taxonID	type	description	language	source
BD7F0CA84213906ECB6FD2FB0566693E.taxon	materials_examined	Type. 1 Cultivated at the Chelsea Physic Garden [in protologue said to " grow naturally in Virginia "], Herb. Miller s. n. (lectotype, designated by Edmonds 1972, pg. 103 [as type]: BM [BM 000617683]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
BD7F0CA84213906ECB6FD2FB0566693E.taxon	description	Description. Annual to short-lived perennial herbs up to 1.5 m tall, subwoody at base. Stems terete or somewhat angled with ridges, older stems often appearing spinescent, not markedly hollow; new growth pubescent with simple, spreading, uniseriate 2 - 8 - celled eglandular trichomes 0.2 - 0.8 mm long, often clustered along the stem angles; older stems glabrescent, with only the trichome bases persisting as pseudo-spines. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.5 - 10.5 cm long, 1.0 - 4.5 cm wide, ovate to elliptic; adaxial surface sparsely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along the lamina and the veins; abaxial surface similar but more densely pubescent; major veins 3 - 6 pairs; base attenuate, decurrent on the petiole; margins entire or occasionally sinuate-dentate; apex acute; petioles (0.3 -) 2.0 - 3.8 (- 4.0) cm long, sparsely pubescent with simple uniseriate trichomes like those on stems. Inflorescences 0.6 - 2.5 cm long, lateral and internodal, unbranched or rarely forked, with (3 -) 4 - 6 (8) flowers (very rarely with many flowers in unusual many-branched inflorescences) clustered near the tips (umbelliform to sub-umbelliform), sparsely pubescent with simple uniseriate trichomes like those on stems; peduncle (0.5 -) 1.0 - 1.8 cm long, delicate; pedicels 3 - 9 mm long, 0.2 - 0.3 mm in diameter at the base and 0.4 - 0.5 mm at the apex, stout, straight and spreading, articulated at the base; pedicel scars spaced 0 - 0.5 mm apart, clustered at the tip of the inflorescence. Buds broadly ellipsoid, the corolla exserted 1 / 3 beyond the calyx lobe tips before anthesis. Flowers 5 - merous, all perfect. Calyx tube 0.8 - 1.3 mm long, the lobes 0.3 - 0.5 mm long, 0.5 - 0.6 mm wide, broadly triangular with obtuse apices, sparsely pubescent with simple uniseriate trichomes like those of the stem. Corolla 3 - 6 mm in diameter, stellate, white with a yellow-green central portion near the base, lobed 1 / 2 - 2 / 3 of the way to the base, the lobes 2.0 - 3.2 mm long, 1.0 - 2.5 mm wide, strongly reflexed at anthesis, later spreading, densely 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 0.5 - 0.8 mm long, adaxially pubescent with tangled uniseriate trichomes; anthers 0.7 - 1.5 mm long, 0.5 - 0.6 mm wide, ellipsoid to almost globose and very plump-looking, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.2 - 2.6 mm long, densely pubescent with 2 - 3 - celled simple uniseriate trichomes 2 / 3 from the base where included in the anther cone, almost included to exserted 0.5 (- 1.0) mm beyond the anther cone; stigma minutely capitate, the surface minutely papillate, green in live plants. Fruit a globose berry, 4 - 9 (- 12) mm in diameter, purplish-black at maturity, opaque, the surface of the pericarp markedly shiny; fruiting pedicels 13 - 18 mm long, ca. 0.7 - 1.0 mm in diameter at the base and 0.8 - 1.0 mm in diameter at the apex, stout, straight and spreading, spaced ca. 1 (- 3) mm apart or tightly clustered, not falling with the fruit, remaining on the plant and persistent on older inflorescences; fruiting calyx lobes not accrescent, the tube less than 1 mm long, the lobes 1 (- 2) mm long, strongly reflexed at fruit maturity. Seeds 30 - 50 per berry, 1.0 - 1.5 mm long, 0.8 - 1.3 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 mostly absent (Australia, South Pacific, and South America), but if present (North America, Mexico, Caribbean, Eurasia and Africa) 2 - 4 (6) per berry, 2 - 4 larger ones> 0.5 mm, and two smaller ones <0.5 mm in diameter. Chromosome number: 2 n = 2 x = 24 (see Saerkinen et al. 2018 for vouchers).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
BD7F0CA84213906ECB6FD2FB0566693E.taxon	distribution	Distribution. (Figure 5) Solanum americanum is a globally distributed weed found throughout the tropics and subtropics; it is not clear where it is native, or if this circumtropical distribution is its native range. In the region treated in this monograph it is commonest in Central America and the Caribbean, but is found around the coasts in North America, especially around the Gulf of Mexico and the Pacific.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
BD7F0CA84213906ECB6FD2FB0566693E.taxon	discussion	Discussion. Solanum americanum is the most widespread and common species of the morelloid solanums (see Saerkinen et al. 2018), and quite possibly the most widely distributed species in Solanum. It has been implicated as the diploid parent in the polyploid events that gave rise to the species of the Old World (e. g., Edmonds 1977; Poczai and Hyvoenen 2011), although this has been disputed (Ma 1995). The name S. americanum has been in common use in North America (e. g., Stebbins and Paddock 1949) for what is now known as S. emulans, but more recently (Schilling 1981) the two taxa have been distinguished and the name S. ptychanthum has been used for the taxon for which the oldest name is S. emulans. The type specimen of S. ptychanthum, however, falls within the variation of S. americanum, so is treated as a synonym here. The application of the name S. americanum to any morelloid species with small anthers from northeastern North America should be viewed with caution. Solanum americanum can be easily recognised in fruit by its shiny black berries with small, strongly reflexed calyx lobes that are held on erect or spreading pedicels. In flower, the species has tiny almost globose anthers 0.8 - 1.5 mm long and short filaments usually less than 1 mm long. It has been often confused with S. emulans and S. nigrescens. Solanum emulans has equally short plump anthers but longer filaments (0.6 - 1 mm versus ca. 0.5 mm long), matte black or green fruits on deflexed pedicels and calyx lobes that are not markedly reflexed in fruit. Ripe berries of S. americanum are shiny black (but that can be difficult to see in herbarium specimens) and in North and Central America and the Caribbean usually have four stone cells in each. Berries of S. emulans have more than five stone cells. When berries ripen in S. americanum they fall from the plant leaving the stout, spreading pedicels behind, while berries drop off with the pedicels in S. emulans leaving only the peduncles behind in herbarium specimens. This can, however, be difficult to see in specimens with only very old inflorescences. Solanum nigrescens differs from S. americanum in having larger anthers always more than 2 mm long, matte black or green fruits that are held on spreading or deflexed pedicels that drop with the berry, and calyx lobes appressed to the berry base in fruit. Berries of S. nigrescens have more than 5 (usually 5 - 6 large and several smaller) stone cells, while plants of S. americanum from this region have 2 (- 4). Inflorescences of S. americanum tend to be more sub-umbelliform in appearance than those of S. nigrescens, and calyx lobes of S. americanum are strongly reflexed and smaller relative to berry size in fruit. D'Arcy (1974 a, b) suggests that S. americanum hybridizes with other diploid species (e. g., S. nigrescens) and that intermediates are common, but did not cite vouchers. Our observations (but see below) are that the two species are usually distinguishable using the characters above. Some geographical trends in the morphological variation within the species can be observed, where populations along the coast of the Gulf of Mexico appear more hairy with duller grey-green leaf coloration, with more narrow, lanceolate rather than ovate leaves, with racemose inflorescences rather than strict umbels, with more rounded calyx lobes that do not always strongly reflex in fruit, and with generally larger fruits. The small anthers combined with stout and spreading pedicels in fruit that remain on the plant after fruits drop off are strong indications that these populations belong to S. americanum and do not represent a distinct species. Variation appears continuous and could be caused by local introgression from the sympatric diploid species S. pseudogracile or S. nigrescens. Collections with forked inflorescences (Nee & McClelland 60259 from Florida; Dancer s. n. from Jamaica) are likely to be isolated aberrant individuals; in other parts of the world populations of S. americanum with highly branched inflorescences occur (e. g., China, type of S. merrillianum T. N. Liou) indicating that plasticity in this character is not unusual. It may also be that plants from cultivated populations in Asia have been brought with rice cultivation to North America. Plants collected as weeds of rice fields in Louisiana (Ma 1995; E. Shilling, pers. comm.) and identified as S. merrillianum are somewhat intermediate between S. americanum and S. nigrescens and could represent recent homoploid hybrids; preliminary data indicate they are members of the clade containing both those taxa. Further studies using molecular markers and carefully comparing Asian and American populations will be necessary to unravel this enigma. Manoko et al. (2007) distinguished S. americanum and S. nodiflorum using AFLP markers; we re-examined the material they used and consider the plants they called S. nodiflorum to be S. americanum as defined here, and plants they called S. americanum represent specimens of S. nigrescens (see Saerkinen et al. 2018: 61). Typification details for the many synonyms of S. americanum can be found in Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
BD7F0CA84213906ECB6FD2FB0566693E.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E2731A0AC8700B825F526C0F56A87F5C.taxon	materials_examined	Type. 2 Mauritius. " Ex ins. Mauritiana ", Herb. Lamarck s. n. (lectotype, designated by Barboza et al. 2013, pg. 242: P [P 00357629]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E2731A0AC8700B825F526C0F56A87F5C.taxon	description	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: 2 n = 2 x = 24 (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E2731A0AC8700B825F526C0F56A87F5C.taxon	distribution	Distribution. (Figure 8) Solanum chenopodioides 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.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E2731A0AC8700B825F526C0F56A87F5C.taxon	discussion	Discussion. Solanum chenopodioides 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. Solanum chenopodioides 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. Solanum chenopodioides 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 Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E2731A0AC8700B825F526C0F56A87F5C.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
87AB3C3975B4B66EFAFA02284EFB4E56.taxon	materials_examined	Type. Cultivated in Vienna [" Hort. Bot. Vindob. "] seeds said to be from Peru, N. von Jacquin s. n. (lectotype, designated by D'Arcy 1970, pg. 559: W [acc. # 0022473]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
87AB3C3975B4B66EFAFA02284EFB4E56.taxon	description	Description. Annual to short lived perennial herbs to 30 - 50 cm tall, subwoody and branching at base. Stems terete, green to straw colour, sprawling, somewhat weak and decumbent, not markedly hollow; new growth nearly glabrous to sparsely pubescent with weak simple, uniseriate appressed 1 - 8 - celled eglandular trichomes, these ca. 0.3 mm long; older stems glabrescent. Sympodial units difoliate or occasionally trifoliate, the leaves not geminate. Leaves simple, 4.5 - 8 cm long, 1.5 - 4 cm wide, ovate-lanceolate, chartaceous to subcoriaceous; both surfaces glabrous or sometimes sparsely ciliate near the base of the winged petiole; major veins 7 - 9 pairs, not clearly evident abaxially in live plants, paler in herbarium specimens; base long-attenuate, decurrent on the petiole; margins entire (in Peru rarely slightly 3 - lobed, Croat 58409); apex acute; petioles 0.5 - 1 cm, glabrous to sparsely puberulent, winged to the base. Inflorescences 2 - 3 cm long, lateral, internodal or opposite the leaves, 4 - 7 times branched, with 20 - 50 (- 60) flowers spaced along the rhachis, nearly glabrous to sparsely pubescent; peduncle 0.1 - 2 cm, straight in fruit; pedicels 2 - 2.5 mm long, less than 0.5 mm in diameter at the base, ca. 0.5 mm in diameter at the apex, spreading, articulated at the base; pedicel scars spaced 1 - 3 mm apart. Buds globose, the corolla about halfway exserted from the calyx tube before anthesis, the tips of the corolla lobes often much more pubescent than the calyx. Flowers 5 - merous, all perfect. Calyx tube 0.5 - 1 mm long, conical or broadly conical, the lobes 0.5 - 0.6 mm long, ca. 0.5 mm wide, broadly triangular, glabrous to very sparsely puberulent with simple, uniseriate trichomes. Corolla 5 - 10 mm in diameter, white or purple, the abaxial surface usually purple, rotate-stellate, the lobes 1 - 2.5 mm long, 1 - 1.5 mm wide, broadly triangular, reflexed at anthesis, later spreading, glabrous adaxially, minutely white-puberulent abaxially on the tips. Stamens equal; filament tube minute; free portion of the filaments ca. 0.2 mm long, adaxially pubescent with simple tangled white trichomes; anthers 0.8 - 1.5 (- 1.8) mm long, ca. 0.5 mm wide, ellipsoid, yellow, somewhat connivent, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style ca. 2 mm long, hardly exserted from the anther cone, pubescent in the lower 2 / 3 with tangled, white uniseriate simple weak-walled trichomes; stigma globose-capitate, minutely papillate, pale green in live plants. Fruit a globose berry, 4 - 6 mm in diameter, orange to red when ripe, opaque, the surface of the pericarp shiny or matte; fruiting pedicels 2 - 3 mm long, ca. 0.5 mm in diameter at base, strongly recurved at the very base, dropping with mature fruits, not persistent; fruiting calyx scarcely accrescent, the tube ca. 1 mm long, the lobes 1 - 1.3 mm long, appressed to the berry. Seeds 20 - 30 per berry, 1.5 - 1.8 mm long, 1.2 - 1.4 mm wide, flattened reniform with a central hilum, light yellow-tan or reddish brown in herbarium material, the surfaces minutely pitted, the testal cells with sinuate margins. Stone cells 2, ca. 1.5 mm in diameter, globose, prominent near the apex of the berry. Chromosome number not known.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
87AB3C3975B4B66EFAFA02284EFB4E56.taxon	distribution	Distribution. (Figure 11) Solanum corymbosum is native to the western slopes of the Andes in Peru, and naturalised in central and southern Mexico, possibly through introduction in colonial times.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
