A revision of the " spiny solanums " of Tropical Asia (Solanum, the Leptostemonum Clade, Solanaceae) Author Aubriot, Xavier Universite Paris-Saclay, CNRS, AgroParisTech, Ecologie Systematique et Evolution, 91190, Gif-sur-Yvette, France & The Natural History Museum, Cromwell Road, London SW 7 5 BD, UK Author Knapp, Sandra https://orcid.org/0000-0001-7698-3945 The Natural History Museum, Cromwell Road, London SW 7 5 BD, UK s.knapp@nhm.ac.uk text PhytoKeys 2022 2022-06-01 198 1 270 http://dx.doi.org/10.3897/phytokeys.198.79514 journal article http://dx.doi.org/10.3897/phytokeys.198.79514 1314-2003-198-1 486F1F1B4F5854D2831AAA341B9A322C 26. Solanum melongena L., Sp. Pl. 1: 186. 1753. Fig. 42 Solanum mexianum Hill, Veg. Syst. 9: 39, pl. 39, fig. 1. 1765. Type. Cultivated "The Rugged Nightshade" (no specimens cited; lectotype, designated here: [illustration] Hill, The Vegetable System 9: Tab 39, f. 1. 1765). Melongena ovata Mill., Gard. Dict., ed. 8, no. 1. 1768. Type. " Melongena ( Ovata ) caule inermi herbaceo, foliis oblong-ovatis tomentosis integris, fructu ovato [ Melongena fructu Solanum oblongo violaceo Tourn. Inst. 151]" (no specimens cited; possibly described from live plants; neotype, designated here: BM [BM000942564]). Melongena teres Mill., Gard. Dict., ed. 8, no. 2. 1768, as " Tereta ". Type. " Melongena ( Tereta ) caule inermi herbaceo, foliis oblong-ovatis tomentose, fructu terete [ Melongena fructu tereti violaceo Tourn. Inst. 151]" (no specimens cited, no material found; probably described from live plants). Melongena incurva Mill., Gard. Dict., ed. 8, no. 3. 1768. Type. " Melongena ( Incurva ) caule inermi herbaceo, foliis oblongis sinuatis tomentosis, fructu incurvo [ Melongena fructu incurvo Tourn. Inst. 152]" (no specimens cited, no material found; probably described from live plants). Melongena spinosa Mill., Gard. Dict., ed. 8, no. 4. 1768. Type. " Melongena ( Spinosa ) Melongena spinosa , foliis sinuatis-lacinitis, fructu tereti, caule herbaceo [ Solanum pomiferum fructu spinoso J.B. 3. 619]" (no specimens cited, no material found; lectotype, designated here: [illustration] " Solanum pomiferum fructu spinoso" Bauhin, Historia plantarum 3: 619. 1651). Solanum zeylanicum Scop., Delic. Fl. Faun. Insubr. 1: 1. 1786. Type. Cultivated in Italy [Pavia] from seeds sent by D. Marsilius (no specimens cited, described from live plants; lectotype, designated here: [illustration] Scopoli, Deliciae Flora et Fauna Insubricae 1: tab. 1. 1786). Solanum album Lour., Fl. Cochinch. 129. 1790. Type. Indonesia. Malaku Islands: Amboina [Ambon Island] (no specimens cited; based on illustration; lectotype, designated here [cited as holotype by Hul and Dy Phon 2014 , pg. 30]: [illustration] "Trongum Solanum agreste spinosum" in Rumphius, Herb. Amboin. 5: 241, tab. 86, fig. 1. 1747). Solanum oviferum Nocca, Ann. Bot. (Usteri) 6: 61. 1793., nom. illeg. superfl. Type. Based on Solanum melongena L. (cited in synonymy) Solanum oviferum Salisb., Prodr. Stirp. Chap. Allerton 134. 1796, nom. illeg. superfl. Type. Based on Solanum melongena L. (cited in synonymy) Solanum esculentum Dunal, Hist. Nat. Solanum 208, tab. 3. 1813, nom. illeg. superfl. Type. Based on Solanum melongena L. (cited in synonymy with the comment [ Dunal 1852 ] "nomen Melongena no accepi, quia sub hoc nomine, species plures confusae suerunt"). Solanum ovigerum Dunal, Hist. Nat. Solanum 210. 1813, nom. illeg. superfl. Type. Based on Solanum melongena L. (cited in synonymy) Solanum pressum Dunal, Hist. Nat. Solanum 217. 1813. Type. Based on "Trongum pra rubrum" of Rumphius (no specimens cited; lectotype, designated here: [illustration] Rumphius, Herbarium Amboinense 5: tab. 86, f. 2. 1747). Solanum melanoxylon Link, Enum. Hort. Berol. Alt. 1: 188. 