Capsicum chinense Jacq., Hort. Bot. Vindob. 3: 38. t. 67. 1776.

Barboza, Gloria E., Garcia, Carolina Carrizo, Bianchetti, Luciano de Bem, Romero, Maria V. & Scaldaferro, Marisel, 2022, Monograph of wild and cultivated chili peppers (Capsicum L., Solanaceae), PhytoKeys 200, pp. 1-423 : 1

11. Capsicum chinense Jacq., Hort. Bot. Vindob. 3: 38. t. 67. 1776.

Fig. 47

Capsicum cerasiforme Mill., Gard. Dict. ed. 8, no. 5. 1768. Type. Cultivated at the Chelsea Physic Garden, London, seeds from Spanish West Indies (no specimens cited).

Capsicum cerasiforme Lam., Tabl. Encycl. 2: 26. 1794, nom. illeg., not Capsicum cerasiforme Mill. (1768). Type. Brazil. "E Brafilia" (lectotype, designated here: P [P00357733]).

Capsicum luteum Lam., Tabl. Encycl. 2: 26. 1794. Type. India. "Ex India. Sonnerat" P. Sonnerat s.n. (lectotype, designated here: P [P00357731]).

Capsicum cerasiforme Willd., Enum. Pl. [Willdenow] 1: 242. 1809, nom. illeg., not Capsicum cerasiforme Mill. (1768). Type. Cultivated in the Berlin Botanic Garden, Germany, “Habitat…..” Capsicum cerasiforme [sheet] 1, herb. Willdenow (lectotype, designated here: B [B-W04427-01-0]).

Capsicum milleri Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 563. 1819, nom. illeg. superfl. Type. Based on Capsicum cerasiforme Mill. (cited in synonymy).

Capsicum indicum Dierb. var. luteum (Lam.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 24. 1829. Type. Based on Capsicum luteum Lam.

Capsicum indicum Dierb. var. humifusum Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 26. 1829. Type. Based on Capsicum cerasiforme Mill. (cited in synonymy).

Capsicum indicum Dierb. var. ochranthum Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 27. 1829. Type. Based on Capsicum cerasiforme Willd. (cited in synonymy).

Capsicum dichotomum Vell., Fl. Flumin.: 61. 1829 ( “1825”); Fl. Flumin. Icon. 2: t. 9. 1831 ( “1827”), as " Capsicum conicum ". Type. Brazil. [Rio de Janeiro]: "Colitur hortis" (lectotype, designated by Knapp et al. 2015, pg. 824: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_012] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 9. 1831).

Capsicum odoriferum Vell., Fl. Flumin.: 61. 1829 ( “1825”); Fl. Flumin. Icon. 2: t. 8. 1831 ( “1827”). Type. Brazil. [Rio de Janeiro]: "Colitur hortis" (lectotype, designated by Knapp et al. 2015, pg. 824: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_011] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 8. 1831).

Capsicum conicum Vell., Fl. Flumin. Icon. 2: t. 9. 1831. ( “1827”), nom. illeg., not Capsicum conicum G.Mey. (1818). Type. Based on Capsicum dichotomum Vell.

Capsicum toxicarium Poepp., Not. Natur- Heilk. 32: 228. 1832. Type. No locality cited (no specimens cited, no original material found). Peru. "Peruvian hispan., Maynas alto", Jan 1830, E.F. Poeppig 2220 (neotype, designated here: W [acc. # 0102196].

Capsicum cereolum Bertol., Hort. Bonon. Pl. Nov. 1: 6. t. 2. 1838. Type. Cultivated in Bologna, Italy "Nascitur in Brasilia unde semina ad nos attulit, et comiter dedit Eq. NUNNEZIUS" (lectotype, designated here [illustration]: Bertoloni, Hort. Bonon. Pl. Nov. 1: 6. t. 2. 1838).

Capsicum cordiforme Mill. var. subangulosum Fingerh., Monogr. Capsic.: 29. 1832. Type. No locality cited (no specimens cited; lectotype, designated here: [Fingerhuth, Monogr. Capsic. Tab. IX d. 1832]).

Capsicum cordiforme Mill. var. majus Fingerh., Monogr. Capsic.: 30. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. IX e. 1832).

Capsicum cordiforme Mill. var. minus Fingerh., Monogr. Capsic.: 30. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. X a. 1832).

Capsicum cordiforme Mill. var. olivaeforme Fingerh., Monogr. Capsic.: 30. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. X b. 1832).

Capsicum cordiforme Mill. var. globosum Fingerh., Monogr. Capsic.: 30. 1832. Type. " Crescit in Indiis et America meridionali" (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. X c. 1832).

Capsicum ustulatum Paxton, Paxton’s Mag. Bot. 5: 197. 1838. Type. Cultivated at Chatwsworth House, England (no specimens cited; lectotype, designated here: [illustration], Paxton, Paxton’s Mag. Bot. 5: [un-numbered plate opposite] 197. 1838).