87AB3C3975B4B66EFAFA02284EFB4E56.taxon	discussion	Discussion. Solanum corymbosum is a member of the Radicans group and is related to species of southern South America (see Saerkinen et al. 2015 b). The distribution of this species in Mexico is highly disjunct from what are presumed native populations in Peru and Mexican populations are thought to represent an introduction of this species in post-Columbian times. It is tempting to speculate on an inadvertent introduction between mining areas, perhaps even in Spanish colonial times. Populations in Mexico show nearly identical haplotypes to those from the coastal regions in Peru (Mitchell 2014), supporting this hypothesis. Mexican populations of S. corymbosum differ from Andean populations in having larger leaves (20 cm 2 Mexico, ca. 9 cm 2 Andes) and larger and fewer berries; an average of ca. 30 berries of 5.5 cm diameter per inflorescence in Mexican specimens versus an average of ca. 50 berries of 3.5 mm in diameter per inflorescence in Andean specimens (Mitchell 2014). This may be due to founder effects in the establishment of the Mexican populations, and Mitchell (2014) has speculated that these populations may be polyploid. Solanum corymbosum can be distinguished from other morelloids occurring in Mexico in its orange to red fruits with two large apical stone cells, its highly branched inflorescences and diminutive flowers with rotate-stellate corollas that are usually white adaxially and purple abaxially. The leaves are thicker than other morelloids from the area, and the petioles are strongly winged. Three collections of Solanum corymbosum in BM [all mounted on a single sheet] 1. " Hort. Paris. L'Heritier 1783 (E Peru Dombey) ", 2. " Hort. Kew. 1785 ", 3. " Peru, Dombey 63, " P-Lam [Morton neg. 8364] are possible isotype material of various of the synonyms. Collections attributed to Dombey from Paris are probably isolectotype material of S. cymosum (see Knapp 2008 b), while those from Kew and the Lamarck herbarium are not type material. It is possible that much of the botanical garden material being described in the late 18 th century came from a few collections and is all genetically the same. Solanum leptanthum is a synonym of S. pubigerum Dunal (a member of the Dulcamaroid clade, Knapp 2013), but variety parviflorum corresponds to S. corymbosum. We have selected the best preserved of the three sheets in the Humboldt and Bonpland herbarium at P (P 00670610) as the lectotype for var. parvifolium.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
87AB3C3975B4B66EFAFA02284EFB4E56.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
5E41D9899EA99E4D72CBD124DE5F8440.taxon	materials_examined	Type. United States of America. California: " Nova California ", D. Douglas s. n. (holotype: G-DC [G 00144189]; isotypes: BM [BM 000838093], K [K 001159712]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
5E41D9899EA99E4D72CBD124DE5F8440.taxon	description	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: 2 n = 2 x = 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).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
5E41D9899EA99E4D72CBD124DE5F8440.taxon	distribution	Distribution. (Figure 14) Solanum douglasii 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.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
5E41D9899EA99E4D72CBD124DE5F8440.taxon	discussion	Discussion. Solanum douglasii is most common west of the Rocky Mountains, along the western coast and southwesternmost United States of America along the Mexican border. Solanum douglasii 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 1974 a, b), but characterised as " ill-defined " by others (e. g. Nee 1999). The description of S. umbelliferum var. 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 Tor rey (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 (NY 00821411) as the lectotype following McNeill (2014). In describing S. profundeincisum Bitter (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.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
5E41D9899EA99E4D72CBD124DE5F8440.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C603910F6F12EB23C4D39B42644DAE2C.taxon	materials_examined	Type. United States of America. " Amer. bor. ", C. S. Rafinesque s. n. [ex Herb. Rafinesque] (neotype, designated here: W [acc. # 0009388]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C603910F6F12EB23C4D39B42644DAE2C.taxon	description	Description. Annual herbs to subwoody perennial shrubs up to 1.0 m tall, branching at base. Stems terete to ridged, green colour, pubescent with simple, appressed, uniseriate eglandular 1 - 5 - celled trichomes, these ca. 0.2 mm long, new growth more densely pubescent. Sympodial units difoliate, not geminate. Leaves simple, 4.5 - 10.5 (- 17.5) cm long, 2.0 - 6.3 (- 8.3) cm wide, ovate, thin membranous, slightly discolorous, green above and purplish tinged underneath, especially so in younger growth; adaxial surface glabrous to sparsely pubescent with appressed translucent, simple, uniseriate trichomes like those on stem scattered mainly along veins; abaxial surface glabrous to sparsely pubescent with trichomes like those of the upper surface on both lamina and veins; primary veins 4 - 6 pairs; base attenuate to acute; margins sinuate dentate, rarely entire; apex acute to acuminate; petiole 1.0 - 5.0 cm long, pubescent with simple uniseriate trichomes like those of the stems. Inflorescences 1.0 - 2.5 cm long, lateral, internodal, unbranched or occasionally forked, with (2) 3 - 6 flowers clustered near the tips (sub-umbelliform), sparsely pubescent with appressed simple uniseriate trichomes like those on stem; peduncle 1.0 - 1.7 cm long, straight; pedicels 8 - 10 mm long, 0.4 - 0.5 mm in diameter at the base and 0.5 - 0.6 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0 - 0.5 mm apart. Buds subglobose, corolla exserted from the calyx to 1 / 3 of its length. Flowers 5 - merous, all perfect. Calyx tube 0.7 - 0.9 mm long, the lobes 0.8 - 2.2 mm long, 0.7 - 1.3 mm wide, ovate to elongate with obtuse apices, sparsely pubescent with appressed hairs like those on stem but shorter. Corolla 8 - 10 mm in diameter, stellate, white with a yellow-green central portion near the base, lobed 1 / 3 to the base, the lobes 3.0 - 4.0 mm long, 1.0 - 1.2 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially along margins and apex with simple uniseriate trichomes like those on stem and leaves but shorter. Stamens equal; filament tube minute, pubescent with spreading uniseriate simple trichomes adaxially; free portion of the filaments 0.6 - 1.0 mm long, pubescent like the tube; anthers (1 -) 1.5 - 1.7 mm long, 0.4 - 0.5 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 3.5 - 4.5 mm long, not exceeding anthers, densely pubescent with 2 - 3 - celled simple uniseriate trichomes along 1 / 3 to 1 / 2 from the base; stigma capitate, minutely papil late, green in live plants. Fruit a globose berry, 6 - 8 mm in diameter, dull purplish-black at maturity, opaque, the surface of the pericarp matte to slightly shiny; fruiting pedicels 8 - 10 mm long, 0.4 - 0.6 mm in diameter at the base, 0.7 - 1.0 mm in diameter at the apex, recurved to reflexed, pedicels spaced 0.5 - 2.5 mm apart, dropping with mature fruits; fruiting calyx somewhat accrescent, the tube less than 1 mm long, the lobes 1.0 - 2.2 mm long, appressed to the surface of the berry or slightly spreading in mature fruit. Seeds 20 - 50 (- 60) per berry, 1.6 - 1.8 mm long, 1.0 - 1.2 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells 6 - 9 (10) per berry, ca. 0.3 mm in diameter. Chromosome number: 2 n = 2 x = 24 (Stebbins and Paddock 1949, as S. americanum; Mulligan 1961, as S. americanum; Soria and Heiser 1961, as S. americanum; Heiser et al. 1965, as S. americanum; Edmonds 1983, as S. americanum; Crompton and Bassett 1976, as S. americanum).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C603910F6F12EB23C4D39B42644DAE2C.taxon	distribution	Distribution. (Figure 17) Solanum emulans is endemic to North America and is the most common species of black nightshade in eastern North America east of the Rocky Mountains from Maine to North Carolina and into Canada. Plants collected near Vancouver (British Columbia) may have been introduced along the railways.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C603910F6F12EB23C4D39B42644DAE2C.taxon	discussion	Discussion. Solanum emulans can be distinguished from other morelloids in North America by the small anthers 1.0 - 1.5 mm long, relatively long filaments 0.6 - 1.0 mm compared to S. americanum, calyx lobes longer than S. americanum and these appressed in fruit rather than strongly reflexed like in S. americanum, pedicel thickened at the apex in fruit (unlike in S. americanum), and pedicels that drop off with mature fruits (pedicels remain on the inflorescence in S. americanum). Solanum emulans has always 4 - 9 (10) stone cells in fruits, while S. americanum either lacks or has maximum of 4 stone cells. Solanum emulans can be distinguished from S. interius and S. nigrescens by its shorter anthers, usually shorter calyx lobes, and usually unbranched inflorescences. When sympatric with the occasionally introduced S. nigrum, S. emulans can be easily distinguished based on anther length and the numerous stone cells in the berries, but S. emulans also generally has thinner leaves that are often purplish tinged beneath. In the Great Plains, the morphologically similar S. interius becomes more common than S. emulans, while along the southern East and Gulf coasts in the United States of America S. americanum becomes more common. Solanum emulans is not known from the Caribbean. Although S. emulans appears to have been in cultivation in European botanical gardens since the 18 th century, it has not escaped and naturalised beyond where it has initially been introduced. The few European specimens are from areas near oil and clothing factories and have apparently not persisted (Polgar 1926). Constantine Rafinesque cited no specific specimens in his many descriptions of new taxa, and any herbarium he kept in North America was widely dispersed after his death and is thought to have been destroyed (Pennell 1944; Warren 2004). A specimen in the Vienna herbarium (W acc. # 0009388) corresponding to the description of S. emulans and labelled " Solanum Virginicum / Amer. Bor. Rafinesque " and the date 1828 may be original material for this name. We have here selected this as a neotype, since there is no evidence in the protologue that this (or any other) specimen was used by Rafinesque to describe S. emulans. The name S. ptychanthum has been used for this species in North America (e. g., Schilling 1981; Voss et al. 1993; Jones 2005), but the type of that name corresponds to a plant of S. americanum (see S. americanum description). Solanum emulans was long ignored, but in the protologue Rafinesque clearly refers to a taxon from " New England and Kentucky " that people were calling " S. virginicum " - probably the Linnaean S. nigrum var. virginicum, not S. virginicum L. (an illegitimate name and orthographic variant of the spiny solanum from India S. virginianum L., see Jarvis 2007) - and his description matches this widespread small-flowered morelloid from eastern North America. The protologue states " NE states, usually mistaken for S. Virg. [virginianum] but smooth smaller, fl. white small, berries pisiform ". Specimens corresponding to this taxon are in the Dillenian herbarium in OXF under the polynomial (Solanum nigrum vulgari simile, caulibus exasperatis Dill. elth. 368, t. 275, f. 256) and correspond to the plate that was the only element cited for Solanum nigrum var. virginicum (Linnaeus 1753). D'Arcy (1974 a) cited " Hungary, Polgar s. n. (MPU) " as the type of Solanum adventitium, but without specifying a locality or number. We do not consider this specific enough to constitute the citation of a single unambiguous specimen and it is likely to be in conflict with the protologue; we therefore lectotypify S. adventitium here. In the protologue of S. adventitium Polgar (1926) cited several of his own collections made at " Meller'schen Oelfabrik " and " Gueterbahnhof " (both in Gyoer, Hungary) between 1915 and 1919, but cited neither numbers nor herbaria. He noted that the plants had disappeared from both localities by October 1919, but again cited no herbarium. His herbarium is kept at BP, and we have selected one of his many collections labelled as S. adventitium in that herbarium collected between 1916 and 1919 from the freight depot in Gyoer as the lectotype (BP- 352743).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C603910F6F12EB23C4D39B42644DAE2C.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
FFDC122C959A208E11486139562F92E7.taxon	materials_examined	Type. Peru? [more likely Chile]. " Cette plante croit au Perou ", J. Dombey [343] (lectotype, first step designated by Edmonds 1972, pg. 107 [as holotype], second step designated by Barboza et al. 2013, pg. 246: P [P 00335357]; isolectotypes: CORD [CORD 00006928], F [v 0043232 F, acc. # 976864], G [G 00359946], G-DC [G 00144483], P [P 00335358]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
