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.

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.

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

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.

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.

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

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.

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).

(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.

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.

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.

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

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.

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.

(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).

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.

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.

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

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.

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.

(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).

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.

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.

(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.

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]).

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.

(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).

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.

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.

(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).

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

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.

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

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.

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.

(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]).

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.

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.

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

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).

Based on Solanum rantonnetii Carrière.

Shrubs 0.5–3 m tall, with multiple stems from the base, these arching and sometimes scandent and sprawling; stems 3–4-angled, the angles yellowish green in live plants and paler than the rest of the stem, sparsely to moderately pubescent with spreading transparent simple uniseriate 1–4-celled trichomes to 0.5 mm long, these occasionally forked or dendritic, glabrescent with age; new growth moderately pubescent with transparent simple uniseriate or occasionally dendritic trichomes like those of the stems; bark of older stems pale greyish brown, prominently angled. Sympodial units unifoliate or more usually difoliate, the leaves usually geminate, if paired the leaves similar in shape and size. Leaves simple; blades of major leaves (1)4–15.5 cm long, (0.5)3.5–7.5 cm wide, ovate, rhombic-elliptic, elliptic or occasionally almost lanceolate, usually broadest at the middle, rarely in the upper half, membranous, concolorous; adaxial surfaces sparsely and evenly pubescent with 1–3-celled simple uniseriate trichomes, these denser along the midrib; abaxial surfaces sparsely to moderately and evenly pubescent with 1–3-celled simple uniseriate trichomes, these denser along the midrib; principal veins 3–7 pairs, more pubescent than the lamina. drying yellowish green abaxially; base attenuate onto the petiole; margins entire or somewhat undulate; apex acute to acuminate; petiole 0.5–2.4(4) cm long, winged from the attenuate leaf base, pubescent with simple uniseriate (or occasionally dendritic) trichomes like those of the stems and leaves; blades of minor leaves similar in size and shape to those of the major leaves, or slightly smaller; petioles 0.5–3 cm long, winged. Inflorescences axillary fascicles with (1)2–7 flowers, pubescent with transparent trichomes like those of the new growth and stems; pedicels 1.2–1.7 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading at anthesis, sparsely to moderately and evenly pubescent with transparent simple (occasionally dendritic) uniseriate 1–3-celled trichomes like those of the stems, articulated at the base; pedicels scars tightly packed in the leaf axils. Buds ellipsoid to fusiform with pointed tips, the corolla more than halfway exserted from the calyx tube before anthesis. Flowers 5-merous, all apparently perfect. Calyx with the tube 1.5–4 mm long, 2.5–4.5 mm wide, openly cup-shaped, with (5)10 linear subulate appendages of variable length 0.25–5.2 mm long, arising ca. 0.25–1 mm from the hyaline rim, usually alternating long and short, sparsely to moderately pubescent with simple trichomes like those of the pedicels. Corolla 1.2–2 cm in diameter, violet with the midveins dark purple and the centre yellow, rotate, lobed less than 1/10 of the way to the base, interpetalar tissue abundantly present, the lobes ca. 1 mm long, ca. 1 mm wide and mere acumens from the rotate corolla, glabrous on both surfaces except for the densely papillate, cucullate tips (acumens). Stamens unequal; filament tube minute; free portion of the filaments of two lengths, three long filaments 2–3 mm long, 2 short filaments 0.8–1.5 mm long, glabrous or adaxially pubescent with tangled weak-walled uniseriate simple trichomes; anthers ellipsoid and slightly curved, orange-yellow, poricidal at the tips, the pores round, distally directed, not elongating to slits with age. Ovary conical, glabrous; style 3.5–5.5 mm long, slightly curved in the same direction as the anthers, glabrous; stigma slightly clavate and bilobed, the surface minutely papillate. Fruit a compressed-ellipsoid or compressed globose berry, 2–3 cm long, 1.3–1.5 cm in diameter (usually absent or smaller and seedless in cultivated plants), yellow or yellowish orange when mature, the pericarp glabrous, thin, shiny and translucent; fruiting pedicels 2.5–4 cm long, ca. 1.5 mm in diameter at the base, ca. 3 mm in diameter at the apex, somewhat woody, spreading or hanging from the weight of the berries; fruiting calyx a plate with the appendages somewhat longer than in flower, spreading and often broken off, stiff and woody. Seeds 20–100 per berry (many fewer in cultivated plants), 2–3.5 mm long, 1.5–3.5 mm wide, rounded and compressed, reddish tan, the surfaces minutely pitted, the testal cells with sinuate margins. Stone cells more than 20 per berry, ca. 0.5–1.5 mm in diameter. Chromosome number: 2n=24 (Gerasimenko and Reznikova 1968 [cited in D’Arcy 1974] as Solanum rantonnetii; Acosta et al. 2005, as L. rantonnei, voucher Moscone 4260 [CORD]).

Lycianthes rantonnetii is widely cultivated in the tropics and subtropics (and even into the temperate zone as a short-lived perennial) worldwide. I have seen no collections from New Guinea, but it has been relatively widely collected in Australia. It is native to southern South America (Argentina, Bolivia, southern Brazil and Paraguay).

In its native range L. rantonnetii is a plant of semi-moist, seasonal forests and open areas; from (sea level) 100 to 2,000 m elevation.

In its native range in Argentina L. rantonnetii is called meloncillo del aire (Barboza 2013).

(IUCN 2020). Not applicable to this species for this region.

Lycianthes rantonnetii is native to South America (Barboza 2013) but widely cultivated in subtropical and temperate areas worldwide. It is the only species occurring in the region treated here that has stone cells in the berries, but in cultivation it rarely sets fruit. It can easily be distinguished from all native species by its rotate corollas with copious interpetalar tissue, orange-yellow anthers that are slightly curved and angled, and somewhat striped stems. In the area treated here I have only seen specimens of L. rantonnetii from Australia, but I would expect it to be in cultivation anywhere in the region.

This species epithet is often seen spelled “rantonnei” but is correctable to “rantonnetii” following Art. 60.9 of the ICN (Turland et al. 2018: Ex. 31), which stipulates that epithets honouring persons where there is an intentional latinisation of the name that involves the omission of a terminal vowel or consonant are not permitted; the epithet in this case honours the French horticulturalist Barthélémy Victor Rantonnet so is correctable to “rantonnetii” even though Carrière (1859) originally spelled in “rantonnei” (using the latinisation Rantonneus).

Dean et al. (2020) cited the type specimens of Solanum urbanum var. foliosum and var. subtomentosum as “holotype”, but Chodat (1916) did not cite a specific herbarium in any part of the protologue thus these names require lectotypification (see McNeill 2014), even though only a single specimen is preserved in the herbarium at G. I have selected the specimens cited as holotypes by Dean et al. (2020) as the lectotypes in both these cases.

Australia. New South Wales: North Coast, 37.5 km from the Bruxner Highway along Long Gully Road, next to Rocky River., 3 Feb 2014, Johnstone 3435 (NSW). Queensland: Wide Bay, cultivated at Nora Creina, Homestead, 1.5km north-northeast of Didcot., 24 Jan 1999, Forster 24067 (BRI); Moreton, Brisbane Mt. Coot-Tha Botanic Gardens, 1 Mar 1984, Whitten s.n. (BRI). South Australia: in creek near Fremont Park, Elizabeth, 10 Feb 1988, Bates 13861 (AD); Rocky River S of Laura, 12 Jan 1993, Bates 30884 (AD); in creek NW below entrance to Angove C[onservation] P[ark], (dedicated 2 weeks ago), 15 Jul 1994, Bates 37051 (AD); Naracoorte, Jul 1940, Cooper s.n. (AD); Region 13, Southeastern Drain L., Oct 1989, Lawson 51 (AD, CANB, MEL, MO); On the roadside at Winding Way, Belair, by vacant block, 18 Feb 1976, Sparrow s.n. (AD); Tusmore, Kenneway St. (rear of No. 20 Lynington St.), 14 Dec 1974, Symon s.n. (CANB). Western Australia: Forrestfield, 1948, Dawson Harrison Ltd s.n. (PERTH); Mingenew tip, 30 Apr 1997, Elson s.n. (PERTH).