1821. Type. Cultivated in Berlin, Germany (no specimens cited; no original material found, likely destroyed). Solanum longum Roxb. ex Wall., Fl. Ind. (Carey & Wallich ed.) 2: 248. 1824. Type. India. West Bengal: Cultivated "HBC" [Hort. Bot. Calcutta], 1821, Without collector s.n. [Wallich cat. 2628D ] (lectotype, designated here: K-W [K001116666]). Solanum ovigerum Dunal var. insanum Blume, Bijdr. Fl. Ned. Ind. 13: 698. 1826. Type. Indonesia. "in hortis frequenter cultum/Terong Pangang" (no specimens cited; no original material found). Solanum pseudo-undatum Blume, Bijdr. Fl. Ned. Ind. 13: 699. 1826. Type. Indonesia. Java. "in locis cultis/nomen Terong Rangu" [ex protologue] Sin. loc., C.L. Blume s.n. (lectotype, designated here: L [L0003642]). Solanum pseudo-undatum Blume var. albiflorum Blume, Bijdr. Fl. Ned. Ind. 13: 699. 1826. Type. Indonesia. "in hortis colitur/Terong lelles" [ex protologue] (no specimens cited; no original material found). Solanum pseudo-undatum Blume var. atropurpurascens Blume, Bijdr. Fl. Ned. Ind. 13: 699. 1826. Type. Indonesia. "cum praecidentibus/Terong Kupa" [ex protologue] (no specimens cited; no original material found). Solanum pseudo-undatum Blume var. leucocarpon Blume, Bijdr. Fl. Ned. Ind. 13: 700. 1826. Type. Indonesia. "montis Salak" [ex protologue] (no specimens cited; no original material found). Solanum edule Schumach. & Thonn., Beskr. Guin. Pl. 125 [145]. 1827. Type. "Guinea" . Sin. loc., cultivated, P. Thonning 141 (lectotype, designated here: C [C10004584]). Solanum melongena L. var. esculentum (Dunal) Walp., Repert. Bot. Syst. (Walpers) 3: 81. 1844. Type. Based on Solanum esculentum Dunal. Solanum aethiopicum L. var. violaceum Dunal, Prodr. [A. P. de Candolle] 13(1): 351. 1852. Type. Based on Solanum aethiopicum sensu Lour., non L. Solanum album Lour. var. richardii Dunal, Prodr. [A. P. de Candolle] 13(1): 362. 1852. Type. Reunion . "Bisom. croix dans les lieux habites" , 1837, A. Richard 186 (holotype: P [00352546]). Solanum album Lour. var. rumphii Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852, nom. illeg. Type. Based on Solanum album Lour. (as "Trongum Solanum agreste "). Solanum edule Thonn. var. multifidum Dunal, Prodr. [A. P. de Candolle] 13(1): 357. 1852. Type. Cultivated "in hort. Tonelle Audiberti cultum" [Audibert nursery in Tonelle, near Tarascon, France], sin. dat., Without collector s.n. (holotype: AV [n.v.]). Solanum serpentinum Desf. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852. Type. Cultivated in Montpellier, France, 17-26 Aug 1838, R.L. Desfontaines s.n. (holotype: MPU [MPU854428]). Solanum esculentum Dunal var. aculeatum Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852. Type. Indonesia. Java: Sin. loc., sin. dat., H. Zollinger 702 (holotype: G-DC [G00131553]; isotypes: BM [BM000778111], K [K000788269]). Solanum esculentum Dunal var. inerme Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852. Type. Brazil. Bahia: sin. loc., sin dat., J.S. Blanchet 368 (lectotype, designated here: G [G00357998]). Solanum esculentum Dunal var. subinerme Dunal, Prodr. [A. P. de Candolle] 13(1): 355. 1852. Type. Brazil. Bahia: sin. loc., 1831, J.S. Blanchet 236 (lectotype, designated here: G-DC [G00131527]). Solanum lagenarium Dunal, Prodr. [A. P. de Candolle] 13(1): 368. 1852. Type. Cultivated "hort. Tonelle cultum" [Audibert nursery in Tonelle, near Tarascon, France], 1824, Anon. s.n. (holotype: AV [n.v.]). Solanum ovigerum Dunal [unranked] sinuatorepandum Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852. Type. Cuba. La Habana: "La Havane" [Havana], 1826, de la Ossa s.n. (lectotype, designated here: G-DC [G00131521]). Solanum ovigerum Dunal [unranked] subrepandum Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852. Type. Cultivated in Geneva, Switzerland "in tepidario DC.", sin. dat., Anon. s.n. (lectotype, designated here: G-DC [G00131523]). Solanum ovigerum Dunal var. violaceum Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852, as Solanum ovigerum [unranked] subrepandum var. Solanum ovigerum violaceum . Type. No specimens or locality cited; reference to Tournefort ("Tourn. inst. 152"). Solanum ovigerum Dunal var. oblongo-cylindricum Dunal, Prodr. [A. P. de Candolle] 13(1): 358. 