Solanum opacum A.Braun & C.D. Bouche , Ind. Sem. Hort. Berol.: 18. 1853

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

9. Solanum opacum A.Braun & C.D. Bouche, Ind. Sem. Hort. Berol.: 18. 1853 Figures 28 , 29

Solanum forsteri Seem., J. Bot. 1: 207. 1863.

Solanum opacum Type. Chile. Región V ( Valparaíso): Easter Island, J.R. Forster & G. Forster s.n. (lectotype, designated here: BM [BM000900372]).

Solanum fauriei H. Lév., Repert. Spec. Nov. Regni Veg. 10: 152. 1911.

Solanum opacum Type. United States of America. Hawaii: Oahu, Kaela, Nov 1909, U. Faurie 861 (lectotype, designated here: BM [BM000846671]; isolectotypes: P [P00315100, P03961997]).

Solanum apopsilomenum Bitter, Repert. Spec. Nov. Regni Veg. 12: 89. 1913.

Solanum opacum Type. New Zealand. Mount [illegible, perhaps Zwart], 28 Sep 1838, E. Schwarz s.n. [Exped. Novara] (holotype: W [0022367]).

Solanum microtatanthum Bitter, Bot. Jahrb. Syst. 55: 63. 1919.

Solanum opacum Type. Papua New Guinea. Kelil, 180 m, F.R.R. Schlechter 16407 (holotype: B, destroyed; lectotype, designated here: E [E00273859]; isolectotypes: G [G00343298], L [L0003621], GH [GH00077834], K [K001080534, K001080535]).

Solanum brachypetalum Bitter, Bot. Jahrb. Syst. 55: 64. 1919.

Solanum opacum Type. Papua New Guinea. "Ssigaun, in Dorfern", 600 m, Jun 1896, C. Lauterbach 2360 (holotype: B, destroyed; lectotype, designated here: WRSA).

Solanum insulae-paschalis Bitter, Nat. Hist. Juan Fernandez & Easter Island 2: 78. 1922.

Solanum opacum Type. Chile. Región V ( Valparaíso): Easter Island, Hanga ho ono (La Perouse), au linem actem Brunnen, C. Skottsberg & I. Skottsberg 663 (lectotype, designated here: GOET [GOET003528]; isolectotype: UPS [UPS104030]).

Solanum nigrum L. var. nihoense F.Br., Bernice P. Bishop Mus. Bull. 81: 36, pl. 16B. 1931.

Solanum opacum Type. United States of America. Hawaii: Leeward Island, Nihoa, E.L. Caum 62 (holotype: BISH; isotypes: K [K000922201], NY [00172289]).

Solanum nigrum L. var. pitcairnense F.Br., Bernice P. Bishop Mus. Bull. 130: 255. 1935.

Solanum opacum Type. Pitcairn Islands: Pitcairn island, 13 Mar 1922, E.H. Quayle 375a (holotype: BISH [BISH1005084]; isotype: BISH [BISH1005083]).

Solanum allanii Polgár, Trans.& Proc. Roy. Soc. N. Z. 69: 278. 1939.

Solanum opacum Type. Cultivated in Györ Hungary from seeds of Miss L.B. Moore #8 (New Zealand, Auckland, Mt. Wellington), Jan 1938, S. Polgár s.n. (lectotype, designated here: BP [acc. # 146410]; isolectotype: BP [acc. # 261051]).

Type.

Cultivated in Germany at the Berlin Botanical Garden from seeds from Australia "New Holland (Listeman) hort. bot. Berlin", Anon. s.n. (lectotype, designated here: HBG [HBG511471]) .

Description.