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

Shrubs or woody vines, 0.3–3 m tall (long); stems terete, densely pubescent with stiff, antrorse simple 4–5-celled uniseriate trichomes to 0.5 mm long, these sometimes from multicellular bases or the bases becoming enlarged with plant growth(i.e., remnants of multicellular bases on old stems but young stems without); new growth sparsely to densely pubescent with stiff simple uniseriate trichomes like those of the stems; bark of older stems brownish grey, glabrescent, corky and warty with persistent multicellular trichome bases. Sympodial units unifoliate (with deciduous minor leaves) or difoliate, the leaves geminate, the leaves of a pair different in size and shape. Leaves simple; blades of major leaves 7–12(17) cm long, 1–5 cm wide, elliptic or more commonly narrowly elliptic, widest at the middle, lower leaves on stems broader than distal ones, discolorous, thick and chartaceous (papery fide Takeuchi 11804), slightly bullate; adaxial surfaces shiny, glabrous, the veins deeply impressed; abaxial surfaces glabrous or with a few antrorse simple uniseriate trichomes like those of the stems along the midrib, sometimes purple-tinged (fide Womersley 4883); principal veins 4–6 pairs, impressed above, prominent below and occasionally sparsely pubescent; base acute; margins entire, revolute; apex abruptly attenuate to a long drip-tip; petioles 0.3–1.2 cm long, sparsely pubescent with antrorse simple uniseriate trichomes to ca. 0.5 mm long like those of the stems; blades of minor leaves 0.3–0.7 cm long, 0.3–0.7 cm wide, orbicular or somewhat heart-shaped, texture and pubescence like that of the major leaves, often deciduous especially on reproductive stems; base truncate or cordate; margins entire, revolute; apex rounded; petioles less than 0.1 cm long, sparsely pubescent. Inflorescences axillary fascicles of 2–4 flowers, usually only a single flower open at a time, densely pubescent like the stems with stiff antrorse simple uniseriate trichomes; pedicels at anthesis 0.8–1 cm long, ca. 0.5 mm in diameter at the base, 1–1.5 mm in diameter at the apex, erect (?) or spreading, sparsely pubescent with stiff antrorse simple trichomes like those of the stems, articulated at the base; pedicel scars tightly packed in the leaf axils. Buds long-ellipsoid, the corolla strongly exserted from the calyx tube just before anthesis, included in early buds. Flowers 5-merous (6-merous in Womersley 4883), apparently all perfect, heterostyly not observed. Calyx tube 2.5–3 mm long, 3–3.5 mm wide, cup-shaped, woody and stiff in dry material, perhaps fleshy in live plants, white or cream-colored, with no appendages, sparsely pubescent with stiff antrorse simple uniseriate trichomes ca. 0.5 mm long. Corolla 1.4–2 cm in diameter, white tinged with purple, purplish blue or purple, deeply stellate, lobed nearly to the base, interpetalar tissue absent, the lobes 8–10 mm long, 1.5–2 mm wide, reflexed or spreading, membranous/fleshy, glabrous on both surfaces, with minutely papillate tips and margins. Stamens equal; filament tube minute; free portion of the filaments 0.75–1 mm long, glabrous or with a few stiff simple uniseriate trichomes abaxially; anthers 5–6 mm long, ca. 1 mm wide, ellipsoid and somewhat tapering at the tips, yellow, poricidal at the tips, the pores round, distally directed, not elongating to slits with age. Ovary conical, glabrous; style 5.5–6 mm long, straight, glabrous (white fide Takeuchi 11804); stigma clavate, the surfaces minutely papillate. Fruit a globose berry, often apically umbonate, 0.7–0.8 cm in diameter, green or dark green (immature?), ripening purple-black, the pericarp thick and somewhat woody (fruits immature?), matte, opaque; fruiting pedicels 1.8–2.1 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, erect or spreading, somewhat woody, the surfaces thickened and rugose; fruiting calyx a stiff, woody plate beneath the fruit, not enclosing the base. Seeds 60–80 per berry, 2–2.5 mm long, 1.5–2 mm wide, flattened reniform with a deep notch, yellowish golden, the surfaces deeply pitted, especially at the thickened margins, the testal cells pentagonal in outline with slightly sinuate walls. Stone cells absent. Chromosome number not known.

(Fig. 47).Lycianthes rostellata is endemic to the island of New Guinea and is relatively widely distributed; it has only been collected in Papua New Guinea (Central, Eastern Highlands, Enga, Oro, Western Highlands).

Lycianthes rostellata grows in montane and submontane forests, as well as riverine regrowth forests, from 740–2,500 m elevation.

Papua New Guinea: baga (Henty et al. NGF-41640)

(IUCN 2020).EOO (94,873 km² - LC); AOO (84 km² - EN). Lycianthes rostellata is known from more than 10 localities, some of which are within protected areas (e.g., Crater Mountain Wildlife Area). It is currently a single-country endemic. This suggests a preliminary threat status of either Least Concern (LC) or Near Threatened (NT).

Lycianthes rostellata is one of the larger-flowered species of Lycianthes on New Guinea. Like L. bambusarum, L. belensis and L. moszkowskii the corolla is ca. 2 cm in diameter; it can be distinguished from L. bambusarum by its pubescent rather than glabrous or minutely papillate new growth, and from L. belensis and L. moszkowskii by its stiff antrorse pubescence of trichomes with multicellular bases. Lycianthes belensis has soft, curled trichomes on the new growth, and L. moszkowskii has antrorse pubescence, but it is sparser than that of L. rostellata and the trichomes never have multicellular bases. Flower buds of L. rostellata are long-ellipsoid and pointed at the tips (the character upon which the specific epithet is based). Lycianthes shanesii of Australia has similar pointed buds, but the corolla lobes are wider relative to their length (2–3 times longer than wide) that are those of L. rostellata (4–5 times longer than wide) and the mature berries of L. shanesii are bright red, while those of L. rostellata are purple-black.

Plants named as Solanum pustulatum (Symon 1985) represent an extreme in trichome morphology with prominent multicellular bases that persist after the uniseriate part of the trichome breaks off, but there is much variation in this character, from no to large multicellular bases as well as in the density of these types of trichomes. The specimen illustrated by Symon (1985: fig. 22,) has unusually wide leaves and persistent minor leaves, most collections of Lycianthes rostellata as circumscribed here have narrower, lance-elliptic leaves and the minor leaves are often deciduous with the sympodial units apparently unifoliate. Plants not in flower also appear to have wider leaves (Symon & Katik 10690, 10677), so there may be some change in leaf shape as stems reach reproductive phase (e.g., Roe 1966).