1852, as Solanum ovigerum [unranked] subrepandum var. Solanum ovigerum oblongo-cylindricum . Type. Based on Solanum ovigerum var. insanum Blume. Solanum trongum Poir. var. sinuatopinnatifidum Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Based on Solanum trongum Poir. sensu Blume [Java. "in agestribus/karnadong", C. Blume s.n. (no specimens cited, based on S. trongum Poir. sensu Blume)]. Solanum requienii Dunal, Prodr. [A. P. de Candolle] 13(1): 363. 1852., as 'Requieni' . Type. Cultivated in France "hort. Tonelle ex Audibert fratr.", 1822, Anon. s.n. (holotype: AV [n.v.]). Solanum sativum Dunal, Prodr. [A. P. de Candolle] 13(1): 360. 1852, nom. illeg. superfl. Type. Based on Solanum pseudo-undatum Blume (cited in synonymy "Nomen Solanum pseudo - Solanum undatum mutavi, quonian semigraecum, semilatinem") Solanum sativum Dunal var. albiflorum (Blume) Dunal, Prodr. [A. P. de Candolle] 13(1): 360. 1852. Type. Based on Solanum pseudo-undatum Blume var. albiflorum Blume. Solanum sativum Dunal var. atropurpurascens (Blume) Dunal, Prodr. [A. P. de Candolle] 13(1): 360. 1852. Type. Based on Solanum pseudo-undatum Blume var. atropurpurascens Blume. Solanum sativum Dunal var. leucocarpon (Blume) Dunal, Prodr. [A. P. de Candolle] 13(1): 361. 1852. Type. Based on Solanum pseudo-undatum Blume var. leucocarpon Blume. Solanum melongena L. var. giganteum Alef., Landw. Fl. 136. 1866, as " gigantea ". Type. Cultivated (no specimens cited; no original material found). Solanum melongena L. var. leucoum Alef., Landw. Fl. 136. 1866. Type. Cultivated (no specimens cited; no original material found). Solanum melongena L. var. stenoides Alef., Landw. Fl. 136. 1866. Type. Cultivated (no specimens cited; no original material found). Solanum melongena L. var. stenoleucum Alef., Landw. Fl. 136. 1866, as " stenoleuca ". Type. Cultivated (no specimens cited; no original material found). Solanum melongenum St.-Lag., Ann. Soc. Bot. Lyon 7: 135. 1880, nom. illeg. superfl. Type. Based on Solanum melongena L. (cited in synonymy). Melongena esculenta (Dunal) Grecescu, Consp. Fl. Romaniei 423. 1898. Type. Based on Solanum esculentum Dunal, nom. illeg. Solanum melongena L. var. inerme (Dunal) Hiern, Cat. Afr. Pl. (Hiern) 1(3): 748. 1898. Type. Based on Solanum esculentum var. inerme Dunal Solanum melongena L. var. viride Dikii, Trudy Prikl. Bot. 88: 104. 1984, as " viridis ". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from Asia Minor. 4 Jul 1971, L. Studentsova s.n. [VIR cat. # 249 ] (holotype: WIR [n.v.]). Solanum melongena L. subsp. agreste Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as " Solanum agrestis ". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India, 7 Aug 1975, N. Frantskevich s.n. [VIR cat. # 713 ] (holotype: WIR [n.v.]). Solanum melongena L. var. angustum Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as " angusta ". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India "c.v. R.R.I.", 7 Aug 1975, N. Frantskevich s.n. [VIR cat. # 2333 ] (holotype: WIR [n.v.]). Solanum melongena L. var. cylindricum (Filov) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as " cylindrica ". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India, 7 Aug 1975, N. Frantskevich s.n. [VIR cat. # 116 ] (holotype: WIR [n.v.]). Solanum melongena L. var. racemiflorum Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as " racemiflora ". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India "c.v Purple Cluster", 27 Sep 1958, V. Barkovskaya s.n. [VIR cat. # 2515 ] (holotype: WIR [n.v.]). Solanum melongena L. var. racemosum (Filov) Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 105. 1984, as " racemosa ". Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from India, 4 Jul 1971, L. Studentsova s.n. [VIR cat. # 709 ] (holotype: WIR [n.v.]). Solanum melongena L. var. globosi Dikii, Trudy Prikl. Bot. Genet. Selek. 88: 106. 1984. Type. Russia. Cultivated at Maikop Experiment station of VIR (Maikop, Krasnodar Krai, Republic of Adygea), originally from Vietnam, 14 Jul 1971, L. Studentsova s.n. [VIR cat. # 2582 ] (holotype: WIR [n.v.]). Type . Cultivated in Uppsala , Sweden , Anonymous s.n. ( lectotype , designated by Schoenbeck-Temesy 1972 , pg. 70: LINN [acc. # 248.28]) . Description. Erect annual herb, 0.2-0.5 m, unarmed or more rarely prickly. Stems erect, terete, unarmed or occasionally prickly, moderately stellate-pubescent to glabrescent; prickles to 3 mm long, less than 0.5 mm wide at the base, straight, acicular, yellow-orange to dark brown, glabrous; pubescence a mixture of minute simple hairs and porrect-stellate trichomes, the stellate trichomes sessile to short-stalked, the stalks to 0.2 mm long, the rays 8-15, 0.3-0.7 mm long, the midpoints ca. same length as the rays or elongated to 1 mm; new growth moderately to densely stellate-pubescent, whitish green; bark of older stems glabrescent, green-brown to dark brown. Sympodial units difoliate, not geminate. Leaves simple, moderately lobed, the blades 10-23 cm long, 9-15 cm wide, 1.5-2 times longer than wide, ovate, chartaceous, drying discolorous, unarmed; adaxial surface yellow-green to red-brown, sparsely to moderately stellate-pubescent, occasionally glabrescent; abaxial surface glabrescent to moderately stellate-pubescent usually more pubescent than the adaxial surface, the trichomes porrect, sessile to short-stalked, the stalks to 0.2 mm long, the rays 5-8, 0.3-1 mm long, the midpoints ca. same length as the rays; major veins 4-7 pairs; base cordate to obtuse; margins lobed, the lobes 2(-3) on each side, 0.5-2 cm long, deltate, apically rounded, the sinuses extending 1/4-1/3 to the midrib; apex acute; petiole 1-10 cm long, 1/4-1/3(-2/3) of the leaf blade length, moderately stellate-pubescent to glabrescent, unarmed or prickly with 1-2 prickles. Inflorescences 6-15 cm long, apparently terminal or lateral, unbranched, with 1-8 flowers, 1-3 flowers open at any one time, moderately stellate-pubescent to glabrescent, with a mix of sessile and short-stalked stellate-porrect trichomes like those of the stems, unarmed; peduncle 0-80 mm long, unarmed; pedicels 0.8-3.5 cm long, 1-1.8 mm in diameter at the base, 1-2 mm in diameter at the apex, erect to pendent, unarmed or more rarely prickly with up to 5 prickles, moderately stellate-pubescent to glabrescent with porrect-stellate trichomes like those of the stems, articulated at the base; pedicel scars spaced 3-5 mm apart. Buds ovoid, the corolla ca. halfway exerted from the calyx before anthesis. Flowers 4-8-merous, heterostylous and the plants andromonoecious, with the lowermost flower long-styled and hermaphrodite, the distal flowers short-styled, staminate and usually somewhat smaller. Calyx with the tube 3-8 mm long in long-styled flowers, urceolate to deeply cup-shaped, 2-3 mm long in short-styled flowers, cup-shaped to elongate cup-shaped, the lobes in long-styled flowers 5-17 mm long 3-5 mm wide, in short-styled flowers 3.5-5 mm long, 2-2.5 mm wide, deltate to narrowly deltate, apically acute to long-acuminate, unarmed or more rarely prickly with up to 30 prickles, moderately stellate-pubescent, with porrect-stellate trichomes like