FFDC122C959A208E11486139562F92E7.taxon	description	Description. Annual or perennial herbs to 1.0 m tall, erect to lax, subwoody at base, sprawling to ca. 2 m across. Stems terete or ridged, green to purple tinged, not markedly hollow sparsely pubescent with simple, uniseriate 1 - 5 - celled eglandular trichomes 0.1 - 0.5 mm long; new growth sparsely to densely pubescent with similar simple, uniseriate 1 - 5 - celled eglandular trichomes; older stems sparsely pubescent to glabrescent, pale yellowish brown. Sympodial units difoliate, the leaves not geminate. Leaves simple, (1.5 -) 4.0 - 8.0 (- 2.0) cm long, (0.6 - 2.2 - 4.6 (- 6.5) cm wide, ovate to rhomboidal, green above, slightly paler beneath; adaxial surface sparsely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along lamina and veins; abaxial surface more densely pubescent; major veins 4 - 6 pairs; base cuneate to acute, the two sides slightly unequal, decurrent on the petiole; margins sinuate-dentate or entire; apex acute; petioles 1.0 - 3.5 cm long, sparsely pubescent with simple uniseriate trichomes like those on stem. Inflorescences (1.0 -) 1.5 - 3.0 (- 4.0) cm long, lateral, internodal, forked or more rarely unbranched, with 6 - 14 flowers clustered at the tips (sub-umbelliform) or evenly spaced along the rhachis, sparsely pubescent with simple uniseriate trichomes like those on stem; peduncle (1.0 -) 1.5 - 2.0 cm long; pedicels 4.0 - 7.5 mm long, 0.2 - 0.3 mm in diameter at the base and 0.3 - 0.4 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0.2 - 2.5 mm apart. Buds subglobose, the corolla exserted 1 / 3 - 1 / 2 from the calyx tube before anthesis. Flowers 5 - merous, all perfect. Calyx tube 2 - 3 mm long, conical, the lobes 0.8 - 1.5 mm long, 0.6 - 1.0 mm wide, rectangular to narrowly obovate with obtuse to shortly acute apices, pubescent with simple uniseriate trichomes like those on stem but shorter. Corolla 12 - 20 mm in diameter, white to lilac with a green or yellow-green central portion near the base, this sometimes purplish near the lobe midvein, stellate, lobed 1 / 3 - 1 / 2 of the way to the base, the lobes 5.5 - 7.0 mm long, 2.8 - 5.5 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially with 1 - 4 - celled simple uniseriate trichomes, especially along the margins and apex, these shorter than the trichomes of the stems and leaves. Stamens equal; filament tube minute; free portion of the filaments 0.9 - 1.6 (2) mm long, adaxially pubescent with tangled uniseriate 4 - 6 - celled simple trichomes; anthers 2.3 - 3.3 (- 3.6) mm long, 0.8 - 1.0 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 6.0 - 6.5 mm long, densely pubescent with 2 - 3 - celled simple uniseriate trichomes in the lower 1 / 2 - 2 / 3, exserted 2 - 3 mm beyond the anther cone and somewhat curved; stigma capitate, minutely papillate, yellow or green in live plants. Fruit a globose berry, 6 - 9 mm in diameter, dull green to purple at maturity, opaque, the surface of the pericarp matte; fruiting pedicels 7 - 12 mm long, 0.2 - 0.4 mm in diameter at the base, 0.5 - 1.0 mm in diameter at the apex, strongly reflexed, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube 1.0 - 2.0 mm long, the lobes 1.5 - 2.5 mm long, appressed against the berry. Seeds 30 - 40 per berry, 1.8 - 2.0 mm long, 1.4 - 1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow-brown, the surface minutely pitted, the testal cells pentagonal in outline. Stone cells 6 - 14 per berry, 0.8 - 1.0 mm in diameter. Chromosome number: 2 n = 6 x = 72 (Stebbins and Paddock 1949; Edmonds 1982, 1983; Chiarini et al. 2017).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
FFDC122C959A208E11486139562F92E7.taxon	distribution	Distribution. (Figure 20) Solanum furcatum is native to Chile (incl. the Juan Fernandez Islands) and adjacent Andean Argentina. It is probably locally introduced and naturalised along the west coast of the United States of America, Australia and New Zealand. Only a few specimens have been seen from California and Oregon, but in those areas the species is clearly naturalised. Wiggins (1980) recorded S. furcatum from Baja California (Mexico), with no specimen citations; all specimens we have seen identified as S. furcatum from Baja California are plants of S. douglasii.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
FFDC122C959A208E11486139562F92E7.taxon	discussion	Discussion. Solanum furcatum can be distinguished from the similar and sympatric S. douglasii in its usually forked inflorescences, globose buds from which the style is often exserted, ellipsoid anthers on distinct filaments, and berries with usually more than 10 stone cells. Solanum furcatum is not sympatric with S. nigrescens but differs from it in the same set of characters and in its style that is exserted for as long as the anther cone (e. g., exserted portion of the style equal to the length of the anther cone). Solanum furcatum was considered to be an introduction from Chile by Stebbins and Paddock (1949) and is well-established but not common in coastal areas from Oregon to central California. Other Chilean species have similarly become established along the west coast of the United States [such as Nicotiana acuminata (Graham) Hook., see Goodspeed 1954; Knapp in press]. Details of typification for the synonyms of S. furcatum can be found in Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
FFDC122C959A208E11486139562F92E7.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
3E657D97531BAC49217F74BC90CC95A6.taxon	materials_examined	Type. United States of America. Nebraska: Hooker County, on Middle Loup River, near Mullen, 20 Jul 1893, P. A. Rydberg 1385 (lectotype, designated here: NY [NY 00138953] isotypes: GH [GH 00077424], H [acc. # 1087075], NDG [NDG 45091], NEB [NEB-V- 0000607], NY [NY 00138952], US [US 00027625, acc. # 210385; US 02828882, acc. # 210353]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
3E657D97531BAC49217F74BC90CC95A6.taxon	description	Description. Annual herbs to subwoody perennial shrubs up to 1.0 m tall, branching at base. Stems terete to ridged, pale straw colour, sparsely pubescent with simple, appressed, uniseriate (2) 4 - 8 - celled trichomes, these ca. 0.6 mm long, the new growth more densely pubescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 4.5 - 11.2 cm long, 2.3 - 6.8 cm wide, ovate to broadly ovate, membranous, green on both sides; adaxial surface sparsely pubescent with appressed translucent, simple, uniseriate trichomes like those on stem scattered along veins and lamina; abaxial surface more densely pubescent with trichomes like those of the upper surface across both lamina and veins; primary veins 4 - 6 pairs; base attenuate; margins sinuate dentate, especially so up to 2 / 3 from the base, to occasionally entire; apex acute to acuminate; petiole 0.5 - 3.5 cm long, pubescent with simple uniseriate trichomes like those of the stems. Inflorescences 2.4 - 3.5 cm long, lateral, internodal, unbranched or rarely forked, with (2) 3 - 8 flowers clustered near the tips (sub-umbelliform) or less commonly the distal flowers spaced along the rhachis, the lowermost flower distant from the rest, sparsely pubescent with appressed simple uniseriate trichomes like those on stem, rhachis 2 - 10 mm long when present; peduncle 1.0 - 2.0 cm long, straight; pedicels 5 - 8 mm long, 0.3 - 0.4 mm in diameter at the base and 0.4 - 0.5 mm in diameter at the apex, spreading, the terminal pedicels articulated at the base, but the lowermost flower with the pedicel articulated in the basal 1 / 4 to 1 / 3; pedicels spaced 0 - 1.0 mm apart. Buds globose, corolla exserted from the calyx 1 / 5 to 1 / 3. Flowers 5 - merous, all perfect. Calyx tube 1.0 - 1.5 mm long, lobes irregularly unequal, the longest 1.7 - 4.5 mm long, 0.6 - 0.7 mm wide, lanceolate with acute to acuminate apices, sparsely pubescent with appressed hairs like those on stem but shorter. Corolla 6 - 12 mm in diameter, stellate, white with a yellow-green central portion near the base, lobed 1 / 2 to 2 / 3 to the base, the lobes 4.0 - 5.0 mm long, 2.0 - 3.0 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially with simple uniseriate trichomes like those on stem and leaves but shorter. Stamens equal; filament tube minute; free portion of the filaments 0.7 - 1.0 mm long, pubescent with tangled uniseriate simple trichomes; anthers 1.8 - 2.5 mm long, 0.6 - 0.9 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 3.5 - 4.5 mm long, exserted 0 - 1 mm beyond anther cone, densely pubescent with 2 - 3 - celled simple uniseriate trichomes along 2 / 3 from the base; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 10 - 14 mm in diameter, purple-black at maturity, opaque, the surface of the pericarp shiny; fruiting pedicels 6 - 10 mm long, 0.4 - 0.6 mm in diameter at the base, 0.6 - 1.0 mm in diameter at the apex, recurved to reflexed, pedicels spaced 0.5 - 2.5 mm apart, dropping with mature fruits, occasionally not dropping; fruiting calyx not accrescent, the tube 1.5 - 2.0 mm long, the lobes (2.0 -) 3.0 - 4.0 mm long with the apices spreading to strongly reflexed in fruit. Seeds 20 - 40 per berry, 1.8 - 2.0 mm long, 1.5 - 1.6 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells 2 - 4, 0.8 - 1.0 mm in diameter, white or cream coloured. Chromosome number: 2 n = 2 x = 24 (Heiser et al. 1965).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
3E657D97531BAC49217F74BC90CC95A6.taxon	distribution	Distribution. (Figure 23) Solanum interius is endemic to North America, and the most common species of morelloid in the Great Plains. It is less common west of the Rocky Mountains, and although it does extend to Arizona and western New Mexico, S. interius does not occur on the Gulf Coast where it is replaced by S. nigrescens. Records of S. interius for Canada (Saskatchewan, Harms 2006) have not been verified with voucher specimens, although it is to be expected there. Specimens annotated by Harms as S. interius in SASK are of S. emulans (e. g., Child s. n., collected 29 July 1941)	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
3E657D97531BAC49217F74BC90CC95A6.taxon	discussion	Discussion. Solanum interius can be distinguished from other North American morelloids by its inflorescence with apparently uneven branches, one with several flowers and the other apparently with a single flower that is actually the basal flower with the articulation ca. 1 / 4 to 1 / 3 of the way up the pedicel, very like the pedicel articulation in wild potatoes. Other distinguishing features to be used in combination with this are the medium-sized anthers 1.8 - 2.5 mm long and relatively long rectangular calyx lobes with rounded apices. Solanum nigrescens has more regularly spaced flowers, occasionally branched inflorescences with more than one flower per branch and is more common along the Gulf Coast but distinguishing the two species without locality information can be difficult. The seeds of S. interius are much larger than those of S. nigrescens (1.8 - 2 mm versus 1.2 - 1.5 mm long) or any other of the diploid morelloids occurring in the area. Stone cell number can also be used as a distinguishing character; S. interius has 2 - 4 stone cells in each berry while S. nigrescens has more than 6 and often as many as 12. Solanum interius is sympatric with S. emulans and can be distinguished from that species in its longer anthers (1.8 - 2.5 mm long versus 1.5 - 1.8 mm long), its rounded calyx lobe apices, and its larger berries (10 - 14 mm in diameter versus 6 - 8 mm in diameter) with larger seeds (1.8 - 2.0 mm long versus 1.6 - 1.8 mm long). The flowers of S. emulans are usually smaller than those of S. interius. Nee (on label of Nee 61337) says that living plants of the two species are quite distinct, and that S. interius is a perennial growing in the shade of single trees. In describing S. interius Rydberg (1905) cited a collection but no herbarium. We have selected the sheet in the herbarium (NY) where he worked (NY 00138953) that is annotated " Type " as the lectotype of this species following best practice as described in McNeill (2014).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
3E657D97531BAC49217F74BC90CC95A6.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
73FBEB8934E430622BCB9F5A6FAD33CE.taxon	materials_examined	Type. Venezuela. Aragua: Colonia Tovar, Sep 1847, J. W. K. Moritz 1643 (holotype: B, destroyed; lectotype, designated by D'Arcy 1974 a, pg. 737: P [P 00336967]; isolectotypes: B [destroyed, F. neg. 2669], BM [BM 000617678], F [v 0073325 F, acc. # 612111], HBG [HBG- 511459], K [K 000585559]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
73FBEB8934E430622BCB9F5A6FAD33CE.taxon	description	Description. Perennial herb, woody at the base, 0.7 - 2 m tall, perhaps occasionally annual or only persisting for a few years. Stems terete or angled with spinescent processes, often described as " viney ", arching and scrambling over other vegetation, often drying blackish grey; young stems densely pubescent with somewhat antrorse, simple uniseriate eglandular trichomes 0.5 - 1 mm long, the trichomes drying white, soon glabrescent; new growth densely white pubescent like the young stems, glabrescent; bark of older stems green to greenish brown. Sympodial units difoliate or unifoliate, the leaves not geminate. Leaves simple, occasionally with a few dentate teeth near the base, (2) 4 - 10 (12) cm long, (0.8) 1.8 - 4.5 (5.5) cm wide, elliptic to narrowly obovate, sometimes thick (described as succulent), but more often membranous; adaxial surfaces sparsely pubescent with simple 3 - 4 - celled uniseriate trichomes or almost glabrous, the trichomes denser on veins and midrib; abaxial surfaces sparsely pubescent to glabrous like the adaxial surfaces, but the trichomes denser along the veins; principal veins 5 - 7 pairs, drying paler abaxially; base abruptly attenuate along the petiole; margins entire to sparsely toothed near the base; apex acute to narrowly acute; petiole 0.5 - 2.5 cm, sparsely pubescent with antrorse simple uniseriate trichomes like those of the stems and leaves. Inflorescences internodal or very occasionally leaf-opposed, 0.7 - 4 cm long, unbranched, with 2 - 3 (7) flowers clustered in the distal part of the rhachis (sub-umbelliform), sparsely pubescent with simple uniseriate trichomes like those of the stems and leaves; peduncle 0.5 - 4 cm long; pedicels 1 - 1.3 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, tapering gradually and appearing relatively stout, often described as reddish purple or purple, spreading at anthesis, sparsely pubescent or glabrous, articulated at the base; pedicel scars tightly packed in the distal portion of the inflorescence, less than 0.5 mm apart or occasionally the lowermost scar to 2 mm apart. Buds broadly ellipsoid to subglobose, the corolla long-exserted from the calyx tube before anthesis. Flowers 5 - merous, perfect. Calyx tube 1 - 1.5 mm long, conical, the lobes 0.5 - 0.8 (1) mm long, 0.5 - 1 mm wide, broadly deltate with acute apices, sparsely pubescent with simple uniseriate trichomes like those of the pedicel or almost glabrous. Corolla 10 - 20 mm in diameter, white to lilac or tinged with lilac, the central portion yellowish green, stellate, lobed 1 / 2 to 2 / 3 of the way to the base, the lobes 4 - 6 mm long, 1.5 - 3 mm wide, triangular, reflexed or spreading at anthesis. Stamens equal; filament tube minute and barely visible, the free portion of the filaments 1 - 2 mm long, pubescent with tangled simple uniseriate trichomes adaxially; anthers (2.7) 3 - 4 mm long, 1 - 1.5 mm wide, ellipsoid, bright yellow, the surfaces smooth, poricidal at the tips, the pores elongating to slits with age. Ovary glabrous; style 5 - 6 mm long, densely pubescent with tangled simple uniseriate trichomes in the basal half where included in the anther cone, exserted from the anther cone; stigma capitate or minutely capitate, bright green, the surface densely papillate. Fruit a globose berry, 0.8 - 1 cm in diameter, green turning to black when ripe, or occasionally green when ripe (Nee & Whalen 16839), opaque, the pericarp thin, more or less shiny but not brilliantly so; fruiting pedicels 15 - 17 mm long, tapering from a base 0.7 - 1 mm in diameter to an apex 1.5 - 2 cm in diameter, somewhat woody, strongly deflexed (very occasionally appearing spreading due to herbarium specimen preparation), dropping with mature fruits or occasionally remaining on the inflorescence rhachis; fruiting calyx not accrescent, the tube 1 - 1.5 (2) cm long, appressed to the berry, the lobes 0.5 - 1 mm long, appressed or spreading at the tips. Seeds (10) 30 - 50 per berry, 1.2 - 1.5 mm long, 0.8 - 1 mm wide, tear-drop shaped, tan to reddish brown, the surfaces minutely pitted, the testal cells pentagonal, more elongate and rectangular near the hilum. Stone cells (2) 4 - 5 (6) per berry, 0.5 - 0.7 mm in diameter. Chromosome number: 2 n = 2 x = 24 (Heiser 1955, as S. leonii), 2 n = 6 x = 72 (Heiser 1963, as. S. paredesii).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