Papua New Guinea. Central: Boridi, 1,524 m, 3 Oct 1935, Carr 14357 (BM, K, NY); subdsitrict Port Moresby, east slope of Lake Myola No. 1, 1,920 m, 17 Sep 1973, Croft & Lelean NGF-34800 (E, K, L, LAE, MO, QRS, US); Efogi environs, Owen Stanley range,, 1830 m, 14 Sep 1970, Schodde 5705 (A, K, L, LAE); Murray Pass, near the summit, 2,650 m, 13 Jun 1984, Symon 13874 (K, L, LAE, MO); Murray Pass, near crest of pass, 2,800 m, 13 Jun 1984, Symon 13875 (K, L, LAE, MO); Murray Pass, “Mur Mur Pass”, 2,700 m, 13 Jun 1984, Symon 13876 (K, L, LAE), Symon 13877 (K, L, LAE, MO). Eastern Highlands: above Fatima River, Marafuga, Sub-dist. Goroka, 2,100 m, 13 Nov 1968, Millar NGF-40737 (K, L, LAE); Daulo Pass, top of Daulo Pass, 2,320 m, 22 Jun 1977, Symon & Katik 10677 (K, L, LAE); near summit of Daulo, 22 Jun 1977, Symon 10678 (MO, US); Crater Mountain Wildlife Area, E of Haia village along Wara Sename, Chimbu (Simbu) province, Crater, 740 m, 16 Mar 1997, Takeuchi 11804 (K, L, LAE, US); Nondugl, Al River Valley, 2,134 m, 7 Apr 1953, Womersley 4883 (A, BM, K, L, LAE). Enga: between Mount Hagen and Wabog, 42 km from Mount Hagen, 29 km after turnoff to Wabog, 13 km before Waterfall Village, 2,520 m, 23 Jun 1977, Symon 10687 (K, L, LAE, MO); Wahgi-Sepik Divide, from Banz to Tabibunga, 4 km after crest and 39 km before Tabibunga, 27 Jun 1977, Symon 10704 (K, L, LAE, MO). Oro: Alola, 1,829 m, 4 Dec 1935, Carr 13611 (BM, K, NY), 1,981 m, 11 Dec 1933, Carr 13737 (K), 11 Dec 1935, Carr 13738 (BM, K, NY); eastern side lake Myola No.1 (Northern District), Kokoda subdistrict, 2,000 m, 23 Jul 1974, Croft et al. LAE-61993 (A, K, LAE). Sanduan: Bulindip, W of Oksapmin, Telefomin subdistrict, 1,981 m, 19 Oct 1968, Henty et al. NGF-41640 (A, K, L, LAE). Southern Highlands: Onim Hill, Mt. Gilwe Timber area, Mendi subdistrict, 2,500 m, 19 May 1975, Argent 8/ 19 (K); Mount Giluwe, track from Onim to SW summit [Eastern Highlands on label], 2,290 m, 19 Jul 1976, van Royen 11511 (K, L, LAE); between Nol and Mendi, 24 km from Mendi [georef to Mendi], 24 Jun 1977, Symon & Katik 10690 (K, L, LAE, MO). Western Highlands: Wapalepa, Kepaka, Upper Kaugel Valley, Hagen [Mt Hagen?], 2,652 m, 13 Jul 1969, Bowers 796 (L, LAE, US); Waghi & Jim Divide, Minz subprovince, 10 Aug 1981, Kerenga & Croft LAE-77644 (K, L, LAE); Kundip, Mt. Hagen subdistrict, 2,134 m, 10 Sep 1963, Millar & Garay NGF-18664 (GH, LAE); Wabag Road, 1.5 miles from turn-off, Mount Hagen subdistrict, 2,469 m, 30 Sep 1968, Vandenberg et al. NGF-39883 (A, K, L, LAE).

Based on Solanum shanesii F.Muell.

Shrubs or small trees 2–8 m tall, with single trunks to 10 cm diameter at breast height; stems terete, glabrous, with prominent white lenticels; new growth minutely puberulent with simple uniseriate golden 2–4-celled trichomes less than 0.5 mm long; bark of older stems pale greyish brown, blistering and peeling on older stems. Sympodial units difoliate, the leaves geminate, the leaves of a pair similar in shape, differing only somewhat in size. Leaves simple; blades of major leaves 7–12 cm long, 2.5–7.2 cm wide, elliptic to elliptic-obovate, broadest in the upper half or rarely at the middle, membranous, concolorous; adaxial surfaces completely glabrous; abaxial surfaces completely glabrous; principal veins 5–6 pairs, these sometimes purple abaxially (fide Clarkson 4586); base attenuate onto the petiole; margins entire (undulate fide Clarkson 4586); apex acute to acuminate; petiole 1–2.5 cm long, winged from the attenuate leaf base, with a few golden trichomes less than 0.5 mm long near the base; blades of minor leaves 2.5–7.5 cm long, 1–5.3 cm wide, like the major leaves in shape, texture, and pubescence; petioles 0.5–1.5 cm long. Inflorescences axillary fascicles with (1)2–3 flowers, minutely puberulent with a few golden simple uniseriate trichomes like those of the new growth; pedicels 1.2–1.7 cm long, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, spreading at anthesis, almost completely glabrous, but often with a few golden simple uniseriate trichomes like those of stems along their length, articulated at the base; pedicels scars tightly packed in the leaf axils. Buds ellipsoid to fusiform with pointed tips, the corolla ca. halfway exserted from the calyx tube before anthesis, in young buds the calyx completely closed. Flowers (4)5-merous, heterostylous, the plants possibly andromonoecious (fide Symon and Clarkson 1985). Calyx with the tube ca. 3 mm long, ca. 4 mm wide, openly cup-shaped, with no appendages, the hyaline calyx rim ca. 0.5 mm wide, often irregularly torn and apparently 4–5-lobed, glabrous or with a few scattered simple trichomes like those of the pedicels. Corolla 1.6–2 cm in diameter, dark purple with the midveins darker than the surrounding tissue, stellate, lobed ca. 3/4 of the way to the base, thin edge of interpetalar tissue present, the lobes 7–7.5 mm long, 2.5–3 mm wide, spreading, glabrous on both surfaces except for the densely papillate, cucullate tips. Stamens equal or slightly unequal (fide Symon and Clarkson 1985); filament tube minute; free portion of the filaments 1–1.2 mm long, glabrous; anthers 4–4.5 mm long, 1–1.25 mm wide, ellipsoid and slightly tapering at the tips, yellow, poricidal at the tips, the pores tear-drop shaped, elongating to slits with age. Ovary conical, glabrous; style in long-styled flowers ca. 6 mm long, straight, glabrous, in short-styled flowers ca. 1.5 mm long, glabrous; stigma minutely capitate, the surface minutely papillate. Fruit a globose berry, 1.3–1.5 cm in diameter, bright red when mature, the pericarp glabrous, thin, shiny and somewhat translucent; fruiting pedicels 1.8–3.5 cm long, 1–2 mm in diameter at the base, 2–4 mm in diameter at the apex, somewhat woody, spreading or slightly hanging from the weight of the berries; fruiting calyx a plate with an undulate margin (appearing “ruffly”) subtending the berry, somewhat stiff and woody, the margins appearing lobed. Seeds 50–100 per berry, 3.5–4.5 mm long, 3–3.5 mm wide, flattened reniform, pale yellowish tan, the margin darker and thickened and the seed looking almost winged, the body of the seed shallowly pitted, the margins with the testal cells deeper, the testal cells with sinuate margins. Stone cells absent. Chromosome number: n=12 (Symon and Clarkson 1985; voucher Clarkson 4585).

(Fig. 50).Lycianthes shanesii is endemic to Australia from central eastern Queensland and the York Peninsula.

Lycianthes shanesii grows in deciduous and semi-deciduous forests (“closed forest pockets” fide Symon and Clarkson 1985) and monsoon forests and is relatively common where it occurs; from sea level to 450 m elevation.

None recorded.