those of the pedicels. Corolla 2.5-5 cm in diameter in long-styled flowers, 2.4-4 cm in diameter in short-styled flowers, white to mauve or purple, stellate, lobed 1/4-1/2 of the way to the base, the lobes 10-15 mm long, 8-13 mm wide in long-styled flowers, 5-8 mm long and 8-13 mm wide in short-styled flowers, broad-deltate, spreading at anthesis, not opening fully in long-styled flowers, sparsely stellate-pubescent abaxially, the trichomes porrect, sessile or stalked, the stalks to 0.2 mm, the rays 4-8, 0.2-0.7 mm long, the midpoints ca. same length as the rays. Stamens equal; anthers 5.5-7.5 mm long in long-styled flowers, 5.5-6 mm long in short-styled flowers, ca. 2 mm wide, connivent, tapering, yellow, glabrous, poricidal at the tips, the pores not elongating to slits with drying; filament tube 2-3 mm long, glabrous; free portion of the filaments 1.2-3 mm long, glabrous. Ovary conical, stellate-pubescent in upper part near the style base; style in long-styled flowers ca. 9 mm long, broad and straight, moderately stellate-pubescent in the lower half to 3/4, in short-styled flowers 2-3 mm long, pubescent at the base; stigma globose capitate to somewhat bilobed, bright green in live plants, the surface minutely papillate. Fruit a globose to ovoid, ellipsoid, or oblong to variously curved berry (many cultivars with a wide variety of fruit shapes), 1 (rarely> 1) per infructescence, 3-20 cm long, 3-7 cm wide, green, sometimes mottled or striped, white, pink, mauve, purple, or black when young, usually white or maroon at maturity, the pericarp smooth, glabrous; fruiting pedicels 2.5-8 cm long, 2-4 mm in diameter at the base, 5-10 mm in diameter at the apex, unarmed or sparsely prickly, woody, pendent and deflexed from the weight of the berry; fruiting calyx lobes elongating to 12-50 mm long, 1/4-1/3 the length of the mature fruit, often cup-shaped around the fruit in some cultivars, unarmed or sparsely prickly. Seeds usually aborted in cultivars. Chromosome number: n = 12 ( Doganlar et al. 2002 ; Barchi et al. 2019 ; Chapman 2019 ; Gramazio et al. 2019 ; Wei et al. 2020 ). Figure 42. Solanum melongena L. A herbarium specimen collected in India ( Mace s.n. , P00058105) B habit (field photograph, unvouchered, Thailand) C detail view of a hermaphroditic flower ( Meeboonya et al. RM 294 , Thailand) D detail view of an oblong and white fruit ( Meeboonya et al. RM 250 , Thailand) E detail view of a round green fruit marbled with white ( Meeboonya et al. RM 248 , Thailand). Photograph credits: A CC-BY, Museum national d'Histoire naturelle, Paris B-E X. Aubriot. Distribution. Solanum melongena is cultivated worldwide in tropical and subtropical areas (in the temperate zone under glass); the greatest diversity of landraces and cultivars is found in Asia (India, China and southeast Asia), with secondary centres in the Middle East and around the Mediterranean. The origin of Solanum melongena is in Asia, but the exact place of domestication in not clear (see Meyer et al. 2012 ; references in Knapp et al. 2013 ). We have not provided a distribution map as herbarium specimens do not accurately reflect the range of the species in cultivation. Ecology and habitat. Solanum melongena is only known from cultivation (see below). Common names and uses. China. qie ( Zhang et al. 1994 ), qie zi ( Lester and Hawkes 2001 ). India. brinjal [Hindi] ( Lester and Hawkes 2001 ); Bihar: baigan, bhanta ( Varma 1981 ); Karnataka: advibadani, doorla ( Singh 1988 ); Tamil Nadu: kathiri (Tamil, Matthew 1983 ); kaththiri ([Tamil] Henry et al. 1987 ). Japan. nasu ( Lester and Hawkes 2001 ). Indonesia. Java: enchung, terong ( Burkill 1935 ); Sumatra: tiung, chung ( Burkill 1935 ). Malaysia/Singapore. tĕrong (with many qualifiers indicating varieties, Burkill 1935 ). Solanum