Solanum nitidibaccatum Bitter, Repert. Spec. Nov. Regni Veg. 11: 208. 1912

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

8. Solanum nitidibaccatum Bitter, Repert. Spec. Nov. Regni Veg. 11: 208. 1912 Figures 25 , 26

Solanum styleanum Dunal, Prodr. [A. P. de Candolle] 13(1): 44. 1852.

Solanum nitidibaccatum Type. Chile. Sin. loc., J. Styles s.n. (holotype: G-DC [G00144016]).

Bosleria nevadensis A.Nelson, Proc. Biol. Soc. Washington 18(30): 175. 1905.

Solanum nitidibaccatum Type. United States of America. Nevada: Washoe County, Pyramid Lake, 9 Jun 1903, G.H. True s.n. (holotype: RM [RM0004387]).

Solanum nitidibaccatum Bitter forma integrifolium Blom, Acta Horti Gothob. 24: 118. 1961.

Solanum nitidibaccatum Type. Sweden. Göteborg, som ogräs i Göteborgs Botaniska Trädgård, 10 Sep 1948, C. Blom s.n. (lectotype, designated here: GB [GB-0146965]; isolectotype: E [E00593447]).

Solanum patagonicum C.V.Morton, Revis. Argentine Sp. Solanum 146. 1976.

Solanum nitidibaccatum Type. Chile. Región XII (Magallanes): Río Paine, 100m, 15 Jan 1931, A. Donat 415 (holotype: BM [BM000617673]; isotypes: BA, BAF, GH [GH00077732], K, SI [SI003331, SI003332], US [00027733, acc. # 2639758]).

Solanum physalifolium Rusby var. nitidibaccatum (Bitter) Edmonds, Bot. J. Linn. Soc. 92: 27. 1986.

Solanum nitidibaccatum Type. Based on Solanum nitidibaccatum Bitter

Type.

Chile. Sin. loc., 1829, E.F. Poeppig s.n. (lectotype, designated by Edmonds 1986, pg. 27: W [W0004151]; isolectotype: F [V0073346F, acc. # 875221]) .

Description.

Annual prostrate or spreading herbs to 20 cm tall, branching at base. Stems decumbent or ascending, terete, green, not markedly hollow; new growth densely viscid-pubescent with simple, spreading, uniseriate, translucent, glandular trichomes, these 2-8(10)-celled, 1.5-2.0 mm long, with a glandular apical cell; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 2.0-5.5(-9.5) cm long, 1.5-5.0(-6.5) cm wide, ovate to broadly ovate, rarely elliptic, membranous, green, concolorous, without smell; adaxial surface sparsely pubescent with spreading 2-4-celled translucent, simple, uniseriate gland-tipped trichomes like those of the stem, these denser along the veins; abaxial surface more evenly densely pubescent on the lamina and veins; major veins 3-6 pairs, not clearly evident abaxially; base attenuate to cuneate, at times asymmetric, decurrent on the petiole; margins entire or sinuate-dentate; apex acute to obtuse; petioles 0.5-2.7(-4.5) cm long, sparsely pubescent with simple uniseriate glandular trichomes like those of the stems and leaves. Inflorescences 1.0-2.0 cm long, generally internodal but occasionally leaf-opposed, simple, the flowers spaced along the rhachis, with 4-8(-10) flowers, sparsely pubescent with spreading trichomes like those on stems and leaves; peduncle 0.6-1.3 cm long, straight; pedicels 4-12 mm long, 0.1-0.2 mm in diameter at the base and 0.2-0.4 mm at apex, straight and spreading, articulated at the base; pedicel scars spaced 0.3-1 mm apart. Buds subglobose, the corolla only slightly exserted from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 1-2 mm long, conical, the lobes 1.7-2.5 mm long, less than 1 mm wide, triangular with acute to obtuse apices, sparsely pubescent with 1-4-celled glandular trichomes like those of the pedicels. Corolla 4-6 mm in diameter, white with a yellow-green central eye with black “V” or “U” shaped margins in the lobe sinuses, rotate-stellate, lobed 1/3 of the way to the base, the lobes 2.3-3.2 mm long, 2.5-3.7 mm wide, spreading at anthesis, sparsely papillate-pubescent abaxially with 1-4-celled simple uniseriate trichomes, especially along tips and midvein. Stamens equal; filament tube minute; free portion of the filaments 1.5-2.0 mm long, adaxially sparsely pubescent with tangled uniseriate 4-6-celled simple trichomes; anthers 1.0-1.4 mm long, 0.5-0.8 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5-3.0 mm long, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted 0.2-1.0 mm beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 4-13 mm in diameter, brownish-green and marbled with white (this not easily visible in herbarium specimens) at maturity, the surface of the pericarp usually shiny; fruiting pedicels 4-13 mm long, ca. 0.2 mm in diameter at the base, spaced 1-3 mm apart, reflexed and slightly curving, dropping with mature fruits, not persistent; fruiting calyx accrescent, becoming papery in mature fruit, the tube ca. 3 mm long, the lobes 2.5-3.5(-4) mm long, 3-4 mm wide, appressed against the berry, but the berry clearly visible. Seeds 13-24 per berry, 2.0-2.2 mm long, 1.2-1.4 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells usually (1-)2-3 per berry, occasionally absent, ca. 0.5 mm in diameter. Chromosome number: 2 n =2 x =24 ( Ellison 1936; Henderson 1974; Saarisalo-Taubert 1967; Edmonds 1977, 1981, 1982, 1983, 1984a, 1986; Randell and Symon 1976; Symon 1981; Olet et al. 2015).

