Perennial herbs, shrubs, vines, lianas or trees, sometimes epiphytic. Stems terete or angled, glabrous or pubescent with simple (unbranched), forked, dendritic or stellate trichomes, these usually eglandular, but sometimes glandular. New growth usually with papillae, these sometimes glandular. Sympodial units unifoliate or difoliate, if difoliate the leaves geminate and often differing in both size and shape (anisophyllous). Leaves simple, entire, glabrous or pubescent with simple (unbranched), forked, dendritic or stellate trichomes, these usually eglandular, but sometimes glandular; petioles well-developed or not. Inflorescences axillary or adnate to the stems and caulescent (L. kaernbachii only), fasciculate or with a short rhachis; pedicels articulated at the base. Flowers 4–6-merous, usually 5-merous, but some species (e.g., Lycianthes banahaensis (Elmer) Bitter of the Philippines) consistently 4-merous, perfect or heterostylous, long- and short-styled flowers borne on the same or different plants (in Australia, New Guinea and the Pacific probably dioecious). Calyx with a truncate rim, with various numbers (usually multiples of five, but sometimes fewer) of appendages protruding from the calyx tube below or just at the rim, or without appendages; appendages small bumps to linear subulate in shape. Corolla rotate to deeply stellate, white to deep purple (yellow in L. banahaensis), often with the midvein of the lobes darker and the centre paler or yellow-green, interpetalar tissue present or absent, the lobes minute (rotate corollas) or long-triangular, spreading, cupped or reflexed at anthesis. Stamens equal or unequal, if unequal due to anther and/or filament differences; anthers plumply ellipsoid and obovate to tapering at the tips, usually dehiscing by apical pores, these sometimes opening to longitudinal slits with age. Ovary conical or globose, glabrous; style straight or curved, the stigma minutely capitate, clavate or strongly bifid with diverging lobes. Fruit a berry, globose to ellipsoid to ovoid, green, orange, red or purple, sometimes with stone cells in the mesocarp. Seeds few to many, usually flattened, sometimes winged (e.g., L. moszkowskii). Chromosome number: n=1224 (few species have chromosome counts).

Species of Lycianthes are found in the Americas, Asia, Australia, New Guinea and the islands of the Pacific. Species richness is concentrated in Mexico and Central America. No Lycianthes species are native to Africa, Europe or North America north of Mexico.

By far the greatest species diversity in Lycianthes occurs in the Americas (see Dean et al. 2020), but significant diversity occurs on the island of New Guinea (treated here). The description above attempts to cover variation across the distribution, both in the Americas and in the eastern Hemisphere. Ongoing work on the species of Asia (Japan, Philippines and Indonesia to India) may reveal new character states not included above.

As discussed above, the species treated here are mostly (excepting the widespread and weedy L. biflora) not found elsewhere in Asia, and although they may not be a phylogenetically distinct group, they are geographically logical to treat as a unit. The species treated here are, with a few exceptions, very rarely collected and there are many gaps in our knowledge of both their distribution and morphology. Field observations indicate this is not just due to under-collecting, but that these plants are rare where they do occur; this is often the case for large woody lianas and epiphytes of primary forests (e.g., Orejuela 2021).

Based on Solanum bambusarum Bitter.

Figure 4. 

Lycianthes bambusarum (Bitter) Bitter. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 23, as S. umbonatum Symon). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Shrubs, scrambling shrubs, lianas or epiphytes, to 3.5 m tall (long); stems terete, glabrous; new growth minutely puberulent with tiny usually single-celled papillate trichomes less than 0.1 mm long, soon glabrous; bark of older stems pale beige, somewhat corky and peeling. Sympodial units unifoliate, the leaves not geminate. Leaves simple; blades 4–11 cm long, 2–4.5 cm wide, narrowly elliptic to lanceolate, less commonly elliptic (i.e., Craven & Schodde 1258), slightly discolorous, membranous to chartaceous; adaxial and abaxial surfaces glabrous, the midrib somewhat keeled adaxially; principal veins 4–5 pairs, prominently anastomosing in arches before the margins; base acute to more commonly attenuate; margins entire; apex acuminate; petiole 0.6–1.2 cm long, glabrous. Inflorescences axillary fascicles of 1–4 (rarely to 8–10, e.g., Hartley 11434) flowers, only 1–2 open at a time, completely glabrous; pedicels (0.6 in bud) 1–1.2 cm long at anthesis, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, glabrous or minutely papillate near the base, articulated at the base; pedicel scars tightly packed in leaf axils. Buds globose, becoming ellipsoid, the corolla exserted halfway from the calyx tube just before anthesis. Flowers 4–5-merous, unisexual and heterostylous, the flowers on individual plants apparently all either short-styled or long-styled and the plants dioecious (needs field testing). Calyx with the tube 2.5–3 mm long, 4–5 mm in diameter, cup-shaped, glabrous or minutely papillate, apparently fleshy, purple, with 4–5 small umbonate appendages to ca. 0.5 mm long or the appendages absent, the rim entire and extending for 0.25–0.5 mm beyond the appendages, the appendages more prominent in buds. Corolla 0.8–1.2 cm in diameter, purple, stellate, lobed 3/4 of the way to the base, interpetalar tissue absent, the lobes 4–5 mm long, 1.5–2 mm wide, erect to spreading, fleshy (stiff and woody in dried specimens), glabrous abaxially and adaxially but densely papillate on tips and margins, the tips cucullate. Stamens equal; filament tube minute; free portion of the filaments ca. 0.1 mm long, glabrous; anthers 4–4.5 mm long, 1.5–2 mm wide, ellipsoid, somewhat tapering at the tips, bright yellow, poricidal at the tips, the pores directed distally, not elongating to slits with age. Ovary conical, vestigial in short-styled flowers, glabrous; style in short-styled flowers less than 1 mm long, in long-styled flowers 4–6 mm long, straight, purple, glabrous; stigma bilobed, the surfaces minutely papillate. Fruit a globose berry, 0.6–0.7 cm in diameter, green (immature? – in Streimann 9635 remnants of the style still at apex), the pericarp glabrous, thin, matte, opaque; fruiting pedicels 1–1.3 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, erect or spreading, somewhat woody, purple or green; fruiting calyx a cup-like spreading plate beneath the berry, somewhat thickened and warty (fleshy in live plants?). Seeds 50–70 per berry, ca. 2.5 mm long, ca. 2.5 mm wide, round with a deep notch at the hilum, yellowish tan, the surfaces deeply pitted, the testal cells pentagonal in outline. Stone cells absent. Chromosome number not known.

Figure 5. 

Lycianthes bambusarum herbarium specimen. Papua New Guinea. Morobe: Conn & Kairo 142 (A). Courtesy of the Herbarium of the Arnold Arboretum of Harvard University, reproduced with permission.

(Fig. 6). Lycianthes bambusarum is endemic to the island of New Guinea; collected only from Papua New Guinea (Chimbu, Morobe, Madang [destroyed type only]).

Figure 6. 

Distribution of Lycianthes bambusarum.

Lycianthes bambusarum is a plant of montane forests with bamboo, Pandanus and/or Nothofagus, between 450 and 2,400 m elevation.

None recorded.

(IUCN 2020). EOO (7,346 km² - VU); AOO (44 km² - EN). Lycianthes bambusarum is known from three localities; in light of this and the threats to forest habitats on New Guinea more generally, I propose a preliminary threat status of Endangered (EN [B1,2ab (iii, iv)]) for L. bambusarum.

Lycianthes bambusarum was distinguished by Bitter (1917) on the basis of its unifoliate sympodia, narrow leaves and 4-merous flowers; Symon (1985) recognised no other collections as this species, and described L. umbonata (as S. umbonatum) on the basis of its 5-merous flowers with distinct “umbos” on the calyx. The calyx appendages in L. bambusarum are very evident in bud (as in the type of S. umbonatum) but in flower are very small to non-existent (as shown in Bitter 1917: 92). Based on the otherwise widely overlapping descriptions I am here treating L. umbonata as a synonym of L. bambusarum.

Lycianthes bambusarum is similar to L. rostellata but differs from that species in being almost completely glabrous on stems and leaves, having unifoliate sympodia, and having heterostylous flowers. Lycianthes rostellata is distinctly pubescent, especially on the stems, with stiff, antrorse simple trichomes that often have multicellular bases, has small trowel-shaped minor leaves, and appears to have all bisexual flowers. The two taxa share narrow leaves and berries with numerous deeply notched seeds.

The type of Lycianthes bambusarum (as S. bambusarum) was collected by C.L. Ledermann in his explorations of the central mountain ranges of what is now Papua New Guinea (Ledermann 1919; Veldkamp et al. 1988; Takeuchi and Golman 2002). I have found no duplicates of this collection (Ledermann 12129), the original of which was cited as being in the herbarium in Berlin (Bitter 1917). These areas have rarely been accessed since Ledermann’s collecting, and I have seen no specimens from Madang that correspond to L. bambusarum. The collection (Takeuchi et al. 11704) selected here as a neotype matches the protologue, is from a similar elevation in the same mountain range and has a number of duplicates that are widely distributed, including in Papua New Guinea (where the neotype sheet is held in LAE).

Papua New Guinea. Morobe: near Blue Point, Mt. Kaindi, Wau, 800 m, 24 Apr 1977, Conn & Kairo 142 (A, K); near Wengomanga, via Oiwa, Aseki Patrol Area, 11 Apr 1966, Craven & Schodde 1258 (A, K, L, LAE, US); Wau, Mount Kaindi, contour trail in forest, 2,390 m, 10 Jul 1977, Fallen 374 (L, LAE, MO); Tymne-Wago track, 457 m, 18 Mar 1963, Hartley 11434 (A, K, L, LAE); Aseki-Spreader Div., Menyamya subdistrict, 1,800 m, 8 Jan 1972, Stevens LAE-54766 (A, K, L, LAE, US); Ekuti Divide, Bulolo-Aseki road, 33 km WSW of Bulolo, 2,250 m, 17 Oct 1982, Streimann 9635 (A, E, K, L, LAE); Angabena Ridge, ca. 3 km from Aseki-Menyamya Rd., Menyamya subdistrict, 1,675 m, 7 Jan 1972, Streimann & Stevens LAE-53892 (A, K, L, LAE); Aseki road from Bulolo, subdistrict Wau, 2,300 m, 29 Jul 1977, Symon 10632 (K, L, LAE, MO); Mount Kaindi, upper slopes of Mt. Kaindi, 2,000 m, 30 May 1984, Symon & Katik 13822 (K, L, LAE, MO); Aseki road below the crest, 31 May 1984, Symon & Katik 13829 (K, L, LAE, MO).

Based on Solanum belense Merr. & L.M.Perry.

Figure 7. 