Capsicum cerasiforme Willd. var. maurocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 422. 1852. Type. No locality cited (no specimens cited; lectotype designated here: [illustration] " Solanum Capsicum dictum perenne minus Capsicum erectum " Weinemann, Phytanthoza Iconogr. 4: 349, t. 930 c. 1745).

Capsicum oxycarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 426. 1852. Type. Brazil. Rio de Janeiro, 1831, M. Blanchet 90 (holotype: G-DC [G00200070]; isotype: MPU [MPU023041]).

Capsicum cordiforme Mill. var. subsulcatum Dunal, Prodr. [A. P. de Candolle] 13(1): 427. 1852. Type. No locality cited (no specimens cited; lectotype designated here: [illustration] Fingerhuth, Monogr. Capsic. Tab. IX c. 1832).

Capsicum annuum L. var. chinense (Jacq.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum chinense Jacq.

Capsicum annuum L. var. luteum (Lam.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum luteum Lam.

Capsicum annuum L. var. milleri (Roem. & Schult.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum milleri Roem. & Schult.

Capsicum grossum Willd. var. cerasiforme (Lam.) C.B.Clarke, Fl. Brit. India 4: 239. 1883. Type. Based on Capsicum cerasiforme Lam.

Capsicum annuum L. var. cerasiforme (Mill.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 92. 1898. Type. Based on Capsicum cerasiforme Mill.

Capsicum frutescens L. var. cerasiforme (Mill.) L.H.Bailey, Gentes Herbarum 1: 129 1923. Type. Based on Capsicum cerasiforme Mill.

Capsicum assamicum J.Purkay. & Lok.Singh, Ozean J. Appl. Sci. 5(1): 56. 2012. Type. India. Assam: Tezpur, 157 m, 25 May 2008, Purkayastha & Singh, DRLT 12 (holotype: CAL [by error, BSI in protologue, n.v.]; isotype: CAL [by error, DRLT and BSI in protologue, n.v.]).

Type.

Cultivated in Vienna, Austria ["Hort. Bot. Vindob."], N.J. von Jacquin s.n. (lectotype, designated here: W [acc. # 0080115]) .

Description.

Low, erect, short-lived subshrubs or rarely shrubs, 0.5-1.5 (-2.5) m tall, with the main stem (0.5-) 0.8-1.5 cm in diameter at base, few to much branched from near the base. Young stems 4-angled, fragile, green or greenish-brown, glabrous to glabrescent, with spreading, simple, uniseriate, (2-) 4-5-celled, eglandular trichomes 0.07-0.7 mm long; nodes solid, green or purple; bark of older stems light brown or green with light brown stripes, glabrous to sparsely pubescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous, concolorous or slightly discolorous, dark green above, pale green below, glabrescent to sparsely pubescent, with simple, 4-9-celled, eglandular trichomes 0.6-1.2 mm long, especially along the mid-vein or with a tuft of trichomes in the basal vein axils abaxially; blades of major leaves (4-) 5.25-10 (-15.5) cm long, 2.4-4 (-7) cm wide, ovate to elliptic, the major veins 4-5 (-6) on each side of mid-vein, the base attenuate or truncate and rather unequal, the margins entire, the apex short-acuminate, acuminate or acute; petioles 0.5-3 (-3.5) cm, glabrous to sparsely pubescent; blades of minor leaves similar in shape, 4-5.5 cm long, 2-2.5 cm wide; petioles 0.7-1.5 cm long, with the same pubescence as the major leaves. Inflorescences axillary, 2-4 (-5) flowers per axil, occasionally solitary flower; flowering pedicels 12-20 (-30) mm long, angled, erect or slightly spreading, geniculate at anthesis (wild forms) or pendent and non-geniculate (domesticated forms), green or green with purple lines, glabrous to moderately pubescent, the eglandular trichomes short, antrorse; pedicels scars conspicuous, slightly corky. Buds ellipsoid, cream or greenish-white. Flowers 5-6-merous. Calyx 1-2.5 (-3) mm long, 3-4 mm wide, cup-shaped, green, pentagonal or hexagonal in outline, the main veins strongly marked, the calyx appendages absent or with 5-6 mucro-like appendages, glabrous to moderately pubescent with similar short or long eglandular trichomes as the stems. Corolla 5 (-6)-merous, (5-) 6.5-8 mm long, 10-15 (-20) mm in diameter, dull white or greenish-white, occasionally with purple spots outside and within, stellate with interpetalar membrane, lobed nearly 2/3 of the way to the base, glabrous adaxially and abaxially, the tube (2-) 2.5-3 mm long, the lobes 3.5-5 mm long, 2-3.5 mm wide, triangular, spreading, the margins papillate, the tips acute, cucullate, papillate. Stamens 5 (-6), equal; filaments 1-1.3 mm long, white, cream or purple, inserted on the corolla 1-1.5 mm from the base, with auricles fused to the corolla at the point of insertion; anthers (1-) 1.38-2.05 mm long, blue or bluish-grey, very rarely yellow or greenish-white, broadly ellipsoid or ellipsoid, connivent or not connivent at anthesis, the connective sometimes wide and clearer. Gynoecium with the ovary 2-2.5 mm long, 2.5-3.5 mm in diameter, 2-3 (-4)-carpelar, light green, subglobose; ovules more than two per locule; nectary ca. 0.5 mm tall; styles heteromorphic, 3-4.1 mm, included, at the same level to the stamen length or exserted ca. 2 mm beyond the anthers, lilac or white, cylindrical; stigma ca. 0.15 mm long, ca. 0.5 mm wide, minute, discoid, pale green or pale yellow. Berry <10 mm in diameter, subglobose and orange to red (in wild populations), highly variable in size, shape and colour (in semi-domesticated or domesticated cultivars): subglobose or triangular, 10-20 mm in diameter, to long-triangular or campanulate, 30-60 (-100) mm long, 20-30 mm in diameter, some blocky, with the apex pointed, blunt or long-acuminate and upcurved and the base obtuse or truncate, green, yellow, brown or purple when immature, pale yellow, yellow, dark brown, orange, red or vermilion-scarlet at maturity, deciduous or persistent, very pungent (sometimes non-pungent), the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 15-45 (-55) mm long, thick, angled, strongly widened distally, erect and rigid (wild) or pendent and curved (domesticated); fruiting calyx 5-10 mm in diameter, persistent, not accrescent, discoid or shallowly cup-shaped, sometimes reflexed, with a strong annular constriction at junction with the pedicel (wild and domesticated), sometimes the margin ripped, green. Seeds 14-35 per fruit, (2.7-) 3-4 mm long, (2.5-) 3-3.5 mm wide, C-shaped or subglobose, pale yellow or nearly white, the seed coat smooth (SM), cerebelloid (SEM), the cells irregular in seed body, polygonal at margins, the lateral walls sinuate in the seed body, straight at margins; embryo imbricate.