Distribution

(Figure 27 ). Native to southern South America but sporadically to widely adventive in Europe, North America, Australia and New Zealand.

Ecology.

Grows along roadsides, in disturbed and cultivated areas in the shade of trees and shrubs, in rocky and sandy areas; between sea level and 2,000 (-2,700) m elevation in its native range, between sea level and 2,400 m in the Old World primarily as a weed in cultivations.

Common names.

Australia: cherry nightshade ( Walsh and Entwisle 1999); Austria: Glanzbeeren-Nachtschatten ( Fischer et al. 2005); Denmark: storbægret natskygge ( Hartvig 2015); Finland: kehtokoiso ( Hämet-Ahti et al. 1998); France: morelle à feuilles de coqueret ( Jauzein and Nawrot 2011); New Zealand: hairy nightshade ( Webb et al. 1988); Sweden: bägarnattskatta ( Mossberg et al. 2003); United Kingdom: Argentinian nightshade, green nightshade ( Stace 2010).

Uses.

None recorded; a weed of agriculture and actively controlled.

Preliminary conservation status

( IUCN 2016). Solanum nitidibaccatum is a relatively weedy species that is invasive where introduced; in its native range it is widespread and can be assigned a preliminary status of LC (Least Concern; Table 7 ). The EOO is relatively large even if considering only the native American range (1,591,444 km2) and the assessment status does not change.

Discussion.

Solanum nitidibaccatum is morphologically similar to S. sarrachoides and has been treated under that taxon in many previous treatments (e.g. Schilling 1981, Schilling and Heiser 1979). Edmonds (1986) clarified the distinction between the two taxa, together with a discussion of the problems surrounding its correct identification and its complex synonymy and lectotypification. Solanum nitidibaccatum was treated as a subspecies of S. physalifolium ( Edmonds 1986). Solanum physalifolium is not known from the Old World and is endemic to South-Central Andes of Argentina, Bolivia and Peru to elevations between 2,000 and 2,900 m; distinguishing features of that species can be found in the key in Barboza et al. (2013).

Solanum nitidibaccatum is a diploid species native to the south-eastern parts of South America and within the Old World is morphologically most similar to S. sarrachoides , with which it has been confused. The two species are best distinguished by the complete inclusion of buds in calyx before anthesis, the larger stone cells in berries and the more erect habit of S. sarrachoides (see key in Barboza et al. 2013). Solanum nitidibaccatum has shorter and more ovoid anthers compared to S. sarrachoides , where anthers are more elongate-ellipsoid and the fruiting calyx of S. nitidibaccatum does not cover the berry as much as does that of S. sarrachoides.

