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 douglasii Dunal, Prodr. [A. P. de Candolle] 13(1): 48. 1852Figure 12, 13 Solanum umbelliferum Eschsch. var. trachycladum Torr., Pacific Railr. Rep. Parke, Bot. 7(3) [preprint]: 17. 1856. Type. United States of America. California: Ventura County, San Buenaventura Ranch, 16 Feb 1855, T. Antisell s.n. (lectotype, designated here: NY [NY00821411]). Solanum arizonicum Parish, Proc. Calif. Acad. Sci., ser. 3, 2: 165. 1901. Type. United States of America. Arizona: Copper Basin, J.W. Toumey 397 (holotype: US [acc. # 211749, US00027460; isotype: UC n.v.). Solanum extusviolascens Bitter, Repert. Spec. Nov. Regni Veg. 11: 7. 1912. Type. Mexico. Sin. loc., J.G. Schaffner 654 (holotype: B, destroyed; no duplicates found). Solanum profundeincisum Bitter, Repert. Spec. Nov. Regni Veg. 12: 80. 1913. Type. Mexico. Baja California: Guadalupe Island, cañon near beach, 1875, E. Palmer 61 (lectotype, designated here: UPS [UPS-V-851402]; isolectotypes: BM [BM001007201], MO [MO-568699, acc. # 5510874], NY [NY00139024, NY00828776], YU [YU065318]). Type.

United States of America. California: “Nova California”, D. Douglas s.n. (holotype: G-DC [G00144189]; isotypes: BM [BM000838093], K [K001159712]).

Description.

Perennial, subwoody herbs or shrubs, erect to ascending, up to 2 m tall. Stems terete, green or purple-tinged, moderately to densely pubescent with simple, uniseriate 4–10-celled spreading eglandular trichomes, 0.5–1 mm long; new growth more densely pubescent. Sympodial units difoliate, not geminate. Leaves simple, 3–10(–17) cm long, 1.3–5(–7.5) cm wide, (broadly) ovate to lanceolate, green or marked with purple, green above, paler greyish-green below; adaxial surface moderately to densely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along the lamina and veins; abaxial surface more densely pubescent than the abaxial surface; primary veins 4–6 pairs, clearly evident abaxially; base abruptly contracted to attenuate, at times asymmetric, decurrent on the petiole; margins sinuate-dentate to toothed, rarely entire; apex acute; petiole 1–4(–7) cm long, moderately to densely pubescent with simple, uniseriate like those on stem. Inflorescences 1.5–4.5 cm long, lateral, internodal, unbranched to occasionally forked, with (3–)6–14 flowers spaced along the rhachis, moderately to densely pubescent with simple, uniseriate trichomes like those on stems; peduncle 1.5–4 cm long; pedicels 10–41 mm long, 0.3–0.4 mm in diameter at the base and 0.4–0.6 mm in diameter at the apex, straight and spreading, articulated at the base, spaced ca. 0.5–1 mm apart. Buds ovoid and narrower at the tips, the corolla exserted 1/5 of its length beyond the calyx tube. Flowers 5-merous, all perfect. Calyx tube 1–2 mm long, the lobes (1-)1.5–2.9 mm long, 0.7–1.5 mm wide, lanceolate to broadly triangular with obtuse to acute apices, moderately to densely pubescent with simple, uniseriate trichomes like those on stem. Corolla 13–15(–20) mm in diameter, stellate, white to lilac with a yellow-green central eye with black coloration at the base, lobed 1/3 to the base, the lobes 4.5–7 mm long, 2–4 mm wide, strongly reflexed at anthesis, sparsely pubescent abaxially with 1–4-celled simple uniseriate trichomes like those on stems and leaves but shorter. Stamens equal; filament tube 0.3–1 mm long; free portion of the filaments 0.1–0.5(1) mm long, sparsely pubescent with spreading uniseriate 4–6-celled simple trichomes adaxially; anthers (2.5-)3–4.5 mm long, 0.9–1.2 mm wide, ellipsoid and slightly tapered towards the tips, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 6.5–7.5 mm long, exserted 1.7–2.3 mm beyond the anther cone, densely pubescent with 2–3-celled simple uniseriate trichomes to 1/2–2/3 from the base; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 6–14 mm in diameter, black at maturity, opaque, the surface of the pericarp matte; fruiting pedicels 8–11 mm long, 0.4–0.5 mm in diameter at the base, 0.5–0.6 mm in diameter at the apex, spaced 1–3 mm apart, spreading to reflexed, dropping with mature fruits, very occasionally remaining on the inflorescence rhachis; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes 1.2–3 mm long, appressed against the berry. Seeds usually >50 per berry, 1.5–1.9 mm long, 1.2–1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells (2-)6–8 per berry, rather large, 0.5–0.7 mm in diameter. Chromosome number: 2n=2×=24 (Henderson 1974; Heiser et al. 1965 as S.amethystinum; Edmonds 1982, 1983; Stebbins and Paddock 1949; Heiser 1955 (as S.amethystinum); Soria and Heiser 1961 (as S.amethystinum and S.douglasii).