Distribution.

Capsicum chinense wild forms were thought to originally occur in the north-central Amazon Basin lowlands (Brazil), where domestication is thought to have occurred ( Pickersgill 1971; Eshbaugh 1993). Pickersgill (1984) stated that wild C. chinense is confined to the lowlands of the Amazon, Orinoco and eastern Brazil, while Moses et al. (2014) suggested that populations in Central America and the Caribbean may have been primarily derived from progenitors from the Upper Amazon Region and later diverged through geographical isolation. Wild populations can still be found in the nature, but they are difficult to locate or are restricted to remote areas. Wild C. chinense (indigenous name: Pimi’ró) have been found in Roraima State (Brazil) ( Barbosa et al. 2002, 2008; Busso et al. 2003; Nascimento-Filho et al. 2007; Romero da Cruz and Forni Martins 2015), which confirms the occurrence of probable ancestors in the eastern lowlands of Amazonia (Fig. 48 ).

Capsicum chinense has been introduced into United States of America ( Bosland and Votava 2000), Mexico through Cuba ( González Estrada et al. 2010; Ruiz-Lau et al. 2011; López Castilla et al. 2019) and Central and South America where is found cultivated or escaped from cultivation; it has also been introduced outside the Americas (Eastern Europe, Africa, Asia: China, Japan, Taiwan, India and others) mainly by Portuguese explorers ( Eshbaugh 1983; Andrews 1993; Meghvansi et al. 2010).

Ecology.

Capsicum chinense grows in wet tropical and subtropical forests where native communities cultivate it around their chakras or in-home gardens, between 100 and 800 (-1,800) m elevation. Wild populations in northern Brazil grow in lowlands and disperse spontaneously in fallow agriculture areas, before the beginning of the rainy period ( Barbosa et al. 2002).

Phenology.

Flowering and fruiting all year in most parts of its range.

Chromosome number.

n = 12 ( Pickersgill 1966; Pozzobon and Schifino-Wittman 2006); 2n = 2x = 24 ( Pickersgill 1977; Moscone et al. 2003, 2007; Pozzobon et al. 2006; Romero da Cruz and Forni Martins 2015).

Common names.