Solanum nitidibaccatum has been introduced extensively to other parts of the world where it has become a prolific and successful weed of disturbed sites usually associated with agriculture of the wool trade. Trade with South America - particularly the importation of grain, seeds and the spreading of wool ‘shoddy’ - has been largely responsible for its introduction into Europe, with one of its common European names being the Argentinian nightshade. The taxon is now a widespread adventive in Europe where it is rapidly becoming naturalised and often forms extensive populations. It has been introduced into Australia on a number of occasions, where it persists as a weed of cultivation and is sparingly established in all States ( Henderson 1974; Symon 1981). The species is locally abundant throughout North America ( Ogg et al. 1981) where it is particularly widespread in the Pacific States and the West ( Bohs in press). It was introduced into the South Island of New Zealand around 1968 where it rapidly became established and is now also found in northern parts of North Island ( Healy 1974). It also has been sparingly introduced into equatorial regions of Africa (see Appendix 2).

In parts of eastern England, S. x procurrens A.C.Leslie ( Leslie 1978), a sterile tetraploid hybrid between S. nitidibaccatum and S. nigrum , is locally established where the two species grow together ( Stace 2010: 531; Stace et al. 2015). The hybrid has also been recorded in New Zealand ( Webb et al. 1988). These plants are intermediate between the two species, with glandular pubescence and black berries with somewhat accrescent calyx lobes. A good description and distribution map for this local hybrid can be found in Stace et al. (2015). We have not seen convincing material from elsewhere in Europe of this hybrid, but it potentially occurs wherever S. nitidibaccatum and S. nigrum co-occur. As it is sterile, however, it does not spread or persist.

The amateur Swedish botanist Carl Blom rarely cited herbaria for his names; we have selected the sheet in GB, where he worked, as the lectotype of his var. Solanum nitidibaccatum integrifolium .

Selected specimens examined.

Australia. Australian Capital Territory: CSIRO Black Mountain site, Acton, Canberra , 9 Jan 1995, Lepschi 1729 (AD, CANB, HO,NSW) ; New South Wales: Canyonleigh via Marulan , May 1976, Cooper s.n. (NSW); White Rock, 8 km upstream Bathurst, 9 May 1982, Dellon s.n. (NSW) ; Queensland: The Summit c. 5 miles NNE of Stanthorpe, 8 Feb 1972, Henderson & Parham 1241 (AD, BRI, MEL, NSW) ; South Australia: Adelaide Hills , 20 Mar 1986, Jackson 5976 (AD, CANB); Tea Tree Gully, 12 Mar 1974, Spooner 3341 (AD); Mitcham, 18 Jan 1980, Winn s.n. (AD) ; Tasmania: Murphy’s Flat Reserve, 2010 ( Bush Blitz ), 25 Mar 2010, Baker 2203 (HO); 12 Jan 2001, Buchanan 15827 (HO) ; Victoria: Australian Alps , 19 Jan 1981, Beauglehole 68690 (MEL); Melbourne; Ripponlea Estate, Hotham Street, Elsternwick, 6 Mar 1984, Clarke 1682 (NSW); Milne’s property, 27 Jan 1966, Shepherd 215 (CANB); Riverina , Jan 1975, Without Collector s.n. (MEL) ; Western Australia: Badgerup Road opposite Jambaris Road, Wanneroo , 15 May 1997, Burt s.n. (PERTH); Pinjarra , 17 Nov 1987, Sheldow s.n. (PERTH) .

Austria. Burgenland: Nordburgenland, WSW von Eisenstadt , S von Müllendorf, 2 Aug 2007, Barta s.n. (BM); Seewinkel, zw. Apetion u Frauernkirchen, knapp W v E-Teil d. Fuchslochlacke, W v Feldweg, 12 Oct 2008, Walter 7085 (W) ; Nieder-Österreich: Marchfeld, nahe Gänserndorf, Ackerrand neben der Strasse 1.5-1.7 km W der Kirche von Weikendorf, 28 Sep 2010, Barta s.n. (BM); Wiener Becken, Steinfeld, 7.5 km NNE Wiener Neustadt, 1.7 km NW Eggendorf, knapp WNW, 19 Sep 1998, Walter 4189 (K, W) ; Wien: 10 Bezirk, in Kurpark Oberlau, 22 Oct 2012, Barta 1219 (W); 22 Bezirk, knapp S der Ostbahngleitstrasse nahe der U2-Haltestelle Aspern-Nord, 6 Aug 2014, Barta 3270 (W) .

