Capsicum annuum L. var. annuum

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

1a. Capsicum annuum L. var. annuum

Figs 20 , 21

Capsicum grossum L., Mant. Pl.: 47. 1767. Type. "Habitat in India … H.U. " HU [Horto Upsaliensis]: Fructu vario crasso. Caulis biennis, Herb. Linn. N° 249.5 (lectotype, designated here: LINN [LINN-HL249-5]).

Capsicum cordiforme Mill., Gard. Dict. ed. 8, no. 2. 1768. Type. Cultivated at the Chelsea Physic Garden (no specimens cited; no original material located).

Capsicum tetragonum Mill., Gard. Dict. ed. 8, no. 3. 1768. Type. Cultivated at the Chelsea Physic Garden (no specimens cited; no original material located).

Capsicum angulosum Mill., Gard. Dict. ed. 8, no. 4. 1768. Type. Cultivated at the Chelsea Physic Garden (no specimens cited; no original material located).

Capsicum olivaeforme Mill., Gard. Dict. ed. 8, no. 6. 1768. Type. Cultivated at the Chelsea Physic Garden, seeds from “Barbadoes” (no specimens cited; no original material located).

Capsicum pyramidale Mill., Gard. Dict. ed. 8, no. 7. 1768. Type. Cultivated at the Chelsea Physic Garden, seeds from Egypt (no specimens cited; no original material located).

Capsicum conicum Lam., Tabl. Encycl. 2: 26. 1794. Type. "Ex Indiis" Herb. Lamarck s.n. (lectotype, designated here: P-LAM [P00357734]).

Capsicum bicolor Jacq., Fragm. Bot. 66, tab 99, fig. 1. 1809. Type. "Patriam ignoro" (no specimens cited; lectotype, designated here: Jacquin, Fragm. Bot.: 66, tab 99, fig. 1. 1809).

Capsicum grossum Willd., Enum. Pl. [Willdenow] 1: 241. 1809, nom. illeg., not Capsicum grossum L. (1767). Type. " Habitat in India orientali " Capsicum grossum [sheet] 2, Herb. Willdenow (lectotype, designated here: B [B-W04425-02-0]).

Capsicum sphaericum Willd., Enum. Pl. [Willdenow] 1: 241. 1809. Type. “Habitat…. " (lectotype, designated here: B [B-W04426-01-0, F neg. 2886]).

Capsicum nigrum Willd., Enum. Pl. [Willdenow] 1: 242. 1809, nom. illeg. superfl. Type. Based on Capsicum bicolor Jacq. (cited in synonymy).

Capsicum purpureum Vahl ex Hornem., Hort. Bot. Hafn. 1: 224. 1813. Type. [Denmark]. Hort. Haf., 1802, Herb. Vahl s.n. (lectotype, designated here: C [C10019148]).

Capsicum ovatum DC., Cat. Pl. Horti Monsp.: 86. 1813. Type. “Habitat….” (no specimens cited; no original material located; Capsicum ovatum , Anonymous s.n. (neotype, designated here: G-DC [G00200072]).

Capsicum longum DC., Cat. Pl. Horti Monsp.: 86. 1813. Type. "Hab... in hortis frequens" (no specimens cited; lectotype, designated here [illustration]: "Piper Calecuticum sive Capsicum oblongius, Bauhin et al., Hist. pl. 2: 943, f. I. 1651).

Capsicum globiferum G.Mey., Prim. Fl. Esseq.: 113. 1818. Type. "In plantationibus", no specimens cited; [Guyana]. Río Essequibo, E.K. Rodschied 29 (lectotype, designated here: GOET [GOET003420]).

Capsicum purpureum Roxb., Fl. Ind., ed. Carey & Wall. 2: 259. 1824, nom. illeg., not Capsicum purpureum Vahl ex Hornem. (1813). Type. "Most likely from the Molucca Islands" (no specimens cited; neotype, designated here: " C. purpureum , H.B.C." [Horto Botanici Calcutta]: K [K001132446]).

Capsicum indicum Dierb. var. vulgatum Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 22. 1829, nom. illeg. superfl. Type. Based on Capsicum annuum L. (cited in synonymy).

Capsicum indicum Dierb. var. longum (DC.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 23. 1829. Type. Based on Capsicum longum DC.

Capsicum indicum Dierb. var. tetragonum (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 23. 1829. Type. Based on Capsicum tetragonum Mill.

Capsicum indicum Dierb. var. angulosum (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 25. 1829. Type. Based on Capsicum angulosum Mill.

Capsicum indicum Dierb. var. cordiforme (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 25. 1829. Type. Based on Capsicum cordiforme Mill.

Capsicum indicum Dierb. var. grossum (L.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 26. 1829. Type. Based on Capsicum grossum L.

Capsicum indicum Dierb. var. sphaericum (Willd.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 27. 1829. Type. Based on Capsicum sphaericum Willd.

Capsicum indicum Dierb. var. ovatum (DC.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 27. 1829. Type. Based on Capsicum ovatum DC.

Capsicum indicum Dierb. var. pyramidale (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 28. 1829. Type. Based on Capsicum pyramidale Mill.