Argentina: Ají bolita (Salta, Hilgert 2060). Belize: Jabanero pepper (Corozal, Balick 2211); Bolivia: Ají (Beni, Williams 937; La Paz, Williams 774), Aji mote (Santa Cruz, Krapovickas & Schinini 32488), Ají soliman (Cochabamba, Thomas 1093), Ají trompillo (Cochabamba, Thomas 704), Ají grande rojo (Beni, Rivero 428); Brazil: Baianinha ( Rondônia, Teles Walter 579), Canaimé, Chumbinho, Malaguetão, Morangão, Moranguinho, Murici, Murupi, Pimentãozinho, Peão-amaerlo, Peão-vermelho, Peixe-boi, Pimenta amarela, Pimenta-moranga, Pimenta murici, Pimenta ornamental, Pimenta-roxa, Chifre-de-carneiro, Chifre-de veado, Cumari-do-Pará, Esporão-de-galo, Olho-de-cará, Olho-de-carangueijo, Olho-de-mutum, Olho-de-peixe (falsa), Olho-de-peixe (verdadeira), Pimenta-vermelha longa, Unha-de-gato, Pimenta-de-cherio amarela (ardosa), Pimenta-de-cheiro longa, Pimenta-de mesa-ardosa (Roraima, Barbosa et al. 2006), Pimenta ( Goiás, Amaral 139; São Paulo, Hoehne s.n.), Habanero vermelha ( São Paulo, Lélis et al. 12), Pimenta açaí ( Amapá, Pereira et al. 1918), Pimenta acerola ( Amapá, Pereira et al. 1816), Pimenta-biquinho (Mina Gerais, Goiás, Carvalho et al. 2006), Pimenta boliviana (Rondonia, Teles Walter et al. 572), Pimenta camapú ( Amapá, Pereira et al. 2011), Pimenta cambuí ( Bahía, Medeiros et al. 196), Pimenta-doce ( Rondônia, Teles Walter 567), Pimenta habanero ( Carvalho et al. 2006), Pimenta-murupi ( Região Norte, Carvalho et al. 2006), Pimenta queimosa ( Amapá, Pereira et al. 1905), Pimenta do Curupira (Roraima, Barbosa et al. 2002; Nascimento Filho et al. 2007), Pimenta amarela ardosa ( Amapá, Pereira et al. 1908), Pimenta de bode ( Região Centro-Oeste, Carvalho et al. 2006), Pimenta de cheira (Pickersgill RU72-175), Pimenta de cheiro ( Bahía, Medeiros et al. 182; Rio de Janeiro, Gonçalves & Kaketru 2; Paraíba, Gadelha Neto 1748), Pimenta queimosa amarela ( Amapá, Pereira et al. 1904), Pimenta roxa alongada ( São Paulo, Hoehne s.n.), Pimenta de cheiro amarela ( Amapá, Pereira et al. 1901), Pimenta de cheiro legítimo ( Río de Janerio, Freire de Carvalho s.n.), Pimenta de cheiro redondinha ( Pereira et al. 2011), Pimenta de cheiro tacacá ( Amapá, Pereira et al. 2011), Pimenta-de-cheiro vermelha ( Amapá, Pereira et al. 1831; Roraima, Barbosa et al. 2006), Pimenta-dedo-de-papagaio ( Bahía, Medeiros et al. 200), Pimenta de cheiro de panela ( Amapá, Pereira et al. 2011); Colombia: Ají (Meta, Quevedo et al. 1816, Santander, Rosado 01), Trompito (Vichada, García Guzmán s.n.), Aji amarillo (Amazonas, Cárdenas et al. 9439), Ají pecas ( Caquetá, Cárdenas et al. 9311), Aji trompito ( Caquetá, Cárdenas et al. 9356), Ojito de Lucía ( Caquetá, Cárdenas et al. 9331), Ají de babilla (Amazonas, Cárdenas et al. 9431); Costa Rica: Chile, Chile chiricano, Chile panameño ( Bohs 2015); Ecuador: Ají (Napo, Ríos 429; Morona-Santiago, Villegas & Meneses 16), Ají botoncillo (Guayas, Holmgren & Heilborn s.n.); French Guiana: Péppé (Cayenne, Fleury 124); Jamaica: Scotch bonnet (Jamaica, Bosland and Votava 2000); Mexico: Habanero (Quintana Roo, Palma Gutiérrez 85-69), Rosita ( Yucatán, López Castilla et al. 2019), Chile cimarrón ( México, Hinton 4336), Chile costeño (Campeche, Ramírez A. 56), Chile habanero ( Yucatán, Ucan 4652), Chile xalapeño (Quintana Roo, Palma Gutiérrez 85-25), Chile de árbol ( Michoacán, Soto Núñez & Silva R. 3842); Panamá: Ají Chombo ( Bosland and Votava 2000); Peru: Ají (Lambayeque, Llatas Quiroz 3454; San Martín. Plowman & Schunke V. 11695), Arequipeño (Lima, Velarde Núñez 8), Charapita (Iquitos, Bosland and Votava 2000), Chinchi-uchu (Peru, Andrews 1984), Esticana (Cuzco, Ferreyra 16384), Limos (Lima, Velarde Núñez 16), Pucunuchu (San Martín, Plowman & Schunke 11497), Ají amarillo (Pasco, Huamán et al. 0317), Ají bravo (Loreto, Plowman 2457), Aji Charapa (Iquitos, Bosland and Votava 2000), Ají Dulce (Loreto, Plowman 2458), Ají mutecillo (Amazonas, Campos & Vargas 6945), Ají rojo o ají chanchamayo (Pasco, Oxapampa, Huamán et al. 0320), Ají Panca ( Bosland and Votava 2000), Ají Pucomucho ( Bosland and Votava 2000), Ajuju aji (Loreto, Hormia 2194), Colorado Cuzco (Lima, Velarde Núñez 4), Colorado Trujillo (Lima, Velarde Núñez 9), Munición uchu (Loreto, Williams 5225), Rojo Arequipa (Lima, Velarde Núñez 12), Ají corazón de pollo (Madre de Dios, Pesha Baca 95); Puerto Rico: Rocotillo ( Bosland and Votava 2000); Trinidad and Tobago: Congo ( Bosland and Votava 2000); United Stated of America: Datil (Florida, Bosland and Votava 2000), Rocotillo (Texas, Andrews 1984).