73FBEB8934E430622BCB9F5A6FAD33CE.taxon	distribution	Distribution. (Figure 26) Solanum macrotonum occurs from Guatemala to northern South America and in the Antilles on the islands of Hispaniola and Jamaica.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
73FBEB8934E430622BCB9F5A6FAD33CE.taxon	discussion	Discussion. Solanum macrotonum is broadly sympatric with S. nigrescens across its entire range, and in Mexico and northern Central America with S. douglasii. It is similar to them in having usually 4 to 5 stone cells per berry and black fruits that are more or less shiny. It can be distinguished from S. nigrescens in having longer anthers (to 4 mm rather than to 2.5 mm) and in having more robust, longer fruiting pedicels that are strongly deflexed. It differs from S. douglasii in having strictly ellipsoid anthers (versus the slightly tapering anthers of S. douglasii) on longer filaments, and similarly in the strongly deflexed fruiting pedicels. Many annotations in herbaria have been done based on elevation (see comments in Bohs 2015) so care must be taken with these determinations. Measurement of anthers is the best way to determine identifications unambiguously. In general, S. macrotonum does occupy slightly higher elevations than does S. nigrescens, and appears to be confined to cloud forests, but S. nigrescens has a wide elevation range and ecological tolerance. Solanum macrotonum is one of few morelloids with differing chromosome counts across its range. D'Arcy (1974 a) reported a chromosome number of " n = 36 " for S. macrotonum as a personal communication from J. M. Edmonds; the chromosome count in Edmonds (1972) is not new, and we presume it is a reference to the count (" numero de cromosomas - 36 "; Heiser 1963) given in the protologue of S. paredesii, which Edmonds (1972) placed in tentative synonymy with S. macrotonum. Some other chromosome vouchers of S. macrotonum at IND, however (e. g., Heiser 4854) are noted as having " n = 24 " on the label; Heiser (1963) did not cite these in the description of S. paredesii. Chromosome counts for S. leonii, here treated in synonymy with S. macrotonum, indicate it is diploid, with n = 12 (Heiser 1955). Chromosome number variation within a species is known in Solanum (e. g., in the potatoes, see Spooner et al. 2014), and sometimes occurs sporadically at the edges of species ranges. It will be important to assess this across the range of S. macrotonum, because we cannot find any morphological characteristic that distinguishes vouchers with different chromosome counts. Bitter (1912 b) cited a single specimen from B in the protologue of S. macrotonum that is no longer extant. The sheet at P (P 00336967) that has a label with all of the details cited in the protologue was chosen by D'Arcy (1974 a) as the lectotype of S. macrotonum. Solanum megalophyllum was described from cultivated specimens from the herbarium of Aylmer B. Lambert seen by Bitter (1913) in the herbarium in Vienna. Of the two specimens that are clearly duplicates of this collection, we have selected that with the more complete annotation in Bitter's hand as the lectotype (W- 1889 - 0291427). Heiser (1955, 1963) described both S. leonii and S. paredesii citing " IND " as type. Heiser 5001 (type of S. paredesii) is only represented by a single sheet, but there are two sheets of Heiser 3597, the type of S. leonii, labelled " sheet 1 " and " sheet 2 ". We interpret this as a two-sheet holotype (see Turland et al. 2018, Art. 8, Ex. 7). Sheet 2 (IND- 0136010) is better material with more leaves and flowers. It is possible that S. frutescens A. Braun & C. D. Bouche is an older name for this taxon; that name has never been used, however, and it has been proposed for suppression (see Knapp et al. 2018; Doubtful Species).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
73FBEB8934E430622BCB9F5A6FAD33CE.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C1FA30475967674FD05DAAACA79A0C59.taxon	materials_examined	Type. Mexico. Oaxaca: " Cordillera " [" aux bords des ruiseaux de la cordillera de Yavezia "], Nov-Apr 1848, H. Galeotti 1238 (lectotype, designated by D'Arcy 1974 a, pg. 737: P [P 00337261]; isolectotypes: BR [BR 000000825045, BR 0000008250483], W [acc. # 0022312, acc. # 1889 - 0291397]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C1FA30475967674FD05DAAACA79A0C59.taxon	description	Description. Perennial herbs to 3 m tall, sometimes epiphytic. Stems terete or more usually angled to ridged, green or sometimes tinged purplish green, usually lax and somewhat scrambling, glabrescent to sparsely pubescent with antrorse simple eglandular uniseriate trichomes to 1 mm long, these white when dry and usually somewhat curved, occasionally on older stems the trichome bases enlarged and forming spinescent processes; new growth more densely pubescent. Sympodial units difoliate, geminate or not, the leaves if paired of similar size and shape. Leaves simple, (1.5) 4 - 10.5 (15) cm long, (0.5) 2 - 5 (7.5) cm wide, elliptic to elliptic ovate, membranous; surfaces sparsely to moderately pubescent with simple eglandular uniseriate trichomes to 1 mm long, these denser on the veins and abaxially; principal veins 5 - 6 pairs; base abruptly attenuate, usually decurrent on the petiole; margins entire to sinuate or dentate, the teeth irregular and unevenly spaced, often larger in the basal half of the lamina; apex acute or occasionally acuminate; petiole 0.5 - 2 cm long, sparsely pubescent like the stems and leaves. Inflorescence 1 - 3.5 cm long, lateral and internodal, unbranched to occasionally forked, with (2) 5 - 10 flowers clustered at the tip (sub-umbelliform) or spaced along the rhachis (depending on inflorescence age), sparsely pubescent with antrorse simple eglandular trichomes like the stems; rhachis 0.3 - 1 cm long; peduncle 1 - 2.5 cm long, slender, spreading; pedicels 0.4 - 0.7 cm long, slender and threadlike, spreading at anthesis, ca. 1 mm in diameter at the base, ca. 0.5 mm in diameter at the apex, sparsely pubescent like the inflorescence axis. Buds ellipsoid with blunt tips, the corolla strongly exserted from the calyx tube long before anthesis. Flowers 5 - merous, all perfect. Calyx tube 1 - 1.2 mm, conical, the lobes 0.5 - 0.8 (1) mm long, 0.5 - 1 mm wide, broadly deltate to deltate, the apices acute or occasionally somewhat rounded. Corolla 8 - 10 mm in diameter, white or less often pale purple, with a green or yellow-green (very occasionally dark purple) central portion near the base of the lobes, stellate, lobed ca. 3 / 4 of the way to the base, the lobes 3 - 4 mm long, 1.5 - 2 mm wide, narrowly triangular, reflexed or spread ing, densely papillate abaxially, the papillae ca. 0.1 mm long, denser at the tips and margins. Stamens equal; filament tube minute; free portion of the filaments 0.5 - 2 mm long, densely pubescent adaxially with tangled simple trichomes; anthers 2 - 2.8 (3) mm long, 1 - 1.1 mm wide, yellow, ellipsoid or narrowly ellipsoid, sagittate at the base, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 3.5 - 5 mm long, usually somewhat curved, often exserted from the bud before anthesis, densely pubescent in the basal 2 / 3 (the portion inside the anther cone), exserted from the anther cone; stigma minutely capitate, the surface papillose. Fruit a globose berry, 6 - 8 mm in diameter, dull green to purplish black at maturity, opaque, the pericarp thin and usually matte but sometimes slightly shiny; fruiting pedicels 10 - 12 mm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, not markedly woody, spreading, dropping with mature fruits or occasionally remaining on the inflorescence rhachis; fruiting calyx not accrescent, tube ca. 1 mm long, the lobes 0.5 - 1.1 mm long, spreading and appressed to the berry, very occasionally somewhat reflexed. Seeds (5) 10 - 50 per berry, 1.2 - 1.5 mm long, 1 - 1.1 mm wide, tear-drop shaped, pale brown to yellow, the surfaces minutely pitted, the testal cells square or pentagonal in shape, becoming elongate and rectangular near the subapical hilum. Stone cells 4 - 13, mostly commonly 5 or 6, rather large ca. 0.5 mm in diameter. Chromosome number: 2 n = 2 x = 24 (Heiser 1955 as S. costaricense; Heiser et al. 1965 as S. amethystinum).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C1FA30475967674FD05DAAACA79A0C59.taxon	distribution	Distribution. (Figure 29) Solanum nigrescens is a widespread species ranging from the southeastern United States of America through Central America, northern South America, and the Caribbean; in the southeastern United States of America it is found along the Gulf Coast and slightly inland but does not extend to the Great Plains.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C1FA30475967674FD05DAAACA79A0C59.taxon	discussion	Discussion. Solanum nigrescens is one of the commonest and most widely distributed of all morelloid species in Central America of America and the Caribbean. It is very variable morphologically, perhaps due to its wide ecological tolerance and occurrence in many different habitats. It is sympatric or occurs parapatrically with S. americanum, S. douglasii (in Mexico), S. interius and S. pseudogracile. It may hybridize with S. americanum in the southeastern United States (see discussion under S. americanum). Distinguishing features of each of those taxa can be found in the discussions of those species. Solanum nigrescens is a perennial and has been reported to be epiphytic (D'Arcy 1974 a, b). Where it and S. americanum occur in sympatry, the matte berries with appressed to spreading calyx lobes of S. nigrescens are distinct from the shiny berries with strongly reflexed tiny calyx lobes of S. americanum; anther length also differs (0.7 - 1.5 mm in S. americanum versus 2 - 2.8 (3) mm in S. nigrescens). In central Mexico, were S. nigrescens and S. douglasii co-occur, anther length (3 - 4 mm in S. douglasii versus 1.8 - 2.5 mm in S. nigrescens) is a good distinguishing feature; in fruit, these two taxa can be almost impossible to tell apart. Nee (1993) considered S. nigrescens to occupy wetter forest types than does S. douglasii but did not have not enough material from Veracruz to make the distinction. We find that the two species occur in very similar habitats, but that S. nigrescens is a more Caribbean species on the eastern side of the Sierra Madre and around the Gulf of Mexico, while S. douglasii occurs along the Pacific coast and into central Mexico, but also does occur in the Chihuahuan Desert biome. Specimens from Quintana Roo identified as S. nigrum in Sousa and Cabrera (1983) are S. nigrescens (Cabrera 875, 1151). Like most of these morelloid species, it is very weedy and occupies a wide range of disturbed and undisturbed habitats. In the southeastern United States (e. g., Texas) the distributions of S. nigrescens and S. interius are very close if not interdigitating. Solanum nigrescens can be distinguished from S. interius in its smaller seeds, more numerous stone cells in the berry and usually acute calyx lobe apices. The unusual pedicel articulation of the basal flower (in the lower third of the pedicel) in the inflorescences of S. interius has not been seen in S. nigrescens. Solanum nigrescens also appears to occur in more mesic and coastal habitats than S. interius, which is a species of the Great Plains. Material identified as S. americanum by Manoko et al. (2007) represents specimens of S. nigrescens (see Saerkinen et al. 2018: 61). Bitter (1914) reported large numbers of stone cells in the berries of many of the names we consider synonyms of S. nigrescens. In general, S. nigrescens has more stone cells in its berries than does the similar S. douglasii, but these can be difficult to see as some of them are very tiny. Dunal (1852) cited several specimens in describing S. nodiflorum var. puberulum, all from the Candolle herbarium at G: " Carolina meridionali, Fraser ", " Florida, Mich. f. ", " Mexico circa Bejar, Berlandier 1904 " and " China, Staunton ". Of these, the Fraser collection is a mixed collection composed of two tiny fragments of S. nigrescens (G 00144217) and one larger fragment of Capsicum annuum L. (G 00144268), mounted on the same sheet is a tiny fragment of S. americanum attributed to Michaux filius (G 00144264), and the Staunton sheet (G 00144265) is of a plant of S. americanum. Edmonds inadvertently (see Prado et al. 2015) selected as the lectotype for this name the sheet of Berlandier 1904 held in G-DC (G 00144231) by citing it as " holotype "; this is fortunate because it is unambiguous, was cited by Dunal (1852), and has duplicated in several other herbaria. When Berlandier was collecting, southern Texas was part of Mexico, and Bejar was the name for what today is San Antonio, the capital of Bexar County. D'Arcy (1974 a) lectotypified Solanum caribaeum citing a specimen from Jamaica in " G-DC ex Kew ". In the protologue of S. caribaeum Dunal (1852) cited " In insulis Caribaeis, Jamaica, Guadalupa (ex h. DC) " suggesting that more than one specimen was consulted. In G-DC there is a single specimen with the label " S. caribearum Nob. 1835 " (G 00144199) that is the only element of unambiguous material we have seen, and we designate this sheet in a second stage lectotypification. It is clear that more than this single sheet was used in the preparing the description, G 00144199 has only fruits and the description has details of both flowers and fruits. Kuntze (1891) equated his S. nigrum var. amethystinum with " S. nigrum subsp. genuinum Sendtn. " a name not validly published (see Saerkinen et al. 2018) and distinguished it by its violet flowers. He did not explicitly cite specimens but did cite the locality " Costa Rica. Irazu ". We have selected the specimen in Kuntze's herbarium held at NY (NY 00688134) as the neotype for this name. Both S. prionopterum (Bitter 1912 a) and S. gollmeri (Bitter 1912 b) were described from material collected by J. Gollmer around Caracas, in two separate publications. It is likely that the seed collection grown in Berlin in 1859 that was used to describe S. gollmeri was derived from the same material collected in 1854 in Venezuela. The holotypes in B for both these names were destroyed; the photograph of the holotype of S. gollmeri (F neg. 2689) has a tiny leaf fragment attached, but no such material is associated with the photograph taken of the holotype of S. prionopterum (F neg. 2699). We designate this fragment (F- 621268) as the lectotype of S. gollmeri; it is not, however, original material for S. prionopterum, and we hope a duplicate of the Gollmer