Lycianthes belensis (Merr. & L.M.Perry) A.R.Bean. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 9, as S. belense Merr. & L.M.Perry). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Small shrubs to 1 m tall; stems terete, moderately pubescent with tiny glandular papillae and simple uniseriate 1–5-celled trichomes ca. 0.5 mm long, the trichomes transparent, weak-walled and collapsing, somewhat tangled; new growth densely pubescent with simple uniseriate trichomes like those of the stems, glabrescent; bark of older stems brown, glabrescent. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing only in size, not in shape, the minor leaves often deciduous. Leaves simple; blades of major leaves 7–14 cm long, 3–6.5 cm wide, elliptic, widest in the middle, concolorous, membranous to chartaceous; adaxial surfaces glabrous; abaxial surfaces often purplish green, with the lamina glabrous and scattered simple uniseriate trichomes ca. 0.5 mm long on the veins and midrib; principal veins 4–6 pairs, puberulent beneath; base narrowly acute, margins entire; apex acute to abruptly acuminate; petiole 0.6–1(1.5) cm long, glabrous or with a few scattered trichomes like those of the stems; blades of minor leaves 2.5–4 cm long, 1.3–1.7 cm wide, in shape and pubescence like the major leaves; petioles 0.5–0.7 cm long. Inflorescences axillary fascicles of ca. 4 flowers, 1–2 open at a time, moderately pubescent like the stems; pedicels 0.7–2 cm long at anthesis, lengthening considerably at anthesis, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, glabrous, articulated at the base; pedicel scars closely spaced in leaf axils. Buds ellipsoid, the corolla included in the calyx tube and exserted halfway just before anthesis. Flowers 5-merous, apparently unisexual and heterostylous, individual specimens with either short-styled or long-styled flowers, the plants possibly dioecious. Calyx tube 2.5–3 mm long, 2.5–3 mm in diameter, cup-shaped, glabrous with the rim minutely papillate, occasionally with up to 3 very small appendages, the rim transparent and ruffly, extending ca. 0.75 mm beyond the appendages (if present). Corolla ca. 2.4 cm in diameter, white or purple, stellate, lobed ca. 2/3 of the way to the base, a thin rim of interpetalar tissue present, the lobes ca. 3 mm long, ca. 0.9 mm wide, spreading, membranous, glabrous except for the densely papillate tips, margins and adaxial midvein. Stamens equal; filament tube minute; free portion of the filaments ca. (0.5) 3 mm long, glabrous; anthers 2.5–3 mm long, ca. 1.5 mm wide, ellipsoid and tapering, yellow, poricidal at the tips, the pores directed distally and not elongating with age. Ovary conical, glabrous; style in short-styled flowers less than 1 mm long, in long-styled flowers ca. 4 mm long (only seen in buds), straight, glabrous; stigma strongly bifid with diverging lobes ca. 0.25 mm long, the surfaces minutely papillate. Fruit a globose berry, ca. 1 cm in diameter, green (immature), the pericarp glabrous, relatively thin, matte, opaque; fruiting pedicels ca. 1.5 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, green (?), erect or spreading, not markedly woody; fruiting calyx a spreading plate beneath the berry, somewhat fleshy except for the thinner rim. Seeds and stone cells not seen. Chromosome number not known.

Figure 8. 

Lycianthes belensis herbarium specimen. Indonesia. Papua: Brass 11223 (isotype of S. belense, BM000778128). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

(Fig. 9). Lycianthes belensis is endemic to New Guinea; found in Papua New Guinea (Chimbu, Eastern Highlands) and Indonesia (Papua).

Figure 9. 

Distribution of Lycianthes belensis.

Lycianthes belensis is rarely collected but appears to be a plant of Fagaceae and Castanopsis forests, from sea level to around 2,300 m elevation.

None recorded.

(IUCN 2020). EOO (10,187 km² - VU); AOO (16 km² - EN). Lycianthes belensis is known from only three widely spaced localities, all in forested areas; it is very rarely collected and has not been collected recently. I propose a preliminary threat status of Endangered (EN [B2 a(iii, iv)]) for L. belensis.

There are very few collections of Lycianthes belensis and the range of variation needs further examination with additional material. The type collection (Brass 11223) has somewhat larger flowers on longer pedicels than other material, but leaf shape, pubescence and seed morphology suggest these all correspond to the same taxon. Specimens here included in L. lucens from the islands in the Louisiade Archipelago have been identified as L. belensis. Lycianthes belensis differs from L. lucens in its pubescent (versus glabrous) stems and its calyx with a thin, translucent ‘ruffly’ rim and 0–3 tiny appendages (versus a less obviously translucent rim with 3–5 triangular appendages emerging at right angles to the calyx tube).

Lycianthes belensis is a plant of high elevations while L lucens, also a plant of montane forests, is found at lower elevations. Their phylogenetic relationship has not been tested.

Indonesia. [type only]

Papua New Guinea. Chimbu: Chimbu valley, Gembogl, Yei nigl, 2,700 m, 31 Jan 1981, Sterly 80-469 (L); Eastern Highlands: Kassam, [Kassam Pass], 1,370 m, 3 Nov 1959, Brass 32400 (LAE, US); Mount Gahavisuka, 2,250 m, 16 Mar 1984, Cruttwell 2602 (L).

Based on Solanum biflorum Lour.

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.

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).

Figure 11. 

Lycianthes biflora herbarium specimen. Papua New Guinea. Morobe: Floyd NGF-7509 (BM001018933). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

(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.

Figure 12. 

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

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.

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

(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).

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.

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).

Based on Solanum bitterianum Symon.

Figure 13. 

Lycianthes bitteriana (Symon) A.R.Bean. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 5, as S. bitterianum Symon). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Large woody herbs to shrubs ca. 2 m tall; stems terete, densely pubescent with uniseriate dendritic 5–10-celled trichomes to 0.5 mm long, the branches short and congested (“tannenbaumartig”), drying yellowish tan; new growth densely pubescent with uniseriate dendritic trichomes like those of the stems, drying yellowish tan, not markedly glabrescent; bark of older stems dark brown, somewhat glabrescent. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing only in size, not in shape. Leaves simple; blades of major leaves 9–14 cm long, 4–6 cm wide, elliptic to broadly elliptic, widest in the middle, discolorous, membranous to chartaceous; adaxial surfaces moderately and evenly pubescent with dendritic trichomes with congested branches like those of the stems, these denser along the veins; abaxial surfaces more densely pubescent with dendritic trichomes, but the lamina still visible; principal veins 8–9 pairs, yellowish tan abaxially; base acute to truncate, oblique; margins entire; apex acuminate; petioles 1.6–3 cm long, densely dendritic-pubescent; blades of minor leaves 2.5–5.5 cm long, 1.3–3 cm wide, similar in shape and pubescence to the major leaves; petioles 0.5–1 cm long. Inflorescences axillary, the flowers borne on a woody axis with 3–4 short branches to 0.8 cm long, with 10–15 flowers, densely dendritic-pubescent with trichomes like those of the stems and leaves; pedicels 0.9–1 cm long at anthesis, ca. 0.5 mm in diameter at the bae, ca. 1 mm in diameter at the apex, densely pubescent with dendritic trichomes with congested branches, articulated at the base; pedicel scars tightly packed along the woody axes. Buds narrowly ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis. Flowers 5-merous, apparently perfect. Calyx tube 2–2.5 mm long, 2–2.5 mm wide, cup-shaped, densely dendritic-pubescent, with 4–5 linear, awl-shaped appendages 0.5–1 mm long, these varying in length within individual flowers, the rim extending ca. 0.1 mm beyond the appendages. Corolla 1–1.2 cm in diameter, white or “whitish blue”, stellate, lobed ca. halfway to the base, interpetalar tissue present, the lobes 3–3.5 mm long, ca. 2.5 mm wide, spreading or slightly reflexed, membranous, glabrous with densely papillate tips and a few dendritic trichomes along the midvein adaxially. Stamens equal; filament tube minute; free portion of the filaments ca. 0.5 mm long, glabrous; anthers ca. 2.5 mm long, ca. 1 mm wide, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary conical, glabrous; style ca. 6.5 mm long, glabrous; stigma minutely capitate and slightly bilobed. Fruit a globose berry, 0.6–0.7 cm in diameter, black or purple-black when ripe, the pericarp glabrous, thin, matte, opaque; fruiting pedicels 1.1–1.3 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, green (?), erect to spreading, densely dendritic-pubescent; fruiting calyx a spreading cup beneath the berry, dendritic-pubescent. Seeds ca. 100 + per berry, ca. 1.5 mm long, ca. 1.5 mm wide, flattened with a deep notch at the hilum, yellowish tan, the surfaces minutely pitted, the testal cells sinuate in outline. Stone cells absent. Chromosome number not known.

Figure 14. 

Lycianthes bitteriana herbarium specimen. Papua New Guinea. Morobe: Streimann & Kairo NGF-25854 (A). Courtesy of the Herbarium of the Arnold Arboretum of Harvard University, reproduced with permission.

(Fig. 15). Lycianthes bitteriana is endemic to the island of New Guinea; it has only been collected in Papua New Guinea (Morobe).

Figure 15. 

Distribution of Lycianthes bitteriana.

Lycianthes bitteriana is a plant of secondary forests at mid-elevations, from ca. 1,000 m.

None recorded.

(IUCN 2020). EOO (22 km² - CR); AOO (12 km² - EN). Lycianthes bitteriana is known from three localities less than 5 km apart, but could be more widely distributed, given its apparently secondary forest nature. I propose a preliminary threat status of Critically Endangered (CR [B1, 2a(iv)]) for L. bitteriana, but field assessment of its distribution is a priority.

Lycianthes bitteriana is a distinctive species with its congested branched trichomes that look like tiny Christmas trees (“tannbaumartig” of Seithe 1962) and shiny black berries. Unlike many of the other endemic New Guinea Lycianthes, L. bitteriana is a coarse herb or small shrub, in habit very similar to the widely distributed L. biflora. These two species can be easily distinguished by mature berry colour (black in L. bitteriana versus bright red in L. biflora), inflorescence morphology (many flowers on a short axis in L. bitteriana versus few-flowered fascicles in L. biflora), and branched trichomes morphology (L. bitteriana with congested branches versus the loosely branched trichomes of L. biflora). In addition, the corolla of L. bitteriana has abundant interpetalar tissue and is divided ca. halfway to the base, while that of L. biflora is deeply stellate with little or no interpetalar tissue.

Lycianthes dendropilosa has similar branched trichome with congested branches but differs from L. bitteriana in number of flowers per inflorescence (many in L. bitteriana, 1–3 in L. dendropilosa), calyx appendages (ca. 10 in L. bitteriana, absent in L. dendropilosa) and adaxial leaf morphology (evenly pubescent in L. bitteriana, glabrous and shiny in L. dendropilosa).

Given the nature of the secondary habitat in which Lycianthes bitteriana occurs, it is likely to be more widely distributed across the island of New Guinea.

Papua New Guinea. Morobe: Hump L.A. 5 mi SE Bulolo, Wau subdistrict, 1,067 m, 15 Mar 1971, Streimann & Kairo NGF-25854 (A, K, LAE); Mun. Bulolo District, Bulolo, 14 Jan 1957, Wells NGF-7565 (K, L, LAE).

Based on Solanum cladotrichotum Bitter.

Figure 16. 

Lycianthes cladotrichota (Bitter) Bitter A flowering branch B inflorescence with long-styled flowers C long-styled flower D short-styled flowers E fruiting branch F berry G, H trichomes from stem I trichome from calyx. (A, G, H Streimann & Katik NGF-30479 B, I Takeuchi 22832 C Crutwell 2310 D photograph of James et al. SAJ-1385 E, F Takeuchi 12379). Drawing by Lucy Smith. Scale bars: 4 cm (A, E); 7 mm (B, C); 7.5 mm (D, F); 0.9 mm (G–I).