Annual or short-lived sprawling to erect perennial herbs to 1 m tall, subwoody and branching at base. Stems spreading to decumbent, terete or sometimes slightly ridged, green to yellow-green, older stems greenish-grey, not or occasionally somewhat hollow; new growth densely to sparsely pubescent with simple, antrorse, uniseriate, translucent, eglandular or sometimes glandular trichomes, these 4-6-celled, 0.5-1 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5-8.0(-17) cm long, 1-4 -(9) cm wide, elliptic to slightly ovate, very variable in size, membranous, green, concolorous, without smell; adaxial surface glabrous or sparsely and evenly pubescent with simple uniseriate ca. 4-celled trichomes to 0.5 mm long; abaxial surfaces glabrous or sparsely pubescent with simple uniseriate trichomes along the veins; major veins 3-8 pairs, not prominent; base cuneate, decurrent on the petiole; margins entire or shallowly toothed, if present the teeth acute; apex acute to acuminate; petioles 0.5-3 cm long, sparsely pubescent with antrorse simple uniseriate trichomes like those of the stems. Inflorescences 1-2 cm long, internodal, unbranched but very rarely furcate, umbelliform to sub-umbelliform, with 3-7 flowers clustered near the tip of the rhachis, sparsely pubescent with antrorse simple uniseriate 3-4-celled trichomes like those of the stems; peduncle 1-3 cm long, straight and stout; pedicels 0.4-0.8 cm long, <0.3 mm in diameter at the base, ca. 0.3 mm in diameter at the apex, filiform, nodding, pubescent like the peduncle, articulated at the base; pedicel scars clustered near the tip of the rhachis, often the lowest flower ca. 0.5 mm spaced from the rest. Buds ellipsoid, the corolla strongly exserted from the calyx tube long before anthesis. Flowers 5-merous, all perfect. Calyx tube 1-2 mm long, conical, the lobes often unequal, the lateral two largest 1-1.2 mm long, 0.5-0.6 mm wide, the top and lowermost 0.4-1.0 mm long, 0.3-0.5 mm wide, long triangular often with a rounded tip, glabrous or sparsely pubescent with antrorse simple uniseriate trichomes ca. 0.5 mm long. Corolla 6-10 mm in diameter, white or white with a purplish tinge, stellate, lobed ca. 1/2 way to the base, the lobes 3.0-4.2 mm long, 1.0-1.2 mm wide, spreading to reflexed, densely papillate on tips and margins. Stamens equal; filament tube <0.1 mm long; free portion of the filaments 0.5-1.0 (-1.5) mm long, glabrous or adaxially pubescent with tangled simple uniseriate trichomes; anthers 1.2-1.6 mm long, 0.7-1.0 mm wide, ellipsoid, yellow, somewhat sagittate at the base, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 3-4 mm long, strongly curved in the distal 1/4, densely pubescent with simple uniseriate trichomes 0.2-0.5 mm long, these tangled in the basal 1/2 to 2/3 of the length, not exserted beyond anthers and only the stigma visible outside the anther cone; stigma capitate, the surfaces minutely papillate. Fruit a globose berry, 4-10 mm in diameter, green or bluish-black at maturity, the pericarp thin, matte; fruiting pedicels 0.7-1.5 cm long, ca. 0.5 mm in diameter at the base and the apex, erect or spreading, becoming yellow, falling with the mature fruits, not persistent; fruiting calyx lobes not accrescent, the tube less than 1 mm long, the lobes 1.2-1.7 mm long, appressed to the basal quarter of the berry, occasionally somewhat spreading, never strongly reflexed. Seeds 50-100 per berry, 1.2-2.2 mm long, 0.7-1.8 mm wide, flattened reniform, pale yellowish-tan, the surfaces minutely pitted, thin and the embryo clearly visible, the testal cells rectangular to pentagonal in outline. Stone cells (0-)2(-4) per berry,>0.5 mm in diameter, brown. Chromosome number: 2n =6x=72 ( Henderson 1974; Symon 1985).

Distribution

(Figure 30 ). Native in the Pacific islands, from Australia, Papua New Guinea and New Zealand, to Taiwan and Hawaii, west to the Pitcairn Islands and Rapa Nui (Easter Island).

Ecology.

Grows in disturbed areas, along roadsides and field edges; between sea level and 2,600 m elevation in a wide variety of habitats.

Common names.