Capsicum indicum Dierb. var. olivaeforme (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 28. 1829. Type. Based on Capsicum olivaeforme Mill.

Capsicum indicum Dierb. var. nigrum (Willd.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 29. 1829. Type. Based on Capsicum nigrum Willd.

Capsicum axi Vell., Fl. Flumin.: 61. 1829 ( “1825”); Fl. Flumin. Icon. 2: t. 6. 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_009] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 6. 1831).

Capsicum silvestre Vell., Fl. Flumin. 60. 1829 ( “1825”); Fl. Flumin. Icon. 2: t. 1. 1831 ( “1827”). Type. Brazil. [Rio de Janeiro]: "Ad declivium Alpium Fluminensium" (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_004] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 1. 1831).

Capsicum annuum L. var. rugosulum Fingerh., Monogr. Capsic.: 13. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II b. 1832).

Capsicum annuum L. var. acuminatum Fingerh., Monogr. Capsic.: 13. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II c. 1832).

Capsicum annuum L. var. subangulosum Fingerh., Monogr. Capsic. 13. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II d. 1832).

Capsicum annuum L. var. ovoideum Fingerh., Monogr. Capsic.: 14. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II e. 1832).

Capsicum annuum L. var. abbreviatum Fingerh., Monogr. Capsic.: 14. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II f. 1832).

Capsicum annuum L. var. olivaeforme Fingerh., Monogr. Capsic.: 14. 1832. Type. "Crecit in America meridionali et India oriental" (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II g. 1832).

Capsicum bicolor Jacq. var. purpureum (Vahl ex Hornem.) Fingerh., Monogr. Capsic.: 16. 1832. Type. Based on Capsicum purpureum Vahl ex Hornem.

Capsicum strictum Fingerh., Monogr. Capsic.: 21. 1832. Type. “Patria…..” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. V a. 1832).

Capsicum grossum Capsicum grossum Willd. var. Capsicum grossum pomiforme Fingerh., Monogr. Capsic.: 22. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. V c. 1832).

Capsicum grossum Willd. var. ovatum Fingerh., Monogr. Capsic.: 22. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. V d. 1832).

Capsicum grossum Capsicum grossum Willd. var. Capsicum grossum cordatum Fingerh., Monogr. Capsic.: 22. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VI a. 1832).

Capsicum grossum Capsicum grossum Willd. var. Capsicum grossum angulosum Fingerh., Monogr. Capsic.: 22. 1832. Type: "Patria India orientalis (Herb. Wight et Herb. Hamilt.)" (no specimens found; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VI d. 1832).

Capsicum ceratocarpum Fingerh., Monogr. Capsic.: 22. 1832. Type. “Patria….” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VI c. 1832).

Capsicum longum DC. var. incrassatum Fingerh., Monogr. Capsic.: 24. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VII a. 1832).

Capsicum longum DC. var. latum Fingerh., Monogr. Capsic.: 25. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VII b (as - - [ Capsicum longum ] Capsicum luteum ). 1832).

Capsicum longum DC. var. rectum Fingerh., Monogr. Capsic.: 25. 1832. Type. "Cresit in Indiis et America meridionali" (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VII c. 1832).

Capsicum pendulum Willd. var. torulosum Fingerh., Monogr. Capsic.: 26. 1832. Type. [Indonesia] "in Amboina" (no specimens cited; lectotype, designated here [illustration]: Capsicum rubrum minus Rumphius, Herbarium Amboinense 5, Tab. LXXXVIII, fig. 1, 1747, cited in synonymy).

Capsicum angulosum Mill. var. conicum Fingerh., Monogr. Capsic.: 28. 1832. Type. No locality cited (no specimens cited, no original material located).

Capsicum angulosum Mill. var. ovale Fingerh., Monogr. Capsic.: 28. 1832. Type. “Patria….?” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VIII b. 1832).

Capsicum hamiltonii G.Don, Gen. Hist. 4: 447. 1838. Type. [Caribbean Islands] "Native of the Island of Nevis, in gardens" (no specimens cited, no original material located).

Capsicum annuum L. var. longum (DC.) Sendtn., Fl. Bras. (Martius) 10(6): 144. 1846. Type. Based on Capsicum longum DC.

Capsicum annuum L. var. grossum (Willd.) Sendtn., Fl. Bras. (Martius) 10(6): 147. 1846. Type. Based on Capsicum grossum Willd.

Capsicum annuum L. var. cordiforme (Mill.) Sendtn., Fl. Bras. (Martius) 10(6): 148. 1846. Type. Based on Capsicum cordiforme Mill.

Capsicum abyssinicum A.Rich., Tent. Fl. Abyss 2: 96. 1850. Type. [Ethiopia] "Abyssinia, Ouedjerate", R. Quartin Dillon s.n. (lectotype, designated here: P [P00329903]; isolectotypes: P [P00329904, P00329905]).

Capsicum annuum L. var. oblongum Dunal, Prodr. [A. P. de Candolle] 13(1): 412. 1852. Type. " Capsicum annuum α Capsicum oblongum fructibus rubris", 1844, Herb. Dunal (lectotype, designated here: G-DC [G00131768]).