Shrubs or more commonly described as woody climbers, to 10 m tall (long); stems terete, moderately to densely pubescent with stiff uniseriate dendritic trichomes to 1.5 mm long, the branches antler-like and well-spaced, occasionally also mixed with with stiff uniseriate simple trichomes ; new growth densely pubescent with stiff dendritic trichomes like those of the stems; bark of older stems pale brown, not markedly glabrescent, but becoming somewhat corky and peeling. Sympodia units unifoliate (minor leaves deciduous?) or difoliate and the leaves geminate, the leaves of a pair very different in size and shape; minor leaves often only present on new non-reproductive shoots. Leaves simple; blades of major leaves 7–19 cm long, 3.5–10 cm wide, elliptic to broadly elliptic (the type with narrowly elliptic leaves), widest in the middle, somewhat discolorous, chartaceous to coriaceous, thick and possibly rubbery-fleshy in live plants; adaxial surfaces shiny, glabrous but densely to moderately dendritic-pubescent along the veins and keeled midrib; abaxial surfaces sparsely and evenly pubescent with stiff antler-like dendritic trichomes ca. 0.5 mm long on the lamina, these denser along the veins and midrib; principal veins 8–12 pairs, prominent and impressed adaxially; base acute to truncate; margins entire; apex acute to abruptly acuminate with an elongate drip-tip; petiole 1–1.5 cm long, moderately pubescent with stiff dendritic trichomes like those of the stems and leaves; blades of minor leaves (if present) 1–2.2 cm long, 1–1.2 cm wide, orbicular and somewhat clasping the stem, bullate, pubescence like that of the major leaves; base truncate or cordate; margins entire; apex obtuse or rounded; petioles minute, to 0.5 cm long. Inflorescences dense axillary fascicles of 10–20 flowers, many flowers open at once, densely dendritic-pubescent with antler-like trichomes from the adjacent stems; pedicels at anthesis 0.7–1.5 cm long, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading to erect, possible held below the leaves, sparsely to moderately pubescent with stiff antler-like dendritic trichomes like those of the stems and leaves, articulated at the base; pedicel scars tightly clustered in the leaf axils. Buds ellipsoid, the corolla halfway exserted from the calyx tube before anthesis. Flowers 5-merous, heterostylous and apparently unisexual on separate plants, individual specimens either all long-styled and with fruit (e.g., Takeuchi 22892) or short-styled (e.g., Carr 15946). Calyx tube ca. 2 mm long, 3–4 mm in diameter, woody and stiff in dry specimens, fleshy in live plants, cup-shaped or slightly urceolate, sparsely pubescent with stiff dendritic trichomes like those of the stems and pedicels, mixed with some simple uniseriate trichomes of similar stiffness and size, without appendages, the rim slightly constricted. Corolla 1–1.3 cm in diameter, white or pale purple, stellate, lobed nearly to the base, interpetalar tissue absent, the lobes ca. 5 mm long, ca. 2 mm wide, spreading or perhaps reflexed, thick (fleshy in live plants), densely papillate on tips and margins, the tips slightly cucullate. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, densely pubescent with tangled weak-walled simple uniseriate trichomes; anthers ca. 3 mm long, 1.25–1.5 mm wide, yellow, poricidal at the tips, the pores distally directed, not splitting with age. Ovary conical, glabrous; style in short-styled flowers less than 0.5 mm long or absent, in long-styled flowers 4–4.5 mm long, straight, glabrous; stigma broadly bilobed, the surfaces minutely papillate. Fruit a globose berry, 0.5–0.6 cm in diameter, yellow-green (immature?) or fleshy blue-violet (fide Bitter 1917 in description of S. patellicalyx), the pericarp glabrous, thin, matte, opaque; fruiting pedicels 1–1.4 cm long, ca. 1 mm in diameter at the base, 1.5–2 mm in diameter at the apex, spreading or pendent, somewhat woody, with a few dendritic trichomes; fruiting calyx a spreading cup beneath the berry, woody in dry specimens, fleshy in live plants (?), often with a few dendritic trichomes, but these usually absent and the calyx somewhat corky or warty. Seeds 80–100 per berry, 2–2.5 mm long, 1.2–1.5 mm wide, tear-drop shaped with no distinct notch, the hilum apical, tan, the surfaces deeply pitted (less so in centre of seeds), the testal cells pentagonal in outline. Stone cells absent. Chromosome number not known.

Figure 17. 

Lycianthes cladotrichota herbarium specimen. Papua New Guinea. Oro: Carr 15946 (BM001018981). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

(Fig. 18). Lycianthes cladotrichota is endemic to the island of New Guinea; it has been collected in Papua New Guinea (Central, Eastern Highlands, Milne Bay, Oro, Sanduan, Western) and Indonesia (Papua).

Figure 18. 

Distribution of Lycianthes cladotrichota.

Lycianthes cladotrichota is a plant of lowland rainforest, occurring between 50 and 1,400 m elevation.

Papua New Guinea. Oro: ahara (Orokaira language, Hoogland 3979).

(IUCN 2020). EOO (254,412 km² - LC); AOO (48 km² - EN). Lycianthes cladotrichota is known from six localities across the island of New Guinea, but in spite of its apparently large distribution it has a relatively small area of occupancy (due to lack of collecting generally). Given the small AOO and its primary forest habitat, I suggest a preliminary threat status of Vulnerable (VU [B2a(iii, iv)]) for L. cladotrichota. It does occur in at least one protected area (Crater Mountain Wildlife Area).

Lycianthes cladotrichota a relatively widely distributed species of wet, lowland forests. It is distinctive in its stiff, antrorse pubescence of dendritic trichomes, thick, shiny major leaves, and tiny, orbicular minor leaves. The sympodia often appear unifoliate, but this is due to the abscission of the minor leaves – on young stems they are uniformly present (Fig. 2B). The thick, slightly urceolate calyx tube (Fig. 2F) is somewhat like that of L. oliveriana, but that species is glabrous, and the minor leaves are not tiny and deciduous. The fruiting calyx in L. cladotrichota is a spreading cup, while that of L. oliveriana cups the lower part of the berry, making it look at bit like an acorn. Lycianthes impar has similar small, heart-shaped or orbicular minor leaves, but can easily be distinguished by its simple rather than dendritic pubescence and in its inflorescence with a distinct axis; L. cladotrichota flowers are strictly fasciculate in the leaf axils.

The holotype of Lycianthes cladotrichota (Ledermann 12606) was in Berlin and is no longer extant; fortunately, unlike many other Ledermann collections, several duplicates exist. Lycianthes patellicalyx was described (Bitter 1917) from collections made a few months earlier than those used to describe L. cladotrichota, the principal distinguishing characteristics of the former were its slightly broader and less pubescent leaves. Lycianthes cladotrichota was described from flowering plants, and L. patellicalyx from fruiting material; in these plants that are possibly dioecious these differences can seem striking in the absence of more specimens. I have found duplicates of neither of the collections cited in the protologue of L. patellicalyx (Ledermann 11272, 11483), both from Mount Hunstein (Veldkamp et al. 1988). Symon (1985) placed L. patellicalyx (as S. patellicalyx) in synonymy with L. cladotrichota based on the similarity in descriptions, I maintain that here, and await further collections to neotypify the former name.

Indonesia. Papua: Mimika Regency, Mount Jaya, PT-Freeport Indonesia Concession Area, Kuala Kencana, near Ecological Plot 5, 65 m, 25 Jan 1998, Johns et al. 8900 (A, K, MO).

Papua New Guinea. Central: Kairuku-Hiri District, Mt. Gerebu, Trail towards summit ridge, 489 m, 5 Nov 2013, James et al. SAJ 1385 (BISH, BM, LAE). Eastern Highlands: M[oun]t Otto, south slopes,, m, 6 Aug 1959, Brass 30845 (K, L, LAE, US); Mount Gahavisuka, 2,250 m, Apr 1983, Cruttwell 2310 (K, L, LAE); Daulo Pass, top of Daulo Pass, 2,320 m, 22 Jun 1977, Symon, & Katik 10676 (K, LAE, MO, US); Crater Mountain Wildlife Area, ridge above Hauneäbäbo, 1,920 m, 21 Jul 1998, Takeuchi 12379 (K, US). Milne Bay: Raba Raba subdistrict, junction Ugat and Mayu Rivers, near Mayu Island (Mt Suckling complex), 400 m, 25 Jul 1972, Streimann & Katik NGF-34091 (E, K, L, LAE, US); Oro: Isuarava [Isurava], 1,067 m, 4 Mar 1936, Carr 15946 (BM, G, K, L, NY, P), 15 Mar 1936, Carr 16109 (BM, G, K, L, NY); “Northern Div.” near Pitoki village, ca. 3 km S of Kokoda station, 23 Sep 1953, Hoogland 3979 (A, LAE); Kokoda, ‘Northern District, Papua’, 400 m, 28 Jul 1964, Millar NGF-23549 (K, L, LAE). Western: Baia River (Expedition Bivouac 3) survey track B, 300 m, 13 Feb 2008, Takeuchi et al. 22892 (A, K, LAE, MO).

Based on Solanum dendropilosum Symon.

Figure 19. 

Lycianthes dendropilosa (Symon) A.R.Bean. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 11, as S. dendropilosum Symon). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Straggling shrubs or root climbers, 1–1.5 m tall; stems terete, densely glandular-papillate and densely pubescent with uniseriate dendritic trichomes to 0.75 mm long, the branches short and congested (“tannenbaumartig” sensu Seithe 1962), these drying yellowish tan; new growth densely papillate and pubescent like the stems, the trichomes tangled and interwoven; bark of older stems pale brown, becoming somewhat corky and glabrescent. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and shape. Leaves simple; blades of major leaves 2.5–4 (8) cm long, 1–2.5 cm wide, elliptic to narrowly elliptic (lanceolate in Symon & Katik 10691), widest in the middle, discolorous, coriaceous or membranous (Symon & Katik 10691); adaxial surfaces shiny, sparsely and unevenly pubescent with dendritic trichomes with congested branches like those of the stems, these denser along the midrib and principal veins; abaxial surfaces densely dendritic pubescent (lamina barely visible in Hoogland & Schoode 7291), yellowish tan; principal veins 4–6 pairs, densely pubescent, the midrib keeled adaxially; base acute; margins entire, strongly revolute in Hoogland & Schoode 7291; apex acute or more often acuminate; petiole 0.4–0.6 cm long, densely dendritic-pubescent like the stems; blades of minor leaves 0.6–0.7 cm long, 0.6–0.7 cm wide, orbicular or heart-shaped, similar in texture and pubescence to the major leaves; base cordate or rounded; margins entire or revolute; apex rounded; petiole absent or less than 0.1 cm long, densely dendritic-pubescent. Inflorescences axillary fascicles of 1–3 flowers, only one open at a time, densely dendritic-pubescent; pedicels at anthesis 1–1.2 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, erect or spreading, densely pubescent with golden dendritic trichomes with congested branches like those of the stems and leaves, articulated at the base; pedicel scars tightly packed in the leaf axils, Buds long-ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers 5-merous, probably heterostylous and unisexual, no styles seen in the only flowering specimen seen (Hoogland & Schoode 7291), the plants possibly dioecious. Calyx tube ca. 3 mm long, 3–3.5 mm in diameter, cup-shaped, densely golden dendritic-pubescent with trichomes like those of the pedicels, thick and coriaceous (fleshy in live plants?), without appendages, the rim slightly thickened. Corolla 1.5–2 cm in diameter, purple (mauve), deeply stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 6–9 mm long, 2–3 mm wide, spreading, thick and fleshy, adaxially glabrous, abaxially moderately pubescent with simple uniseriate trichomes 0.25–0.5 mm long, if these branched then merely forked, the branches not congested, the tips and margins densely papillate, the tips cucullate. Stamens equal; filament tube minute; free portion of the filaments 1–1.5 mm long, glabrous; anthers 5–7 mm long, 1–1.2 mm wide, long-ellipsoid and tapered at the tips, yellow, poricidal at the tips, the pores distally directed, not elongating with age. Ovary not seen. Fruit and seeds not known. Chromosome number not known.