melongena is a worldwide food crop; after tomato ( S. lycopersicum L.) it is the most widely cultivated fruit crop in the Solanaceae ( Daunay and Janick 2007 ; Knapp et al. 2019b ). Recent work to improve eggplants in the face of climate change has seen increased interest in wild relatives and crosses with wild species across the spiny solanums (e.g., Garcia-Fortea et al. 2019 ). See section on Uses (pp. 18-20) for further details. Discussion of the many aspects of eggplant biology can be found in Chapman (2019) . Preliminary conservation status ( IUCN 2019 ). Solanum melongena is known primarily from cultivation, and as such we have not assigned it a threat status using the IUCN criteria. Discussion. Solanum melongena is known to cross easily with its wild progenitor S. insanum ( Davidar et al. 2015 ; Mutegi et al. 2015 ) and distinguishing the two taxa can be difficult especially in older specimens or in feral populations. We have chosen to recognise the two as distinct species rather than as infraspecific taxa (e.g., Lester and Hasan 1991 ; Daunay and Hazra 2012 ) because cultivated and wild plants are on distinct evolutionary trajectories. Wild plants are subject to natural selection, while cultivated plants are commensal with human beings. We recognise that this can lead to difficulties in identifying individual specimens, particularly those from old collections where details of provenance are not clear. Distinguishing the cultivated S. melongena from wild S. insanum can be particularly difficult in the absence of fruits. Keys in previous publications ( Knapp et al. 2013 ; Ranil et al. 2016 ) can be used for identification, but in general specimens we assign to S. insanum are pricklier, with smaller flowers and fruit, and more ruderal than those we assign to S. melongena . Flowers of S. melongena are often fasciated (with more than 5 parts), but usually are not in local cultivars. Local cultivars of S. melongena in tropical Asia can be especially difficult to distinguish from S. insanum . The only original material we have found for Sir John Hill's Solanum mexianum is his illustration ( Hill 1765 ) that clearly represents S. melongena ; we lectotypify the name with this illustration. Miller (1768) recognised the pre-Linnaean genus Melongena , in which he segregated those species treated as Solanum by Linnaeus (1753) as S. melongena . His species were distinguished based mainly on fruit size and shape; he considered colour variants to all belong to his Melongena ovata . He did not cite any specimens in his treatment, he cited instead pre-Linnaean polynomials mostly from Tournefort (1700) , none of which are illustrated in that work. Original material is thus apparently lacking for these names. From Miller's (1768) discussion of Melongena he had obviously cultivated all of the taxa he recognised at the Chelsea Physic Garden. A single specimen of S. melongena from the Miller herbarium has been found in BM, labelled on the verso side with "Hort. Chelsea, Miller" and " Melongena ovata " in Solander's handwriting, indicating it was so filed in Miller's herbarium (J. Wajer, person. comm.). We designate this here as the neotype for Melongena ovata , as it is not clear it is original material; all of these species of Melongena are certainly described from living material, as were most of the plants in the "Gardener's dictionary" ( Miller 1768 ). We have not designated neotypes for Melongena teres and M. incurva ; they should perhaps be sought in the Tournefort herbarium in Paris. Melongena spinosa was described ( Miller 1768 ) with reference to Bauhin's (1651) "Historia plantarum" (" Solanum pomiferum fructu spinoso J.B. 3. 