Indigenous names.

Bolivia: Bidó (Tacana, La Paz, Williams 1177), Cheti (Trinitario, Cochabamba, Thomas 704), Ta’ (Beni, Davis 1066), Hchetgi (Trinitario, Cochabamba, Thomas 1092), Tata (Yuqui, Cochabamba, Martínez et al. 246), Uchu (Quechua, La Paz, Williams 772), Winno (Yuracare, Cochabamba, Thomas 1093), Winno manera (Yuracare, Cochabamba, Thomas 704); Brazil: Pimi’ró (Macuxi, Roraima, Barbosa et al. 2002; Nascimento Filho et al. 2007); Colombia: Aati (Curripaco, Guainía, Espina et al. 182); Chamouiju (Colono Huitoto, Caquetá, Cárdenas et al. 9331), Doehoe (Amazonas, Cárdenas et al. 9439), Duaó ( Caquetá, Cárdenas et al. 9385), Gubsovia (Makuna, Amazonas, Cárdenas et al. 9431), Kukunarí (And, Amazonas, Castro & Matapí 565), Jade deé (Mui, Amazonas, Castro & Ramírez 132), Kukunari (Amazonas, Castro & Matapí 565), Fakiki deé (Mui, Amazonas, Castro & Rodríguez 309), Lole-ra (Norte de Santander, Solange 8), Tada tarre ( Sáliba, Casanare, Camargo 020), Wijichi-taku (And, Amazonas, Castro & Matapí 561); Ecuador: Atalba ucho (Kichwa, Sucumbíos, Moya & Reyes 207), Ampi (Shuar, Zamora-Chinchipe, Santín et al. 102), Ccoma ( Cofán, Sucumbios, Cerón 20848), Giimo (Oncaye, Napo, Davis & Yost 993), Giimo (Huaorani, Napo, Miller & Yépez 651), Giimoñabu (Huaorani, Napo, Miller & Yépez 651), Guimo (Huaorani, Orellana, Naranjo & Freire 329), Giimohue (plant) (Huaoroni, Napo, Miller & Yépez 651), Guimuwe (Huaorani, Orellana, Freire & Naranjo 612), Hueapia (Secoya, Cerón 62013), Imiá (Shuar Jívaro, Pastaza, Lewis et al. 13960), Jinia (Shuar, Morona Santiago, Pujupet RBAE 1005), Juminialinae ([Guahibo?] Casanare, Camargo 020), Khoma ( Kofán, Napo, Pinkley 230), Munisión (Kichwa, Orellana, Carrillo & Reyes 434), Ochabia (Siona, Sucumbíos, Miranda & Moya 437), Piujimia (Orellana, Herrera & Guerrero 186), Ucho (Quechua, Napo, Miller et al. 2404), Úchu (Achuar Jívaro, Pastaza, Lewis et al. 13960), Aji ucho (Quechua, Orellana, Reyes & Carrillo 773), Anya bia (= snake pepper) (Napo, Vickers 178), Arara uchu (Quechua, Orellana, Kohn 1613), Araya ucho (Quechua, Sucumbíos, Reyes & Moya 234), Araray-uchu (Quechua, Napo, Balslev & Irvine 4598), Biruti uchu (Quechua, Orellana, Kohn 1611), Butun uchu (Quechua, Napo, Ríos & Oña 430), Chipiri khuma ( Kofán, Napo, Pinkley 256), Cua Khoma ( Kofán, Napo, Pinkley 544), Palanda uchu (Quechua, Orellana, Kohn 1612), Puca uchu (Quechua, Orellana, Kohn 1896), Sasi uchu (Quechua, Orellana, Kohn 1611), Tiupi tio (Cayapa, Esmeraldas, Kvist 40566), Tɛsi khuma ( Kofán, Napo, Pinkley 257), thothao khuma ( Kofán, Napo, Pinkley 259), Tota cu’ma ( Kofán, Napo, Cerón et al. 5849), Uchu muyu (Quechua, Napo, Ríos 429), Wea bia (Siona, Napo, Vickers 115), Bula muyu uchu (Quechua, Orellana, Kohn 1910), Dio tape fin chuno (Cayapa, Esmeralda, Kvist 40586); Guyana: Há-ka-pu-tá (Cuyuni-Mazaruni, Tillett & Tillett 45424); Tor-tor-oi-ma (Cuyuni-Mazaruni, Tillett & Tillett 45627); Mexico: Habanero ik (Maya, Quintana Roo, Gutiérrez 24), Maax-ik (Maya, Quintana Roo, Gutiérrez 388), X-Kat ik (Quintana Roo, Palma Gutiérrez 85-24); Peru: Imia (Achual Jívaro, Loreto, Lewis et al. 11207), Tots (Yanesha, Pasco, Huamán et al. 0317), Yaájima (Amazonas, Kayap 652); Surinam: Ah-se-se (Kaxuyana, Kwamala, Plotkin 139), Pepra (Sranam, Kwamala, Plotkin 139), Pom-we (Tirio, Kwamala, Plotkin 139).