Solanumdouglasii Dunal A Habit B detail of adaxial leaf surface C detail of abaxial leaf surface D floral bud E dissected flower F fruiting branch G maturing fruit (A–GCarter et al. 2149). Drawing by R. Wise.

Solanumdouglasii Dunal A Habit B young and maturing inflorescences with buds and flowers C flowers at full anthesis D fully mature matte black fruits with appressed calyx lobes (A–DOchoterena et al. 979). Photos by S. Knapp.

Distribution.

(Figure 14) Solanumdouglasii occurs in North America from California east to Arizona and south to Nicaragua; it is the most common black nightshade in the southwestern United States of America and northern Mexico.

Distribution of Solanumdouglasii Dunal

Ecology.

Open areas and disturbed habitats in a wide variety of vegetation types, from xerophytic to mesophytic forests and oak-pine woodlands between (sea level-) 600 and 3,400 m elevation.

Common names.

United States of America. Arizona nightshade (Martin and Hutchins 1980), Douglas’ horse-nettle (NatureServe 2017), Douglas’s nightshade (Peck 1941; Martin and Hutchins 1980; Nee 2012), Greenspot nightshade (USDA Plants 2017). Mexico. Hierba (yerba) mora (many sources, [Chihuahua] Chichequelite (Pennington 42), [Chiapas] Moen (Tzeltal, Shilom Ton 9185), Mora wamul (Tzeltal, Gómez López 426), [Guerrero] Moradito (Kruse 1656), [Hidalgo] Tomaquilit (Villa Kumel 53), [México] Tomatillo (la Cruz Bolaños Adec-12), [Oaxaca] Pchfux-yaas (Zapotec, Hunn OAX-1547), Skelemal ch’aben (Tzeltal, López Pérez 326), [Puebla] Teconchichi (Tlapa & Ubierna 105), [Sonora] Chichicalite (Guizar N. et al. 4260).

Uses.

United States of America. [California] Leaves used as a potherb (Luiseño people of Orange County, Sparkman 1908); juice of berries used as wash for inflamed eyes and in tattooing or for dye (Luiseño people, Sparkman 1908; Cahuilla people of the Sonoran Desert, Bean and Saubel 1972). Mexico. Leaves used as a potherb (“quelite”). See also section on Uses.

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

Least Concern (LC). Solanumdouglasii is widespread and weedy in the southwestern United States of America and throughout Mexico. For EOO see Table 6.

Discussion.

Solanumdouglasii is most common west of the Rocky Mountains, along the western coast and southwesternmost United States of America along the Mexican border. Solanumdouglasii can be distinguished from the morphologically similar and sympatric S.nigrescens by its longer, slightly tapering anthers (greater than 3 mm long and in North America usually 4–4.5 mm long) and the minute free portion of the filaments. Both species are morphologically highly variable and sympatric through much of Mexico and Central America, often growing in the same areas; detailed studies are needed to establish whether interbreeding occurs between particular areas/populations in areas of sympatry. The two species have been put in synonymy by other authors (e.g., Edmonds 1972; D’Arcy 1974a, b), but characterised as “ill-defined” by others (e.g. Nee 1999).

The description of S.umbelliferumvar.trachycladum cites “Santa Inez and San Buenaventura Ranch” and “Flowers apparently white, about as large as in S.nigrum” (Torrey 1856) with no collector or date. The plants collected in the several expeditions ordered by the United States Government to plan a railway leading across the Rocky Mountains to the Pacific were variously described by Asa Gray (Harvard) and John Torrey (New York). Thomas Antisell collected between the Rio Grande River and southern California; his collections are described in Volume VII of the Reports (Brendel 1880) by Torrey. We have only found a single specimen collected by Antisell and annotated by Torrey with this name; it has the locality “San Buenaventura Ranch/Feb 16/Dr Antisell”. We select this sheet (NY00821411) as the lectotype following McNeill (2014).