Australia: currant, green-berry nightshade ( Symon 1981; Harden 1993), dark nightshade ( Walsh and Entwisle 1999), tamatha, tamada; Fiji: mau, mboro laukana, murimuri; Indonesia: kenkene; Malaysia: tutan; Papua New Guinea: bek, kepa, kepilam, ongkara, tagur/tagura, tuskombuk, wietat; Pitcairn Islands: o brew.

Uses.

Leaves eaten as greens throughout the species range.

Preliminary conservation status

( IUCN 2016). Solanum opacum is a relatively widespread species across the Pacific; it can be assigned a preliminary status of LC (Least Concern; Table 7 ) on a global scale, but local country assessments may differ. The species has a very scattered distribution in the Pacific and, although its overall range is very large, the land area it occupies is small.

Discussion.

Solanum opacum is extremely similar to S. americanum ; both species have minute flowers with anthers usually less than 1.5 mm long. They co-occur across the Pacific, but can be distinguished easily, particularly in fruit. Solanum opacum has matte berries that are green or greyish-purple at maturity, with appressed or slightly spreading calyx lobes, while S. americanum has very shiny berries that are black or dark purple at maturity and calyx lobes that are very strongly reflexed. The pedicels of S. americanum remain on the plant as fruits fall, while those of S. opacum drop with the mature berries. In flower, the relative length of filament to anther is a useful distinguishing character; S. opacum has filaments that are as long as or slightly longer than the anthers, while the filaments of S. americanum are always shorter than the anthers.

Our circumscription of S. opacum encompasses a much wider distribution than that of Henderson (1974), who treated only the Australian populations. The treatments of "Solanum nigrum" or "Solanum americanum" in the Pacific could apply to either S. opacum or S. americanum . Degener (1945) records the use of morelloid species in Hawaii for both food and medicine; both species occur there.

Solanum opacum is hexaploid and, based on network analysis of arbitrary amplified DNA markers, Poczai and Hyvönen (2011) suggested that Australian populations of S. retroflexum were a possible tetraploid parent of S. opacum . The putative diploid parent is S. chenopodioides , S. nitidibaccatum or some other closely related species ( Poczai and Hyvönen 2011). Morphologically, S. retroflexum somewhat resembles S. opacum , but the shared molecular markers could also come from a common progenitor of the two polyploid species.

Henderson (1974) neotypifed S. opacum with material collected by Robert Brown on the voyage of the Discovery (Bird Sound, 19 Sep 1802, R. Brown s.n. (neotype, designated by Henderson 1974, pg. 39: NSW [NSW125341]; isoneotypes: BM [BM000900380], K, MEL)) because he could find no original material corresponding to the collector or locality cited in the protologue ("E Nova Hollandia. Semina communicavit Listemann") and presumed it to have been in Berlin and subsequently destroyed. However, recent digitisation of historical material in European herbaria has uncovered original material not seen by Henderson (http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.hbg511471) that was annotated as “lectotype” in 1974 by J.M. Edmonds, but never published as such. The fact that original material has been found means that Henderson’s (1974) neotypification must be set aside ( McNeill et al. 2012, Art. 9.19) and we designate the sheet in HBG (HBG511471) as the lectotype of S. opacum . It matches the wild-collected plants selected by Henderson exactly.

Solanum forsteri was named in honour of the Forster’s (father Johann and son Georg, who accompanied Captain Cook on the HMS Resolution from 1772 to 1775) and we have selected the sheet collected by them on Easter Island (BM000900372) as the lectotype. This specimen is mounted together with a collection of S. americanum that is on different paper (white rather than the blue associated with Forster’s material), showing the difficulty that early botanists (and this holds true still today) had in distinguishing these two species with minute anthers that co-occur across the Pacific.

In describing S. microtatanthum , Bitter (1919) cited a specimen in the Berlin herbarium, now destroyed; we have selected the most complete of the duplicates we have seen as the lectotype (E00273859). The protologue of S. fauriei cited no herbaria and we have selected the best preserved of the duplicates we have seen as the lectotype (BM000846671). We have selected the sheet with better material for both fruits and flowers (BP No. 146410) as the lectotype of S. allanii .

Selected specimens examined.