(IUCN 2020).EOO (349,347 km² - LC); AOO (196 km² - EN). Lycianthes shanesii is known from more than 10 localities and has quite a wide distribution on the eastern coast of the Cape York Peninsula and on islands in the Torres Strait; populations occur in lands managed by aboriginal peoples for conservation. This suggests a preliminary threat status of Least Concern (LC) but documentation of threats to the dry forests of northern Australia may cause this to be modified.

Lycianthes shanesii was described based only on fruiting collections and not included in Bitter (1919) nor in Symon (1981a); Bitter, writing between the two World Wars in Europe, would not have had access to the specimens held in Australia, and Symon (1981a) thought it was a species of Capsicum. Later, Symon and Clarkson (1985) described it in full (as S. shanesii) from Clarkson’s flowering Queensland collections, where the poricidal anthers clearly showed it was not Capsicum. Symon and Clarkson (1985) realised it was distinct from the taxa native to nearby New Guinea and suggested it was closely related to L. synanthera and L. heteroclita (Sendtn.) Bitter of Central America, rather than to other species from southeast Asia or New Guinea. They (Symon and Clarkson 1985) discounted long-distance dispersal for the occurrence of L. shanesii (as S. shanesii) in Australia.

Lycianthes shanesii was included in the Solanaceae-wide phylogeny of Särkinen et al. (2013) and resolved as sister to the clade of Lycianthes that included all the Asian species sampled (L. biflora and L. lysimachioides (Wall.)Bitter) plus L. synanthera and other American species. Resolution of the relationship between Lycianthes and Capsicum has proved difficult (see Spalink et al. 2018), and none of the New Guinea or other southeast Asian taxa have ever been included in a phylogenetic analysis.

Like Lycianthes vitiensis, L. shanesii is a tree with a single trunk, unlike most of the New Guinea species that are shrubs or woody vines. Lycianthes shanesii grows in deciduous or semi-deciduous forests, rather than the wet, mossy forests inhabited by the other taxa treated here. It can be distinguished from L. vitiensis by its sparse, golden pubescence on the new growth and young stems; L. vitiensis has dense, tangled, reddish brown pubescence on stems and young leaves. The inflorescence of L. shanesii is strictly axillary and few-flowered while that of L. vitiensis has a short axis and is many-flowered.

The flower buds of Lycianthes shanesii resemble those of L. rostellata of New Guinea in being long-ellipsoid and somewhat beaked and the anthers are large (4–4.5 mm long in L. shanesii and 5–6 mm long in L. rostellata) and similarly tapered at the tips. Lycianthes shanesii is easily distinguished from L. rostellata in its distribution and habitat, and also by the wider corolla lobes (3–4 times longer than wide in L. shanesii, 4–5 times longer than wide in L. rostellata). Lycianthes shanesii also clearly differs in the pubescence of young stems; L. rostellata has dense, stiff antrorse pubescence on young stems while L. shanesii has sparse, golden trichomes on the young stems.

Symon and Clarkson (1985) cited Symon (1981b) as the designation (“proposal”) of the lectotype for Solanum shanesii. The citation of lectotype in Symon (1981b) is ambiguous and could refer to either of the two syntypes (O’Shanesy 6 ser. 1 or Dallachy 435), thus I consider the explicit citation of the O’Shanesy collection as lectotype in Symon and Clarkson (1985) as the effective lectotypification of this name.

Australia. Queensland: Mount Stuart, 9 km S of Townsville, 14 Dec 1991, Bean 3865 (AD, BRI); South Kennedy, Hazelwood Gorge, 13 km SSW of Eungella, 15 Dec 1992, Bean 5271 (BRI); Wigton Island, c. 50km NE of Mackay, 28 Jun 2000, Bean & Champion 16706 (BRI); Cook, 11.8 km N of the Palmer River on the Peninsula Development Road, 450 m, 23 Dec 1981, Clarkson 4217 (AD, BRI, DNA, K, MBA, MEL, QRS); 11.7 km N of the Palmer River on the Peninsula Development Road, ca. 500 NE of the road, 450 m, 14 Mar 1983, Clarkson 4585 (AD, BRI, CANB, K, MBA, MEL, MO, NSW, QRS), 14 Mar 1983, Clarkson 4586 (AD, BRI, K, MBA, QRS), 31 Jan 1984, Clarkson 5131 (AD, BRI, CANB, K, QRS); 3.4 km N of Spear Creek on the Peninsula Development Road, 11.3 km N of the Palmer River Crossing, 4 Mar 1987, Clarkson & McDonald 6674 (BRI, QRS); 12 km from the East Normanby River crossing on the Lakeland Downs to Cooktown road, 6 Mar 1987, Clarkson & McDonald 6768 (BRI, QRS); Dauan Island, Mount Cornwallis, 17 Feb 1993, Clarkson 7813 (AD, BRI, CANB, DNA, MO, NSW); Mt White, c. 2 km SSW of Coen, 20 Dec 1989, Clarkson 8205 (AD, BRI, MBA, MEL, QRS); North Kennedy, Granite Ironbark Hill, high range army training area, 31 Dec 1996, Cumming 15486 (BRI); Mt Fox, 18 Feb 1997, Cumming 15841 (BRI); Rockhampton, 17 Mar 1863, Dallachy 345 (MEL, P); Cook, Hannibal Island, near Shelburne Bay, ca. 16 km W of Helby H, 3 Jul 1969, Done s.n. (BRI); Browns Peak, 75.4km ENE of Lakefield Homestead, Starcke Pastoral Holding (GR 7868-661813) RF site 49, Cape York Peninsula, 9 May 1993, Fell et al. 3232 (BRI); Cape Melville National Park NP 4 Eumangin/Temple Creek catchment, 11.5km NW of Barrow Point, 79.2 km NNE of Lakefield Ranger Base, 4 May 1994, Fell 4314 (BRI); Altanmoui Range, Cape Melville National Park, 1.6 km E of Flat Hill, 62.6 km NE of Lakefield Ranger Base, 4 May 1994, Fell & McDonald 4354 (BRI); Pulu Islet, off western shore of Mabuiag Island, Torres Strait, 14 Apr 2009, Fell 10000 (BRI, CNS, DNA); South Kennedy, Mount Bella Vista, 17 Dec 1992, Fensham 579 (BRI); Leichhardt, ‘Clifton’, northern Boomer Range, 7 Feb 1993, Fensham 718 (BRI); Back Creek, “Killarney”, Connors Range, 1 Mar 1993, Fensham 749 (AD, BRI); Leichhardt, ‘Fort Cooper’, 7 Apr 1993, Fensham 823 (BRI); Cook, NPR166, Black Mountain, Helenvale Road, Site 17, 12 Mar 2001, Ford & Holmes 2647 (BRI); Port Curtis, Camoo Caves, 11 Jun 1989, Forster & Tucker . 5108 (BRI); SF 471, Mount Coulston, 4 Oct 1989, Forster & Bean 5808 (BRI); Pine Mountain, State Forest 79, 21 Apr 1991, Forster 8012 (BRI, MEL); Cook, Possum Scrub, 22 Jun 1994, Forster & Tucker 15280 (BRI); Bolt Head, Temple Bay, 26 Jun 1996, Forster 19385 (BRI); 1.5 mls E of the highway, Byerstown Range, May 1980, Godwin C-881 (QRS); Kings Plains Station, Barron Range SW of Cooktown, 7 Jun 1983, Godwin C-2416 (QRS); Maitland Downs Holding, 560 m, 26 Jan 1989, Gray 4972 (DNA, MO, QRS), Gray 4974 (QRS); Yam Island, Torres Strait, 8 Feb 1989, Gray 4981 (QRS); Kum Kum Range, Nychum Station, 6 Apr 1996, Gray 6706 (QRS); OK Mine Road, 11 km from Burke Developmental Road, 2 Apr 2002, Gray 8068 (QRS); Cook, Great Barrier Reef, Restoration Rock, near Cape Weymouth, Portland Roads, 24 Jul 1969, Heatwole s.n. (BRI); Great Dividing Range, S of Byerstown, 10 Jun 1971, Hyland 5222 (QRS); Lankelly Creek Road, 6 Apr 1976, Hyland 8713 (QRS); Mutee Head, 26 May 1981, Hyland 11071 (QRS); Maitland Downs Holding, Parish of Byerstown, 13 Dec 1988, Hyland 13764 (QRS), 11 Mar 1993, Hyland, B. 14694 (QRS); Mt White, Coen, 22 Apr 1993, Hyland 14781 (QRS); Haggerstone Island, 10 May 1994, Hyland. 15119 (QRS); About 10 km north of the Palmer River Roadhouse, 12 Feb 1997, Hyland 15534 (BRI, QRS); Mt White, Coen, 20 Jan 2000, Hyland 16345 (BRI, QRS); Maitland Downs Holding, 29 Sep 1988, Hyland 25551 RFK, (QRS), 13 Dec 1988, Hyland 25640 RFK (BRI, QRS); Cook, Cooktown Development Road opposite Bonnie Glen Station turnoff, 8 Jun 1996, Jago 4014 (BRI); Unnamed basalt-capped hill on Springvale Station, adjacent to Plum Tree Creek, 13 Mar 2017, Kerrigan 1343 (CNS); North Kennedy, Upper slopes of high ridge, ca. 3.5km SW of Earlando, Dryander National Park, 2 Jun 1994, McDonald & Champion 5862 (BRI); 3.4 km N of Spear Creek on the Peninsula Development Road. 11.3 km N of the Palmer River crossing, 4 Mar 1987, McDonald 6674 (AD); 12 km from the east Normanby River crossing on the Lakeland Downs to Cooktown road, 6 Mar 1987, McDonald 6768 (AD); Rocky knob just S of the divide on the Palmer River road, 23 Nov 1972, Nicholson AFO-4776 (QRS); Port Curtis, Mt Archer, Rockhampton, 12 Mar 2003, Nicholson NJN 455 (BRI); Rockhampton, 1 Feb 1869, O’Shanesy s.n. (MEL); Cook, King’s Plain Stn, Mt Emily, 9 Jan 2014, Roberts KRM-15160 (BRI); Mt Pinnacle, 4 Mar 1987, Sankowsky 615 (QRS); Port Curtis, over hanging path to caves, Olsen’s Capricorn Caves, Olsen’s caves Rd, The Caves, 12 Feb 2014, Shapcott & Howard MGH-40 (BRI); Colosseum Cave, Mt Etna Caves NP, N of Rockhampton, 1 Apr 1993, Thomas 8951 (BRI); 18 km NNW of Gladstone, 2 Mar 1997, Thompson & Turpin GLA-22 (BRI); Middle Percy Island, 5 Mar 1906, Tryon s.n. (BRI); Olsens’ Caves, 1 Feb 1989, Vavryn s.n. (BRI); Cammoo Caves, 1 Jan 1987, Vavryn 17 (BRI); Cook, Cooktown Developmental Road-Sackleys Hill, 6 Jan 2002, Wannan & Jago 2328 (BRI, NSW); Seisia Village, Northern Peninsula Area, 17 Jan 1998, Waterhouse 4806 (BRI, DNA), 12 Mar 1999, Waterhouse 5114 (CANB), Waterhouse 5114 (BRI, DNA); tributary of Mossman River, 25 km S of Laura, 8 Mar 2017, Worboys 1344 (CNS).