Belgium. Wallonia: Vesdre , Sep 1958, Lousley s.n. (K) .

France. Grand Est: Bas-Rhin, Strasbourg, Graffenstaden , Oct 1961, Patzak s.n. (W); Nouvelle Aquitaine: Gironde, Bordeaux, Bassens , 24 Jul 1927, Duffour 5538 (BM, MA); Gironde, Bassens, 9 Sep 1931, Jallu 1038 (H, P) .

Germany. Hessen: Frankfurt am Main, Frankfurt-am-Main, Hessen-Nassau , Sep 1911, Peipers s.n. (BM) ; Niedersachsen: Leer, Ostfriesl, Blamgelände, 10 Oct 1955, Klimmek s.n. (W) .

Ireland. Leinster: Kilkenny, Rosbercon Port , 5 Oct 1995, Reynolds s.n. (BM) ,

Netherlands. Gelderland: Nijmegen, prov. Gelderland , 23 Sep 1928, Kern & Reichgelt 14998 (BM, H) .

New Zealand. North Island: Waikato, Pukekawa, 18 Dec 1986, Dawes s.n. (AK); Auckland, Arikikapakapa Golf Course, Rotorua Ecological District, Northern Volcanic Plateau Ecological Region , 15 Mar 2013, Hobbs 13297 (AK) . South Island: Canterbury, Christchurch, waste land behind Fendalton Mall buildings, Fendalton , 2 Feb 1999, Healy 99/ 34 (AK, CHR) .

Sweden. Prov. Halland, 26 Sep 1923, Jungner s.n. (BM); Haisenborg, vid fabrikenb Kärnan, 6 Oct 1938, Lange s.n. (BM); Skåne, Saxtorp, Flygeltofta, 28 Aug 1958, Nilsson s.n. (BM); Götaland: Halland, Halmstad, Gustavsfält, 22 Sep 1955, Blom s.n. (BM); Halland, pr. urben Halmiam, 26 Apr 1923, Jungner s.n. (BM); Halland, prope usbem Halmiam, 26 Sep 1923, Jungner s.n. (K); Halland, Halmstad, 26 Sep 1923, Jungner 1375 (K); Västra Götaland, Göteborg, Ringon, 29 Sep 1951, Blom s.n. (K) .

United Kingdom. Channel Isles: Jersey , N of St. Ouen’s Bay, Sep 1996, Dupree s.n. (BM) ; England: Essex, vice county 18, Dagenham dumps, 1938, Airy Shaw s.n. (K); Bedfordshire , Flitton , 14 Oct 1950, Doug 1401 (K); Bedfordshire, nr Flitwick, 16 Sep 1988, Hanson 499 (BM); Hampshire, Blackmoor, 24 Oct 1959, Lousley s.n. (BM); Berkshire, Reading, 25 Nov 1999, Rutherford s.n. (BM); Gloucestershire, Wapping Wharf, Bristol Harbour, 14 Sep 1942, Sandwith s.n. (BM) ; Scotland: Highlands, Auldearn, Broomhill Farm , 27 Oct 1967, McCallum-Webster s.n. (BM); Midlothian, Railway Tip, Borthwick, 20 Jul 1963, McCallum-Webster s.n. (BM); Midlothian, Borthwicts, 20 Jul 1963, McCallum-Webster 8768 (K) ; Wales: Vale of Glamorgan, Barry Dock , 29 Sep 1923, Melville s.n. (BM) .

Figures

Figure 25. Solanum nitidibaccatum A Habit B Fruiting habit C Fruiting habit showing leaf variation D Detail of adaxial leaf surface E Detail of abaxial leaf surface F Bud G Dissected flower H Fruit (A, C, F Henning 14; B, D-E, H Blake 186; C Arnow 740). Drawing by R. Wise.

Figure 26. Solanum nitidibaccatum A Habit B Young inflorescence with flower buds C Flowers at anthesis D Maturing fruits (A-D Saerkinen et al. 4076). Photos by T. Saerkinen.

Figure 27. Distribution of Solanum nitidibaccatum in its non-native range in the Old World.