Capsicum pyramidale Mill. var. longicorne Dunal, Prodr. [A. P. de Candolle] 13(1): 414. 1852. Type. [Indonesia] Java, 1843, H. Zollinger 489 (lectotype, designated here: G-DC [G00131841]; isolectotypes: G [G00390281], LE).

Capsicum bicolor Jacq. var. purpureum (Vahl ex Hornem.) Dunal, Prodr. [A. P. de Candolle] 13(1): 414. 1852. Type. Based on Capsicum purpureum Vahl ex Hornem.

Capsicum testiculatum Vis. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 424. 1852. Type. In Hort. Montpellier [seeds sent by R. de Visiani], 1837, Anonymous s.n. (lectotype, designated here: G-DC [G00200067]; isolectotype: MPU [MPU023039]).

Capsicum angulosum Mill. var. macrocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 426. 1852. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VIII a (as Capsicum angulosum M.). 1832).

Capsicum leucocarpon Dunal, Prodr. [A. P. de Candolle] 13(1): 429. 1852. Type. Cultivated in England " Capsicum americanum latifolium , fructu oblongo erecto candido" ( Miller 1752) (no specimens cited, no original material located).

Capsicum dulce Dunal, Prodr. [A. P. de Candolle] 13(1): 428. 1852. Type. Cultivated in Montpellier, France, "In hortis botanicis cultum" (no specimens cited; no original material located).

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

Capsicum annuum L. var. angulosum (Mill.) Alef., Landw. Fl.: 132. 1866, as ' angulatum '. Type. Based on Capsicum angulosum Mill.

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

Capsicum annuum L. var. globiferum (G.Mey.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum globiferum G.Mey.

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

Capsicum annuum L. var. tetragonum (Mill.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum tetragonum Mill.

Capsicum annuum L. var. tetragonum (Mill.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum tetragonum Mill.

Capsicum annuum L. var. purpureum (Roxb.) Alef., Landw. Fl.: 134. 1866. Type. Based on Capsicum purpureum Roxb.

Capsicum annuum L. var. ceratocarpum (Fingerh.) Alef., Landw. Fl.: 134. 1866. Type. Capsicum ceratocarpum Fingerh.

Capsicum annuum L. var. bicolor (Jacq.) Alef., Landw. Fl.: 134. 1866. Type. Capsicum bicolor Jacq.

Capsicum fasciculatum Sturtev., Bull. Torrey Bot. Club 15(5): 133. 1888. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: " Tenjikumamori , Capsicum annuum L. ( Solaneae)", Tanaka & Motoyoshi, Sô-Mokou-Zoussets, vol. 3, Tab. 38. 1874).

Capsicum annuum L. var. longum (DC.) Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. Based on Capsicum longum DC.

Capsicum annuum L. var. erectum Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. "Java, cult." (no specimens cited, no original material located).

Capsicum annuum L. var. grossum (L.) Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. Based on Capsicum grossum L.

Capsicum annuum L. var. fasciculatum (Sturtev.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 68, pl. 9, f. 4. 1898. Type. Based on Capsicum fasciculatum Sturtev.

Capsicum frutescens L. var. lanicaule Greenm., Proc. Amer. Acad. Arts 39: 88. 1903. Type. Mexico. Jalisco: along Ave. Vallarta in Ciudad Granja, on the western outskirts of Guadalajara, 31 Dec 1886, E. Palmer 639 (lectotype, designated here: US [00816554, acc. # 92534], isolectotype: BM [BM000775827]).

Capsicum velutinum De Wild., Pl. Bequaert. 1: 413. 1922. Type. [Democratic Republic of the Congo]. Basankusu, Mar 1913, O. Lamboray 22 (lectotype, designated here: BR [BR000000649909]).

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

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

Capsicum annuum L. forma erectum Makino, J. Jap. Bot. 3(8): 29. 1926, as " Capsicum annuum var. fasciculatum f. erectum ". Type. "Hab. JAPAN, cultivated" (no specimens cited, no original material located).

Capsicum annuum L. forma pendulum Makino, J. Jap. Bot. 3(8): 29. 1926, as " Capsicum annuum var. fasciculatum f. pendulum ". Type. "Hab. JAPAN, cultivated, rare" (no specimens cited, no original material located).

Capsicum petenense Standl., Publ. Carnegie Inst. Wash. 461(4): 84. 1935. Type. Guatemala. Distr. Peten, La Libertad, Jun 1933, C. L. Lundell 3754 (holotype: F [v0072800F, acc. # 685329]; isotypes: CORD [CORD00101764 fragment ex MICH], MICH [1109873]).

Capsicum sonitpurense J.Sarma & G.Dutta, Bangladesh J. Pl. Taxon. 24(2): 215. 2017. Type. India. Assam, Sonitpur, Tezpur, 49 m, 22 Oct 2016, J. Sarma & G. Dutta 394 (holotype: ASSAM [acc. # 95893, sheet 394A]; isotypes: TUH [Tezpur University Herbarium, 3 sheets 394 B, C, D]).

Description.