Based on Solanum vitiense Seem.

Trees to 15 m tall, with single trunks to 30 cm diameter at breast height; stems terete, glabrous or minutely puberulent (especially near the nodes) with weak-walled, 10–15-celled simple uniseriate trichomes to 0.25 mm long, these drying golden reddish brown, soon glabrescent; new growth densely puberulent-papillate with weak-walled 10–15 celled simple uniseriate trichomes like those of the stems near the nodes; bark of older stems pale grey or whitish grey. Sympodial units difoliate, the leaves geminate, the leaves of a pair, differing in size but not in shape. Leaves simple; blades of major leaves 8–28 cm long, (2.5)4–12.5 cm wide, elliptic to broadly elliptic, widest at the middle, somewhat discolorous, membranous to thick and chartaceous (rubbery in live plants?); adaxial surfaces shiny, completely glabrous; abaxial surfaces glabrous; principal veins 7–10 pairs, prominent beneath, drying reddish brown; base acute; margins entire; apex acute to acuminate; petioles 2–4.5 cm long (or perhaps even longer in larger leaves), glabrous; blades of minor leaves 3.8–11.5 cm long, 2.5–6.5 cm wide, shape, texture and pubescence like than of the major leaves; base acute; margins entire; apex acute to acuminate, sometimes rounded; petioles 0.7–1.5 cm long, glabrous. Inflorescences axillary 4–10 flowered fascicles with several flowers usually open at once, sometimes developing a woody axis or axes to 0.6 cm long, pubescent with floccose reddish brown trichomes like those of the new growth; pedicels at anthesis 1.5–2 cm long, 0.5–0.75 mm in diameter at the base, 1.5–2 mm in diameter at the apex, spreading, glabrous, articulated at the base; pedicel scars tightly packed in the leaf axils or along the short inflorescence axes. Buds ellipsoid, the corolla exserted ca. halfway from the calyx tube before anthesis. Flowers 5-merous, heterostylous and unisexual, specimens have only short-styled flowers or long-styled flowers and fruits, the plants probably dioecious (described as androdioecious, with staminate and hermaphroditic flowers, by Reinecke 1898, as Brachistus feddei). Calyx tube 3–3.5 mm long, 3–5 mm wide, openly cup-shaped, chartaceous or somewhat fleshy, often purple or flushed with purple, sparsely puberulent with floccose trichomes like those of the stems, without appendages, the rim hyaline and irregular (“ruffly”). Corolla 1–1.8 cm in diameter, white, sometimes fragrant, stellate, lobed ca. 3/4 of the way to the base, interpetalar tissue present, the lobes 5–8 mm long, 2.5–3 mm wide, spreading to somewhat reflexed, adaxially glabrous, abaxially glabrous or densely papillate, the tips and margins always densely papillate. Stamens equal; filament tube minute; free portion of the filaments 1–1.5 mm long, glabrous; anthers 2.5–4 mm long, 1–1.5 mm wide, ellipsoid and slightly tapering at the tips, white, cream-colored or yellow, poricidal at the tips, the pores tear-drop shaped and markedly splitting to slits with age, some plants apparently with longitudinally dehiscent anthers and no pores (see figure 5 in Smith 1991). Ovary conical, glabrous, vestigial in short-styled flowers; style in short-styled flowers to 1.5 mm long, in long-styled flowers 5.5–6 mm long, straight, glabrous; stigma clavate, occasionally somewhat bilobed, the surfaces minutely papillate. Fruit a globose berry, 0.5–1.5 cm in diameter, green to orange when immature and bright red when ripe (description as “atroviridi” by Reinecke 1898 possibly refers to dry material; blue fide Kajewski 1863), the pericarp glabrous, shiny, translucent; fruiting pedicels 2–2.5 cm long, ca. 1.5 mm in diameter at the base, 2–2.5 mm in diameter at the apex, spreading, glabrous, woody, corky and rugose; fruiting calyx a cup with a spreading thinner rim beneath the fruit, the rim thinner than the woody calyx cup, the rim sometimes reflexed. Seeds (10)30–50 per berry, 4–5 mm long, 3.5–4 mm wide, flattened and rounded with a prominent notch, yellowish tan, the surfaces deeply pitted at the thickened margins with pentagonal to rectangular testal cells, the centre of the seed smooth or only shallowly pitted with testal cells pentagonal in outline. Stone cells absent. Chromosome number not known.