619"). In that work there is an illustration labelled " Solanum pomiferum fructu spinoso" that is clearly S. melongena , with two fruits, one an ovoid shape with a prickly calyx and the other a different elongate shape with a non-prickly calyx. We designate this illustration as the lectotype for M. spinosa , it is the only original material we have found. In describing S. zeylanicum Scopoli (1786) provided an extremely detailed description of the plant and an excellent illustration both clearly referrable to S. melongena , although to a prickly relatively small-fruited variety. Scopoli (1786) stated that the fruit pulp was white ( "albida" ) and the illustration shows large, pendent fruits. In describing S. album , Loureiro (1790) referred only to "Trongum Solanum agreste . Album. Rumph. Amb. l. 8. c. 48. P. 241" and refers to its use "Virtus radices Ondontologica. Bacca edulis". In Rumphius's "Herbarium Amboinense" (Rumphius 1747) "Trongum Solanum agreste Solanum album /Trongum pra" comprises two white-flowered taxa, one with green fruits marbled with white and the other with red fruits, but the use in dentistry cited by Loureiro (1790) is part of the description of the previous taxon, "Trongum Solanum agreste /Trongum udan" (Rumphius 1747: 240). Hul and Dy Phon (2014) cited as "holotype" of S. album the illustration of "Trongum Solanum agreste spinosum" (Rumphius 1747: tab. 86, f. 1), a taxon described as part of "Trongum Solanum agreste ", not "Trongum Solanum agreste Solanum album ". No botanical materials have survived from Loureiro's stay in Indochina ( Merrill 1935 ), so a holotype is out of the question; we have designated as the neotype the illustration cited by Loureiro (1790) that was clearly available to him at the time he described S. album (see Turland et al. 2018 , Art. 8.1). In his interpretation of the "Herbarium Amboinense", Merrill (1917) cited a specimen that "almost certainly represents " Solanum agreste Solanum album " Rumph." ( Robinson 286 , US acc. # 654604). This is a specimen of S. lasiocarpum Dunal and does not correspond with the illustrations in Rumphius (1747). Dunal (1813) based his S. pressum entirely on Rumphius' (1747) "Trongum pra rubrum", citing no other material and taking his description entirely from that of Rumphius. He took his epithet from the suggestion that "Trongum Solanum agreste Solanum album " was translatable to Latin as "Trongum Solanum pressum " (Rumphius 1747). We have selected the one element cited by Dunal (1813) as the lectotype of S. pressum ; the illustration shows an unarmed plant with solitary flowers and berries. Rumphius' (1747) use of the word "rubrum" may indicate the berries were red, thus perhaps making this name a synonym of S. aethiopicum L., but this is not mentioned in the description. In the description of S. longum Roxburgh (1824) states that he distinguishes this from S. melongena by its elongate fruit form, in all other respects said to be exactly like the brinjal eggplant. Roxburgh (1814) first used the name Solanum longum in his "Hortus Bengalensis", which is a list of the plants growing in the Calcutta Botanical Garden; we have not found any unambiguous original material, nor any specimens with fruits as those in the description, the description may have come from live plants. We have selected as a neotype for S. longum specimen in the Wallich herbarium at Kew of a plant cultivated in the Calcutta Botanical Garden in 1821 (Wallich cat. 2628D) that has an immature fruit that looks like it might be of the elongate form (K001116666). Unravelling the identities of the various taxa referrable to S. melongena described by Blume (1826) is complicated by the lack of specimens of these cultivated plants. We here typify only those names for which we have found unambiguous original material; from Blume's descriptions the identity of the plants in question is clear. In