Annual herbs or short-lived, compact, low subshrubs, 1-1.5 m tall, the main stem 0.5-1 cm in diameter at base, branched from near the base. Young stems 3-4-angled, fragile, green to brownish-green, sometimes with purple lines, glabrous, glabrescent to moderately pubescent, rarely densely pubescent, with appressed-antrorse, simple, uniseriate, (5-) 8-13)-celled, eglandular trichomes 0.5-1 (-2) mm long; nodes green or with purple spots; bark of older stems light brown or brown, glabrous to sparsely pubescent; lenticels absent or few. Sympodial units difoliate, the leaves geminate; leaf pair similar in size and shape. Leaves membranous, concolorous, pale to dark green, glabrous to moderately pubescent on both sides, especially on the main veins abaxially, the trichomes similar to those of the stems; blades of all leaves 3-7 (-15.5) cm long, 2.5-5 (-8) cm wide, ovate to elliptic, the major veins (3-) 5-8 on each side of mid-vein, the base truncate to cordate or cuneate to attenuate, the margins entire, the apex acuminate or long-acuminate; petioles (0.5-) 4-7 (-10) cm, with the same pubescence as the stems. Inflorescences axillary, 1 (- 2) flowers per axil, rarely more; flowering pedicels (6-) 10-40 mm long, angled, erect and geniculate at anthesis or pendent and non-geniculate, green or purple, glabrous to moderately pubescent, the eglandular trichomes usually short, antrorse; pedicels scars inconspicuous. Buds globose, white or purple. Flowers 5-7-merous. Calyx 1-4 mm long, 3-5 mm wide, cup-shaped, green, strongly 5-10-nerved, glabrous to moderately pubescent with similar short or long eglandular trichomes as the stems, the calyx appendages usually 5 (-7), minute, 0.3-0.5 mm long. Corolla 8-15 mm long, (8-) 10-22 mm in diameter, entirely white, rarely entirely pale yellow or purple, stellate with narrow interpetalar membrane, lobed ca. halfway or 2/3 of the way to the base, glabrous adaxially and abaxially, the tube 3-8 mm long, the lobes 5-7 mm long, 3.5-5.5 mm wide, ovate, spreading, the margins finely ciliate, the tips acute, papillate. Stamens 5-7, equal; filaments 1-3 mm long, white or cream, sometimes purple, inserted on the corolla 1-1.5 mm from the base, with auricles fused to the corolla tube at the point of insertion; anthers 2-3 mm, ellipsoid or ovoid, pale blue to purplish, very rarely yellow, connivent or not connivent at anthesis. Gynoecium with ovary 1.5-3 mm long, 1.2-2.5 mm in diameter, ovoid or globose, green; nectary ca. 0.5 mm tall, pale green; style heteromorphic, short style 2.2-2.5 mm, not exceeding the anthers, medium style nearly the same height as the anthers, long style 3-5.1 mm, exserted 1.3-2.3 mm beyond the anthers, cylindrical, white or purple; stigma 0.1-0.2 mm long, ca. 0.4 mm wide, discoid or capitate, pale green or yellow. Berry highly variable in shape, size and colour, usually blocky or elongate, less commonly globose, up to 300 mm long, 6-65 mm in diameter, green, yellow or purple when immature, yellow, red, brown, purple or purple-black at maturity, persistent, pungent or non-pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 25-50 (-70) mm, erect or pendent, rigid, angled, uniformly widened, green; fruiting calyx 15-25 mm in diameter, slightly accrescent, discoid or rather cup-shaped, green. Seeds more than 50 per fruit, 3.8-4.4 mm long, 3.2-3.6 mm wide, C-shaped, pale yellow, the seed coat smooth to slightly reticulate (SM), cerebelloid (SEM), the cells irregular in shape, the lateral walls sinuate; embryo imbricate.

Distribution.

Capsicum annuum var. annuum is the most extensively cultivated pepper worldwide.

Ecology.

Capsicum annuum var. annuum is found in diverse habitats throughout its wide distribution and is well adapted to the highlands environments (0-2,600 m elevation).

Phenology.

Flowering and fruiting all year.

Chromosome number.

2n = 2x = 24 ( Pickersgill 1971, 1977, 1991; Moscone et al. 2007).

Common names.