(Fig. 53).Lycianthes vitiensis is a mostly Pacific species, known from New Guinea (Bougainville Island), the Solomon Islands, Samoa, Fiji and Tonga. It is very common in Fiji (Smith 1991).

Lycianthes vitiensis is a plant of many forest types, and has been collected both in primary and secondary rainforests and along streams; from 30 to 1,200 m elevation.

There are no standard common names (Smith 1991), but several different names have been recorded across its range. Fiji: malawatha (Smith 5828), nandata (Degener 14832), nggalinggwa (Smith 5270), teleniva (Smith 5250), tungasele (Smith 1244), vualiku (Smith 5250). New Guinea (Bougainville): em-wuye (Kajewski 1863), kosiou (Waterhouse 88), neketigo (Kajewski 1800), nua (Waterhouse 272B). Samoa: koriele (Guppy 160), uagagi (Christopherson 2697), u’agani (Bristol 12146)]. No uses have been recorded.

(IUCN 2020).EOO (1,758,710 km² - LC); AOO (288 km² - EN). Lycianthes vitiensis is known from more than 10 localities on many islands in the western Pacific and is often described as a “common tree” on herbarium labels. This suggests it should be assessed as Least Concern (LC), despite its island occurrence.

Lycianthes vitiensis is a relatively common slender tree; both Seemann (1863) and Reinecke (1898) recognised it as having long- and short-styled flowers. Bitter (1919) did not include this species in his treatment of Lycianthes, perhaps because in German herbaria it was often cited as a species of Brachistus Miers (see below) due to longitudinal anther dehiscence. Symon (1985) suggested that L. vitiensis (as S. vitiense) showed intermediate characters between Lycianthes (as subgenus Lycianthes) and Solanum, citing the short inflorescence axis and calyx that sometimes appears five lobed.

Lycianthes vitiensis is not easily confused with any other species of Lycianthes treated here. Like L. shanesii it is a small tree; it differs from that species in its many-flowered inflorescence with a short axis (versus few-flowered strictly axillary inflorescences in L. shanesii) and in the dense, tangled reddish brown pubescence of 10–15-celled trichomes on young stems and inflorescences (versus sparse golden pubescence of 2–3-celled trichomes of L. shanesii). Lycianthes vitiensis is not sympatric with any other Lycianthes in Asia, Australia or the Pacific. It is most similar to L. banahaensis (Elmer) Bitter of the Philippines (not treated here) with which it shares floccose reddish brown pubescence of stems and new growth, but differs from that species in its white or violet (versus yellow) flowers and red (versus orange) mature berries.

Lycianthes vitiensis is unusual in Lycianthes in its anther dehiscence by both apical pores and more commonly by longitudinal slits. Many species of Solanum with ellipsoid anthers have tear-drop shaped apical pores that split longitudinally with age and drying (see Särkinen et al. 2018; Knapp et al. 2019). The longitudinally dehiscent anthers of L. vitiensis, in contrast, do not appear to be poricidal at the outset (see figure 5 in Smith 1991: 24). This difference does not correlate with the sex expression of individual flowers, and its occurrence merits further study.

Flowers of Lycianthes vitiensis are sometimes described as fragrant (e.g., Mauriasi et al. BISP-13952), other times as with no smell (e.g., Mauriasi et al. BISP-8491); this may be due to time of day or degree of anthesis,

In the protologue of Solanum vitiense, Seemann (1863) only cited his collection number “340” but no herbarium. I have selected the specimen at Kew (K000759477) as the lectotype, because it is well-preserved and has a clear label with both the number and the locality. Other specimens at Kew cited as types by Symon (1985) are not clearly duplicates of this number. Smith (1991) cited two sheets at Kew as “holotype”, but this does not constitute effective lectotypification.

In the protologue of Brachistus feddei, Reinecke (1898) cited three of his own Samoan collections: Reinecke 58 (male), Reinecke 58a (male) and Reinecke 78 (hermaphrodite). Reinecke’s own collections are housed in WRSL. Of the collections located there Reinecke 78 is the best preserved and has bisexual flowers, therefore the duplicate of this collection at WRSL is here selected as the lectotype. The duplicate of Reinecke 78 held at BISH has both the locality and date of collection matching the protologue and is and isolectotype (BISH1005071). A note on the specimen from A.C. Smith dated June 1988 indicates the holotype is at B; this was a common mistaken assumption for many names coined by German botanists. Nowhere in Reinecke’s protologue does he indicate material in Berlin; his personal herbarium is held in WRSL (now in Poland). Duplicates of Reinecke’s collections held at K (Reinecke 58, K000759467; Reinecke 58a K000759466) cited by Symon as types, have different dates than those in the protologue (September in the protologue and October on the Kew specimens) and are thus I do not strictly consider them isolectotypes.