collection from 1854 used to describe S. prionopterum will eventually be found. Bitter (1913) described S. pruinosum var. phyllolophum from living material originally sent by David Fairchild of the USDA as No. 32065; he cited no specimens. In the Germplasm Resources Information Network of the USDA (USDA 2017) the germplasm accession PI- 32065 is recorded as " S. nigrum " collected by C. A. Purpus in Puebla, Mexico. Fairchild (1912) recorded the exact locality as " Esperanza, Puebla, 2,700 m [9,850 feet] ". We have not yet seen a collection made by Carl Purpus with this exact information, hence do not designate a neotype until we have searched more exhaustively for a collection with the correct locality. It may be Purpus only collected seeds, and not herbarium specimens. Bitter (1913) described S. subelineatum from living material original sent from the USDA as No. 32067 grown in the Bremen botanical garden; he cited no specimens. In the Germplasm Resources Information Network of the USDA (USDA 2017) the germplasm accession PI- 32067 is recorded as " S. nigrum " collected by C. A. Purpus in San Luis Potosi, Mexico. Fairchild (1913) recorded the exact locality as " Rascon, San Luis Potosi, 400 to 500 m [1,300 to 1,650 feet] ". As with S. pruinosum var. phyllolophum we refrain from lectotypifying this anticipating encountering a duplicate. In describing S. oligospermum Bitter (1913) cited two specimens of Pringle 4948 at " herb. Haussknecht.!, Turic.! " (today JE and Z). Edmonds (1972) inadvertently lectotypified this name with the Z sheet by stating " (Z holotype!) " (see Prado et al. 2015). We here specify the individual sheet (Z 000033841) in that herbarium that is the lectotype. There are many well-preserved duplicates of this collection (see synonymy), all with flowers and fruit. Solanum approximatum, S. durangoense, and S. purpuratum were described (Bitter 1913) from B sheets that are now destroyed. We have selected lectotypes for these names from the large number of extant duplicates, using the best-preserved sheets with both flowers and fruits (if possible). Many specimens of S. nigrescens from Venezuela in US were annotated as " S. jahnii Bitter " by C. V. Morton, a designation not validly published (nomen nudum) based on Jahn 588. That collection corresponds to S. interandinum Bitter, a taxon not known from Central America or Mexico. Heiser (1955) cited only IND as the type in the protologue of S. costaricense, two sheets in IND are labelled " Type ". The sheets are clearly labelled as " sheet 1 " and " sheet 2 " and we interpret them as a two-sheet holotype (see Turland et al. 2018, Art. 8, Ex. 7). IND- 1000068 is the better material, with flowers and complete vegetative material.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
C1FA30475967674FD05DAAACA79A0C59.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
9898A1C6A8CF2616CB85925928E5FFA2.taxon	materials_examined	Type. 3 " Habitat in Orbis totius cultis " [sheet marked with Θ, meaning central part of Asia = Middle East], Without collector s. n. (lectotype, designated by Henderson 1974, pg. 19: LINN [LINN 248 - 18]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
9898A1C6A8CF2616CB85925928E5FFA2.taxon	description	Description. Annual or short-lived perennial herbs to 1.0 m tall, branching 10 - 30 cm from the base. Stems terete to sharply angled and ridged, green, the ridges often spinescent, not markedly hollow; new growth sparsely to densely pubescent with simple, spreading, uniseriate 1 - 6 - celled trichomes 0.5 - 0.6 mm long, these eglandular and / or glandular; older stems glabrescent, the trichome bases persisting as pseudospines. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.8 - 7.2 (- 14.5) cm long, 2.5 - 5.0 (- 9.5) cm wide, broadly ovate, green; adaxial surface sparsely pubescent with spreading, simple, uniseriate trichomes like those on stem evenly scattered along veins and lamina; abaxial surface more densely pubescent along veins and sparsely along lamina with eglandular and / or glandular trichomes like those of the stems; major veins 5 - 7 pairs; base obtuse to truncate, somewhat attenuate; margins sinuate-dentate, especially in the lower 2 / 3, to occasionally entire or deeply toothed; apex acute; petioles 0.5 - 3.0 cm long, pubescent with simple uniseriate glandular and eglandular trichomes like those of the stems. Inflorescences 0.8 - 2.0 cm long, lateral, internodal, unbranched (occasionally forked), with (3 -) 4 - 10 flowers spaced along the rhachis, pubescent with spreading simple uniseriate trichomes like those on stem; peduncle 0.5 - 1.5 cm long, straight; pedicels 3 - 5 mm long, 0.2 - 0.3 mm in diameter at the base and 0.2 - 0.3 mm at the apex, spreading, articulated at the base; pedicel scars spaced 0.3 - 0.7 mm apart. Buds subglobose, the corolla approximately halfway exserted from the calyx before anthesis. Flowers 5 - merous, all perfect. Calyx tube 0.8 - 1.0 mm long, the lobes 0.5 - 0.8 mm long, 0.6 - 0.8 mm wide, triangular with acute or somewhat rounded apices, pubescent with spreading simple uniseriate eglandular and glandular trichomes like those of the pedicels. Corolla 10 - 12 mm in diameter, white with a yellow-green central portion near the base, stellate, lobed 1 / 2 to 2 / 3 of the way to the base, the lobes 4.0 - 5.0 mm long, 2.0 - 2.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate-pubescent abaxially with simple uniseriate eglandular trichomes. Stamens equal; filament tube very short to minute; free portion of the filaments 0.5 - 0.7 mm long, adaxially pubescent with spreading uniseriate simple trichomes; anthers 1.8 - 2.5 mm long, 0.8 - 1.0 mm wide, ellipsoid, very slightly wider at base, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5 - 3.5 mm long, densely pubescent with tangled 2 - 3 - celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted 0 - 1 mm beyond anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 6 - 10 mm in diameter, purple-black or green to yellowish green at maturity, opaque, the surface of the pericarp matte or slightly shiny; fruiting pedicels 10 - 12 mm long, 0.4 - 0.5 mm in diameter at the base, 1.0 - 1.1 mm in diameter at the apex, generally spreading to occasionally recurved, spaced 1.0 - 2.0 mm apart, dropping with mature fruits, usually not persistent but occasionally remaining on the rhachis; fruiting calyx not accrescent, tube 0.7 - 1.5 mm long, the lobes 1.0 - 2.0 mm long, spreading to reflexed in fruit. Seeds (15 -) 20 - 40 per berry, 1.8 - 2.0 mm long, 1.5 - 1.6 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent (North America and Europe) but usually 2 (- 8) per berry in other areas (Asia), ca. 0.5 mm in diameter, brown. Chromosome number: 2 n = 6 x = 72 (for vouchers and references see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
9898A1C6A8CF2616CB85925928E5FFA2.taxon	distribution	Distribution. (Figure 32) Solanum nigrum is native to Eurasia and has been sporadically introduced and locally naturalised in temperate North America. It is possible that populations from eastern and western areas have different origins (see below).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
9898A1C6A8CF2616CB85925928E5FFA2.taxon	discussion	Discussion. Solanum nigrum is probably introduced in temperate North America; populations on the eastern seaboard best match European populations in overall morphology. Populations from the western coast (e. g., British Colombia) are morphologically more similar to Eurasian plants and it is possible that they are the result either of introductions from that region over the Pacific or are relictual native plants that came across the Bering Straits during warm periods in the Quaternary (e. g., DeChaine 2008; Ickert-Bond et al. 2009). Molecular population genetic analysis of these populations has not been done but should shed some light on the status of S. nigrum in North America. Solanum nigrum can be distinguished from other North American species (e. g., S. americanum, S. douglasii, S. emulans, S. interius, S. nigrescens) in the character combination of thicker peduncles and pedicels, larger seeds and fruits lacking stone cells. It has longer anthers (2.5 - 3 mm) than S. emulans and S. americanum, both of which have tiny anthers ca. 1.5 mm long. Like S. nigrescens, it has inflorescences with the flowers spaced along the rhachis, but S. nigrescens has prominent stone cells in the berries and smaller seeds. Solanum interius has similarly large seeds but has fewer flowers per inflorescence and distinctive basal flower pedicel position (articulation above the join with the rhachis). Michael Nee (pers. comm.) has observed its spread and increase in the New York City area over the last decade; it is possible that more collections will be made throughout North America in the coming years. For typification details of the many synonyms of S. nigrum see Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
9898A1C6A8CF2616CB85925928E5FFA2.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E5C74AA121F6C0B12066940AE45E40D7.taxon	materials_examined	Type. 4 Chile. Sin. loc., 1829, E. F. Poeppig s. n. (lectotype, designated by Edmonds 1986, pg. 27: W [acc. # 0004151]; isolectotype: F [v 0073346 F, acc. # 875221]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E5C74AA121F6C0B12066940AE45E40D7.taxon	description	Description. Annual herbs to 20 cm tall, prostrate and spreading to 30 cm in diameter or more. Stems terete, green, not markedly hollow; new growth densely viscid-pubescent with translucent simple, uniseriate 2 - 8 (10) - celled spreading trichomes 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; 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, lateral, generally internodal but in new growth appearing leaf-opposed, unbranched, with 4 - 8 (- 10) flowers clustered at the tip (sub-umbelliform) or spread along a short rhachis, sparsely pubescent with spreading trichomes like those on stems and leaves; peduncle 0.6 - 1.3 cm long; pedicels 4 - 12 mm long, 0.1 - 0.2 mm in diameter at the base and 0.2 - 0.4 mm in diameter at the 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 edges 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, translucent, 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; fruit ing calyx accrescent, becoming papery in mature fruit, the tube ca. 3 mm long, the lobes 2.5 - 3.5 (- 4.0) mm long and 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 (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E5C74AA121F6C0B12066940AE45E40D7.taxon	distribution	Distribution. (Figure 35) Solanum nitidibaccatum has an amphitropical distribution in temperate South America and temperate western North America, including northern Baja California. A single collection (Hammel et al. 6964) is known from the high elevation regions of Chiriqui, Panama (D'Arcy 1987). Solanum nitidibaccatum has often been recorded as adventive in North America, but the large number of early herbarium collections far from ports of entry suggest it is native (see also S. triflorum) at least from the Rocky Mountains westwards.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E5C74AA121F6C0B12066940AE45E40D7.taxon	discussion	Discussion. Solanum nitidibaccatum is morphologically similar to and has been treated as S. sarrachoides in most previous treatments of North American morelloids (e. g., Schilling 1981; Schilling and Heiser 1979); it is also often identified as S. physalifolium Rusby. Edmonds (1986) showed that S. nitidibaccatum and S. sarrachoides were distinct morphologically, and phylogenetic results (Saerkinen et al. 2015 b) confirm this; molecular sequence data also show these two taxa are not closely related despite their overall similarity. Solanum nitidibaccatum has also sometimes been treated at subspecific rank within S. physalifolium, an Andean endemic (see Saerkinen and Knapp 2016), but the species are distinct although preliminary data suggest they are closely related (see Saerkinen and Knapp 2016). Solanum nitidibaccatum is usually thought to be native to the southeastern parts of South America, from which it has been introduced extensively to other parts of the world where it has become a prolific and successful weed of disturbed sites. The species is locally abundant throughout North America (Ogg et al. 1981) and is perhaps native there west of the Rockies (see Distribution above). Solanum nitidibaccatum can be distinguished from S. sarrachoides in its shorter, plumper anthers, the blackish purple markings in the centre of the corolla on the margins of the central star, and in its fruits that are shiny at maturity, marbled with white (not usually visible on herbarium sheets) and not completely enclosed in the accrescent calyx. In addition, the mature inflorescences of S. nitidibaccatum are always internodal while those of S. sarrachoides are usually leaf-opposed. Details of typification of the synonyms of S. nitidibaccatum can be found in Barboza et al. (2013) and Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
E5C74AA121F6C0B12066940AE45E40D7.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
271A31962CD6B2F79800D3331FB1BCAC.taxon	materials_examined	Type. Mexico. " Circa Mexico ", J. Berlandier 751 (holotype: G [G 00418346]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