Argentina: Ají balita (Jujuy, Moscone 204), Pimiento (Corrientes, Anzótegui & Benitez 237; Salta, Hunziker 25498), Serrano (Salta, Hunziker 25492), Pimiento Calahorra ( Córdoba, Hunziker 29428); Bolivia: Ají (Beni, Balderrama 10; Santa Cruz, Saldías P. 759), Urubibi (Beni, Ticona & Saravia May 10), Pimentón colorado (Santa Cruz, Libreros et al. 2014), Brazil: Pimentão (Roraima, Barbosa et al. 2006; São Paulo, Duth s.n.), Pimenta-americana, Pimentão-vermelho, Pimentão-indigena, Pimenta-de-mesa, Pimenta-ornamental, Pimentão-bola, Pimenta-doce (Roraima, Barbosa et al. 2006), Pimenta-jalapeño, Pimenta-cayenne, Pimenta-serrano ( Carvalho et al. 2006); Chile: Ají ( Concepción, Junge 5755); Colombia: Ají (Amazonas, Cordero P. 683; Chocó, La Rotta & Martínez 737; Guainía, Espina et al. 189; Vichada, Rodríguez 165), Pimentón (Amazonas, Torres & Morales 139; Caquetá, Cárdenas et al. 9310; Cundinamarca, Correa N. 006), Ají amarillo ( Caquetá, Cárdenas et al. 9375, Vichada, Rodríguez 1), Ají bravo (Amazonas, Posada 2580), Ají dulce (Amazonas, Cárdenas et al. 9438; Cundinamarca, Duque-Jaramillo 3555), Ají grande (Amazonas, Cárdenas et al. 9432), Ají pajaa (Vaupes, Rodríguez 86), Ají picante (Amazonas, Posada 2581), Ají pimentón (Amazonas, Torres et al. 4011), Ají yunga ( Nariño, de Benavides 4673), Ají yuquitania (Vaupes, Plowman 11986), Pimentón dulce ( Caquetá, Cárdenas et al. 9360), Pimentón quisquis (Meta, Álvarez & Montañez 1), Ají de agua (Vaupes, Rodríguez 102), Ají de blanco (Amazonas, Henao & Kuiru 172), Ají de curripaco ( Vaupés, Rodríguez 105), Ají de gente (Amazonas, Cárdenas et al. 9424), Ají mas picante (Vaupes, Rodríguez 83), Diente de chucha ( Caquetá, Cárdenas et al. 9319), Pipí de perro ( Caquetá, Cárdenas et al. 9306), Ají largo de blanco (Amazonas, Henao 316), Ecuador: Ají (Chimborazo, Lara s.n.; Pichincha, Mejía 001), Pimiento (Guayas, Bonifaz & Cornejo 4158; Pichincha, Narváez 018), Ají colorado (Tungurahua, Acosta Solís 8848), Ají manzana (Guayas, Valverde 392), Ají patateño (Chimborazo, Ganchozo 007), Manzanita de Eva (Guayas, Valverde 35); El Salvador: Chile de relleno (San Salvador, Calderón 523); Guatemala: Chile (Quezaltenango, Steyermark 34462); Honduras: Chile fuerte ( Morazán, Molina R. & Molina 34535), Chile picante ( Copán, Molina R. & Molina 33572; Morazán, Molina R. 34008), Chile de gallina (Cortes, Chevez 40); Mexico: Chile (Chiapas, Calzada et al. 3774; Michoacán, Miranda et al. 1519B; Tamaulipas, Rodríguez & Lira 63; Veracruz, Baizabal & Zola B. 12), Picante (Veracruz, Baizabal & Zola B. 11), Pimentón ( Michoacán, Bye et al. QD 246), Chile ancho (Guanajuato, Benítez 689; Tamaulipas, Rodríguez & Lira 63b), Chile bolita (Zacatecas, Benítez 738), Chile camote (Tamaulipas, Hernández 1944), Chile cimarrón ( México, Hinton 4336), Chile chaua ( Yucatán, Simá 610), Chile cora (Zacatecas, Benítez 734), Chile costeño (Campeche, Ramírez A. 56), Chile chilaca (Guanajuato, Benítez et al. 386), Chile delgado (Quintana Roo, Gutiérrez 85-27), Chile dulce ( Michoacán, Bye et al. 90; Tabasco, Ortíz 01; Yucatán, Simá 606), Chile gordo (Oaxaca, Hernández Ortega 484; Veracruz, Vázquez 673), Chile guajon (Zacatecas, Benítez 724), Chile kat ( Yucatán, Ucan et al. 3519), Chile largo (Quintana Roo, Gutiérrez 85-41), Chile pableño (Guanajuato, Vieyra s.n.), Chile pasilla (Tamaulipas, Rodríguez & Lira 63ª), Chile pimienta (Mexico, Rodin 40), Chile pimiento (Mexico, Linares 846), Chile serrano (Hidalgo, Villa 71; Oaxaca, Martínez Calderón 1643; Veracruz, Zola & Baizabal 1439), Chile uñepicho (Veracruz, Díaz Rico 48), Chile verde, chile de huerta ( Michoacán, Soto Núñez et al. 5442), Chile xalapeño, Calzada 2367), Fruto azul (Guerrero, Díaz Rico 221), Pimiento grande (Oaxaca, Bamonte 77), Chile de agua (Oaxaca, Acosta Castellanos 9417), Chile de árbol ( Michoacán, Soto N. 14302), Chile de vida o chilar (Oaxaca, Moreno 29), Chile mira parba (Tabasco, Ortíz 07), Chile pico paloma (Tabasco, Ortíz 20), Chile de árbol de bola ( Michoacán, Soto Núñez 14047), Chile de uña de perro (Veracruz, Vázquez 998); Nicaragua: Chile ( León, Guzmán et al. 1011); Panamá: Ají (Canal Zone, Standley 28523), Sweet pepper, pimiento morrón, ají (Canal Zone, Standley 29880); Peru: Encarnado (Lima, Velarde Nuñez 20), Pimiento (Lima, Vilcapoma S. 84), Aji amarillo (Lima, Vilcapoma S. 85), Ají cerezo (Lambayeque, Libreros et al. 2013), Ají dulce (Loreto, Hormia 2228), Ají limón (Trujillo, Plowman 14541), Ají tomate ( Junín, Ridoutt 11714), Bobo panca (Lima, Velarde Nuñez 19), Cerezo triangular (Lambayeque, Libreros et al. 2013), Conico amarillo (Lima, Velarde Nuñez 26), Cónico panca (Lima, Velarde Nuñez 18), Tambo Tacna (Lima, Velarde Nuñez 22), Aji Acarí Moquegua (Lima, Velarde Nuñez 7); United States of America: Black Chile or Chile negro (New Mexico, Wooton 48), Red Chile or Chile rojo (New Mexico, Wooton 49). Venezuela: Ají caribe (Portuguesa, Aymard 5108).