Figure 20. 

Lycianthes dendropilosa herbarium specimen. Papua New Guinea. Western Highlands: Hoogland & Schodde 7291 (isotype of S. dendropilosum, BM000886135). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

(Fig. 21). Lycianthes dendropilosa is endemic to New Guinea; it has only been collected in Papua New Guinea (Southern Highlands, Western).

Figure 21. 

Distribution of Lycianthes dendropilosa.

Lycianthes dendropilosa has been collected in open areas near villages (Symon & Katik 10691 from “near old garden site”), from 2,400 to 2,700 m elevation.

None recorded.

(IUCN 2020). EOO (0 km² - CR); AOO (8 km² - CR). Lycianthes dendropilosa is known from only two localities and has been very rarely collected. It could be assessed as Data Deficient (DD), but given the threats to New Guinea forests more broadly I suggest it warrants assessment as Critically Endangered (CR [B1,2a,b(iii,iv)]); more collecting and a better understanding of its distribution are priorities.

Lycianthes dendropilosa is a distinctive species with small leaves, branched trichomes with densely congested branches, and 1–3 flowers per leaf axil. It shares trichome type with L. bitteriana, but that species is a coarse herb rather than a straggling shrub, has many flowers on a short inflorescence axis rather than 1–3 flowers per axil, and smaller flowers with copious interpetalar tissue (1–1.2 cm in diameter with interpetalar tissue in L. bitteriana, 1.5–2 cm in diameter without interpetalar tissue in L. dendropilosa). Fruits of L. dendropilosa are not yet known. Other than the type, that has thick, coriaceous leaves with somewhat revolute margins, the only other specimen I have seen that is attributable to this species (Symon & Katik 10691) is sterile; the leaves on this specimen are much thinner in texture and more elongate, indicating this may represent a juvenile pre-flowering individual.

Papua New Guinea. Southern Highlands: between Nol and Mendi, 24 km from Mendi, just after crest [“and o”], 2,000 m, 24 Jun 1977, Symon & Katik 10691 (L, LAE).

Based on Solanum impar Warb.

Figure 22. 

Lycianthes impar (Warb.) Bitter A flowering branch B inflorescence with bud C short-styled flower D berries E seeds. (A Utteridge et al. 119, B van Leeuwen 11108, C Sands 7329, D, E Puradyatmika et al. 10428). Drawing by Lucy Smith. Scale bars: 4 cm (A); 5 mm (B, C); 0.75 cm (D); 1.4 mm (E).

Small trees or climbers, to 3.5 m tall; stems terete, sometimes with adventitious roots from the nodes (Kloss s.n., 22 Nov 1912), moderately pubescent with transparent, antrorsely curled simple uniseriate 2–3-celled trichomes to 0.5 mm long, the basal cells sometimes enlarged; new growth densely papillate and pubescent with antrorse trichomes like those of the stems; bark of older stems pale grey-brown and somewhat corky and peeling. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and shape. Leaves simple; blades of major leaves 8.5–17 cm long, 2.9–6 cm wide, elliptic, discolorous, chartaceous or coriaceous; adaxial surfaces shiny, completely glabrous; abaxial surfaces glabrous; principal veins 8–10 pairs, prominent and pale yellow of reddish tan beneath; base acute; margins entire; apex acuminate; petiole 0.5–1 cm long, with a few simple antrorse trichomes like those of the stems at the very base; blades of minor leaves 0.8–1.5 cm long, 0.8–1.1 cm wide, orbicular to obcordate but quite variable in shape even within a single plant, similar in texture and pubescence to the majors; base cordate or truncate; margins entire; apex acute or rounded; petiole absent to less than 0.5 cm long, glabrous to absent. Inflorescences axillary with a short axis 0.4–1 cm long, 10–12-flowered, hanging under the leaves, only 1–2 flowers open at a time, glabrous and corky; pedicels at anthesis 0.6–0.9 cm long, ca. 0.5 mm in diameter at the base, ca. 1.25 mm in diameter at the apex, spreading or nodding, glabrous, articulated at the base pedicel scars closely packed, from the very base of the axis. Buds ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis. Flowers 5-merous, only short-styled flowers seen, the plants possibly dioecious. Calyx tube 2.5–3 mm long, ca. 3 mm wide, elongate cup-shaped, thick and probably somewhat fleshy in live plants, purple, glabrous, without appendages, the rim slightly thinner and sparsely papillate. Corolla ca. 1.2 cm in diameter, purple or violet with the lobes white (fide Sands et al. 7329), stellate, lobed nearly to the base, interpetalar tissue absent, the lobes ca. 5 mm long, ca. 1.5 mm wide, spreading, thick and fleshy, both surfaces glabrous, densely papillate on tips and margins. Stamens equal; filament tube minute; free portion of the filaments 0.75–1 mm long, glabrous; anthers ca. 3 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores round, directed distally, not elongating to slits with age. Ovary conical, glabrous, vestigial (only seen in short-styled flowers); style and stigma not seen. Fruit an elongate berry, 0.7–1 cm long, ca. 0.6 cm wide, bright blue or purple-blue when ripe, whitish blue to almost white when immature (fide Utteridge et al. 119), the pericarp glabrous, thin, matte, opaque, the fruit flesh purple (fide Utteridge et al. 119); fruiting pedicels 1–1.2 cm long, ca. 0.75 mm in diameter at the base, ca. 2 mm in diameter at the apex, purple, pendent and hanging below the leaves; fruiting calyx a “cupule-like structure” (fide Utteridge et al. 119) subtending the fruit, bright purple or white (fide Takeuchi 9181), thick and probably fleshy in live plants. Seeds 10–20 per berry, ca. 4 mm long, ca. 1.5 mm wide, reniform, not markedly flattened, white (“white, kidney-shaped” in live plants fide Utteridge et al. 119) or pale tan, the surface minutely pitted, the testal cells rectangular in outline. Stone cells absent. Chromosome number not known.

Figure 23. 

Lycianthes impar herbarium specimen. Indonesia. Papua: Brass 6796 (A). Courtesy of the Herbarium of the Arnold Arboretum of Harvard University, reproduced with permission.

(Fig. 24). Lycianthes impar is endemic to the island of New Guinea; it has been collected in Papua New Guinea (Southern Highlands, Western) and Indonesia (Papua, Papua Barat).

Figure 24. 

Distribution of Lycianthes impar.

Lycianthes impar occurs in lowland rainforests on alluvium, between 10 and 210 m elevation.

None recorded.

(IUCN 2020). EOO (257,568 km² - LC); AOO (48 km² - EN). Lycianthes impar is known from more than seven localities in both Indonesia and Papua New Guinea, some of which are in protected areas (Parsch et al. 2022). It is a mostly lowland forest plant that is rather rarely collected; I suggest a preliminary threat status of Vulnerable (VU [B2a,b(iii, iv)])) based on the AOO, the number of localities and the threats for lowland forests more generally (Parsch et al. 2022).

Lycianthes impar is a small tree or vine with strikingly differently shaped major and minor leaves. The orbicular or heart-shaped minor leaves are like those of L. cladotrichota and L. peranomala, but L. cladotrichota differs in leaves that are pubescent with dendritic trichomes; both L. impar and L. peranomala have glabrous leaves. Stem trichomes of L. impar are soft and somewhat tangled, while those of L. peranomala are stiff and markedly antrorse. Lycianthes impar differs from both those species in having a short, sometimes forked, inflorescence axis; both L. peranomala and L. cladotrichota have strictly fasciculate flowers. Lycianthes oliveriana is similarly glabrous, but Warburg’s description of Solanum impar clearly mentions a “peduncle” (inflorescence axis), distinguishing it clearly from L. oliveriana that lacks any axis. The soft, ovoid berries of L. impar are quite distinct from the woodier globose berries of L. oliveriana. The striking contrast in colour (purple/white) between the fleshy calyx cup and berry during fruit ripening mentioned on labels (e.g., Utteridge et al. 119, Takeuchi 9181) suggests the fruits are bird dispersed.

Symon (1985) mistakenly synonymised Solanum ridleyanum with Lycianthes moszkowskii (as S. moszkowskii) from which it differs in pubescence type (soft and tangled versus stiff and antrorse), inflorescence morphology (with a small sometimes forked axis versus strictly fasciculate), fruit shape and colour (purplish blue and elongate-ovoid versus globose and bright cherry red) and seed morphology (narrowly kidney-shaped versus strikingly winged in L. moszkowskii).

The type collection of Solanum impar was made by Otto Warburg (Warburg 21244) in the McCluer Gulf area on the N coast of the Bomberai Peninsula in Papua Barat, where he made a short stop in January of 1889 on his world tour (van Steenis-Kruseman 1985). I have found no duplicates of his gathering, nor have I found any specimens collected from the same area. A collection made further southwest of the type locality near the town of Oeta in Papua by Aët (an Indonesian collector who collected widely in conjunction with various Dutch botanists, including on the Lundquist expedition, see van Steenis-Kruseman 1985) is a good match for the protologue, has flowers and fruit and is from similar habitat. I here designate Aët 407 (BO acc. # 1579909) as the neotype for L. impar.

In the herbarium at Naturalis (L) Georg Bitter annotated herbarium specimens of Lycianthes impar with designations he never published “Lycianthes amblycarpa” (Versteeg 1351) and “Lycianthes radicans” (Lam 706) (see Names not validly published). Another name not validly published “Lycianthes fistulosa” (with no attribution) is written on the duplicate of Versteeg 1351 at BO (acc. # 1588531).

