PlantaeSolanalesSolanaceaeKnappSandraBarbozaGloria E.BohsLynnSärkinenTiinaA revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the CaribbeanPhytoKeys3052019123114410.3897/phytokeys.123.31738 Solanum chenopodioides Lam ., Tabl. Encycl. 2: 18. 1794Figures 6, 7 Solanum sublobatum Willd. ex Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 664. 1819. Type. Argentina. Buenos Aires, Anon. s.n. [probably P. Commerson] (Herb. Willdenow 4336) (lectotype, designated by Edmonds 1972, pg. 105 [as type ex photo]: B [B-W04336-01-0]). Solanum besseri Weinm., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 593. 1819. Type. “In America” [cultivated in Europe?], Anon. s.n. (no specimens cited; no original material located; neotype, designated by Särkinen et al. 2018, pg. 65: G-DC [G00144198]). Solanum subspatulatum Sendtn., Fl. Bras. (Martius) 10: 45, tab. 4, fig. 16–18. 1846. Type. Brazil. Sin. loc., F. Sellow s.n. (holotype: B, destroyed, F neg. 3183; lectotype, designated by D’Arcy 1974a, pg. 735 [as type]: P [P00384051]; isolectotype: F [v0361921F, acc. # 621700, fragment]). Witheringia chenopodioides (Lam.) J.Rémy, Fl. Chil. [Gay] 5: 69. 1849. Type. Based on Solanumchenopodioides Lam. Solanum isabellei Dunal, Prodr. [A. P. de Candolle] 13(1): 153. 1852. Type. Uruguay. Montevideo, Lat. aust. 34°45'08", 1838, A. Isabelle s.n. (lectotype, designated by Särkinen et al. 2018, pg. 65: G-DC (G00145645); isolectotypes: F [v0073298F, acc. # 680251; v0073299F, acc. # 680253], K [K000585686], P [P00384071], W [acc. # 1889-115034]). Solanum chenopodiifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 44. 1852. Type. Argentina/Uruguay. “Buenos Aires et Montevideo”, P. Commerson s.n. (lectotype, designated Edmonds 1972, pg. 108 [as holotype], second step designated by Särkinen et al. 2018, pg. 65: P [P00384081]). Solanum crenatodentatum Dunal var. ramosissimum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852. Type. United States of America. Louisiana: “Basse Louisiane”, 1839, G.D. Barbe 82 (holotype: P [P00362535]). Solanum gracile Dunal, Prodr. [A.P. de Candolle] 13(1): 54. 1852, nom. illeg., not Solanumgracile Sendtn. (1846). Type. Brazil. Rio de Janeiro: “Rio de Janeiro”, 1831–1833, C. Gaudichaud 520 (lectotype, designated by Henderson 1974, pg. 46: G-DC [G00144391]; isolectotypes: G [G00343457], P [P00384052, P00384053]). Solanum gracile Dunal var. microphyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852. Type. Argentina/Uruguay. “Circa Buenos Ayres et Montevideo”, P. Commerson s.n. (lectotype, designated by Morton 1976, pg. 151: P [P00384061, Morton neg. 8207]; possible isolectotype: F [v0073283F, acc. # 976485, fragment only]). Solanum nodiflorum Jacq. var. microphyllum Hassl., Repert. Spec. Nov. Regni Veg. 9: 118. 1911. Type. Paraguay. Estrella: Mar, É. Hassler 10271 (holotype: G?, Morton neg. 8612). Solanum vile Bitter, Repert. Spec. Nov. Regni Veg. 11: 221. 1912. Type. Brazil. Rio de Janeiro: Restinga do Harpoador, E. Ule 4310 (lectotype, designated by Särkinen et al. 2018, pg. 66: CORD [CORD00004277]; isolectotype: HBG [HBG-511507]). Solanum gracilius Herter, Rev. Sudamer. Bot. 7: 266. 1943. Type. Based on (replacement name for) S.gracile Dunal Solanum ottonis Hyl., Uppsala Univ. Årsskr. 7: 279. 1945. Type. Based on (replacement name for) Solanumgracile Dunal Type.<sup><xref ref-type="fn" rid="en2">2</xref></sup>

Mauritius. “Ex ins. Mauritiana”, Herb. Lamarck s.n. (lectotype, designated by Barboza et al. 2013, pg. 242: P [P00357629]).

Description.