Indigenous names.

Bolivia: Ta (Beni, Ticona & Saravia May 10); Colombia: Aati (Curripaco, Guainía, Espina et al. 189), Aii (Cauca, Plowman & Vaughan 5370), Asi (Piapoco, Vichada, Rodríguez 177), Azi (Piapocos, Vichada, Rodríguez 165), Biaá (Tanimuka, Amazonas, Cárdenas et al. 9406), Coc (Puinabe, Vichada, Rodríguez 169), Curripaati (Tucano, Guainía, Marín & Rodríguez 502), Fecogɨ (Bora, Amazonas, Torres et al. 4020), Fekorai (Huitoto-Mɨnɨka, Amazonas, Henao 167), Jipujou ( Caquetá, Cárdenas et al. 9330), Jumerien (Sukuare, Vichada, Rodríguez 1), Mèe (Colona, Amazonas, Torres & Rodríguez 2021), Munɨ (Huitoto, Amazonas, Posada 2577), Nubata (Andoque, Amazonas, Torres et al. 4047), Pidá ( Emberá, Chocó, La Rotta & Martínez 737), Rɨairai (Huitoto-Mɨnɨka, Amazonas, Henao & Kuiru 172), Arera rɨairai (Huitoto-Mɨnɨka, Amazonas, Henao 316), Yicane ( Miraña, Caquetá, Cárdenas et al. 9375), Jeba gayebá (Mui, Castro et al. 238), Masan via (Amazonas, Cárdenas et al. 9424), Pipita deé (Mui, Amazonas, Castro 305), Viahoracá carunoje (Tanimuka, Amazonas, Cárdenas et al. 9432); Ecuador: Aatyu (Chapalaachi, Yañez et al. 1485), Uchu (Quichua, Napo, Kohn 1225), Ahí bia (Siona & Secoya Indians, Napo, Vickers 211), Suara pia (Siona & Secoya Indians, Napo, Vickers 227), Soa horo bia (Siona & Secoya Indians, Napo, Vickers 200); Mexico: Cants (Huave, Oaxaca, Zizumbo & Colunga 145), Chaunik ( Yucatán, Vargas 66), Guiin-cànár (Zapateco, Oaxaca, Hunn OAX-1345), Guiin-ló-yág (Zapateco, Oaxaca, Hunn OAX-1341), Guiin-ló-ngÚbidz (Zapateco, Oaxaca, Hunn OAX-1343), Guiin-nàl-zhàb (Zapateco, Oaxaca, Hunn OAX-1342), Guiin-txxtlé (Zapoteco, Oaxaca, Hunn OAX-1344), Moo-o-re (Oaxaca, Hernández Ortega 482), Moo-o-qui (Oaxaca, Hernández Ortega 481), Niiy (Oaxaca, Antonio B. GUI 201), Xcatic (Maya, Quintana Roo, Villanueva 591), X-mash ik (Quintana Roo, Gutiérrez 85-26), X-mehen (Quintana Roo, Gutiérrez 85-27), Xkat-ik (Maya, Quintana Roo, Gutiérrez 26), Ya Jimia (Morona-Santiago, Evans 4384), Ya’axik (Maya, Quintana Roo, Gutiérrez 109), Chaua ik ( Yucatán, May 39), Ixa nadun (Guerrero, Wagenbreth 130), Kat ik ( Yucatán, Ucan et al. 3529), Nadam kanc (Huave, Oaxaca, Bamonte 77), Namis kanc (Huave, Oaxaca, Bamonte 79), Yaá dia (Mixteco, Guerrero, Díaz Rico 221), Yak ik (Quintana Roo, Gutiérrez 85-71), Ixe dun xkuiya smidi (Guerrero, Wagenbreth 687); Peru: Iwiá (Mayna Jívaro, Loreto, Lewis et al. 10922), Kistian jima (Amazonas, Ancuash 297), Mun hima (Amazonas, Berlín 1572), Tsitikana ogat-santsakarioni (Machiguenga, Cuzco, Johnson 70).

Uses.