Indonesia. Papua: Rouffaerriiver, Motorbiv, 100 m, Nov 1926, Docters van Leeuwen 11108 (K, L); Utakwa Expedition to Mt. Carstensz. Camp I-III, 22 Nov 1912, Kloss s.n. (BM); reg. flum. Mamberamo, pr. Pioniersbiv [Sungai Mamberano] [Geelvink Bay], 10 m, 23 Jul 1920, Lam 706 (BO, K, L); Mimika Regency, Mount Jaya, PT-Freeport Indonesia Concession Area, between Kali Kopi levee (new E Levee) and the Kopi River, along black water stream flowering into the Kopi River (also black water), 210 m, 9 Mar 1999, Puradyatmika et al. 10428 (A, K, MO); Mimika Regency, Mount Jaya, PT-Freeport Indonesia Concession Area, Kuala Kencana, near PT Freeport Indonesia Office, 50–100 m, 27 Aug 1998, Sands 7329 (A, K); PT-Freeport Indonesia Concession Area, Rimba Irian Golf Course, on outskirts of Kuala Kencana, 10 m, 15 Mar 1999, Utteridge et al. 119 (A, K, L, MO); “fluv. Lorentz” [Lorentz River = Sungai Unir] [Snow Mountains fide Symon 1985], 19 May 1907, Versteeg 1137 (BO, L); fluv. Lorentz, prope ‘Sabang’ [Lorentz River, Sabang van Weel’s camp; Alkmaar bivouac] [Snow Mountains fide Symon 1985], 2 Jul 1907, Versteeg 1351 (BO, L). Papua Barat (West Papua): Sorong, near Klamono, 20 Aug 1948, Pleyte 633 (BO, L).

Papua New Guinea. sin. loc. (“Solanum elegans Zp./N Guinea”), Without Collector s.n. (L). Southern Highlands: Kutubu patrol area, karst limestone NW of Yorokobaiu village, 600 m, 10 Sep 1993, Takeuchi 9181 (A); Western: Fly River, 528 mile camp., May 1936, Brass 6796 (A, BM, BRI, L); Kiunga subdistr., base camp (Ok Tedi river), 700 m, 2 Nov 1969, Foreman & Galore NGF-45764 (L); Kiunga subdistr., Ok Tedi headwaters, near Kennecott field camp, 800 m, 29 Oct 1969, Henty et al. NGF-42805 (L); Kiunga, Kiunga subdistr., 30 m, 9 Aug 1971, Streimann & Katik LAE-51786 (L).

Based on Solanum kaernbachii Lauterb. & K.Schum.

Figure 25. 

Lycianthes kaernbachii (Lauterb. & K.Schum.) Bitter. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 13, as S. kaernbachii Lauterb. & K.Schum.). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Woody climber or large liana, size not recorded; stems terete, densely to moderately pubescent with simple uniseriate 3–7-celled weak-walled, trichomes to 1 mm long, these occasionally forked, drying golden tan; new growth densely pubescent with simple uniseriate trichomes like those of the stems, bright golden tan in dry material; bark of older stems brown, not markedly glabrescent on twigs. Sympodial units difoliate, the leaves geminate, the leaves of a pair different in size and shape. Leaves simple; blades of major leaves 9–21 cm long, 3.5–11 cm wide, elliptic to narrowly elliptic, widest in the middle, discolorous, coriaceous (“greasy” fide Kairo & Streimann NGF-30943); adaxial surfaces shiny, glabrous to sparsely pubescent with simple uniseriate trichomes, these denser along the veins; abaxial surfaces moderately to densely and even pubescent with simple uniseriate trichomes to 1 mm long, these 3–7-celled, weak-walled, drying golden tan, denser along the veins; principal veins 7–10 pairs, the midrib somewhat keeled, the veins impressed above; base acute to somewhat cordate-truncate, oblique; margins entire, slightly revolute; apex acute to abruptly acuminate; petioles 0.5–3 cm long, densely pubescent with simple uniseriate trichomes like those of the stems and leaves; blades of minor leaves 2.5–6.5 cm long, 2–6 cm wide, broadly elliptic to orbicular, texture and pubescence like that of the major leaves; base rounded or cordate; margins entire to slightly revolute; apex rounded or obtuse; petioles absent to 0.6 cm long, pubescence like thatof the major leaves. Inflorescences paired rows of cauliflorous pedicels along the stem between the nodes and in leaf axils with groups of more than 10 flowers, many flowers open simultaneously, pubescence like that of the stems; pedicels 0.8–1.1 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender and spreading, purple or green, glabrous to sparsely to moderately pubescent with golden simple uniseriate trichomes to 0.25 mm long, these less dense than stem pubescence, articulated at the base; pedicel scars closely spaced in rows along the stems. Buds ellipsoid, the corolla more or less strongly exserted from the calyx tube before anthesis. Flowers 5-merous, heterostylous and unisexual, individual specimens with either short-styled or long-styled flowers, the plants probably dioecious. Calyx tube 2–2.5 mm long, 2.5–3 mm in diameter, urn-shaped, thick and woody in dry material (fleshy in live plants?), slightly tuberculate, with scattered to denser simple uniseriate trichomes like those of the pedicels, without appendages, the rim constricted and thickened. Corolla 0.8–0.9 cm in diameter, creamy white to reddish cream (fragrant fide Kairo & Streimann NGF-30943), stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 3–4 mm long, 1.2–1.5 mm wide, spreading or reflexed, thick and fleshy (live plants), glabrous adaxially, minutely puberulent abaxially, the tips and margins densely papillate, the tips strongly cucullate. Stamens equal; filament tube minute; free portion of the filaments 1–1.5 mm long, glabrous; anthers 1.5–2 mm long, ca. 1 mm wide, plumply ellipsoid, yellow, poricidal at the tips, the pores directed distally, not elongating to slits with age. Ovary globose to conical, glabrous, vestigial in short-styled flowers; style in short-styled flowers apparently absent, in long-styled flowers ca. 4.5 mm long, straight, glabrous; stigma strongly bifid, the lobes ca. 1 mm long, the surfaces minutely papillate. Fruit a globose berry, 0.5–0.6 cm in diameter, green, the pericarp glabrous, hard and somewhat woody in dried material, opaque; fruiting pedicels 0.9–1.1 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading, more or less woody, slightly tuberculate; fruiting calyx a cup around the basal third of the berry, woody in dry material (fleshy in live plants?), with a few simple uniseriate trichomes, somewhat tuberculate. Seeds 2–20 per berry, 3–3.5 mm long, 2–2.5 mm wide, flattened reniform, without a deep notch, reddish tan, the surfaces deeply pitted, especially near the margins, the testal cells sinuate in outline. Stone cells absent. Chromosome number not known.

Figure 26. 

Lycianthes kaernbachii herbarium specimen. Papua New Guinea. Morobe: Kairo & Streimann NGF-30943 (A). Courtesy of the Herbarium of the Arnold Arboretum of Harvard University, reproduced with permission.

(Fig. 27). Lycianthes kaernbachii is endemic to the island of New Guinea; it has only been collected in Papua New Guinea (Madang, Morobe) in the eastern Finisterre Range.

Figure 27. 

Distribution of Lycianthes kaernbachii.

Lycianthes kaernbachii is a plant of primary lowland rainforest, between 30 and 700 m elevation.

Papua New Guinea. nigukwaa (Schumann and Lauterbach 1901).

(IUCN 2020). EOO (8,882 km² - VU); AOO (24 km² - EN). Lycianthes kaernbachii is known from five localities, all in the Finesterre range. Its small range, coupled with the lack of collections from protected areas suggest a preliminary threat status of Endangered (EN[B1,2ab(iii,iv)]) for L. kaernbachii.

Lycianthes kaernbachii is probably the most distinctive and unusual of the New Guinea species of Lycianthes. It is a large canopy liana with softly pubescent leaves and is cauliflorous, a character not seen elsewhere in the genus, and one that I have not seen elsewhere in the family. The inflorescence axis is apparently adnate to the stem, and the buds, flowers and fruits are in two parallel rows of varying length between the nodes. Flowers of L. kaernbachii, at less than 1 cm in diameter, are among the smallest (with L. peranomala) amongst New Guinea Lycianthes. Corolla lobes of L. kaernbachii are valvate in bud and thick and fleshy on live plants, becoming somewhat woody on dry specimens and completely lack interpetalar tissue. This type of corolla is shared with L. oliveriana and L. peranomala, but L. kaernbachii is easily distinguished from those taxa by its soft pubescence on abaxial leaf surfaces (L. oliveriana is glabrous and L. peranomala has sparse pubescence of stiff antrorse trichomes only along the veins) and its cauliflory (the other two species have strictly axillary inflorescences).

Millar NGF-23365 is less pubescent than most of the other collections of Lycianthes kaernbachii I have seen and was tentatively identified as L. oliveriana by Symon based on a non-reproductive specimen; he later suggested it might be a mixed collection (Symon 1985). It does, however, have the distinctive linear cauliflory of L. kaernbachii on other duplicates and the pubescence of the new growth indicates it is not a mixed collection but rather an unusual sparsely pubescent plant of L. kaernbachii.

The type specimen of Solanum kaernbachii was held in Berlin and was destroyed in the Second World War, along with the many other types no longer extant (Vorontsova and Knapp 2010); no duplicates have been found. As a neotype (L.2881629) I have selected a collection from the same locality “Sattelberg” in eastern Papua New Guinea (Clemens 1289) that corresponds to the description in the protologue and is held in several herbaria.

The holotype of Solanum schlechterianum was not destroyed along with other New Guinea collections at Berlin; Bitter (1917) acknowledged the similarity of his new species with L. kaernbachii (as S. kaernbachii) and his illustration clearly shows the distinctive caulescent inflorescences.

Papua New Guinea. Madang: “Kaiser Wilhelmsland, Waldern am Djamu”, 700 m, 24 Apr 1908, Schlechter 17339 (P). Morobe: Wareo, 610 m, 1 Jan 1935, Clemens & Clemens 1426 (L); Sankwet L.A., Lae, 60 m, 30 Nov 1967, Kairo & Streimann. NGF-30943 (A, E, K, L, NSW), 30 m, 5 Jan 1968, Kairo & Emos NGF-30983 (A, E, K, L, LAE); Bupu village above Wampit, 762 m, 3 Mar 1964, Millar NGF-23260 (K, L, LAE, NY, US); Bupu village above Wampit, 701 m, 4 Mar 1964, Millar NGF-23365 (A, K, L, LAE, US); midway of Buso River, SE of Buso Camp, 18 Jun 1984, Vinas & Kairo 308 (K).

Like L. vitiensis, but differing in its shrub rather than tree habit, narrowly elliptic rather than elliptic leaves, its fewer-flowered inflorescences that are strictly axillary rather than many-flowered on an elongate axis, presence of triangular calyx appendages versus lack of appendages, anthers that are strictly poricidal versus anthers dehiscing through elongate slits or the pores lengthening with age, and smaller seeds (3 mm versus 4–5 mm long) that lack a distinct notch.

Papua New Guinea. New Ireland: Lihir Island [Niolam Island], Mount Tementa, above Palie Mission, Namatanai subprovince, 710 m, 7 Nov 1984, O. Gideon LAE-57196 (holotype: LAE [acc. # 256314]; isotypes: K [K000922490], L [L.2882045]).

Figure 28. 

Lycianthes lucens S.Knapp A flowering branch B flower bud C flower D flower bud with petals removed to show stamens E fruiting branch F berries G seeds. (A, C, E, G Sands et al. 2230 B Sands et al. 2073). Drawing by Lucy Smith. Scale bars: 4 cm (A, E); 7 mm (B, C); 6 mm (D); 2 cm (F); 2.5 mm (G).