271A31962CD6B2F79800D3331FB1BCAC.taxon	description	Description. Perennial herb, 0.7 - 1 m tall, perhaps occasionally annual or only persisting for a few years. Stems angled to winged, lacking spinescent processes, usually erect, but occasionally lax and somewhat scrambling; young stems densely to sparsely pubescent with glandular, simple uniseriate trichomes 0.5 - 2 mm long, the trichomes (2 -) 4 - 15 celled, the basal cells larger, the trichomes drying translucent; new growth densely glandular pubescent and sticky; bark of older stems greenish brown. Sympodial units difoliate, the leaves not geminate. Leaves simple, occasionally shallowly toothed, 2.5 - 6.5 cm long, 1.2 - 2.8 cm wide, elliptic to ovate, widest in the lower half, membranous; adaxial and abaxial surfaces evenly and densely glandular-pubescent with simple uniseriate trichomes to 2 mm long, these denser abaxially and along the veins; principal veins 4 - 6 pairs, drying paler than the lamina; base attenuate onto the petiole; margins entire to shallowly and irregularly toothed, the teeth mostly in the basal third of the blade, usually with minute glandular papillae with 2 - celled glandular tips that dry dark brown; apex acute to acuminate; petiole 0.5 - 2 cm, narrowly winged from the attenuate leaf base. Inflorescences 0.8 - 2.5 cm long, unbranched, internodal, with 3 - 6 flowers (usually ca. 4) clustered in the distal third or quarter (sub-umbelliform), densely glandular-pubescent like the stems and leaves; peduncle 0.8 - 2.5 cm long; pedicels 0.7 - 0.9 cm long at anthesis, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender and tapering, densely glandular-pubescent with short uniseriate trichomes and glandular papillae, with only a few trichomes to 2 mm long present, spreading at anthesis, articulated at the base; pedicels scars closely packed in the distal part of the inflorescence, with the lowermost ca. 1 mm distant from the rest. Buds globose to broadly ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers 5 - merous, all perfect. Calyx tube 1.5 - 2 mm long, conical to cylindrical, the lobes 0.5 - 1 mm long, 0.8 - 1 mm wide, deltate to triangular, the tips obtuse or rounded, densely glandular-pubescent like the pedicels with uniseriate trichomes and papillae. Corolla 10 - 15 mm in diameter, white or pale purple with a darker brownish purple central star, stellate, lobed 1 / 2 to 2 / 3 of the way to the base, the lobes 3.5 - 5 mm long, 2 - 3 mm wide, triangular, reflexed to spreading at anthesis, the abaxial surfaces densely papillate, the trichomes not glandular. Stamens equal; filament tube to 0.5 mm; free portion of the filaments 0.5 - 1 mm long, glabrous or with a few weak tangled simple uniseriate trichomes adaxially; anthers 2.5 - 3.5 mm long, 0.5 - 1 mm wide, ellipsoid, bright yellow, poricidal at the tips, the pores elongating to slits with age. Ovary conical, glabrous; style 4.5 - 6 mm long, sparsely pubescent with weak tangled trichomes to densely papillate in the lower part where included in the anther cone, only slightly (ca. 0.5 mm) exserted from the anther cone; stigma capitate, densely papillate. Fruit a globose berry, 0.5 - 1 cm in diameter, green to deep purple (red when ripe? Martinez 1211); opaque (mature fruits not seen on live plants but not markedly translucent when dry), the pericarp thin, matte; fruiting pedicels 6 - 9 mm long, enlarging from a base 0.6 - 1 mm in diameter to an apex 1 - 1.5 mm in diameter, not distinctly woody, spreading; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes 1.5 - 2 mm long, appressed to the berry, venation very apparent and thickened. Seeds 10 - 30 per berry, 1 - 1.5 mm long, 1 - 1.2 mm wide, tear-drop shaped, reddish gold, the surfaces minutely putted, the testal cells pentagonal. Stone cells 2 - 4 (- 6) per berry, 0.5 - 0.7 mm in diameter, pale cream. Chromosome number: not known.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
271A31962CD6B2F79800D3331FB1BCAC.taxon	distribution	Distribution. (Figure 38) Solanum pruinosum occurs from the state of Nuevo Leon in Mexico south to the state of Oaxaca, across the central Volcanic Belt (Rzedowski 1978).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
271A31962CD6B2F79800D3331FB1BCAC.taxon	discussion	Discussion. Solanum pruinosum differs from S. nigrescens and S. douglasii, with which it is sympatric, in its glandular pubescence. It is possible that these specimens represent isolated glandular populations of those two taxa, but in the absence of data showing this we elect to recognise these populations at the species level until further work across the range in Mexico is done. The three taxa share numerous stone cells (ca. 5 - 6, more in S. nigrescens) in the ripe berries, and subumbellate to somewhat " racemose " inflorescences. The flowers of S. pruinosum are intermediate in size between S. douglasii (15 - 20 mm in diameter) and S. nigrescens (8 - 10 mm). Label data from Martinez 1211 (MO) note the berries as " rojo " (red), but no other specimens have this data, so we suspect it is either a mistake, or an interpretation of purplish red. The locality on the type specimen of S. pruinosum is only given as " circa Mexico ", but is likely to have been collected in central Mexico; Berlandier was in Mexico City and vicinity from the time of his arrival in Mexico in 1826 until his journey north to the Tamaulipas-Texas borderlands in late 1827 (Lawson 2012). We have not been able to trace any duplicates of the type specimens of S. dasyadenium and var. uberius, both described from material held in Berlin (Bitter 1912 a) and subsequently destroyed. Until we better understand the range of variation in these taxa, and their relationship to S. douglasii and S. nigrescens, we prefer not to neotypify these names at present.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
271A31962CD6B2F79800D3331FB1BCAC.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
2AEAE1BF00E1CF00A4AAAD13533DE710.taxon	materials_examined	Type. United States of America. North Carolina: Onslow County, N of Surf City, North Carolina Hwy. 210, 16 Jul 1960, C. R. Bell 17061 (holotype: IND [IND- 0136007, acc. # 145606]; isotype: IND [IND- 0136008, acc. # 145605], NCU [NCU 00062742]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
2AEAE1BF00E1CF00A4AAAD13533DE710.taxon	description	Description. Subwoody annual herb to perennial shrub up to 1.0 m tall, branching at base. Stems terete or with minute spinescent processes, green-grey to straw colour, sparsely to moderately pubescent with simple, appressed, uniseriate eglandular 4 - 9 - celled trichomes, these ca. 0.8 mm long; new growth more densely pubescent. Sympodial units difoliate, not geminate. Leaves simple, (1.3) 1.8 - 8.3 (- 10.5) cm long, (0.6 -) 1.1 - 3.7 cm wide, ovate-lanceolate to narrowly ovate, slightly discolorous, green above and pale grey underneath; adaxial surface sparsely pubescent with appressed translucent, simple, uniseriate trichomes like those on stem, these denser along the veins; abaxial surface more densely pubescent like those of the upper surface evenly across lamina and veins; primary veins 3 - 5 (6) pairs; base attenuate to acute, slightly unequal; margins entire to occasionally shallowly sinuate dentate; apex acuminate to acute; petiole (0.7 -) 1.0 - 2.4 cm long, pubescent with simple uniseriate trichomes like those of the stems and leaves. Inflorescences 1.2 - 2.0 cm long, lateral, internodal, unbranched or rarely forked, then with rhachis 0.4 - 0.5 mm long, with 3 - 8 flowers spaced along the rhachis, sparsely pubescent with appressed simple uniseriate trichomes like those on stem; peduncle 1.2 - 1.8 cm long, straight; pedicels 5 - 8 mm long, 0.2 - 0.3 mm in diameter at the base and 0.5 - 0.6 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0 - 1 mm apart. Buds ellipsoid, corolla exserted from the calyx to 2 / 3 of its length. Flowers 5 - merous, all perfect. Calyx tube 1.0 - 1.5 mm long, the lobes 0.5 - 1.0 mm long, ca. 1 mm wide, broadly ovate to obovate with obtuse to shortly acute apices, sparsely pubescent with appressed hairs like those on stem but shorter. Corolla 10 - 12 mm in diameter, deeply stellate, white with a yellow-green central portion near the base, some with darker blackish-purple colouration around the central star, lobed 2 / 3 to 4 / 5 to the base, the lobes 4.0 - 5.0 mm long, 1.6 - 3.0 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially with simple uniseriate trichomes like those on stem and leaves but shorter. Stamens equal; filament tube minute; free portion of the filaments 0.6 - 1.0 mm long, adaxially pubescent with tangled uniseriate 4 - 6 - celled simple trichomes; anthers 2.2 - 2.6 mm long, 0.5 - 0.7 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 3.5 - 4.0 mm long, exserted up to (1.0 -) 2.5 mm beyond the anther cone, densely pubescent with 2 - 3 - celled simple uniseriate trichomes at the base; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, (4 -) 8 - 14 mm in diameter, dull purplish-black at maturity, opaque, the surface of the pericarp matte and somewhat glaucous; fruiting pedicels 7 - 10 mm long, 0.3 - 0.4 mm in diameter at the base, (0.6 -) 0.9 - 1.0 mm in diameter at the apex, deflexed, becoming woody, pedicels spaced (0) 0.5 - 3.0 mm apart, dropping with mature fruits; fruiting calyx not accrescent, the tube 1.0 - 1.5 mm long, the lobes 2.5 - 3.0 mm long, lobes reflexed in fruit. Seeds 20 - 50 (- 60) per berry, 1.1 - 1.3 mm long, 0.8 - 0.9 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 (very rarely 2). Chromosome number: 2 n = 2 x = 24 (Heiser et al. 1965; Heiser et al. 1979).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
2AEAE1BF00E1CF00A4AAAD13533DE710.taxon	distribution	Distribution. (Figure 41) Solanum pseudogracile is endemic to the southeastern United States of America from the Atlantic coast of the Carolinas to the Gulf Coast in Florida and Alabama. Although we have seen no collections yet, we expect this species to also occur in the Bahamas.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
2AEAE1BF00E1CF00A4AAAD13533DE710.taxon	discussion	Discussion. Solanum pseudogracile is a species of the eastern North American coastal plain, and usually occurs in coastal areas not far from the sea. It is very similar to the adventive S. chenopodioides and can be very difficult to distinguish from it. Solanum pseudogracile has longer rectangular to obovate calyx lobes that are rounded to acute at the apex and reflexed in fruit, while S. chenopodioides has short triangular calyx lobes that are acute at the apex and always appressed in fruit. In addition, S. pseudogracile has a longer style that extends (1) 2.0 - 2.5 mm beyond the anther cone, compared to S. chenopodioides where the style is exserted only to 1.5 mm beyond the anther cone. The species differs from S. nigrescens in lacking stone cells or rarely having 2, while S. nigrescens always has 4 - 13 stone cells per fruit. In the absence of fruit these two species can be very difficult to distinguish; they are widely sympatric along the Gulf Coast of the southern United States of America. Solanum pseudogracile may be merely a form of S. chenopodioides, with which it shares many characteristics such as appressed white pubescence and absence of stone cells, but further population level work using molecular and other field markers will need to be undertaken. Distinguishing features of these morelloid species often disappear in herbarium specimens (see D'Arcy 1974 b), making analysis using herbarium specimens difficult. Nee (pers. comm.) has seen plants he has identified as the two taxa growing together. The Florida plants identified as S. americanum var. baylisii by D'Arcy (1974 b) fit our concept here of S. pseudogracile, while the type of D'Arcy's variety is a plant of S. chenopodioides collected from New Zealand. Many of the plants identified as S. chenopodioides in the Florida Plant Atlas (http: // florida. plantatlas. usf. edu /) are most likely S. pseudogracile. In describing S. pseudogracile Heiser (1978) cited only IND. There are two sheets of Bell 17061 in IND, one annotated type. We select this sheet (IND- 0136007) as the lectotype for S. pseudogracile; the sheets are not numbered sheet 1 and sheet 2 as was done by Heiser for other species in this group (e. g., S. costaricense, S. leonii).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
2AEAE1BF00E1CF00A4AAAD13533DE710.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
6B2B520D7E0341CBD307E05FDA1448C0.taxon	materials_examined	Type. 5 South Africa. Eastern Cape: Graaff Reinet (" Graafeynet "), 3000 - 4000 ft, 1838, J. F. Drege 7864 b (lectotype designated by Saerkinen et al. 2018, pg. 134: G-DC [G 00144331]; isolectotypes: K [K 000414172], S [acc. # S-G- 5707]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
6B2B520D7E0341CBD307E05FDA1448C0.taxon	description	Description. Annual to perennial herbs to 0.6 m tall, often woody at the base. Stems terete or ridged, 0.3 - 0.6 cm in diameter, green to yellowish-brown, prostrate or erect, the lowermost lateral branches usually spreading, if stems ridged the ridges sometimes spinescent, not markedly hollow; new growth sparsely to densely pubescent with glandular and / or eglandular simple spreading uniseriate 1 - 5 (- 8) - celled trichomes 0.1 - 0.8 mm long; older stems glabrescent, straw coloured. Sympodial units difoliate, the leaves not geminate. Leaves simple, (0.5 -) 1.5 - 7.5 cm long, 1.5 - 5.5 cm wide, rhomboidal to lanceolate, slightly discolorous; adaxial surface green sparsely to densely pubescent with simple uniseriate trichomes like those on stem evenly spread along lamina and veins; abaxial surface slightly paler, more densely pubescent along veins and lamina; major veins 3 - 7 pairs, pairs not strictly opposite, not prominent; base truncate then abruptly attenuate along the petiole; margins shallowly toothed, the teeth rounded; apex acute, the tip sometimes rounded; petioles (0.5 -) 1.5 - 3.5 cm long, sparsely to densely pubescent with simple uniseriate trichomes like those of the stems. Inflorescences 1.8 - 3.0 cm long, internodal, unbranched, with 3 - 7 flowers clustered towards the tip of the rhachis (sub-umbelliform), sparsely to densely pubescent with glandular and / or eglandular simple uniseriate trichomes like those on stems; peduncle 1.5 - 3.5 cm long, erect, green; pedicels 1.0 - 1.5 cm long, 0.3 - 0.6 mm in diameter at the base, 0.4 - 0.6 mm in diameter at the apex, recurving but not fully reflexed, pubescent like the peduncle, becoming woody, green or yellow-brown, articulated at the base; pedicel scars spaced 0 - 0.5 mm apart. Buds globose, the corolla 1 / 3 exserted from the calyx before anthesis. Flowers 5 - merous, all perfect. Calyx tube 1.0 - 1.7 mm long, campanulate, the lobes equal, 1.0 - 1.5 mm long, less than 1 mm wide, oblong with rounded tips, green, sparsely pubescent with simple uniseriate trichomes like of the inflorescence. Corolla 11 - 16 mm in diameter, white, with a yellow basal star, stellate, lobed to 1 / 2 - 2 / 3 towards the base, the lobes 5.0 - 6.0 mm long, 2.5 - 2.7 mm wide, spreading to reflexed, densely papillate-pubescent abaxially with simple uniseriate trichomes, these denser on tips and margins. Stamens equal; filament tube minute; free portion of the filaments 1.2 - 1.5 mm long, glabrous or adaxially pubescent with tangled 6 - 8 - celled simple uniseriate trichomes; anthers 1.3 - 1.8 (- 2.0) mm long, 1.0 - 1.5 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying, the connective becoming brownish in dry material. Ovary rounded, glabrous; style 1.9 - 2.2 mm long, slightly curved, pubescent with simple uniseriate trichomes 0.2 - 0.5 mm long in the basal 1 / 3 where included in the anther cone, exserted 0.5 - 1.5 mm beyond anther cone; stigma capitate, the surface minutely papillate. Fruit a globose to ellipsoid berry, 6 - 10 mm in diameter, purple-black at maturity, opaque, the pericarp thin, matte with a glaucous cast; fruiting pedicels 10 - 15 mm long, 0.4 - 0.6 mm in diameter at the base, 1.0 - 1.2 mm in diameter at the apex, becoming woody, recurving to deflexed, pale green to yellow-brown, persistent, spaced 0 - 0.5 mm apart, not falling with the fruit, remaining on the plant and persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.0 - 1.5 mm long, the lobes 1.5 - 2.0 mm long, strongly reflexed. Seeds (5 -) 12 - 35 per berry, 1.3 - 1.5 mm long, 1.6 - 1.8 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow to brown, the surfaces minutely pitted, the testal cells rectangular to pentagonal in outline. Stone cells absent. Chromosome number: 2 n = 4 x = 48 (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