Uses.

Capsicum chinense has medicinal, ornamental and food uses. Indigeneous communities and rural people widely cultivated this species around their homes to consume the fruits raw, cooked or used as a spice ( Barbosa et al. 2002). The consumption of the fruits of this species has spread globally and are highly appreciated (sweet and pungent) due to its fleshiness and distinct aroma ( Kurian and Starks 2002; Basu and De 2003; Gahungu et al. 2011) in sauces and powders.

In addition to medicinal properties indicated in Table 3 , C. chinense has spiritual uses which have sometimes been recorded by researchers on herbarium specimen labels. For example, in Ecuador (Comunidad Oncaye, Davis & Yost 993), fruits were used by a shaman’s wife in the termination of an ayahuasa session, (Comunidad Bataburo, Freire & Naranjo 612), the entire plant is used to cure “espanto” (fearfulness) and (Comunidad Indillama, Reyes 502) and the plant is used to cure illness caused by the "mal viento", an evil spirit of the air. In Peru (Comunidad Palcazú, Huamán et al. 317), the smoke of a single dried fruit is used to help children when they are tearful. In Brazil (Roraima; Comunidad Macuxi, Barbosa et al. 2002), the common name of the species is an allusion to ‘curupira’, a supernatural being, guardian of the Amazon forests in Tupi mythology ( Gomez Platero and Palma Ehrichs 2011). In addition, outside the Americas, C. chinense has a broad spectrum of ethnopharmacological applications and medicinal properties (e.g. see Meghvansi et al. 2010).

Preliminary conservation assessment.

EOO (18,657,305 km2); AOO (712 km2). Capsicum chinense is a very widespread cultivated species worldwide; we assign the status of Least Concern (LC).

Discussion.

Capsicum chinense belongs to the Annum clade ( Carrizo García et al. 2016). It is most popularly known as Habanero (Mexico and United States of America) or Scotch Bonnet (Caribbean Islands), but it has numerous common and indigenous names in South America (see above and Baba et al. 2016), as well as landrace and cultivar names (e.g. Carolina Reaper, Bhut Jolokia, Red Savina, Congo pepper, NuMex Suave, NuMex Trick-or-Treat). Many of the names refer to differences in the fruit’s pungency, shape or colour ( Bosland and Votava 2000; Votava and Bosland 2004; Bosland and Baral 2007; Bosland et al. 2012; Bosland and Coon 2015).

Ever since Smith and Heiser (1957) first recognised C. chinense as a cultivated species in the genus, its recognition as an independent entity and as a member of the C. annuum complex ( C. annuum - C. frutescens - C. chinense ) has been debated in literature (see Baral and Bosland 2004; Eshbaugh 2012 and references therein), with arguments based on morphological, cytogenetic, biochemical (isoenzymes), phylogenetic and reproductive evidence. Recently, Raveendar et al. (2017) established the phylogenetic relationships of 13 complete chloroplast genome sequences belonging to the Solanaceae (five Capsicum species included). This study showed that the C. chinense cp genome is much closer to C. annuum var. glabriusculum , wild progenitor of C. annuum , than any other sampled Capsicum species ( C. annuum var. annuum , C. frutescens , C. baccatum and C. lycianthoides ). This indicates that members of the C. annuum complex share a common gene pool as was suggested by Eshbaugh (1983), Ibiza et al. (2012) and others.

An unequivocal morphological delimitation for C. chinense is difficult for two reasons: first, intermediate well-established types occur between wild and domesticated forms, probably due to intraspecific hybridisation ( Pickersgill et al. 1979) and second, some specimens are hardly indistinguishable from their closest relatives ( C. frutescens and C. annuum ) at flowering or fruiting stage (Smith and Heiser 1957; Eshbaugh 1976; Baral and Bosland 2004; Pickersgill 2016). Capsicum chinense is a short-lived subshrub or rarely shrub with 2-4 (-5) flowers per node, flowering pedicels that can be erect or spreading and geniculate at anthesis (mostly in wild forms) or pendent and non-geniculate (mostly in domesticated cultivars), a calyx with strongly-marked main veins that lacks appendages or is pentagonal or hexagonal in outline with 5-6 mucro-like appendages, a corolla that is dull white or greenish-white, fruits that are variable in shape, size and colour, deciduous or persistent, a fruiting calyx with a strong annular constriction at the junction with the pedicel and a clearly discoid fruiting calyx. The discoid calyx with a distinct annular constriction is the most conspicuous and consistent feature of C. chinense (Fig. 47 ).