Fiji. ‘Fiejie Islands’, US Exploring Expedition under the command of Capt Wilkes s.n. (NY). Lau Archipelago: Kabara, ‘Kambara’, limestone formation, 2 Mar 1934, Smith 1244 (K, NY, P, US). Ngau: hills E of Herald Bay, inland from Sawaieke, 300–450 m, 15 Jun 1953, Smith 7797 (K, NY, US); Ovalau: sin. loc., 15 Oct 1924, Bryan 608 (US); near summit of main range W of Levuka, 200 m, 26 Jan 1928, Gillespie 4446 (K, US); hills above Levuka reservoir, 400 m, 30 Jan 1928, Gillespie 4517 (NY); hills east of Lovoni Valley, 300–500 m, 11 May 1953, Smith 7343 (K, NY, P, US); slopes of Mt. Koronimiko, vicinity of Thawathi, 250–350 m, 13 Jul 1953, Smith 8082 (K, NY, P, US); Vanua Balavu: ‘Vanua Mbalavu’, northern limestone section, 2 Apr 1934, Smith 1508 (K, NY, P, US); Vanua Levu: Mt. Delaikoro, Macuata, 762–914 m, 21 Aug 1962, Koroiveibau 12807 (K); Cakaudrove, Navonu, Nov 1964, Qoro USDA-14091 (K, NY); southern slope of Korotini Range, below Navitho Pass, Thakaundrove, 300–650 m, 21 Nov 1933, Smith 574 (K, NY, P, US); Yanawai River region, Mount Kasi, Thakaundrove, 300–430 m, 10 May 1934, Smith 1826 (K, NY, US); Viti Levu: Serua District, 152 m, 14 Jun 1961, Bola 41 (K); vicinity of Nandarivatu, TholoNorth, 700–900 m, 16 Mar 1941, Degener 14832 (K, NY, US); Vuninatiambua, Navai, Tholo North, 750–900 m, 4 Feb 1941, Degener 14875 (K, MO, NY, US); Mbulu, near Sovi Bay, Tholo West, 20 Apr 1941, Degener 15032 (K, MO, NY, P, US); Mt. Victoria [Mount Tomanivi], 29 Dec 1968, Degener et al. 32083 (E, NY); S of Naboutini, Serua, 274 m, 17 Dec 1963, Forestry Department 965 (K); Nadarivatu, 823 m, Aug 1907, Gibbs 568 (BM, K); Tamavua woods 8 miles above Suva, 150 m, 6 Aug 1928, Gillespie 2016 (K, NY, P, US); above waterfall near Namuamua, Namosi, 400 m, 2 Oct 1927, Gillespie 3251 (K, NY, US); slopes of Mount Victoria [Mount Tomanivi], 1,000 m, 29 Nov 1927, Gillespie 4082 (NY); Albuggi Levu, Graeffe, E. s.n. (K); Mount Evans [= Mount Koroyanitu], Lautoka, 914 m, 3 Oct 1920, Greenwood 127 (K); Lautoka, Mt. Evans [=Mount Koroyanitu], 750 m, 16 Sep 1945, Greenwood 1151 (K, US); banks of Samaruna [=Samambula?] River, Horne 714 (K); Nadroga-Navosa, Nausori Highlands, 579 m, 20 Jul 1964, Kuruveli 13888 (K); Naboutini, Serua, 9 Sep 1964, Kuruveli & Qoro 14007 (K); Tailevu, Dakuivuna, 13 Nov 1957, Ledua 11018 (K); 9 miles above Suva, Jul 1932, Meebold 17036 (K); prov. Ba, Nadarivatu, 1,067–1,219 m, 24 Oct 1962, Parham & Koroveibau 13051 (K); Naitsiri, Princes [Princess] Road, 21 Apr 1936, Raigiso 3141 (K); sin. loc., Seemann 42 (BM); [Tailevu], 1860, Seemann 387 (K, P); western and southern slopes of Mount Tomonivi (Mt. Victoria) [Mount Tomanivi], Mba (formerly Tholo North), 850–1,150 m, 7 Jul 1947, Smith 5250 (K, NY, P, US); western and southern slopes of Mount Tomonivi (Mt. Victoria), Mba (formerly Tholo North), 850–1,150 m, 7 Jul 1947, Smith 5270 (K, NY, P, US); northern portion of Rairaimatuku Plateau, between Nandrau and Nanga, Nandronga and Navosa (formerly Tholo North), 725–825 m, 4 Aug 1947, Smith 5501 (K, NY, P, US); hills between Nggaliwana and Nandala Creeks, south of Nauwanga, Mba (formerly Tholo North), 725–850 m, 26 Aug 1947, Smith 5828 (K, NY, P, US); hills bordering Wainavindrau Creek, in vicinity of Wainimakutu, Namosi, 150–250 m, 17 Sep 1953, Smith 8894 (K, NY, P, US); hills W of Wainikoroiluva River, near Namuamua, Namosi, 50–200 m, 15 Oct 1953, Smith 8930 (K, NY, P, US); hills N of Ngaloa, in darinage of Waininggere Creek, Serua, 30–150 m, 19 Nov 1953, Smith 9171 (K, NY, P, US); Serua, hills west of Waivunu Creek, between Ngaloa and Korovou, 50–150 m, 23 Nov 1953, Smith 9264 (K, NY, US); Nambavatu [Creek], Tothill, B.H. 556 (K), Tothill 633 (K); path Kalumbo (Suva), 5 Aug 1926, Tothill 636 (K); Nandarivatu [= Nadarivatu], 914 m, 1927, Tothill 638 (K), Feb 1927, Tothill 639 (K).

Papua New Guinea. Bougainville (North Solomons): Bougainville Island, 22 miles NW of Tonlei Harbour, Buin subdistrict, 91 m, 25 Aug 1969, Foreman NGF-45698 (A, K, L, LAE); Kugi-maru, Buin., 150 m, 2 Jun 1930, Kajewski 1800 (BM, G, P); Kugumaru, Buin., 150 m, 12 Jun 1930, Kajewski 1863 (BM, G, P); vicinity of Aku [Aka] village, ca. 10 miles W of Buin Station, 30 m, 21 Sep 1964, Schodde & Craven 4094 (A, G, K, L, LAE); “Melanesia, Siwai, Bougainville”, Sep 1932, Waterhouse 66 (K, NY, US), Sep 1930, Waterhouse 272-B, (K);

Samoa. sin. loc., Horne s.n. (K), Powell 365 (K), Whitmee 52 (K), Aug 1875, Whitmee 185 [a] (K). Savai’i: far inland from Aopo, base line 4 of Division of Forestry 1968 forestry survey, 300 m, 28 Jun 1968, Bristol 2146 (K, LAE, NY, US); above Salailua, 650 m, 22 May 1924, Bryan 173 (K, US); Olo, [Mount Olomanu], 700 m, 4 Aug 1931, Christophersen & Hume 2249 (K, NY, P, US), 9 Aug 1931, Christophersen & Hume 2320 (K, NY, P, US), 9 Aug 1931, Christophersen & Hume 2324 [a] (P, US), 9 Aug 1931, Christophersen & Hume 2354 (NY), 26 Aug 1931, Christophersen & Hume 2518 (NY, P), 26 Aug 1931, Christophersen & Hume 2521 (K), 26 Aug 1931, Christophersen & Hume 2528 (US); above Salailua, 900 m, 22 Sep 1931, Christophersen 2697 (K); Lo To, above Salailua, 750 m, 21 Oct 1931, Christophersen 2895 (K, NY, P);); above Salailua, ca. 1,350 m, 7 Nov 1931, Christophersen 3116 (P), 1,200 m, 5 Nov 1931, Christophersen 3123 (NY); ueber Aopo [A’opo], Sep 1894, Reinecke, F. 58 a (K, WRSL); Olomono, uber Olomono, 600 m, 7 Sep 1905, Vaupel s.n. (MO); Forestry Block 25 inland from Asau, 29 Jun 1972, Whistler 4 (US); above Ologogo, 700 m, 21 Aug 1973, Whistler 520 (US); above Asau in Block 28, 550 m, 16 Oct 1973, Whistler 891 (US). Upolu: Malololelei, near Malololelei, 550 m, 17 Aug 1929, Christophersen 326 (NY, P); ‘Ins. Upolu’, Graeffe s.n. (BM, NY), Mar 1880, Graeffe 1354 (K); sin. loc., 1896, Hufnagel s.n. (US); Tanumalala, 200 m, 11 Aug 1955, MacKee 2995 (K); Upalu [Upolu sin. loc.], von Mueller 13 (BM); Sameaberg [=Samea], Oct 1893, Reinecke 58 (K, WRSL); Olonono [illegible-Bugrange?] [Cape Olonono], 7 Nov 1905, Vaupel 104 (K, US); E of the main road near Tiavi, 700 m, 3 Oct 1973, Whistler 1071 (K, US); just N of Lake Lanoanea, 600 m, 5 Nov 1973, Whistler 1090 (US); near Mt. Le Pu’e, 750 m, 7 Dec 1973, Whistler 1253 (US).