Annual herbs to short-lived perennial shrubs up to 1.0 m tall, subwoody and branching at base. Stems terete, green-grey to straw colour, sprawling, somewhat weak and decumbent, not markedly hollow; new growth pubescent with simple, uniseriate appressed 1–6-celled eglandular trichomes, these 0.1–0.6 mm long; older stems more sparsely pubescent, glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5–5.5(–7.0) cm long, 0.5–3.0(–3.5) cm wide, lanceolate to narrowly ovate, rarely ovate, discolorous; adaxial surface green, sparsely pubescent with appressed 1–4-celled translucent, simple, uniseriate trichomes like those on stem, these denser along the veins; abaxial surface pale grey, more densely pubescent with trichomes like those of the upper surface evenly distributed across lamina and veins; major veins 3–6 pairs, not clearly evident abaxially; base attenuate, decurrent on the petiole; margins entire or sinuate; apex acute to obtuse; petioles (0.5-)1.0–1.5(–3.5) cm long, sparsely pubescent with simple uniseriate trichomes like those of the stems and leaves. Inflorescences 1.0–2.5(–4.0) cm long, lateral, generally internodal but appearing leaf-opposed on young shoots, unbranched or rarely forked, with 3–7(–10) flowers clustered near the tips (sub-umbelliform), sparsely pubescent with appressed 1–2-celled simple uniseriate trichomes; peduncle 1.0–2.3(–4.0) cm long, strongly deflexed downwards in fruit; pedicels 5–10 mm long, ca. 0.5 mm in diameter at the base and 1 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0–1 mm apart. Buds elongate-oblong, the corolla only slightly exserted from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 2–3 mm long, conical, the lobes 0.6–1.2 mm long, less than 1 mm wide, broadly deltate to triangular with acute to obtuse apices, sparsely pubescent with 1–4-celled appressed hairs like those on stem but shorter. Corolla 6–12 mm in diameter, white with a black and yellow-green central portion near the base, the black colour usually distal to the yellow green, deeply stellate, lobed 4/5 of the way to the base, the lobes 3.5–4.0 mm long, 1.5–1.9 mm wide, strongly reflexed at anthesis, later spreading, densely puberulent-papillate abaxially with 1–4-celled simple uniseriate trichomes, these denser on the tips and margins. Stamens equal; filament tube minute; free portion of the filaments 0.6–1.0 mm long, adaxially pubescent with simple tangled uniseriate 4–6-celled simple trichomes; anthers (2.0–)2.3–2.8 mm long, 0.5–0.8 mm wide, narrowly ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying, the connective becoming darker brown with age in dry plants. Ovary globose, glabrous; style 3.7–4.5 mm long, densely pubescent with 2–3-celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted up to 1.5 mm beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 4–9 mm in diameter, dull purplish-black at maturity, opaque, the surface of the pericarp matte and somewhat glaucous; fruiting pedicels 6–13 mm long, 1.2–1.4 mm in diameter at the base, reflexed and slightly curving, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes 1–1.5 mm long, appressed against the berry. Seeds (13–)20–35(–50) per berry, 1.2–1.4 mm long, 1.0–1.2 mm wide, flattened and tear-drop shaped with a subapical hilum, pale yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent. Chromosome number: 2n=2×=24 (see Särkinen et al. 2018).

Solanumchenopodioides Lam. A Habit B detail of adaxial leaf surface C detail of abaxial leaf surface D opening bud E dissected flower F fruiting branch G detail of infructescence H maturing fruit I fully mature fruit (A–EFox s.n.F–IHieronymus s.n.). Drawing by R. Wise (previously published in “PhytoKeys 106”).

Solanumchenopodioides Lam. A Habit B buds C flowers at full anthesis D fully mature matte black fruits with appressed calyx lobes (A–D Nijmegen accession A14750051). Photos by S. Knapp and G. van der Weerden (previously published in “PhytoKeys 106”).

Distribution.

(Figure 8) Solanumchenopodioides is native to southern South America, and has been introduced globally, largely with the wool trade. The species is relatively uncommon in North America, where it is most likely introduced.

Distribution of Solanumchenopodioides Lam.

Ecology.

Solanumchenopodioides is an adventive species in North America and occurs only in sporadic populations close to urban areas and human disturbance between 0 and 2,000 m elevation.

Common names.

None recorded.

Uses.

None recorded.

Preliminary conservation status (<xref ref-type="bibr" rid="B115">IUCN 2017</xref>).

LC (Least Concern). Solanumchenopodioides is a widespread weed of disturbed areas (see Barboza et al. 2013; Särkinen et al. 2018). For EOO see Table 6.

Discussion.