Australia. New South Wales: Botany Bay, 28 Apr 1770, Banks & Solander s.n. (BM); Sans Souci, nr Georges River, 27 Sep 1961, Goode 346 (K, NSW); Kyogle , Toonumbar State Forest 26 km NW of Kyogle, 22 Feb 1972, Henderson 1261 (AD, AD, NSW); Upper Hunter Shire , Ben Halls Gap State Forest , 20 Feb 1991, Hosking 307 (NE); Wollongong, Bulli, 1875, Johnson s.n. (MEL); Wollongong, Mt Keira , 27 Jul 1983, Mills s.n. (WOLL) ; Norfolk Island: sin. loc, Oct 1805, Caley s.n. (BM); Kingston, 24 Aug 1964, Uhe 1126 (K); Northern Territory: Adelaide River, Northern Territory, grown in glasshouse at Indooroopilly, Brisbane, 18 Dec 1972, Henderson 1356 (CANB, K); Queensland: Nr border of Queensland/New South Wales, Main Range National Park, 10 Feb 2006, Bohs et al. 3561 (BM, UT); Birnam Range 7.5 km NE of Beaudesert, Tremayne rd, Jan 2001, Halford Q-3882 (BRI, K, MEL, NSW); c. 13 km WNW of Goondiwindi on rd to St. George , 30 Sep 1975, Henderson 2357 (AD, BRI, K); South Australia: Upper River Murray nr 375 mile peg 4 mi downstream from Little Hunchee Island, on North bank, 16 Sep 1979, Symon 11586 (AD, CANB, K); Kangaroo Island , Kangaroo Island , Waterhouse s.n. (MEL); Tasmania: Tasman, Port Arthur , 1893, Bufton 11 (MEL); sin. loc, 1842, Gunn 51[a] (BM); Latrobe, Harford on Solomans Hill , 6 Nov 1932, Hamilton 163 (CANB, HO); Van Diemen’s Land, Gunn, 1835, Without collector s.n. (K); King Island , Nr Sea Elephant River , 23 Mar 2009, Wapstra et al. 691 (AD); Victoria: Mt Buffalo access rd 12 km WNW of Bright, 1 Jan 2003, Lepschi 4916 (CANB, MEL); Latrobe, 18 Jul 1988, Thompson 157 (AD, AD, MEL); Cedar Creek , Bentleigh district , 12 Feb 1920, White s.n. (NSW); Western Australia: c. 200 mi N of Chesapeake rd 1.5 km E of Mt Chuladup, 7 Dec 2001, Barker 8355 (AD); Eyre, S of Cocklebeddy, 5 Dec 1962, MacDonald 29 (K); Bayswater, Lower Swan River , 20 Mar 1909, Morrison 19014 (K) .

Chile. Región V ( Valparaíso): Easter Island, von Chamisso s.n. (F, NY); Juan Fernández Islands, Masafuera , Quebrada de Veradero , 12 Mar 1917, Skottsberg & Skottsberg 568 (GOET) .

Cook Islands. Mitiaro: just before entering swamp on rd to Atai Foodland , 22 Jul 1991, Luttrell 187 (FHO); rd from the village of Takaue to the marsh, 24 Apr 1985, Whistler 5576 (US) .

Fiji. Yuen Yick’s farm, Nasinu, Naitasiri, 3 Dec 1957, Ledua s.n. (K); sin. loc., 1860, Seemann 344 (BM, K); Fulanga: Limestone Formation, 22 Feb 1934, Smith 1174 (K); Kandavu: Mount Mbuke Levu, 23 Oct 1933, Smith 206 (K); Viti Levu: Mba, summit of Mt. Nanggaranambuluta E of Nandarivatu, 19 Jun 1947, Smith 4849 (K, US); Hills E of Wainikoroiluva River, nr Namuamua, Namosi, 15 Oct 1953, Smith 9070 (K, US).