Solomon Islands. sin. loc., Kajewski D s.n. (BM, P). Central Province: Florida Island, Popinisura [=Nggela Islands, Nggela Sule], Comins 235 (K); Rove Area, Big Nggela [=Nggela Sule], 91 m, 24 Jul 1969, Gafui & collectors BSIP-16787 (K). Choiseul Province: Choiseul Island, East Mbirambira, West Choiseul, valley bottom, 15 m, 17 Jan 1970, Gafui & collectors BISP-18862 (K). Guadalcanal Province: Berande River., 31 Dec 1930, Kajewski 2388 (A, BM, G, P); NW Guadalcanal, Mt. Austen, W side of the trial plot, 304 m, 27 Nov 1964, Kere BSIP-4936 (K), 23 Dec 1964, Kere BSIP-5057 (K); NW Guadalcanal, Mt. Mambulu summit [=Mount Austen], 457 m, 19 Jul 1967, Nakisi BSIP-8013 (K); Guadalcanal Island, Guadalcanal, west fork Tenau River, 30 Sep 1945, Riley 45 (A, NY, US); Guadalcanal, Mt Austen near Honiara, 304 m, 13 Mar 1963, Whitmore BSIP-776 (K); N Guadalcanal, Kukum, 0.5 mile inland for training college, 7 Feb 1964, Whitmore BSIP-2567 (K). Isabel Province: Samusodu, SW Santa Ysabel, 85 m, 2 May 1966, Beer’s collectors BSIP-7225 (K); Korigole Bay, SW Santa Ysabel, 48 m, 13 Jun 1966, Beer’s collectors BSIP-7306 (K); Ysabel Island, Kolokofa R, NW Santa Isabel, 250 m, 5 Apr 1966, Teona BISP-6362 (K, L, LAE, US); Santa Ysabel, near Maringe Lagoon, Molau Village, 670 m, 27 Oct 1963, Whitmore BSIP-2437 (K). Malaita Province: N of Anihonora’a Village, E Malaita, 5 Sep 1969, Gafui & collectors BSIP-16450 (K). Western Province: Shortland Island, May 1884, Guppy 160 (K); SE Kolombangara, W of Vila River, 30 m, 20 Dec 1967, Mauriasi & collectors BSIP-8491 (K); Fauro Island, Haliuna River area, 114 m, 16 Apr 1969, Mauriasi & collectors BISP-13952 (A, K, L, LAE); NW Treasury Island [=Mono Island], Akolea River area, 60 m, 26 Apr 1969, Mauriasi & collectors BSIP-14089 (K); Palusua, SE Mono, 22 m, 11 May 1969, Mauriasi & collectors BSIP-14154 (K); New Georgia Group, Baga Island, 26 Jan 1964, Whitmore’s collectors BSIP-3081 (K).

Tonga. Vava’u: above Ha’alaufuli on northeastern side of island, 60 m, 21 May 1953, Yuncker 16103 (NY, US).

Based on Solanum wollastonii Wernham.

Slender woody climber or epiphyte, to ca. 3 m tall (long); stems terete, glabrous, flushed with purple (fide Mustaqim et al. 2022); new growth minutely papillate, soon glabrescent; bark of older stems pale tan, peeling. Sympodial units difoliate, the leaves geminate, the leaves of a pair differing in size and sometimes in shape. Leaves simple; blade of major leaves 6–11 cm long, 2–4 cm side, elliptic, somewhat oblique, widest at the middle, discolorous, membranous; adaxial surfaces shiny, glabrous, the venation impressed, the midrib keeled; abaxial surfaces paler, glabrous, the veins drying reddish gold; principal veins 4–6 pairs, impressed above, reddish golden below, glabrous; base acute to cuneate; margins entire, somewhat revolute; apex abruptly acuminate with an elongate drip-tip 1–2 cm long; petioles 0.3–0.6 cm long, glabrous; blades of minor leaves 1.9–2.7 cm long, 1.5–2 cm wide, elliptic to narrowly elliptic, texture and pubescence like that of the major leaves; base acute; margins entire; apex acute or rounded; petioles ca. 0.2 cm long, glabrous. Inflorescences axillary 1–3(5) flowered fascicles, with only a single flower open at a time, glabrous; pedicels 1.8–2.2 cm long, ca. 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading and nodding from weight of flowers, green flushed with purple (fide Mustaqim et al. 2022), glabrous, articulated at the base; pedicel scars tightly packed in the leaf axils. Buds narrowly ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers 5-merous, apparently perfect (only long-styled flowers seen). Calyx tube 3–3.5 mm long, 4–4.5 mm in diameter, openly cup-shaped, pale purple, glabrous, with 5 triangular appendages 1–2 mm long emerging 0.75–1 mm below the rim and perpendicular to the calyx tube, the rim entire. Corolla 1.6–2 cm in diameter, white, deeply stellate, lobed nearly to the base, interpetalar tissue present, the lobes ca. 10 mm long, ca. 2.5 mm wide, spreading and slightly cupped, glabrous, minutely papillate at the slightly cucullate tips. Stamens equal; filament tube minute; free portion of the filaments ca. 1 mm long, glabrous; anthers 4.5–5 mm long, ellipsoid, yellow, poricidal at the tips, the pores tear-drop shaped, lengthening to slits with age. Ovary conical, glabrous; style ca. 7.5 mm long, straight, glabrous; stigma bilobed to bifid with the lobes ca. 0.2 mm long, the surfaces minutely papillate. Fruit a globose berry (described from Mustaqim et al. 2022), ca. 0.7 cm in diameter, green (immature?), the pericarp glabrous, matte, opaque; fruiting pedicels to 3 cm long, pendent, bright purple (dull violet) especially towards the enlarged apex, glabrous; fruiting calyx a cup subtending the fruit, tightly adhering to the basal part of the berry, the appendages fleshy and somewhat enlarged, slightly backwards pointing. Seeds and stone cells not seen. Chromosome number not known.

(Fig. 56).Lycianthes wollastonii is endemic to the island of New Guinea; it is known only from Puncak Jaya (Mount Carstenz) in Indonesia (Papua).

Lycianthes wollastonii is a plant of mossy montane forests, growing between 1,500 and 1,800 m elevation.

None recorded.

(IUCN 2020).EOO (0 km² - CR); AOO (8 km² - CR). Lycianthes wollastonii is known from a single locality. Mustaqim et al. (2022) assessed L. wollastonii as Endangered (EN B2ab(iii)) due to its small range in an area threatened by logging and mining. They did not assign a more threatened status due to the number of populations assumed to be two (their collection and the type), however, although the type locality is somewhat imprecise, it can be pinpointed to a small area using the description of the Wollaston expedition itinerary (Kloss 1916) and is thus distinct from the collection locality cited by Mustaqim et al. (2022). I consider the species should be assigned a threat status of Critically endangered (CR) due to its narrow distribution and the threats for human alteration of the environment. This species merits further collecting and population survey.

Until recently (Mustaqim et al. 2022) Lycianthes wollastonii was known only from Cecil Boden Kloss’s type collection, made on the expedition led by Arthur Wollaston to “Snow Mountains” (Kloss 1916). Their collection, made in 2018 on the ascent of Mount Jaya, is the first fruiting record for this species and confirms both its distinctness from other New Guinea Lycianthes and its rarity and restricted distribution. I have described the fruits and associated characters from Mustaqim et al. (2022); colour of mature fruits is still not known. Wernham (1916) described L. wollastonii (as S. wollastonii) as an epiphyte, but Mustaqim et al. (2022) described it as a slender shrub; Kloss could have mistaken the habit due to the dense covering of moss on the stems, or L. wollastonii may indeed sometimes grow on the branches of other plants in its wet cloud forest habitat as do other Solanaceae (e.g., S. gonyrhachis S.Knapp of Bolivia, see Knapp 2002).

Lycianthes wollastonii is somewhat like L. lucens in its glabrous leaves, triangular calyx appendages held perpendicular to the calyx tube (see figure 1 in Mustaqim et al. 2022) and large flowers. It differs from L. lucens in its homostylous (to be confirmed) flowers (heterostylous in L. lucens), anthers 4–4.5 mm long (versus 2–2.5 mm long in L. lucens) and minor leaves differing significantly in shape from the major leaves (similar in shape, but not in size in L. lucens).

The large flowers with narrow corolla lobes of Lycianthes wollastonii are somewhat similar to those of L. rostellata. The two species can be distinguished by stem pubescence (absent in L. wollastonii, of stiff, antrorse trichomes with multicellular bases in L. rostellata) and calyx appendage morphology (absent in L. rostellata, triangular in L. wollastonii).

Indonesia. Papua: Timika Regency, Mount Jaya, Tembagabura, Borobudur, 2,090 m, 21 Nov 2018, Mustaqim & Manurang 2213 (BO).