Capsicum annuum var. annuum is the economically most important member of the genus. The fruits are widely used in international cuisine in a broad spectrum of meals and preparations, because of their aroma, flavour, texture and level of pungency. Some cultivars have good acceptance as ornamental plants due to the colour of the leaves and the brightness of the colourful and usually erect fruits (e.g. Christmas peppers, Bolivian rainbow, Fig. 21 ). There are few instances where medicinal uses have been recorded on herbarium labels (Table 3 ), but the medical and nutritional importance, as well as the pharmacological properties and therapeutic effects of the active compounds, of C. annuum fruits have been extensively highlighted ( Al-Snafi 2015; Srinivasan 2016; Masud Parvez 2017; Sricharoen et al. 2017; Saleh et al. 2018; Sanati et al. 2018; Roman et al. 2020).

Preliminary conservation assessment.

Capsicum annuum var. annuum is not under threat.

Discussion.

The domesticated taxon C. annuum var. annuum belongs to the Annuum clade, together with C. chinense , C. frutescens and C. galapagoense ( Carrizo García et al. 2016). The three domesticated species and their conspecific wild populations constitute the Capsicum annuum primary gene pool ( van Zonneveld et al. 2015).

The vast majority of the modern landraces, varietals and hybrids of chili peppers belong to this variety ( Bosland and Votava 2000) and it is consequently the most intensively studied species of Capsicum with regard to diversity, domestication and genetics ( OECD 2006; Pickersgill 2016; Acquadro et al. 2020 and references therein). Conversely, a full comprehension of its taxonomy has not been achieved in the last 50 years. Research indicates that its domestication could have been initiated in central-east Mexico over 6,500 years ago ( Kraft et al. 2014).

Due to the selective pressure for domestication and diversification, defining a characteristic group of traits for var. Capsicum annuum annuum is difficult; however, the most distinctive features are its herbaceous to shrubby, annual or perennial habit, the solitary axillary flowers (rarely two or more), the strongly 5-10-nerved calyx, the large white (or purple) corollas (up to nearly 22 mm in diameter) and the usually persistent and pendent fruits, which are highly variable in size, form, colour and pungency. Some of these traits contrast with those of var. Capsicum annuum glabriusculum which has a shrubby habit, 5-nerved calyx, smaller corollas (≤ 12 mm in diameter) and small (<10 mm in diameter), globose, ellipsoid or ovoid, erect, red or red-orange, deciduous fruits.

Philip Miller was the curator of the Chelsea Physic Garden in London in the late 18th century. Many of the plants he grew there were new taxa in his “Gardener’s Dictionary" (1768). He described several Capsicum species ( C. cordiforme , C. tetragonum , C. angulosum , C. olivaeforme and C. pyramidale ), based on cultivated specimens obtained from seeds of different provenance. As was the practice at the time, he did not cite specimens and is likely to have based his descriptions on living plants. Most of these plants were described as annuals with white flowers and a variety of fruit sizes, shapes (heart-shaped, angular-obtuse, oval-shaped, pyramidal), colours (yellow, scarlet, red), textures and positions (pendent or upright), characters that are highly variable due to human selection in these domesticated species. Specimens made from plants grown by Miller are found in several different places, mostly at BM and its associated historical herbaria. As these names are almost certainly described from living plants and, thus, will need neotypification, we do not typify them here, but leave that for a separate study when these materials, including any non-digitised specimens, can be studied in detail.

We found a collection in the Lamarck Herbarium with a label indicating that it belongs to C. conicum (P00357734) which we designate here as the lectotype.

Capsicum bicolor was probably described only from living material cultivated in the gardens of Schönbrunn Palace near Vienna (Austria). Jacquin (1809) cited no specimens for C. bicolor and gave no place of origin for his species. The description is quite complete and includes a colourful illustration (Tab. 99, fig. 1); both fall within our circumscription of C. annuum . Since no specimens have been found, we designate the illustration here as the lectotype.

There are two sheets of original material labelled C. grossum in Willdenow’s Herbarium held at Berlin. Both contain reproductive branches; one of these (B-W 04425 -02 0) consists of two fruiting branches that exactly match Willdenow’s description ( Willdenow 1809) and is, therefore, designated the lectotype here.

Willdenow (1809) cited neither specimen nor locality when describing C. sphaericum . We found at B original material labelled " C. sphaericum , Hort. Bot. Berol. W" (B-W04426-01-0) and select it here as the lectotype.

In the protologue of C. purpureum , Hornemann (1813) stated "Herb. Vahlii … Hab. - -", referring to material cultivated at Horto Hafniensis (Hafnia = Haunia = Copenhagen). A specimen held in C [C10019148], with good flowering material and bearing a label (on the verso of the sheet) with data matching the protologue, is here selected as the lectotype. Additional original material is found at C (C10019147, upper stem only), which corresponds to a Herb. Hornemann specimen cultivated in Copenhagen; this specimen is sterile.

De Candolle (1813) described C. ovatum , based on a living specimen of unknown origin cultivated at the Montpellier Botanical Garden (hort Bot. Monspeliensis), but he cited no herbarium material. We found no original material at MPU; Dunal (1852) expanded the description in the Prodromus with specimens seen in the De Candolle Herbarium, now held in G-DC. At G-DC, there are two elements: a small fruiting fragment possibly from Montpellier ( “h.m.” [my herbarium] on the label) and a more complete specimen with a label stating " Capsicum ovatum D.C." in Dunal’s hand; we designate here this latter specimen (G00200072) as the neotype.