Shrubs 0.5–1.3 m tall; stems terete, glabrous or with very sparse simple or forked uniseriate papillae or trichomes, these sometimes to 5-celled, less than 0.2 mm long, drying reddish gold, the papillae possible glandular, a single collection with adventitious roots from the stem, these filamentous (Croft et al. LAE-71421); new growth glabrous or sparsely papillate, the papillae with darker (glandular?) terminal cells; bark of older stems white to whitish grey, somewhat wrinkled and ridged from drying. Sympodial units difoliate, the leaves geminate, sometimes appearing unifoliate due to the minor leaves abscising, the leaves of a pair similar in shape but not in size. Leaves simple; blades of major leaves 9–15 cm long, 3–7 cm wide, narrowly elliptic, discolorous, membranous; adaxial surfaces very shiny, glabrous; abaxial surfaces glabrous, paler; principal veins 4–7 pairs, yellowish green beneath (in dry material), the midrib not keeled, the veins arching but not forming distinct inframarginal loops; base acute or sometimes somewhat rounded; margins entire; apex acuminate, rarely acute; petioles 0.5–1.5 cm long, completely glabrous; blades of minor leaves 3–9 cm long, 1–4.5 cm wide, shape, texture and pubescence like that of the majors; base acute or sometimes somewhat rounded; margins entire; apex acuminate, rarely acute; petioles 0.3–0.5 cm long, glabrous. Inflorescences axillary fascicles or 1–4 flowers (most often 2–3 flowers), only a single flower open at a time, completely glabrous; pedicels at anthesis 2.5–4 cm long, markedly lengthening just before anthesis, 0.5–0.75 mm in diameter at the base, 1–2 mm in diameter at the apex, pale green or white, spreading or perhaps nodding, completely glabrous and shiny, articulated at the base; pedicel scars in a tight cluster in the leaf axils. Buds ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis, completely included in young buds. Flowers 5-merous, heterostylous and unisexual, specimens with either short-styled flowers or long-styled flowers and fruit, the plants perhaps dioecious, the short-styled flowers marginally smaller than long-styled flowers. Calyx tube 2–4 mm long, shorter in short-styled flowers, 2.5–3.5 mm in diameter, openly cup-shaped, white flushed with purple (fide Sands et al. 1966), with (3)4–5 triangular appendages 1–1.5 mm long, ca. 0.5 mm wide, oriented perpendicular to the calyx tube, these usually of differing lengths in a single flower, emerging 0.5–1 mm from the calyx rim, in very young buds the appendages sometimes papillate. Corolla 1.1–2.4 cm in diameter, short-styled flowers at the smaller size end, white or pale purple, stellate, lobed ca. 3/4 of the way to the base, interpetalar tissue present, the lobes 4–8 mm long, 3–4 mm wide, spreading or slightly cupped, membranous, glabrous on both surfaces, the tips and distal portions of the margins densely papillate. Stamens equal; filament tube minute; free portion of the filaments ca. 1.5 mm long, glabrous; anthers 2.5–3 mm long, ca. 1.5 mm wide, ellipsoid, yellow, poricidal at the tips, the pores distally directed, round and not elongating to slits with age. Ovary conical, glabrous, vestigial in short-styled flowers; style in short-styled flowers vestigial, in long-styled flowers 6–6.5 mm long, straight, glabrous; stigma clavate, the surfaces minutely papillate. Fruit a globose berry, 1.2–1.5 cm in diameter, deep red when ripe, green to orange-red to deep red through development, the pericarp glabrous, thin, shiny, translucent; fruiting pedicels 3–4 cm long, longer in mature fruit, 1–1.5 mm in diameter at the base, 3–3.5 mm in diameter at the apex, not markedly woody, pendent (?), pale green; fruiting calyx a spreading plate beneath the fruit, stiff and perhaps fleshy or woody in live plants, the veins leading to the calyx appendages enlarged and prominent. Seeds 50–100 per berry, ca. 3 mm long, 2.5–3 mm wide, flattened and somewhat round with one straight edge in outline, reddish tan, the margins darker, the surfaces deeply pitted, the testal cells pentagonal in outline. Stone cells absent. Chromosome number not known.

Figure 29. 

Lycianthes lucens herbarium specimen. Papua New Guinea. New Ireland: Sands et al. 2030 (K000922448). Courtesy of the Trustees of the Royal Botanic Garden, Kew, reproduced with permission.

(Fig. 30). Lycianthes lucens is endemic to the islands east of the main island of New Guinea; it has been collected on New Ireland, Lihir Island and the islands of the Louisiade Archipelago (Rossel and West Misima Islands).

Figure 30. 

Distribution of Lycianthes lucens.

Lycianthes lucens grows in the shaded understory of montane rainforests, from 700 to 1,350 m elevation.

None recorded.

(IUCN 2020). EOO (70,602 km² - LC); AOO (24 km² - EN). Lycianthes lucens is known from only four localities on widely separated islands; the large EOO consists mostly of uninhabitable ocean. Gold mining on Lihir island is a significant threat for the forest habitat of this species; I suggest a threat status of Endangered (EN [B1,2a, b(iii,iv)]) for L. lucens.

Lycianthes lucens is a beautiful plant, with large flowers, bright red berries and shiny leaves. The species epithet is derived from the shiny, glabrous leaves. Symon (1985) cited Katik et al. LAE-70928 from Rossel Island as L. belensis (as S. belense), and other specimens in herbaria have been annotated as that species by him. Lycianthes lucens resembles L. belensis in its shiny leaves and mostly axillary inflorescences with few, large flowers but it differs in its glabrous stems and adaxial leaf venation (versus stem and abaxial veins softly pubescent in L. belensis) and triangular calyx appendages to 1.5 mm long that are perpendicular to the calyx tube (versus appendages that are either absent or less than 0.75 mm long and not obviously perpendicular to the calyx tube in L. belensis). The calyx rim in L. belensis is thin, transparent and ruffly (see Fig. 8), while that of L. lucens is strictly truncate with distinct teeth and not markedly ruffly (Fig. 28). Lycianthes belensis is a plant of high elevations in the central ranges, while L. lucens is a plant of montane forests only found on the islands to the east of New Guinea proper.

Lycianthes multifolia has shiny leaves somewhat similar to those of L. lucens but it has smaller flowers (1.4–1.6 cm in diameter versus ca. 2.4 cm in diameter in L. lucens) and densely pubescent stems and venation (versus glabrous in L. lucens). Sympodial units of L. lucens are strictly difoliate, while those of L. multifolia often have more than two leaves at a node.

The only other Lycianthes species found as far east in the New Guinea and Pacific area as L. lucens is L. vitiensis from the island of Bougainville to Samoa. Lycianthes lucens differs from L. vitiensis in habit (shrub versus tree) and inflorescence morphology (axillary fascicles versus with a distinct axis), in addition to the other characters mentioned in the diagnosis.

Like other Lycianthes species in the area, L. lucens appears to be dioecious, with individual specimens bearing either short-styled (staminate) flowers or long-styled (pistillate or bisexual) flowers and fruit.

Papua New Guinea. Milne Bay: Rossel Island, Mt. Rossel, S. slopes, 700 m, 19 Oct 1956, Brass 28494 (A, L); Mt. Oiatau, West Misima Island, subprov. Misima, 700 m, 23 Mar 1979, Croft LAE-71421 (A, K, L, LAE); Mount Rossel, Rossel Island, subprov. Misima, 780 m, 18 Mar 1979, Katik et al. LAE-70928 (A, E, K, L, LAE), 19 Mar 1979, Katik et al. LAE-70954 (K, L, LAE). New Ireland: Lihir Island [Niolam Island], Mount Tementa, above Palie Mission, Namatanai subprovince, 710 m, 7 Nov 1984, Gideon LAE-57196 (K, LAE, L); Hans Meyer Range, on steep ridge below camp, ca. 8 km (map distance) WNW of Taron on east coast, Namatanai subprov., 1,260 m, 9 Oct 1975, Sands et al. 1966 (K), 1,075 m, 8 Oct 1975, Sands et al. 2073 (K), 1,350 m, 15 Oct 1975, Sands et al. 2230 (K).

Based on Solanum moszkowskii Bitter.

Figure 31. 

Lycianthes moszkowskii (Bitter) Bitter. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 15, as S. moszkowskii Bitter). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Erect or sprawling (climbing?) shrubs, 0.7–4.5 m tall; stems terete and very lightly ridged with decurrent leaf bases, glabrous or sparsely to densely pubescent with stiff simple uniseriate 2–4-celled trichomes to 1 mm long, these spreading or sometimes antrorse, the bases sometimes enlarged and pustulate; new growth densely pubescent with stiff antrorse simple uniseriate trichomes to 1 mm long, these soon deciduous in nearly glabrous plants; bark of older stems dark brown, glabrescent. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and shape. Leaves simple; blades of major leaves 7–11 cm long, 3–8.5 cm wide, narrowly obovate to obelliptic, widest in the distal half, discolorous, chartaceous or coriaceous, often drying dark brown; adaxial surfaces shiny, glabrous, the more pubescent plants with stiff antrorse trichomes along the somewhat keeled midrib; abaxial surfaces sparsely to moderately and evenly pubescent on veins and lamina with stiff simple uniseriate trichomes like those of the stems, these not markedly antrorse; principal veins 5–6 pairs, impressed above, the midrib slightly keeled; base acute; margins entire, sometimes somewhat ciliate with stiff simple uniseriate trichomes; apex acuminate or abruptly acuminate; petioles 0.5–1 cm long, glabrous or moderately pubescent with stiff simple uniseriate trichomes like those of the stems; blades of minor leaves 2.5–7 cm long, 1–4 cm wide, elliptic, texture and pubescence like that of the major leaves; base acute; margins entire; apex acute to acuminate; petiole absent to 0.5 cm long, glabrous or pubescent like the stems. Inflorescences axillary fascicles of 1–3 flowers, only one open at a time, pubescent with antrorse or spreading stiff uniseriate trichomes like those of the stems; pedicels at anthesis 1.8–2 cm long, ca. 0.5 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading, thick and apparently fleshy in live plants, green (?), glabrous to sparsely pubescent with antrorse simple uniseriate trichomes like those of the stems, articulated at the base; pedicel scars closely and tightly packed in the leaf axils. Buds globose, later broadly elliptic, the corolla included in the calyx tube until just before anthesis. Flowers 5-merous, heterostylous and apparently unisexual, specimens either have all short-styled flowers or all fruit, the plants possibly dioecious. Calyx tube 3–3.5 mm long, 3.5–5 mm in diameter, cup-shaped, thick and coriaceous, apparently somewhat fleshy in live plants, glabrous to sparsely pubescent with stiff antrorse simple uniseriate trichomes like those of the pedicels, appendages absent, the rim slightly thickened. Corolla 2–2.5 cm in diameter, white or purplish cream, stellate, lobed nearly to the base, interpetalar tissue absent or merely a thin margin on the corolla lobe, the lobes 1–12 mm long, 4–4.5 mm wide, spreading or reflexed, thick and fleshy, glabrous on both surfaces but densely papillate at the tips and margins, the tips cucullate. Stamens equal; filament tube minute; free portion of the filaments 1–2 mm long, glabrous; anthers ca. 3 mm long, ca. 2 mm wide, ellipsoid and slightly tapering, yellow, poricidal at the tips, the pores distally directed, not elongating to slits with age. Ovary conical to globose, glabrous; style in short-styled flowers ca. 1 mm long, in long-styled flowers ca. 5 mm long, straight, glabrous; stigma broadly capitate, the surfaces minutely papillate. Fruit a globose berry, 1.3–1.5 cm in diameter, bright red when ripe, the pericarp glabrous, thin, matte to slightly shiny, opaque; fruiting pedicels (2- immature) 3–3.5 cm long, ca. 1 mm in diameter at the base, 2.5–3 mm in diameter at the apex, spreading or pendent, not markedly woody, green (?); fruiting calyx a spreading saucer beneath the fruit, somewhat corky and rugose (fleshy in live plants?). Seeds 30–40 per berry, 5–6 mm long, 5–6 mm wide, round with a prominent wing 1.5–2 mm wide (total measurement includes the wing), yellowish tan or straw-coloured, the surfaces very shallowly pitted, the testal cells more or less sinuate in outline. Stone cells absent. Chromosome number not known.