Morphologically, Baral and Bosland (2004) found C. chinense produces primarily pendent flowers and calyces with a circular constriction, while C. frutescens produces erect flowers and calyces without a constriction. Another character found in literature to distinguish these species is the degree to which the style is exserted beyond the anthers ( D’Arcy 1973; Eshbaugh 2012). It is most difficult to distinguish these two species at the flowering stage. The great number of specimens of C. chinense , analysed in this treatment, showed that the position of the flowering pedicels and whether or not the style is exserted more than 1 mm beyond the anthers can be quite variable and prevents confident species assignment ( Peña-Yam et al. 2009). Calyx characters can be helpful in the identification of flowering specimens. The flowering calyx in both C. chinense and C. frutescens is deeply cup-shaped, but in C. chinense , the main nerves protrude remarkably from the calyx surface (Fig. 47C, J, L ), while in C. frutescens , they are often completely immersed in the calyx surface (Fig. 68C, G ) or may protrude distally. In both species, these nerves may slightly exceed (less than 1 mm) the calyx edge. In fruit, these species differ clearly. Capsicum chinense has erect and uniformly widened pedicels (wild/domesticated forms, Fig. 47F-H ) or pendent pedicels with one or two swellings distally (domesticated/cultivated forms, Fig. 47L-K ), but in both cases, a noticeable circular constriction at the junction with the fruiting calyx is present. In addition, the calyx flattens completely (discoid calyx) and remains appressed or reflexed to the fruit base (Fig. 47H, M ). In contrast, in C. frutescens , the pedicels are usually erect and widen in a similar way to the wild form of C. chinense , but the circular constriction is totally absent and the fruiting calyx remains deeply cup-shaped, housing the narrowed base of the elongate fruit (Fig. 68G, H ).

The differentiation of C. chinense from C. annuum (wild or domesticated specimens) is sometimes also difficult, especially at the fruiting stage. Wild forms of C. chinense with small red fruits and erect pedicels could be confused with some domesticated (or semi-domesticated) specimens of C. annuum var. glabriusculum , which share the same pedicel position and fruit characteristics (size, shape and colour), although wild forms of this variety have smaller fruits than the typical C. chinense . The presence/absence of the constriction in the pedicel/calyx junction (lacking in C. annuum var. glabriusculum ) allows the assignment to one or other taxon. Furthermore, C. chinense is a more robust plant with larger leaves and flowers than C. annuum var. glabriusculum .

During its long history of cultivation, C. chinense fruits have developed a broad range of variation due to active selection by growers, differing cultivation methods and adaptation to the environment ( Barbosa et al. 2010); in addition, there may have been historical and geographical isolation of subpopulations ( Bharath et al. 2013). This variation has been observed along its distribution ( Bharath et al. 2013; Baba et al. 2016; Moreira et al. 2018), where elongate, campanulate and blocky fruit shapes are more predominant than triangular or rounded shapes. Similarly, red fruits, followed by orange or yellow ones, are the most common, with rare occurrences of white, orange-yellow, black or brown fruits.

The pungency and aroma of this species are also remarkable. The "Carolina Reaper" cultivar is reputed to be the world’s hottest chile, which rates at an average of 1,641,183 Scoville Heat Units (SHU) (https://www.guinnessworldrecords.com/world-records/hottest-chili, accessed on 29 October 2019) and is a favourite amongst hot-pepper lovers. At the other extreme, the fruits of the low pungency varieties (pimenta-doce, pimenta-de-cheiro longa, pimenta-de-cheiro amarela) found in Brazil (Roraima) are used by indigenous and non-indigenous communities and are preferentially used in stews or salads, for preparing jellies or liqueurs or for food dish decoration ( Barbosa et al. 2006). The fruits are also distinguished by their distinct volatile profiles (depending on the cultivars), which give them a powerful exotic-fruity aroma ( Rodríguez-Burruezo et al. 2010; Garruti et al. 2013 and references therein; Patel et al. 2016).

Baral and Bosland (2004) provided morphological, phylogenetic (RADP) and reproductive (sexual compatibility) evidence to differentiate C. chinense from C. frutescens , while a study of genetic diversity (SSR and AFLP) ( Ibiza et al. 2012) has shown that, although C. chinense and C. frutescens are closely related, the molecular characterisation obtained is sufficient to differentiate the individual members of these species. Intermediate phenotypes and intermediate genetic accessions were explained by hybridisation, since gene flow could be occurring in sympatric areas of these two species ( Baral and Bosland 2004; Ibiza et al. 2012).

Jacquin (1776) described C. chinense , based on cultivated material. In the protologue, he stated that he saw this species cultivated on the Island of Martinique (Lesser Antilles), a small island in the Caribbean Sea, but he called it C. chinense , alluding to the region to which he thought the plant was native. We found probable original material only at W (W-acc. # 0080115). This sheet has two labels, one smaller at the lower left corner with the script " Capsicum sinense", probably in Jacquin’s filius writing (see D’Arcy 1970) and another at the lower right corner where " Capsicum sinense" and "Hort. bot. Vind." are handwritten by an unknown hand; this sheet is also annotated "Hb. Jacq." in a different hand and clearer ink. We designate this sheet (W-acc. # 0080115) as the lectotype for this species.