In describing S.profundeincisumBitter (1913) cited two collections of Edward Palmer’s from Guadelupe Island, Palmer 60 pro parte and Palmer 61, both from UPS. Palmer 60 is a mixed collection, some parts of which were used to describe S.calvum (a synonym of S.americanum) and some as part of the protologue of S.profundeincisum. The collection Palmer 61 is represented by many duplicates and is not mixed; we select the UPS sheet of Palmer 61 (UPS-V-851402) cited by Bitter (1913) as the lectotype of S.profundeincisum.

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
HendersonRJF (1974) Solanumnigrum L. (Solanaceae) and related species in Australia.Contributions from the Queensland Herbarium16: 178.HeiserJr CBSoriaJBurtonDL (1965) A numerical taxonomic study of Solanum species and hybrids.American Naturalist99(909): 471488. https://doi.org/10.1086/282392EdmondsJM (1982) Epidermal hair morphology in SolanumL.sectionSolanum. Botanical Journal of the Linnean Society 85(3): 153–167. https://doi.org/10.1111/j.1095-8339.1982.tb02583.xEdmondsJM (1983) Seed coat structure and development in SolanumL.sectionSolanum (Solanaceae).Botanical Journal of the Linnean Society87(3): 229246. https://doi.org/10.1111/j.1095-8339.1983.tb00992.xStebbinsGLPaddockEF (1949) The Solanumnigrum complex in Pacific North America.Madrono10: 7081.HeiserCB (1955) The Solanumnigrum complex in Costa Rica.Ceiba4: 293299.SoriaJHeiserCB (1961) A statistical study of relationships of certain species of the Solanumnigrum complex.Economic Botany15(3): 245255. https://doi.org/10.1007/BF02862165MartinWCHutchinsCR (1980) A flora of New Mexico. J. Cramer in A.R. Gantner Verlag, Vaduz.NatureServe (2017) NatureServe Explorer. An online encyclopedia of life [web application] Version 7.1. NatureServe, Arlington [available http://explorer.matureserve.org, accessed 26 Nov 2017]PeckME (1941) A manual of the higher plants of Oregon. Binfords and Mort, Portland.NeeM (2012) Solanaceae. In: Baldwin BG, Goldman DH, Keil DJ, Patterson R, Rosetti TJ, Wilken DH (Eds) The Jepson Manual: vascular plants of California. University of California Press, Berkeley.USDA Plants (2017) USDA- NCRS The PLANTS Database National Plant Data Team, Greensboro [http://plants.usda.gov, accessed 26 November 2017].SparkmanPS (1908) The culture of the Luiseño Indians.University of California Publications in American archeology and Ethnology8(4): 187234. https://www.sacred-texts.com/nam/ca/coli/index.htmBeanJLSaubelKS (1972) Temalpakh (From the Earth): Cahuilla Indian knowledge and usage of plants. Malki Museum Press, Banning.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.pdfSärkinenTPoczaiPBarbozaGEvan 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.106.21991EdmondsJM (1972) A synopsis of the taxonomy of Solanumsect.Solanum (Maurella) in South America.Kew Bulletin27(1): 95114. https://doi.org/10.2307/4117874D’ArcyWG (1974a) [1973]) Solanaceae, Flora of Panama. Part 9.Annals of the Missouri Botanical Garden60: 573780. https://doi.org/10.2307/2395139D’ArcyWG (1974b) Solanum and its close relatives in Florida.Annals of the Missouri Botanical Garden61(3): 819867. https://doi.org/10.2307/2395032NeeM (1999) Synopsis of Solanum in the New World. In: NeeMSymonDELesterRNJessopJP (Eds) Solanaceae IV: advances in biology and utilization.Royal Botanic Gardens Kew, Richmond, 285333.TorreyJ (1856) Explorations and surveys for a railroad route from the Mississippi River to the Pacific Ocean. Botanical report.Pacific Railway Reports7(3): 128. [preprint of an article published as part of the expedition volume of 1857]BrendelF (1880) Historical sketch of the science of botany in North America from 1840 to 1858.American Naturalist14(1): 2538. https://doi.org/10.1086/272473McNeillJ (2014) Holotype specimens and type citations: General issues.Taxon63(5): 11121113. https://doi.org/10.12705/635.7BitterG (1913) Solana nova vel minus cognita X. XXVII. Morellae nova vel criticae.Repertorium Species Novarum Regni Vegetabilium12: 7689. https://doi.org/10.1002/fedr.19130120905