French Polynesia. Austral Islands: Raivavae, Vaiuru, 10 Aug 1934, Fosberg 11756 (US); Rurutu, du N de l’ile, 17 Apr 1981, Hallé 7032 (US); Rurutu, N Moerai, 25 Apr 1981, Hallé 7321 (US); Raivavae, Ahuoivi, 9 Aug 1934, St. John 16070 (US); Gambier Islands: Mangareva, 1833, Le Guillou s.n. (US); Aukena, Koiovao, 29 May 1934, St. John 14659 (US); Society Islands: Tahiti, 6 Apr 1840, Barclay 3309 (BM)Tahiti, plateau de Taravao, sentier du captage de l’Hamoa, 16 Sep 1982, Florence 3853 (US); Raiatea, first valley S of Uturoa, 14 Oct 1926, Moore 212 (US); Me’et’ia, Fatia-po to Fareura, 12 May 1934, St. John 14187 (US).

Indonesia. Bali: Kintamani , 14 Nov 1966, Schwabe s.n. (B) ; East Nusa Tenggara: Flores, nr Mataloko, Verheijen 24 (L); Flores, nr Mataloko, Verheijen 191 (L) ; Papua: Mimika Regency, PT-Freeport Indonesia Concession Area , 9 Aug 1998, Johns et al. 9565 (A, K); Mimika Regency, W and above Tembagapura, 23 Aug 1998, Sands 7296 (K) ; Sulawesi: Central Sulawesi, Sopu valley , 30 May 1975, Balgooy 3555 (A, K) ; West Papua: West-Irian, Eipomek-Tal, bei Malingdam , 24 Feb 1976, Hiepko & Schutze-Motel 1195 (B) .

Japan. Nagasaki Pref., Katamatsuura-gun, Emukae-cho, Okugawachi-men , 17 Aug 1994, Yonekura 3230 (MO) .

Malaysia. Sabah: Kampung Tiung, side of hill Tiung Valley, Tuaran Distr. , 8 Aug 1985, Maikin Lantoh SAN-108976 (K)

Marquesas Islands. Hatutaa Island : vallon de la pointe SW, 10 Jul 1988, Florence & Teikiteetini 9399 (US)

Marshall Islands. Arno Atoll: Ine village , 12 May 1950, Anderson 3672 (L, US) ; Jaluit Atoll: Jabor , 28 Apr 1958, Fosberg 39475 (US) ; Kwajalein Atoll: Kwajalein Islet , 19 Jan 1950, Fosberg 31180 (US)

New Caledonia. sin. loc, Caldwell s.n. (K); NO de la Nouvelle Caledonie, 1867, Krieger s.n. (W); Nord: Ilôt Poudiou, observatory Island, Isle of Pines, Oct 1853, MacGillivray 805 (K); Sud: Nouméa, Îlot Freycinet, Ilot de Freycinet, Aug 1884, Grunow s.n. (W); Noumea, Baie Tina, 24 Jul 1973, MacKee 26964 (K).

New Zealand. Chatham Islands: Rekohu, ( Chatham Island ), Waitangi, Ellice Point , Chatham Islands Ecological Region and District , 1 Dec 2008, de Lange & Horne CH-2032 (AK) . Kermadec Islands: Macauley Island , Southern Kermadec Islands Group, 29 Jun 2006, Barkla M- 26 (AK); Curtis Island, Nov 1900, Shakespear s.n. (AK). North Island : Northland, dunes near Te Arai Forest Sanctuary, Aupouri State Forest , N of Waimahru Stream, Mangonui County, 25 Sep 1985, Bellingham 0144 (AK) ; Taranaki, Sugarloaf Island, off Mangonui, Moturoa Group , off Karikari Peninsula , 14 May 1976, Wright 1252 (AK) . Outlying Islands: Three Kings Islands, Great Island, above cliff S.E. Bay, 19 Apr 1946, Turbott & Bell s.n. (AK). South Island : Motuara, Banks & Solander s.n. (BM) . Tokelau Territory : Fakaofu , Union Islands, Mar 1891, Lister s.n. (K) .

Niue. Niue Island, "71 Polokai", 1901, Smith s.n. (AK).