describing S. ovigerum var. insanum , he is apparently coining the varietal epithet as new, not citing the Linnaean name S. insanum . The citation of this as occurring in cultivation and having oblong, cylindrical fruit makes it a clear synonym of S. melongena , but no original material has been found. Solanum pseudo-undatum was also described from cultivated plants; here we designate a sheet in L collected by Blume and labelled as " Solanum Solanum pseudoundatum " as the lectotype (L 0003642). The three varieties of S. pseudo-undatum , var. Solanum pseudo-undatum albiflorum , var. Solanum pseudo-undatum atropurpurascens and var. Solanum pseudo-undatum leucocarpon , are distinguished purely on flower or fruit colour and common name (see above); we have found no original material for any of these. Schumacher and Thonning ( Schumacher 1827 ) described S. edule from plants cultivated in west Africa. Of the two collections labelled as S. edule in Thonning's herbarium at Copenhagen, the sheet collected by Thonning himself (C10004584) is here selected as the lectotype, the other sheet, collected by P.E. Isert (C10004585) is less well preserved. Vorontsova and Knapp (2016) incorrectly cited the Thonning collection as "holotype" . The protologue of S. esculentum var. inerme ( Dunal 1852 ) cites two Brazilian collections, one of which ( Salzmann 392 in G-DC, G00131550) was questioned as belonging to S. melongena by Dunal himself. We have therefore selected the other collection, Blanchet 368 (G00357998) that bears a label in Dunal's hand as the lectotype. Var. Solanum melongena subinerme similarly was based on two collections; we have selected Blanchet 236 (G00131527) as the lectotype because it has a label in Dunal's hand saying " Solanum esculentum β 1842" and the other syntype, Sieber 308 from Trinidad (represented by two sheets G00357997, G00131564) is not annotated by Dunal. The infraspecific names Dunal (1852) coined under his S. ovigerum are presented in a manner different to most of the other infraspecific names in this work. The main infraspecific divisions, subrepandum and sinuatorepandum , are indicated with capital letters and are thus unranked; greek letters in this work indicate varietal rank. The varieties listed under [unranked] subrepandum are a mixture of polynomials and references to pre-Linnaean works, we do not typify these here. Dunal (1852) cited several collections in the protologue of [unranked] sinuatorepandum , Wallich cat. 2628d , de la Ossa s.n. and Gaudichaud 475 . We have selected de la Ossa s.n. (G00131521) in G-DC as the lectotype, the Wallich specimen in G-DC is of poor quality and the Gaudichaud collection in Paris is a plant of S. aethiopicum . The protologue of [unranked] subrepandum mentions only "v.s. in h. DC", making the identity of original material difficult. We have selected a specimen in G-DC that has a label in Dunal's hand stating "A. folia subrepanda" as the lectotype for this name (G00131523). The varietal names coined for the various fruit-shape and colour forms of S. melongena in Friedrich Alefeld's "Landwirthschaftliche flora" ( Alefeld 1866 ), a compendium of names for all plants cultivated in Germany, are most likely to have been based on living plants, the descriptions are minimal and no localities are cited. We have not typified these; they are more properly considered as cultivar names. In his work on the cultivated flora of the Soviet Union Filov (1958) coined many names, but many of these had neither description nor diagnosis and so are not validly published. The Russian taxonomist S.P. Dikii (1984) apparently thought Alefeld's names were not validly published and re-published them all at the same rank (see Names not validly published). Specimens examined. See Suppl. materials 1-3.