duplicates as a neotype (LAE, acc. # 293351).

Bitter (1917) described Solanum ledermannii from a specimen of short-styled plant with large leaves ("This magnificent species is very similar to S. Oliverianum , it differs from it in its more vigorous growth and its larger, coarser, leathery leaves. Unfortunately, the fruits of S. Ledermannii are missing from the material, knowledge of which is particularly desirable for comparison with those of S. Oliverianum " Bitter 1917: 109, transl. from the original German). The collection was made by C.L. Ledermann (Ledermann 9214) during his trip exploring the April River in East Sepik. I have found no duplicates of Ledermann 9214; I have therefore selected a collection from East Sepik that matches the protologue and is held in several herbaria as the neotype, although it is from a lower elevation than Ledermann’s collection. The neotype specimen is held at LAE (acc. # 89967) in Papua New Guinea.

Specimens examined.

Indonesia. Maluku: Seram Island, "Seran, Hatomete", 4 Dec 1917, Kornassi 649 (K). Maluku Utara: Halmahera , Halmahera , Toliwang , 18 Oct 1951, Idjan Mochtar 348 (K). Papua: "S Nw Guinea" Sg. Aendosa near Oeta [=Uta], 3 m, 1 Jul 1941, Aët 395 (A, K, L); Rouffaer River [=Tariku River, Sungai Tariku], 175 m, Aug 1926, Docters van Leeuwen 9884 (K); Sawia, 'Nova Guinea neerlandica septemtrionalis’, 100 m, 21 Aug 1911, Gjellerup 613 (K); Kabupaten Manokwari, Kecamantan Manokwari, Arfak Mountains , Mupi Dessa, trail from Mupi village to G. Humibou, near Sungai Mupi, between Kali Umera (stream) and K. Ngwes, 770 m, 11 Apr 1995, Sands et al. 6744 (A, K, L); Mount Jaya , PT-Freeport Indonesia Concession Area, new East Levee road about 1 mile from junction of Main Road at Mile 38, 50 m, 9 Apr 1999, Utteridge et al. 287 (A, K, L); Mount Jaya , PT-Freeport Indonesia Concession Area, Main Road above Mile 38, 230 m, 5 Apr 2000, Utteridge et al. 295 (A, K, L, LAE, MO). Papua Barat (West Papua): Momi, subdistr. Manokwari , Vogelkop, Momi (S of Manokwari), 18 Aug 1948, Kostermans 2704 (K); Sorong, near Klamano [Klamano], 20 m, 10 Aug 1948, Pleyte 623 (A, K); Bird’s Head Peninsula, surroundings of Ayawasi, 14 Jul 1995, Polak 651 (K); Bird’s Head Peninsula, surroundings of Ayawasi, 500 m, 5 Sep 1995, Polak 864 (K); Vogelkop Peninsula, Ife River valley , Bamfot village , 850 m, 2 Nov 1961, Royen & van Sleumer 7621 (K, L, LAE); Vogelkop Peninsula, Ife River valley , central part of Tamray Range, S. slope, path from Sudjak village to Mt. Kusemun , Aiwa River , 840 m, 7 Nov 1961, Royen & van Sleumer 7716 (K) .

Papua New Guinea. "Kaiser Wilhelmsland, am Renegia", 150 m, 3 Oct 1908, Schlechter 18319 (P); "Kaiser Wilhelmsland: walden am Renegia", 150 m, 18 Oct 1908, Schlechter 18427 (E, LAE, P); Sankwet River , 15 Jun 1977, Symon 10659 (US). Central: Veiya, 12 Mar 1935, Carr 11670 (BM, K, L, NY); Port Moresby [National Capital District today], Goldie s.n. (MEL). East Sepik : Waskuk Hills, forest edge near Bangwis stream, 35 m, 2 Jan 2005, Takeuchi et al. 17743 (A, E, K, LAE, MO, US); Waskuk Hills, along Garuka-Bangwis track, 30 m, 3 Jan 2005, Takeuchi et al. 17786 (K); Ambunti District , Waskuk Hills, Seringyam near Mt. Musapien bivouac, 360 m, 15 Nov 2007, Takeuchi & Ama 22112 (A, K, LAE); Gulf: Ravikivau, Gulf District , near Ravikivau, Purari delta, 5 m, 18 Feb 1966, Craven & Schodde 848 (K, L, LAE). Madang: Mt Wilhelm [border of Chimbu, Jiwaka and Madang], near Plot 1200C, 1,200 m, 6 Nov 2012, Molino et al. 3060 (LAE, MPU, P); Morobe: Sattelberg, 1,006 m, 13 Feb 1936, Clemens 1821 (A, L); CRA Camp, near Wafi River , east of Tsili Tsili , 200 m, 3 Mar 1985, Conn et al. 1756 (A, BRI, L, LAE, MO); Bewapi Creek , about 4 miles W of Lae, 60 m, 26 Mar 1962, Hartley 10065 (A, L, LAE); Burep River NE of Lae, 30 m, 30 Apr 1962, Hartley 10136 (A, G, K, L, LAE); Tymne-Wago track, 457 m, 18 Mar 1963, Hartley 11428 (A, K, L, LAE); Oomsis Ridge, 609 m, 22 Feb 1965, Millar NGF-23858 (A, K, L, LAE); Bewapi Creek , 0.5 mile upstream from main road crossing, subdist. Lae , 7 Jun 1977, Symon 10655 (K, L, LAE, LAE); above Bupu village on Lae-Bulolo Road, 17 Jun 1984, Symon 13896 (K); Atzera Range, lowland forest near the 9-11 mile settlement, 300 m, Jan 1994, Takeuchi 9307 (E, NY, SING). Oro: Isuarava, 1,067 m, 4 Mar 1936, Carr 15948 (B, K); Sibium Mountains , ridge to E of Akupe Camp , Giegari , 1,303 m, 8 Feb 2013, Damas et al. SAJ-1050 (K, LAE, US); Sanduan: Telefomin District , Hak Valley , lower slopes of Deptabip ridge [Sanduan Province], 825 m, 8 Mar 1992, Frodin et al. 2352 (K); Folongonom, second bush camp below Tamanagabip on track to Busilmin; Telefomin subdist., 2,300 m, 16 May 1975, Vinas & Wiakabu LAE-59477 (A, E, K, L, LAE). Southern Highlands : Deviation Camp (Expedition Bivouac 5), 2,090 m, 8 Apr 2008, Takeuchi et al. 23895 (A, K, L, LAE, MO, SING, US); Tari subdistrict, Tigibi, 1,570 m, 10 Jun 1966, Vink 16847 (A, L, LAE). Western: Juha North, Bivouac 1, survey track 3, 245 m, 27 Mar 1978, Takeuchi et al. 23389 (K); Juha North (Bivouac 1) survey track 2, slope immediately east of [coordinates], 350 m, 28 Mar 2008, Takeuchi et al. 23491 (A). Western Highlands: 4 miles from Kopiago on Koroba Rd., Kopiago subdistrict, 1,466 m, 2 Nov 1968, Womersley et al. NGF-37289 (K, L, LAE) .

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 37. Lycianthes oliveriana (Lauterb. & K. Schum.) Bitter. Drawing by M. L. Szent-Ivany, first published in Symon (1985: fig. 17, as S. oliverianum Lauterb. & K. Schum.). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Figure 38. Lycianthes oliveriana herbarium specimen. Papua New Guinea. Morobe: Hartley 10065 (A). Courtesy of the Herbarium of the Arnold Arboretum of Harvard University, reproduced with permission.

Figure 39. Distribution of Lycianthes oliveriana.