6B2B520D7E0341CBD307E05FDA1448C0.taxon	distribution	Distribution. (Figure 44) Solanum retroflexum is native to southern Africa but introduced to North America as a garden plant. In North America it is mostly cultivated.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
6B2B520D7E0341CBD307E05FDA1448C0.taxon	discussion	Discussion. In its native range S. retroflexum is a species that shows great variation in its indumentum, the trichomes varying from eglandular to glandular and the leaves from nearly glabrous to densely pubescent. In the geographic region treated here, it has only been collected sporadically from cultivation and appears not to escape or naturalise. The species can be distinguished from other morelloids in North America based on a character combination of inflorescences with 1 - 4 flowers, filaments 1.2 - 1.5 mm long, strongly reflexed calyx lobes in fruit, and matte purple berries that lack stone cells and drop without the pedicels. Solanum americanum has similar small anthers and persistent pedicels, but the berries are very shiny and contain stone cells. Solanum retroflexum is a tetraploid of uncertain parentage (see discussion in Saerkinen et al. 2018). The berries are used in a local jam industry in South Africa (Viljoen 2011) and this is the species introduced as the " wonderberry " by Luther Burbank in the 1930 s (Heiser 1969). The story of the mystery surrounding the identity of Burbank's " wonderberry " is told in detail in Heiser (1969); it was variously considered a fraud or a case of mistaken identity. Burbank contended he had created the " wonderberry " through hybridisation of " S. guineense " (= S. scabrum) and " S. villosum " (probably = S. nitidibaccatum). Various correspondents (see Heiser 1969) suggested it was actually the " garden huckleberry " (S. scabrum) or one of the native black nightshades from North America. Stebbins and Paddock (1949) suggested that the true identity of the " wonderberry " was S. nigrum, a species occasionally found in agricultural fields in the western United States. Examination of specimens grown from the original seeds sold by John Childs, the sole distributor of Burbank's " wonderberry ", have shown that these were plants of S. retroflexum, but how Burbank came to grow them is still not known. Typification details for the synonyms of S. retroflexum can be found in Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
6B2B520D7E0341CBD307E05FDA1448C0.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
EB67601B0DE2A923DD752B0EE685FEBD.taxon	materials_examined	Type. 6 Brazil. " Brasilia australis ", F. Sellow s. n. (lectotype, designated by Edmonds 1986, pg. 16: P [P 00371162]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
EB67601B0DE2A923DD752B0EE685FEBD.taxon	description	Description. Annual herbs to 70 cm tall, usually smaller (but very rarely to 1 m), spreading and decumbent with age. Stems terete, green, generally erect, branching and later spreading, not markedly hollow; new growth densely viscid-pubescent with simple, uniseriate, spreading trichomes with a glandular apical cell, the trichomes of two lengths, 1 - 4 - celled trichomes to 0.5 mm long and 5 - 14 - celled trichomes to 2.0 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.0 - 7.5 cm long, 3.0 - 6.0 cm wide, broadly ovate; adaxial and abaxial surfaces sparsely to densely pubescent with spreading, simple, uniseriate glandular trichomes like those of the stem, evenly distributed on lamina and veins; major veins 3 - 4 pairs; base truncate to cordate, sometimes asymmetric; margins entire or regularly sinuate-dentate; apex acute; petioles 0.5 - 3.2 cm long, sparsely pubescent with trichomes like those of the stem and leaves. Inflorescences 0.7 - 1.7 cm long, lateral, usually leaf-opposed but occasionally internodal (always very near the node), unbranched, with 2 - 5 (6 - 7) flowers clustered at the tip (sub-umbelliform), sparsely pubescent with spreading trichomes like those of the stems; peduncle 0.7 - 1.0 cm long; pedicels 5 - 7 mm long, 0.1 - 0.2 mm in diameter at the base, 0.3 - 0.4 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0 (- 1) mm apart. Buds globose, the corolla only slightly exserted from the calyx tube before anthesis, almost completely included within the calyx lobes and only the tip of the corolla showing. Flowers 5 - merous, all perfect. Calyx tube 0.5 - 1.0 mm long, the lobes 1.5 - 2.0 mm long, 1.3 - 1.5 mm wide, lanceolate to narrowly ovate with acute apices, sparsely pubescent with 1 - 4 - celled spreading glandular trichomes like those on the pedicels but shorter. Corolla 5 - 8 mm in diameter, white with a yellow-green central eye, pentagonal-stellate, lobed 1 / 2 - 1 / 3 of the way to the base, the lobes 3.0 - 4.5 mm long, 5.0 - 7.0 mm wide, spreading at anthesis, sparsely papillate-pubescent abaxially with glandular 1 - 4 - celled simple uniseriate trichomes and eglandular papillae, these denser along margins, tips and midvein. Stamens equal; filament tube minute; free portion of the filaments 1.0 - 1.5 mm long, adaxially sparsely pubescent with tangled uniseriate 4 - 6 - celled simple trichomes; anthers 1.2 - 2.0 mm long, 0.4 - 0.8 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 3.0 - 3.5 mm long, densely pubescent with 2 - 3 - celled simple uniseriate trichomes in the lower 1 / 2 - 2 / 3 where included in the anther cone, not usually exserted beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 6 - 9 mm in diameter, green-brownish grey at maturity, opaque, the surface of the pericarp usually matte; fruiting pedicels 5 - 9 mm long, 0.2 - 0.3 mm in diameter at the base, spaced 0 - 1 mm apart, reflexed, dropping with mature fruits, not persistent; fruiting calyx accrescent, becoming papery in mature fruit, the tube 3 - 4 mm long, the lobes 5.5 - 8.0 mm long and 3.5 - 4.0 mm wide, the tips slightly reflexed or spreading. Seeds (23 -) 59 - 69 (- 93) per berry, 1.3 - 1.7 mm long, 1.0 - 1.5 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 4 - 6 per berry, (0.5) 0.8 - 1 mm in diameter. Chromosome number: 2 n = 2 x = 24 (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
EB67601B0DE2A923DD752B0EE685FEBD.taxon	distribution	Distribution. (Figure 47) Solanum sarrachoides is native to southern South America, and is sporadically introduced to North America, where it is much less common than the morphologically similar S. nitidibaccatum.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
EB67601B0DE2A923DD752B0EE685FEBD.taxon	discussion	Discussion. Many accounts of North American black nightshades have treated as Solanum sarrachoides the species whose correct name is S. nitidibaccatum (e. g., Stebbins and Paddock 1949; Schilling and Heiser 1979; Schilling 1981). Records of S. sarrachoides in the North American literature should therefore be dealt with care due to common misidentification of voucher material. The two taxa can be distinguished based using the following suite of characters: S. sarrachoides has generally truncate leaf bases, leaf-opposed mature inflorescences that are umbellate to sub-umbellate with fewer flowers (2 - 5), shorter calyx lobes 1.0 - 1.4 mm long, and a corolla with yellow-green central eye. Solanum nitidibaccatum has cuneate leaf bases, usually internodal mature inflorescences that are racemose with more flowers (4 - 8), longer calyx lobes 1.8 - 2.5 mm long, and corolla with black-purple edged central eye. The accrescent calyx almost completely encloses the matte-surfaced mature berry in S. sarrachoides, while the shiny, marbled berry of S. nitidibaccatum is always ca. halfway exserted from the calyx lobes. Solanum sarrachoides usually has more stone cells in each berry (4 - 6) than does S. nitidibaccatum (1 - 2, or absent). Though morphologically very similar, preliminary data from both nuclear and plastid DNA sequences suggests the two species are not closely related (T. Saerkinen, prelim. data). Typification details of the synonyms of S. sarrachoides can be found in Barboza et al. (2013) and Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
EB67601B0DE2A923DD752B0EE685FEBD.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
4E832D4F6F433FB0D24F99DD3957D2AB.taxon	materials_examined	Type. 7 Cultivated in Chelsea Physic Garden, said in protologue to " grow naturally in North America ", Herb. Miller s. n. (lectotype, designated by Henderson 1974, pg. 61 [as type]: BM [BM 000847083]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
4E832D4F6F433FB0D24F99DD3957D2AB.taxon	description	Description. Annual or short-lived perennial herbs to 1.5 m tall, often woody at the base. Stems terete, ridged, or winged, green to purple, erect or ascending, if ridged or winged the stems later spinescent, usually somewhat hollow; new growth puberulent with simple spreading uniseriate 2 - 8 - celled eglandular trichomes 0.3 - 0.8 mm long; older stems glabrescent, with or without prominent pseudospines. Sympodial units difoliate, the leaves usually not geminate, but if leaves paired then one is usually smaller. Leaves simple, 4 - 15 (20) cm long, 3 - 10 (16) cm wide, broadly ovate to elliptic, very variable in size depending on cultivars and growth conditions, green to dark green above to somewhat purple coloured, slightly paler; adaxial and abaxial surfaces glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem mainly along veins and scattered along lamina; major veins 3 - 6 (- 8) pairs, paler green or often purple tinged; base abruptly acute or truncate, narrowly winged onto the petiole; margins entire or rarely shallowly sinuate; apex rounded to acute; petioles 1 - 5 (8) cm long, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the stem. Inflorescences 1 - 2 (- 4) cm long, internodal, unbranched, forked or many times branched (in cultivars), with 4 - 10 (30 +) flowers clustered towards the tips (sub-umbelliform) or spread along the rhachis, glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem; peduncle 1 - 5 (- 8) cm long, erect and thick, much thickened at the apex, subwoody, green or purple-tinged; pedicels 0.4 - 1 cm long, 0.3 - 0.5 mm in diameter at the base, 0.75 - 0.9 mm in diameter at the apex and abruptly expanding to the calyx tube, stout, erect and / or spreading, green or purple-tinged, glabrous or minutely pubescent like the peduncle, articulated at the base; pedicel scars tightly clustered near the tip of the rhachis, spaced 0 - 2 mm apart, sometimes with short stumps ca. 0.5 - 1.0 mm long. Buds globose to subglobose, the corolla exserted 1 / 2 - 1 / 3 from the calyx tube before anthesis. Flowers 5 - merous or occasionally fasciate and 6 - 7 - merous in cultivars, all perfect. Calyx tube 0.9 - 1.1 mm long, abruptly cup-shaped with a broad base, the lobes slightly unequal, 0.9 - 1.5 mm long, 0.5 - 1.5 mm wide, broadly deltate with a rounded tip, green or purple-tinged, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the pedicels, the margins often drying scarious and white. Corolla 7 - 12 mm in diameter, white, purple-tinged or occasionally lilac to dark purple, with a yellow basal star, stellate, lobed ca. 1 / 2 of the way to the base, the lobes 2.5 - 4 mm long, 1.5 - 3 mm wide, spreading or reflexed, densely papillate on tips and margins. Stamens equal; filament tube very short, to 0.1 mm long; free portion of the filaments 0.5 - 0.8 mm long, glabrous or pubescent with tangled uniseriate simple trichomes; anthers 2 - 3 mm long, ellipsoid or slightly tapering towards the tips, yellow, orange or brown, poricidal at the tips, the pores lengthening to slits with age and drying, the connective often becoming brownish black in dry specimens. Ovary rounded, glabrous; style 2.5 - 5 mm long, densely pubescent with simple uniseriate trichomes 0.2 - 0.5 mm long in the basal 1 / 2 where included in the anther cone, exserted 0 - 1.5 mm beyond the anther cone; stigma capitate, the surface minutely papillate. Fruit a globose to slightly flattened berry, 10 - 20 mm in diameter, purplish black at maturity, opaque, the pericarp thick, shiny; fruiting pedicels 7 - 15 (20) mm long, 0.5 - 1 mm in diameter at the base, 1.1 - 1.5 mm in diam eter at the apex, stout, erect and spreading, purple or brown, usually not falling with the fruit, remaining on the plant and often persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.5 - 2 mm long, usually tearing unevenly, the lobes 2 - 3 mm long, usually with thicker white margins in dry material, appressed or spreading to slightly reflexed. Seeds (20 -) 100 - 150 per berry, 2 - 2.8 mm long, 1.5 - 1.8 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow-brown or purple, the surfaces minutely pitted, thin and the embryo clearly visible, the testal cells rectangular to pentagonal in outline. Stone cells absent. Chromosome number: 2 n = 6 x = 72 (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
4E832D4F6F433FB0D24F99DD3957D2AB.taxon	distribution	Distribution. (Figure 50) Solanum scabrum is native to tropical Africa; introduced worldwide as a cultivated plant.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
4E832D4F6F433FB0D24F99DD3957D2AB.taxon	discussion	Discussion. Solanum scabrum is a species known only from cultivation in North and Central America and the Caribbean. It is the mostly commonly cultivated morelloid species in Africa, and there is used from both its leaves (eaten as spinach) and its fruits. Specimens of S. scabrum occasionally have been collected from areas where enslaved people were brought from western Africa (e. g., Bahia, Brazil), so it is possible it could occur in especially the Caribbean. Solanum scabrum can be distinguished from the somewhat similar S. americanum by the larger anthers (2.5 - 3.0 mm long versus 0.8 - 1.5 mm long) that usually dry a dirty brownish tan. In both these species, as well as S. retroflexum, the berries drop off without the pedicels at maturity, and lack stone cells except in some populations of S. americanum where up to 4 stone cells have been observed (other populations lacking stone cells completely). Both S. scabrum and S. americanum have purple-black, shiny berries, while S. retroflexum has matte, waxy looking purple berries (with a bloom like blueberries, Heiser 1969). Material seen from North America represents only a fraction of the diversity of S. scabrum across its native range in Africa and is largely composed of specimens of large berried cultivars with simple inflorescences. The cultivated plants are sold in the garden trade in United States of America under the names of ' garden huckleberry'. The origin and identity of this garden plant gained huge interest in the 1960 ' s (Soria and Heiser 1959, 1961). Typification details for the synonyms of S. scabrum, and a complete discussion of its morphological variability in its native range can be found in Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