Figure 32. 

Lycianthes moszkowskii herbarium specimen. Papua New Guinea. Morobe: Sayers NGF-21517 (BM001018996). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

(Fig. 33). Lycianthes moszkowskii is endemic to the island of New Guinea; it has been collected in Papua New Guinea (Eastern Highlands, Madang, Morobe, West Sepik, Western Highlands) and Indonesia (Papua).

Figure 33. 

Distribution of Lycianthes moszkowskii.

Lycianthes moszkowskii is a plant of several types of primary rainforests, oak and Castanopsis forests, mossy montane forests and mid-montane forests; it occurs from 600 to 2,000 m elevation.

None recorded.

(IUCN 2020). EOO (165,342 km² - LC); AOO (80 km² - EN). Lycianthes moszkowskii is known from eight localities, mostly in Papua New Guinea with some collections from protected areas in the Bulolo region (McAdams National Park). I propose a preliminary threat status of Vulnerable (VU [B2a, b(iii,iv)]) for L. moszkowskii.

Lycianthes moszkowskii is one of the more widely distributed species of Lycianthes on the island of New Guinea, only L. oliveriana is more widely distributed. The obovoid leaves, large (2–2.5 cm in diameter) flowers, large berries (to 1.5 cm in diameter) on long fruiting pedicels and large seeds (ca. 6 mm in diameter) with a prominent wing are all found only in L. moszkowskii and serve to distinguish it from all other species on New Guinea. Lycianthes moszkowskii is very variable in pubescence density; a few specimens are densely and evenly pubescent over the entire abaxial leaf surface, while others have trichomes confined to along the midrib or lack trichomes entirely; branches and leaves are usually glabrescent with age and very lightly ridged from the somewhat decurrent leaf bases. The trichomes (if present) are stiff and usually spreading, rather than strongly antrorse, as is common on other species with stiff trichomes (e.g., L. peranomala, L. rostellata). The type collection was a branch with immature berries and the protologue of S. moszkowskii does not mention any pubescence (Bitter 1917).

Many specimens of Lycianthes moszkowskii are identified as “Lycianthes cf. S. belense” by D.E. Symon in various herbaria, especially at the Naturalis Biodiversity Center (e.g. Sayers NGF-21517, L.2859575); care should be taken with early determinations of New Guinea Lycianthes on old annotation labels. Symon (1985) suggested the two species were closely related and they certainly are very similar. Lycianthes belensis has similar inflorescences with few, large flowers and calyces with no appendages but differs from L. moszkowskii in its soft, curling pubescence (versus stiff spreading trichomes of L. moszkowskii) and its calyx with tiny appendages and a rim that is transparent and appears ruffly (versus a truncate calyx with no appendages in L. moszkowskii). The flowers of L. moszkowskii are more stellate with less interpetalar tissue than those of L. belensis. Seeds of L. belensis are not known, if they are strongly winged like those of L. moszkowskii this would be another shared character.

Symon (1985) suggested that Lycianthes moszkowskii (as Solanum moszkowskii) had horticultural potential for its “handsome leaves and large red fruits, looking rather like cherries.” He had not seen long-styled (“female”) flowers, but in common with many other New Guinea Lycianthes, they are present on specimens with fruit, but not on specimens with short-styled flowers, suggesting this species is dioecious.

The type of Solanum moszkowskii was collected in the Van Rees range in northern Papua (Indonesia), an isolated range north of the main central spine of New Guinea; they are the lowest of the ten outlier mountain ranges along the north coast (Diamond and Bishop 2021) and have been very rarely explored and the area is sparsely populated. The German zoologist Max Moszkowski lost all his first set of collections from the Van Rees Mountains in and accident that occurred while descending the Edi Falls on the Mamberano River (van Steenis-Kruseman 1985). The specimen seen by Bitter (1917) was collected on the second attempt to ascend the river. His collections were all sent to Berlin and no duplicates of the type collection of S. moszkowskii have been found, despite intensive searches. In the protologue Bitter (1917) compares L. moszkowskii (as S. moszkowskii) to L. impar (as S. impar) and differentiates them based on the sessile, few-flowered inflorescence of L. moszkowskii, but he also states that they cannot be distinguished without flowers. In designating a neotype for S. moszkowskii that is in accordance with the protologue I have selected a collection with glabrous, slightly ridged stems, narrowly elliptic leaves, and that is in fruit (Rau et al. 73, neotype in LAE); it is unfortunate that it is not from nearer the type locality, but selection of Rau et al. 73 in accordance with the morphology described in the protologue and maintains the circumscription of L. moszkowskii as established by Symon (1985). The collection Streimann NGF-52968 is from nearer the type locality (in the province of Sanduan near the Papua border) and has immature fruit with the distinctive winged seeds of L. moszkowskii but is not in accordance with the protologue as the new growth is densely pubescent.

Indonesia. Papua: 15 km. southwest of Bernhard Camp, Idenburg River [=Taritatu River], small clearing in the mossy forest, 1,800 m, Jan 1939, Brass 12290 (A, L, LAE).

Papua New Guinea. Eastern Highlands: Mount Piora, Kainantu subprov., above Habi’ina village on lower slopes of Mt. Piora, 2,125 m, 7 Sep 1995, Sands et al. 1751 (K). Madang: Track between Budemu and Moro villages, S side of Finisterre Range, eastern Madang District., 21 Oct 1964, Pullen 6011 (BM, LAE); Sewe, Saidor subdistrict, 2,286 m, 10 Aug 1964, Sayers NGF-19830 (A, K, L, LAE). Morobe: Sumanzing, via Heickepe suppl., 1,829 m, 21 Oct 1938, M.S. Clemens 9071 (B); Manki ridge road, 8 Jun 1977, Conn & Kairo 444 (A, K); Mount Shungol, about 5 miles S of Wagau, 1,829 m, 12 Dec 1963, Hartley 12523 a, (A, K, L, LAE); Bulolo District, Gumi, above Gumi Village, Lower Watut TRP, 1,950 m, 17 Oct 1992, Höft 29018 (L); Mount Kandi, Wau, 2200 m, 16 Jul 1978, Kairo 62 (A, K, L, LAE); Manki, Bulolo, 600 m, 17 Jan 1976, Kairo, A. 70 (A, L, LAE), 18 Jan 1976, Kairo 73 (A, K, L, LAE), 19 Jan 1976, Kairo 79 (A, K, L, LAE); Wau, north slope of Mt. Kaindi, 2,050 m, 19 Nov 1983, Kerenga & Dao LAE-56629 (L, LAE); Mount Buruman, Wantoat, 28 May 1980, Kerenga LAE-77590 (LAE); Aseki, beside Aseki road, 2,000 m, 27 Jul 1977, Rau et al. 73 (A, K, L, LAE); Wagau, Morobe Dist., T.N.G., 1,219 m, 5 Jan 1965, Sayers NGF-21517 (BM, L, LAE); Tawa Village near Aseki, Menyamya Subdistrict, 1,700 m, 16 May 1968, Streimann & Kairo NGF-27634 (L); Piwi-anga, Menyamya-Kaintiba Road, Menyamya Subdistrict, 1,800 m, 11 May 1968, Streimann & Kairo NGF-35924 (L); Wau, Aseki road from Bulolo, 2,286 m, 29 May 1977, Symon & Crutwell 10631 (K, L, LAE, MO, US); Aseki, Aseki road, below crest, 1,950 m, 31 May 1984, Symon & Katik 13828 (LAE, MO), Symon & Katik 13830 (L, LAE, MO); Mount Missim, lower slopes, 2 Jun 1984, Symon & Kairo 13846 (L, LAE, MO); Bulolo District, Gumi, Gumi area, 1,750 m, 3 Jun 1984, Symon & Vinas 13853 (L, L, LAE), Symon & Vinas 13854 (L, LAE, MO); near Namie Creek, along Edie Creek road, on flanks of Mount Kaindi, above Wau, 1,585–1,615 m, 8 Sep 1968, Webster & Hildreth 15188 (GH). Sanduan: Vanimo hinterland, Vanimo subdistrict, 500 m, 30 Nov 1971, Streimann NGF-52968 (K, L, LAE). Western Highlands: border with Mandang Province, Bismarck range, summit ridge at Camp 2 (Mt. Oibo), 1,830–2,044 m, 10 Oct 1995, Takeuchi 10663 (A, L, LAE).

Based on Solanum multifolium Merr. & L.M. Perry

Figure 34. 

Lycianthes multifolia (Merr. & L.M.Perry) A.R.Bean. Drawing by M.L. Szent-Ivany, first published in Symon (1985: fig. 16, as S. multifolium Merr. & L.M.Perry). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Slender spindly shrubs 0.8–3 m tall, ; stems terete, densely pubescent with stiff, antrorse 1–4-celled simple uniseriate trichomes 0.2–0.7 mm long, these persistent; new growth moderately pubescent with antrorse simple uniseriate trichomes like those of the stems, these denser along the leaf veins; bark of older stems pale tan, not markedly glabrescent. Sympodial units di- or trifoliate, the leaves geminate or with several leaves at a node, the leaves at a node differing in size but not in shape. Leaves simple; blades of major leaves 5.2–9(12) cm long, 1.5–3(5) cm wide, narrowly elliptic or less commonly elliptic, widest at the middle, discolorous, membranous; adaxial surfaces glabrous or with a few antrorse simple uniseriate trichomes along the midrib and scattered stiff 2–3-celled trichomes on the lamina; abaxial surfaces similar; principal veins 6–7 pairs, the midrib keeled above; base acute to attenuate; margins entire; apex acuminate; petiole 0.3–0.6 cm long, sparsely pubescent with trichomes like those of the stems; blades of minor leaves 1–4 cm long, 0.7–2.2 cm wide, similar in shape, texture and pubescence to the major leaves; base attenuate; margins entire; apex acute to acuminate; petiole 0.2–0.5 cm long, sparsely pubescent. Inflorescences axillary fascicles of 3 flowers, only one flower open at a time, pubescent like the stems; pedicels at anthesis 0.7–0.75 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender and nodding, glabrous or with a very few simple uniseriate trichomes near the base, markedly less pubescent than the stems, articulated at the base; pedicel scars tightly packed in the leaf axils. Buds narrowly ellipsoid, the corolla strongly exserted from the calyx tube long before anthesis. Flowers 5-merous, perhaps heterostylous, the one flowering specimen (Kalkman B.W. 3479) with all short-styled flowers. Calyx tube ca. 2 mm long, ca. 3 mm wide, open cup-shaped, winged or ridged from the veins, translucent and papery, glabrous except for a few simple uniseriate trichomes ca. 0.2 mm long, with 5 tiny nub-like appendages less than 0.5 mm long arising from very close to or at the rim. Corolla 1–4-1.6 cm in diameter, white, deeply stellate, lobed nearly to the base, interpetalar tissue present, the lobes 6–7 mm long, 1.5–2 mm wide, spreading, membranous, glabrous on both surfaces, but the attenuate tip densely papillate. Stamens equal; filament tube minute; free portion of the filaments 0.75–1 mm long, glabrous; anthers ca. 1 mm long, 1.1–1.25 mm wide, ellipsoid and slightly tapering at the tips, yellow, poricidal at the tips, the pores round, directed distally, not elongating with age. Ovary and style not seen in short-styled flowers. Fruit a globose berry, ca. 0.8 cm in diameter, red when ripe, the pericarp glabrous, thin, shiny, translucent; fruiting pedicels 1.4–1.5 cm long, ca. 0.75 mm in diameter at the base, ca. 1.25 mm in diameter at the apex, not markedly woody, orientation not known; fruiting calyx a spreading saucer at the base of the fruit, not markedly woody or rugose. Seeds 10–20 per berry, 4–4.5 mm long, 2.5–3 mm wide, flattened reniform, reddish brown, the surfaces deeply pitted, the testal cells pentagonal in outline. Stone cells absent. Chromosome number not known.