Norfolk Island. Rocky Point Reserve , 6 Oct 1989, Gardner 5866 (AK) .

Papua New Guinea. Boridi, 6 Sep 1935, Carr 12985 (BM, K); Wiligimaan, Baliem, Div. Hollandia, 28 Jun 1961, Versteegh BW-12516 (A, L); Bougainville Island: Namatea, NW Bougainville, 7 Mar 1932, Waterhouse 691 B (K, L); Crown Prince Mountain , Oct 1960, Womersley NGF-13346 (A); Central: E side lake Myola No. 2, Distr. Central, Subdistr. Port Moresby, 13 Sep 1973, Croft NGF-34532 (E, K, L); Kokoda trail, eastern side lake Myola , 22 Jul 1974, Croft LAE-61910 (A, K, QRS); Madang: Saidor, Moro, Naho-Rawa Division, Finisterre mountains , 19 Nov 1964, Perumal 21473 (BM); Morobe: Mount Kaindi , Wau, 4 Oct 1977, Conn & Kairo 491 (CANB, K); Bulldog rd, nr Edie Creek , Wau, 13 Aug 1963, Millar & Holttum NGF-15825[b] (A, US); Southern Highlands : Hagen-Mendi rd, Mendi Subdistrict, Southern Highlands District , 21 Sep 1968, Vandenberg et al. NGF-40062 (K); Western Highlands : nr Ampyak Highlands, Ecological Site 1, 15 Dec 1964, Flenley ANU-2192 (K); nr Tomba village , S slope of Mount Hagen Range, 27 Aug 1956, Hoogland & Pullen 6014 (BM, US); Western Province: Western, 19 Jun 1979, Sohmer LAE-75542 (K); Kubor Range, Uinba, Nona-Minj Divide, 20 Aug 1963, Vink 16311 (K) .

Philippines. Luzon: Cordillera (CAR), Mount Pulag , 26 Jan 1968, Jacobs 7173 (K) .

Pitcairn Islands. Oeno Island : Close to hut, 23 Jun 1934, St. John & Fosberg 15195 (US) ; Pitcairn Island: Bounty Bay , baie de la Bounty, Cap Est, 22 Apr 1991, Florence 10774 (US); Bounty Bay, 15 Jun 1934, Fosberg & Clark 11337 (US) .

Samoa. Apia: Apia , 6 May 1907, Vaupel Sol-3 (K) ; Savai’i: E of Olo , 8 Aug 1931, Christerpherson & Hume 2311 (K) .

Solomon Islands. Guadalcanal: north central Guadalcanal, Tina River , 14 Sep 1967, Maurisi BSIP8109 (K) .

Taiwan. Pingtung County: Wutain Hsiang, Tawu village , 13 Mar 1999, Shu-hui Wu 1161 (MO) ; Taoyuan County: Kuanyin Hsiang, Hsinpo , 28 Dec 1999, Ching-I Peng, 17879 (MO) .

Tonga. Tofua: sin. loc, Jan 1967, Scarth-Johnson s.n. (K) ; Tongatapu: Kologa , Jun 1926, Setchell & Parks 15374 (US) .

United States of America. Hawaii: Leeward Island, Nihoa, 17 Jun 1923, Caum 62 (US); Oahu, Kaela, Nov 1909, Faurie 861 (BM, P); Kauai, Kaholuamanoa, above Waimea, 1 Oct 1895, Heller 2867 (AK, BM, E, US); Pearl and Hermes Atoll, Southeast Island , midway between pond and tower, 18 Aug 1964, Young 112 (US) .

Figures

Figure 28. Solanum opacum. A Habit. B Detail flower (A-B Symon s. n. [ADW 40795]). Drawing by M. L. Szent-Ivany, first published in Symon (1981), courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Figure 29. Solanum opacum A Habit B Shoot with flowers and maturing fruits C Detail of maturing fruits with appressed calyx lobes (A Nijmegen acc. 904570211, B Butaud 3075, C Nijmegen acc. 884570223). Photos by J. - F. Butaud and G. van der Weerden.

Figure 30. Distribution of Solanum opacum.