4E832D4F6F433FB0D24F99DD3957D2AB.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
90ABC0F28AD8E67E2C30154DCC95B8B1.taxon	materials_examined	Type. 8 United States of America. North Dakota: nr. Fort Mandan, Anon. [Lewis & Clark] s. n. (lectotype, designated by Barboza et al. 2013, pg. 260: PH [PH 00030496]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
90ABC0F28AD8E67E2C30154DCC95B8B1.taxon	description	Description. Annual herbs to 40 cm tall, much branched at the base, to 70 cm in diameter. Stems terete, green, decumbent and prostrate, forming adventitious roots at the nodes, not markedly hollow; new growth glabrous to sparsely pubescent with eglandular simple, uniseriate (3 -) 4 - 10 - celled spreading trichomes 0.5 - 2.0 mm long, occasionally with a few glandular trichomes with a 1 - many-celled apical gland; older stems glabrescent. Sympodial units difoliate or trifoliate, the leaves not geminate. Leaves simple and shallowly lobed to deeply pinnatifid, (0.5 -) 2.0 - 4.0 (- 5.0) cm long, 0.2 - 2.9 cm wide, narrowly elliptic to oblong or ovate-elliptic, fleshy in texture, green to dark green; adaxial surface glabrous to sparsely pubescent with simple, uniseriate trichomes like those on stem, scattered along lamina and more densely along the veins; abaxial surface more densely pubescent on veins and lamina; major veins 3 - 6 pairs, not clearly evident abaxially; base cuneate, decurrent on the petiole; sinuate-lobate to deeply pinnatifid to near-pinnate, with 3 - 6 linear to triangular pairs of lobes; apex acute; petioles (0.5 -) 1.0 - 2.0 (- 2.4) cm long, pubescent with simple uniseriate trichomes like those of the stems. Inflorescences 1.0 - 2.0 cm long, internodal, unbranched, with 1 - 5 (- 6) flowers clustered near the tips (sub-umbelliform), glabrous to sparsely pubescent with spreading trichomes like those of the stems; peduncle 0.8 - 3.5 cm long, often with apical leafy " bracteoles " (small, leaf-like structures amongst the pedicels); pedicels 3 - 12 mm long, 0.4 - 0.5 mm in diameter at the base and 0.4 - 0.5 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced 0 (- 0.5) mm apart. Buds narrowly ellipsoid or occasionally narrowly ovoid, the corolla exserted 1 / 5 - 2 / 5 from the calyx tube before anthesis. Flowers 5 - merous, all perfect. Calyx tube 1.0 - 1.5 mm long, conical, the lobes 2.5 - 3.5 (- 7.0) mm long, 0.8 - 1.0 (- 4.0) mm wide, triangular-oblong with acute apices, densely pubescent with simple, uniseriate eglandular trichomes like those of the stem. Corolla 10 - 14 mm in diameter, white to lilac with a yellow-green central eye with black-purple coloration at the base, deeply stellate, lobed 1 / 2 - 3 / 4 of the way to the base, the lobes 4.0 - 5.0 mm long, 1.8 - 2.2 mm wide, reflexed at anthesis, densely pubescent abaxially with short simple uniseriate eglandular trichomes like those on stems and leaves. Stamens equal; filament tube minute; free portion of the filaments 0.6 - 1.0 mm long, adaxially sparsely pubescent with tangled simple, uniseriate trichomes; anthers 2.8 - 3.1 (- 4) mm long, 0.4 - 0.5 mm wide, narrowly ellipsoid, pale yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5 - 3.5 mm long, densely pubescent with 2 - 3 - celled simple uniseriate trichomes to 1 / 2 from the base, not exserted beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 8 - 10 (- 20) mm in diameter, dark green at maturity, opaque, the surface of the pericarp usually shiny; fruiting pedicels 12 - 17 mm long, 0.5 - 1.0 mm in diameter at the base, 1.0 - 1.5 mm in diameter at the apex, spaced 0 - 0.5 (- 1.0) mm apart, reflexed and becoming woody, dropping with mature fruits, not persistent; fruiting calyx elongating in fruit, but not becoming papery nor covering the entire fruit, the tube 2.5 - 3.0 mm long, the lobes (4.0 -) 4.5 - 5.5 (- 8.0) mm long and 2.2 - 3.5 mm wide, strongly reflexed to spreading. Seeds 40 - 60 per berry, 2.0 - 2.5 mm long, 1.7 - 2.0 mm wide, subglobose, yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells 13 - 30, 1.0 - 1.5 mm in diameter. Chromosome number: 2 n = 2 x = 24 (South American populations only, see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
90ABC0F28AD8E67E2C30154DCC95B8B1.taxon	distribution	Distribution. (Figure 53) Solanum triflorum is native to the Americas with a disjunct (amphitropical) distribution between temperate South and North America. In North America it occurs in the United States of America from New Mexico and California north to Canada. The species has been introduced outside its native range in temperate areas of Europe, South Africa and Australia (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
90ABC0F28AD8E67E2C30154DCC95B8B1.taxon	discussion	Discussion. Solanum triflorum is a distinctive species with a prostrate habit, fleshy, usually pinnatifid, leaves, and deeply stellate flowers with long, thin anthers. The inflorescences usually have a small bracetole at the apex, and berry size varies from small (ca. 10 mm) to very large (ca. 20 mm), but usually a given plant has either small or large berries. Numerous stone cells are found in the berries, sometimes almost outnumbering seeds, and large berries can have as many as 30 stone cells. Pubescence of S. triflorum is quite variable (e. g., Subils 1983), and some plants have a few glandular trichomes, but for the most part plants from North America are either glabrous or very sparsely pubescent with spreading and often somewhat tangled simple trichomes. Solanum triflorum has a classic American Amphitropical Distribution (Gray and Hooker 1880; Raven 1963; AAD sensu Simpson et al. 2017), with populations oc curing in North and South America, but not between (see also S. nitidibaccatum). Because of its weedy nature, it is often assumed to be introduced to North America (e. g., https: // plants. usda. gov / core / profile? symbol = SOTR), but the amphitropical distribution pattern is found in other Solanaceae native to both regions such as Lycium L. (Levin et al. 2007), and groups of solanums such as the Carolinense (subsection Lathyrocarpum G. Don, Wahlert et al. 2015, as " section ") and Elaeagnifolium (Knapp et al. 2017) clades. Solanum elaeagnifolium Cav. (Elaeagnifolium clade, Knapp et al. 2017) has an almost identical amphitropical distribution (sensu Simpson et al. 2017) AAD, and is similarly weedy; it has also been assumed to be introduced. Distribution of these disjunct groups is more likely to be the result of long distance dispersal than of vicariance (Guilliams et al 2017), with dispersal after being eaten and passed through an animal's gut (endozoochory) being less common than disperal via attachment to an animal's fur or feathers (epizoochory) (Schenk and Saunders 2017); soft juicy berries make endozoochory more likely as a distribution mechanism, although there is no information on frugivores or fruit dispersal for S. triflorum. The distribution of S. triflorum in temperate areas, but also at higher elevations in deserts and into the more boreal regions of North America places it in the temperate AAD category of Simpson et al. (2017); annuals like S. triflorum predominate in this category. Amongst temperate AAD species the most common direction for distribution is from North to South America, but we suspect that like Verbenaceae (Frost et al. 2017) and Lycium (Levin et al. 2007), most Solanum disjunctions will have a South America to North America directionality. To date, only North American populations of S. triflorum have been included in molecular phylogenetic studies (Saerkinen et al. 2015 b). Typification details for the synonyms of S. triflorum can be found in Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
90ABC0F28AD8E67E2C30154DCC95B8B1.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
600B2207CAF577250B758CEC23C731CF.taxon	materials_examined	Type. 9 Cultivated in Chelsea Physic Garden from " Barbados where it is supposed to grow naturally ", Herb. Miller s. n. (lectotype, designated by Henderson 1974, pg. 54: BM [BM 000942575]).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
600B2207CAF577250B758CEC23C731CF.taxon	description	Description. Annual to short lived perennial herbs, much branched at base and usually bushy in form, up to 0.5 m tall. Stems terete to ridged, green to purple, ascending, not hollow; new growth densely pubescent with eglandular and / or glandular, simple, spreading, uniseriate 3 - 10 - celled trichomes 0.2 - 2.0 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5 - 5.0 (- 10.0) cm long, 0.7 - 2.5 (- 6.5) cm wide, broadly to narrowly ovate to elliptic, green; adaxial surfaces sparsely to densely pubescent with spreading, simple, uniseriate eglandular and / or glandular trichomes like those on stem evenly along veins and lamina; abaxial surfaces more densely pubescent on veins and lamina; major veins 4 - 6 pairs; base acute to truncate, short-attenuate, often asymmetric; margins sinuate-dentate to rarely entire; apex acute; petioles 0.5 - 3.0 (- 4.5) cm long, pubescent with simple uniseriate glandular and / or eglandular trichomes like those on stems. Inflorescences 0.4 - 2.0 cm long, lateral, internodal, unbranched, with (2 -) 3 - 5 (- 8) flowers clustered at the tips (sub-umbelliform, young inflorescences only) or more commonly spaced along the rhachis, pubescent with spreading simple glandular and / or eglandular uniseriate trichomes like those of the stems; peduncle 0.4 - 1.5 cm long, straight; pedicels 4 - 7 mm long, 0.2 - 0.3 mm in diameter at the base and 0.4 - 0.5 mm in diameter at the apex, spreading, articulated at the base; pedicel scars spaced 0 - 1.0 mm apart. Buds globose, the corolla exserted ca. 1 / 5 from the calyx before anthesis. Flowers 5 - merous, all perfect. Calyx tube 1.2 - 1.5 mm long, conical, the lobes 0.8 - 1.5 mm long, 0.5 - 0.8 mm wide, elliptic to triangular with obtuse thickened apices and paler (almost scarious) sinuses, pubescent with spreading simple uniseriate eglandular and / or glandular trichomes like those on stem. Corolla 8 - 15 (- 20) mm in diameter, white with a yellow-green central portion near the base and occasionally with purple stripes along lobe midveins abaxially, stellate, lobed 1 / 2 way to the base, the lobes 2.5 - 4.5 mm long, 2.0 - 3.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate-pubescent abaxially with simple uniseriate eglandular trichomes. Stamens equal; filament tube minute, pubescent with spreading uniseriate simple eglandular trichomes adaxially; free portion of the filaments 1.0 - 1.3 mm long, pubescent like the tube; anthers 1.8 - 2.2 (- 2.4) mm long, 0.5 - 0.7 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.8 - 3.5 (- 4.0) mm long, densely pubescent with 2 - 3 - celled simple uniseriate trichomes in the lower half, exserted 0 - 1 mm beyond anther cone; stigma capitate, the surface minutely papillate, green in live plants. Fruit an ellipsoid berry, usually somewhat longer than broad, 8.5 - 10 mm long, 8.0 - 9.5 mm wide, (red-) orange to yellow at maturity, translucent, the pericarp shiny; fruiting pedicels 8 - 14 mm long, 0.4 - 0.5 mm in diameter at the base, 0.7 - 1.5 mm in diameter at the apex, strongly reflexed, becoming woody, spaced 1.0 - 2.0 mm apart not falling with the fruit, remaining on the plant and always persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.0 - 1.5 mm long, the lobes 2.0 - 3.0 mm long and 1.0 - 2.0 mm wide, strongly reflexed in fruit. Seeds 20 - 40 per berry, 1.8 - 2.2 mm long, 1.5 - 1.7 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent, but occasionally 1 - 2 found in North African and Arabian material. Chromosome number: 2 n = 4 x = 48 (see Saerkinen et al. 2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
600B2207CAF577250B758CEC23C731CF.taxon	distribution	Distribution. (Figure 56) Solanum villosum is native to Europe where it is very common around the Mediterranean basin, the Arabian Peninsula to dry sub-tropical Asia, and eastern Africa (where it is cultivated for its fruit); the species is locally introduced but not persistent and not yet naturalised in North America.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
600B2207CAF577250B758CEC23C731CF.taxon	discussion	Discussion. Solanum villosum is an occasional non-native species to North America; the first known specimen was collected in 1899 by Curtiss from Pensacola, thought to have arrived in ships' ballast from Europe (D'Arcy 1974 b). The species has clearly not persisted or spread in North America despite several introductions (as seen in the few specimen records from widely distant sites). The many synonyms for S. villosum (see Saerkinen et al. 2018 for complete synonymy) reflect the broad morphological variation observed within the species, especially in its native range, where indumentum can range from nearly absent (the plants glabrous) to densely pubescent with eglandular and / or eglandular trichomes, and where cultivation and incipient domestication have resulted in a great range of inflorescence morphologies. Solanum villosum can be easily distinguished from all other morelloid species in North America by its orange-red berries that are often slightly elongated in shape and that lack stone cells. We have been unable to verify with specimens the citation of S. villosum from Texas (Correll and Johnston 1970); this is likely to be a misidentification. The Mexican species S. corymbosum, which also has orange-red berries, has branched inflorescences, minute flowers and berries with two large apical stone cells. All other species of the group in North America have rounded black or green berries. In bud and flower the calyx sinuses in S. villosum are much thinner than the lobes, and in dry specimens are white or transparent below the actual sinus, so that there appears to be a " window " in the calyx tube. In general aspect, plants are similar to those of S. retroflexum, but that species has matte purple berries with a distinct white glaucous bloom (also without stone cells, so herbarium specimens can be difficult to distinguish in the absence of good label information), and the leaves of S. retroflexum are more rhomboid in shape than those of S. villosum. Both of these introduced species are tetraploid and have glandular and eglandular forms in their native ranges. The typification details for the 45 heterotypic Old World synonyms of S. villosum can be found in Saerkinen et al. (2018).	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
600B2207CAF577250B758CEC23C731CF.taxon	materials_examined	Specimens examined. See Suppl. materials 1 and 3.	en	Knapp, Sandra, Barboza, Gloria E., Bohs, Lynn, Saerkinen, Tiina (2019): A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144, DOI: http://dx.doi.org/10.3897/phytokeys.123.31738, URL: http://dx.doi.org/10.3897/phytokeys.123.31738