Figure 35. 

Lycianthes multifolia herbarium specimen. Indonesia. Papua: Brass 12907 (isotype of S. multifolium, BM000778109). Courtesy of the Trustees of the Natural History Museum, London, reproduced with permission.

(Fig. 36). Lycianthes multifolia is endemic to the island of New Guinea; found in Papua New Guinea (Sanduan) and Indonesia (Papua).

Figure 36. 

Distribution of Lycianthes multifolia.

Lycianthes multifolia grows in rainforests from 50 to 1,150 m elevation.

None recorded.

(IUCN 2020). EOO (1,954 km² - EN); AOO (12 km² - EN). Lycianthes multifolia is known from only two widely separated localities; it clearly merits a threat status of Endangered (EN [B1,2a, b(iii, iv)]). The lowland forests where it occurs are within land use concessions (Parsch et al. 2022) in Indonesian Papua.

Lycianthes multifolia is similar morphologically to L. belensis but has smaller flowers (1.4–1.6 cm versus ca. 2.4 cm in diameter). In addition, the calyx of L. multifolia is relatively strongly winged or angled at the veins and has appendages arising from very near the rim, rather than from ca. 0.75 mm below the rim as in L. belensis. In general L. multifolia is a more delicate plant than many other New Guinea Lycianthes. The species name comes from the several leaves at each node seen on the type collection, however, Kalkman BW-3479 has clearly difoliate geminate sympodial units. It is not clear how consistent leaf number per node is across the range of L. multifolia as there are very few collections.

Indonesia. Papua: Res. Hollandia [Jayapura], Nemo, 5 m, 1 Apr 1956, Kalkman BW-3479 (A, K, L, LAE).

Papua New Guinea. Sanduan: near Daunda Bridge, Bewani Highway, subdistrict Vainimo, West Sepik, 120 m, 9 Sep 1977, Wiakabu & Mamalai LAE-70476 (A, E, K, LAE).

Based on Solanum oliverianum Lauterb. & K.Schum.

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.

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.

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.

(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]).

Figure 39. 

Distribution of Lycianthes oliveriana.

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.

None recorded.

(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).

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.

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. Aëndosa 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 & Schodde848 (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).

Based on Solanum peranomalum Wernham ex Ridl.

Figure 40. 

Lycianthes peranomala (Wernhamam ex Ridl.) A.R.Bean. Reproduced from Ridley (1922: pl. 3062, as S. peranomalum Wernham ex Ridl.). Courtesy of permission of NHM Library and Archives, reproduced with permission.

Shrub to 3 m, or a woody climber with length not recorded; stems terete, sparsely pubescent with appressed stiff antrorse simple uniseriate 1–6-celled trichomes to 1 mm long, the basal cell of each enlarged and sometimes remaining as a pustule or bump; new growth densely stiff-pubescent, the trichomes simple, uniseriate and strongly antrorse; bark of older stems brown, glabrescent, somewhat rugose and corky. Sympodial units unifoliate or difoliate, if difoliate the leaves geminate, the leaves of a pair different in size and sometimes shape. Leaves simple; blades of major leaves 9–16 cm long, 4–7 cm wide, elliptic, widest at the middle, the two sides occasionally uneven in size with the basiscopic half narrower, concolorous or somewhat discolorous, chartaceous or coriaceous; adaxial surfaces bullate (fide Takeuchi 11204), glabrous, the midrib keeled; abaxial surfaces glabrous, the veins prominent; principal veins 8–9 pairs, yellowish abaxially; base acute, oblique; margins entire; apex acuminate with an elongate drip-tip; petiole 0.7–1.1 cm long, sparsely pubescent with a few scattered stiff antrorse simple uniseriate trichomes on the adaxial surfaced and near the base; blades of minor leaves 1–2.5(4) cm long, 1–2.2 cm wide, elliptic to orbicular or heart-shaped, often apparently clasping the stem (“retrorsely directed” fide Takeuchi 11204), similar in texture and pubescent to the major leaves; base cordate or rounded; margins entire, usually revolute, sometimes markedly so; apex abruptly acute; petioles 0.1–0.2 cm long, glabrous or sometimes with a few simple trichomes like those of the stems. Inflorescences axillary fascicles of 8–10 flowers, several open at once, sparsely pubescent with stiff antrorse trichomes to 0.5 mm long like those of the stems; pedicels at anthesis 0.6–0.7 cm long, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading (?), white or pale purple, sparsely pubescent with stiff antrorse simple uniseriate trichomes like those of the stems, ca. 0.5 mm long; pedicel scars clustered in the leaf axils; buds ellipsoid, the corolla ca. halfway exserted from the calyx tube before anthesis. Flowers 5-merous, heterostylous and unisexual, specimens with either short-styled flowers or long-styled flowers and fruit, the plants possibly dioecious. Calyx tube 2.5–3 mm long, 2–2.5 mm wide, urn-shaped, the rim somewhat constricted, thick and woody (dry) or fleshy (live plants?), with no appendages, sparsely pubescent with stiff trichomes like those of the pedicels, these deciduous. Corolla 0.6–0.8 cm in diameter, purple, stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 3–4 mm wide, 1.2–2 mm wide, reflexed, thick and fleshy, adaxially glabrous with a prominent ridged midvein, abaxially glabrous or sparsely papillate, densely papillate on tips and margins. Stamens equal; filament tube minute; free portion of the filaments ca. 1 mm long, glabrous; anthers 1.5–2 mm long, 1–1.5 mm wide, plumply ellipsoid, yellow, poricidal at the tips, the pores distally directed, circular, not elongating to slits with age. Ovary conical, glabrous; style in short-styled flowers ca. 0.5 mm long, in long-styled flowers 4–4.5 mm long, glabrous; stigma capitate or minutely bilobed, the surfaces minutely papillate. Fruit a globose berry, 0.5–0.9 cm in diameter, colour at maturity not known, the pericarp brittle in dry material, glabrous, matte, opaque, fruiting pedicels 0.5–0.8 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, stiff and somewhat woody and tuberculate, spreading; fruiting calyx a cup at the base of the fruit, covering less than 1/4 of the berry, somewhat tuberculate, with a few stiff trichomes, but these usually deciduous. Seeds 10–20 per berry, ca. 2 mm long, ca. 1.5 mm wide, flattened reniform or slightly tear-shaped, reddish brown, the surfaces deeply pitted especially on the thickened margins, the testal cells pentagonal in outline. Stone cells absent. Chromosome number not known.

Figure 41. 

Lycianthes peranomala herbarium specimen. Papua New Guinea. Chimbu: Takeuchi 11204 (A). Courtesy of the Herbarium of the Arnold Arboretum of Harvard University, reproduced with permission.

(Fig. 42). Lycianthes peranomala is endemic to the island of New Guinea; known from Papua New Guinea (Chimbu, Madang, Oro) and Indonesia (Papua).

Figure 42. 

Distribution of Lycianthes peranomala.

Lycianthes peranomala occurs in lowland rainforest and riverine forests, between 50 and 640 m elevation.

None recorded.

(IUCN 2020). EOO (189,160 km² - LC); AOO (16km² - EN). Lycianthes peranomala is known from four localities, some of which are within protected areas. Based on EOO alone, it would merit a status of Least Concern, but given the few collections, its forest habitat and general lack of knowledge about the species, I propose a preliminary threat status of Vulnerable (VU [B2a, b(iii,iv)]) for L. peranomala.

Lycianthes peranomala is vegetatively similar to L. impar in its orbicular to heart-shaped minor leaves and relatively large, elliptic major leaves with prominent venation. They differ in trichome morphology, L. peranomala has sparse stiff, strongly antrorse trichomes on stems, veins near the leaf base and calyces, while those of L. impar are soft and curling and found on stems only. Inflorescences of L. impar are elongate with paired pedicel scars, while those of L. peranomala are strictly axillary.

Like Lycianthes kaernbachii and L. oliveriana the flowers of L. peranomala are usually less than 1 cm in diameter with fleshy corolla lobes lacking any interpetalar tissue and the plants are possibly dioecious with short-styled flowers and long-styled flowers (and fruit) on different plants. Lycianthes peranomala can be distinguished from those taxa in its minor leaves that are very dissimilar in shape to the major leaves; both L. kaernbachii had L. oliveriana have minor leaves that are different in size but not so strongly dissimilar in shape.

The name Solanum peranomalum was proposed by Herbert Fuller Wernham, an assistant in the Botany Department of the British Museum, for plants collected by Cecil Boden Kloss on the 1912–1913 expedition led by the ornithologist Arthur Wollaston in a second attempt to reach the high peaks of the “Snow Mountains” (Mount Jaya) in the central range (Ridley 1916). His publication of S. peranomalum was superseded by Nicholas Ridley’s publication of the same name, with an illustration (Fig. 40) but a slightly different and less complete description a few months earlier. Ridley perhaps was inpatient for the Transactions to appear or had little regard for Wernham at the Museum, who had a tragic career cut short by mental health issues and alcoholism (Stearn 1981). It is strange that neither of the two other species described by Wernham (1916) were published by Ridley (S. ridleyanum and S. wollastonii).

Indonesia. [type only]

Papua New Guinea. Chimbu: Crater Mountain Wildlife Management Area, Vicinity of Haia, along the Wara oo streamcourse (first river E of Mt. Widau), 640 m, 16 Sep 1996, Takeuchi 11204 (A, BM, K, L, LAE, US). Madang: Ohu Village, 100 m, 7 Aug 2008, Ctvrtecka 1644 (US). Oro: Kokoda, 365 m, 22 Mar 1936, Carr 16195 (BM, K, NY).