Solanumchenopodioides is a weedy, ruderal species occurring mainly in coastal parts of North America. The species has distinct grey-green appearance due to the pubescence of appressed, eglandular white trichomes. It is morphologically similar to S.pseudogracile and some populations of S.americanum around the coast of the Gulf of Mexico. Solanumchenopodioides can be distinguished from S.pseudogracile only with difficulty, but the short-triangular calyx lobes with acute apices that remain appressed to the berry at fruit maturity, as opposed to the longer, rectangular calyx lobes with rounded to acute apices that are reflexed in fruit of S.pseudogracile, are characters that distinguish the taxa. In flower, the extension of style beyond the anther cone is a good character to separate S.chenopodioides from S.pseudogracile; the style remains almost completely inside the anther cone in S.chenopodioides (exserted to 1–1.5 mm) and is clearly exserted in S.pseudogracile (exserted to (1)2.0–2.5 mm). Many specimens annotated as S.chenopodioides from around the Gulf of Mexico (e.g., Florida) are actually plants of S.pseudogracile.

Solanumchenopodioides can be distinguished from S.nigrescens by the lack of stone cells in fruit, while S.nigrescens has always 4–13 stone cells per fruit. Anthers in S.chenopodioides are always much longer (2.0–2.8 mm) than in S.americanum (0.8–1.5 mm). The strongly deflexed peduncle and pedicels in fruit are distinctive in S.chenopodioides but are not always obvious in herbarium specimens.

Typification details for the synonyms of S.chenopodioides can be found in Särkinen et al. (2018).

Specimens examined.

See Suppl. materials 1 and 3.

Preliminary conservation assessments for morelloid species from the Caribbean and North and Central America. For details see Materials and Methods and individual species treatments. Preliminary assessments are based on EOO only (see Materials and Methods) and have been calculated for worldwide ranges for each species. The EOO and conservation status of species known to be solely cultivated, introduced or adventive in the region has been assessed in Särkinen et al. (2018).

SpeciesPreliminary conservation assessment (IUCN 2017)EOO (km2) [worldwide range]
Solanumamericanum Mill. LC 444,094,992
Solanumchenopodioides Lam. LC 77,207,558
Solanumcorymbosum Jacq. LC 1,621,244 (all); 148,300 (Mexico and Central America only)
Solanumdouglasii Dunal LC 6,419,607
Solanumemulans Raf. LC 5,394,300
Solanumfurcatum Dunal LC 209,035,647 (North America only 4,169 – EN)
Solanuminterius Rydb. LC 4,506,327
Solanummacrotonum Bitter LC 3,804,650
Solanumnigrescens M.Martens & Galeotti LC 15,340,166
Solanumnigrum L. LC 78,076,619
Solanumnitidibaccatum Bitter LC See Särkinen et al. 2018
Solanumpruinosum Bitter LC 294,305
Solanumpseudogracile Heiser LC 1,048,309
Solanumretroflexum Dunal LC See Särkinen et al. 2018
Solanumsarrachoides Sendtn. LC 100,440,077
Solanumscabrum Mill. LC See Särkinen et al. 2018
Solanumtriflorum Nutt. LC 91,711,478
Solanumvillosum Mill. LC See Särkinen et al. 2018

Specimens cited in pdf format (traditional format; only specimens from the region cited)

Data type: PDF file

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Sandra Knapp, Gloria E. Barboza, Lynn Bohs, Tiina Särkinen

Searchable CSV file of all specimens examined for this treatment, including Old World and South America

Data type: CSV file

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Sandra Knapp, Gloria E. Barboza, Lynn Bohs, Tiina Särkinen

Types based on American specimens only, for full synonymy see Särkinen et al. (2018: 65–66)

EdmondsJM (1972) A synopsis of the taxonomy of Solanumsect.Solanum (Maurella) in South America.Kew Bulletin27(1): 95114. https://doi.org/10.2307/4117874SärkinenTPoczaiPBarbozaGEvan der WeerdenGMBadenMKnappS (2018) A revision of the Old World black nightshades (Morelloid clade of Solanum L., Solanaceae).PhytoKeys106: 1223. https://doi.org/10.3897/phytokeys.106.21991D’ArcyWG (1974a) [1973]) Solanaceae, Flora of Panama. Part 9.Annals of the Missouri Botanical Garden60: 573780. https://doi.org/10.2307/2395139HendersonRJF (1974) Solanumnigrum L. (Solanaceae) and related species in Australia.Contributions from the Queensland Herbarium16: 178.MortonCV (1976) A revision of the Argentine species of Solanum. Córdoba: Academia Nacional de Ciencias.BarbozaGEKnappSSärkinenT (2013) Grupo VII. Moreloide. In Anton AM, Zuloaga FO (Eds), Barboza GE (coord.) Flora Argentina vol. 13, Solanaceae. IBODA-IMBIV, CONICET: Buenos Aires & Córdoba, Argentina, 231–264.IUCN (2017) Guidelines for Using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. http://www.iucnredlist.org/documents/RedListGuidelines.pdf