Capsicum cerasiforme was published by Lamarck (1794) with only the type locality "E Brafilia" cited. At P, we found a collection in Lamarck’s Herbarium with a label indicating that it belongs to " Capsicum cerasiforme " (P00357733) which we designate as the lectotype.

For C. luteum , Lamarck (1794) cited in the protologue "Ex India. Sonnerat" and the vernacular names 'piment jaune’ and 'Le piment de Mozambique’. At P, we found a sheet in Lamarck’s Herbarium with original material and the same data (P00357731). The label has a description and pencil drawings that likely were used in preparing the description. We select this sheet as the lectotype.

There are three sheets of original material labelled C. cerasiforme in Willdenow’s Herbarium held at Berlin. All of them contain reproductive branches; one of these (B-W04427-01-0) consists of a fruiting branch that matches Willdenow’s description exactly ( Willdenow 1809) and is, therefore, designated the lectotype.

In describing C. toxicarium , Poeppig (1832) cited no herbarium material. Poeppig’s Herbarium is housed at W ( Stafleu and Cowan 1983) where two specimens from Peru with the handwritten name C. toxicarium are held. On one of the specimens is a note referring to the poisonous effect of this plant known as "aji de veneno", a vernacular name cited in the protologue. This sheet consists of four fruiting branches and a fifth sterile branch (acc. # 0102196). The second sheet is a poorer specimen (one branch with only one fruit, acc. # 0102195) and has a different collection date from the other sheet. We designate W acc. # 0102196 as the neotype of C. toxicarium .

Bertoloni (1838) described C. cereolum , based on material cultivated in the Botanical Garden of Bologna (Italy), from Brazilian seeds. We could find no specimens, but Bertoloni provided a colour plate for C. cereolum that consists of a flowering and a fruiting branch; both description and plate match with our concept of C. chinense and we designate this illustration as the lectotype.

Paxton (1838) described C. ustulatum , based on cultivated plants grown at Chatsworth (England) obtained from seeds of unknown origin, sent by J. Bateman, Esq., of Knypersly, under the name "True Chili Capsicum ". Paxton is not known to have made herbarium material ( Stafleu and Cowan 1983), but he provided a plate opposite page 197 in Paxton (1838) labelled only as " Capsicum " although clearly associated with the protologue; this plate is here designated as the lectotype.

In the protologue of C. cerasiforme var. maurocarpum , Dunal (1852) cited no specimens, but referred this variety to a pre-Linnaean colour figure published by Weinemann (1745), which is here selected as the lectotype. Fruit characters depicted on the plate leave no doubt that this variety corresponds to C. chinense .

Dunal (1852) based C. cordiforme var. subsulcatum on Fig. IX c (as C. cordiforme ) published by Fingerhuth (1832); as Dunal did not mention any specimens, we designate Fingerhuth’s figure as the lectotype.

The original material of C. assamicum could not be found and is apparently lost. According to the protologue ( Purkayastha et al. 2012: 56) "Voucher specimens of the taxon are lodged at the Herbarium of Defence Research Laboratory (DRDO), Tezpur, Assam (DRLT, unregistered herbarium acronym), India and was also sent to the Herbarium of Botanical Survey of India, Eastern Circle, Shillong (BSI), India". The authors were wrong when they assigned the acronym BSI to the Eastern Circle, Shillong; it is should be ASSAM. BSI stands for Botanical Survey of India, Western Circle, Pune, India ( Thiers 2021). An annotation in The International Plant Name Index (https://www.ipni.org/n/60460283-2, accessed on 13 December 2019) "Purkayastha (pers. comm.) admitted that the citation of BSI was an error for CAL". After consulting the respective CAL, BSI and ASSAM curators (Barboza in litt.), neither holotype nor isotypes have been located in these Herbaria.

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 47. Capsicum chinense (A-H from wild plants I-M from domesticated plants) A plant B flower buds C flower on geniculate pedicel D flower, in front view E flower, in full anthesis F immature fruit G fruiting pedicel H mature fruit I flowering branch J flowers on pendent pedicels K, L immature fruits M mature fruit A-H no specimen vouchers (cult. in Banco de Germoplasma de Hortalicas, Embrapa / Hortalicas, Brasilia-DF, Brazil), photos taken in situ by L. Bianchetti I-M no specimen vouchers (cult. in Pairumani, Cochabamba-Bolivia), photos taken in situ by G. E. Barboza.

Figure 48. Distribution of C. chinense.

Figure 68. Capsicum frutescens A plant B flower buds C flower showing the calyx D flower, in lateral view E flower, in front view (note the connivent anthers) F flower after anthers dehiscence G immature (green) and mature (red) fruits H fruit, in longitudinal section showing the seeds A, B, E-G from Barboza & Leiva Gonzalez 4829 C, D, H from Barboza et al. 2041. Photos by G. E. Barboza.