When coining the name C. longum , De Candolle (1813) cited in synonymy pre-Linnean works, some with illustrations. We examined the illustrations and the one from J. Bauhin et al. (1651) best illustrates and corresponds to the original diagnosis and, therefore, is selected here as the lectotype.

Meyer (1818) based his description of C. globiferum on collections made by Ernst Karl Rodschied in what is now Guyana. In Göttingen, where Meyer worked, we found two Rodschied specimens labelled as C. globiferum , both from Rio Essequibo, that are certainly original material. One (GOET003420) has what appears to be a collecting number, Rodschied 29, while the other lacks any indication of a number (GOET003419). We designate here the former and most complete specimen (Rodschied 29) as the lectotype.

Capsicum purpureum is based on a single plant found in the Botanic Garden of Calcutta (India), whose exact origin is unknown, but Roxburgh (1824) suggested the seeds came from the "Molucca Islands". A specimen at Kew (K001132446), labelled as " Capsicum purpureum " and with a faint annotation of “H.B.C.” at the bottom of the sheet, is possible original material, but we cannot be sure it was used by Roxburgh or when it was prepared. We, therefore, designate this sheet as the neotype of Capsicum purpureum .

Dierbach (1829) referred to his C. indicum var. vulgatum as " Capsicum annuum Auctor. plurimor." making this name superfluous. His species was characterised by its red, oblong and straight fruits, one of the most common forms of the domesticated C. annuum .

Fingerhuth (1832) described many domesticated chili pepper taxa (see above); he did not mention specimens and, if he made specimens, the fate of his herbarium is unknown ( Stafleu and Cowan 1976). Fingerhuth provided a set of ten plates, each one with figures in colour that illustrated almost all of the taxa included in his monograph. These figures are of good quality and match the protologues, which allowed us to assign the species to which they belong and to use them as lectotypes for Fingerhuth’s names. The names Capsicum strictum , C. ceratocarpum , C. angulosum (var. Capsicum angulosum macrocarpum and Capsicum ovale ), some varieties of C. annuum (var. Capsicum annuum rugosulum , Capsicum acuminatum , Capsicum subangulosum , Capsicum ovoideum , Capsicum abbreviatum , Capsicum olivaeforme ), C. grossum (var. Capsicum grossum pomiforme , Capsicum ovatum , Capsicum cordatum , Capsicum angulosum ) and C. longum (var. Capsicum longum incrassatum , Capsicum latum , Capsicum rectum ) are here lectotypified, based on the Fingherhuth figures indicated above.

In the protologue of C. abyssinicum , Richard (1850) cited two collections from “Abyssinia” (Ethiopia), one made by León Richard Quartin Dillon and the other by Antoine Petit, botanists on the Lefebvre expedition to the mountains of Africa. Quartin Dillon’s collection is housed at P and consists of three sheets (P00329903, P00329904, P00329905), all of them with complete flowering and fruiting branches. We were unable to find the second collection at P, but a duplicate of the original Petit collection is at MEL (MEL 2442182) and is also a well-preserved specimen. We designate here the best-preserved collection and that which Richard is likely to have seen and used (P00329903) as the lectotype.

Dunal (1852) coined C. annuum var. oblongum with a direct citation to a polynomial and illustration in "Fingerh. l.c. t. 2 f.a" ( Fingerhuth 1832); he also stated "v.s. in h. DC". A sheet in G-DC (G00131768) is labelled " Capsicum annuum α Capsicum oblongum fructibus rubris Fingerh." from "Herb. Dunal 1844", both in Dunal’s hand; we select this specimen as the lectotype.

In his description of C. pyramidale var. longicorne , Dunal (1852) cited the collection Zollinger 489 that he had seen in "h. Boiss. et h. DC." (now G and G-DC). Both specimens consist of fertile branches, but that in G-DC (G00131841) has an immature fruit which confirms the identity of this name and is selected here as the lectotype.

In the protologue of Capsicum testiculatum , Dunal (1852) stated "v.s. in h. Dc. et herb. meo". The original material came from plants grown in the Botanical Garden in Montpellier from seeds sent by R. de Visiani. The sheet in G-DC (G00200067) is a more complete fruiting branch which we select here as the lectotype.

In the protologue of C. angulosum var. macrocarpum , Dunal (1852) cited no specimens, but referred his variety to the illustration "tab. 8, fig. a" provided by Fingerhuth (1832) which is selected as the lectotype.

Dunal based C. leucocarpum on Miller’s (1752) polynomial " Capsicum americanum Capsicum latifolium , fructu Capsicum oblongo erecto candido", referring to a sort of white-coloured C. annuum fruit; Fingerhuth (1832) also transcribed Miller’s polynomial exactly, but he did not provide a formal name, as appears in Index Kewensis ( Capsicum leucocarpon Fingerh., accessed on 20 April 2020) or IPNI ( Capsicum leucocarpon Mill. ex Fingerh. (accessed on 20 April 2020).

Sturtevant (1888b) described C. fasciculatum , based on his own cultivated living material known as Bouquet rouge (French garden name) or Red Cluster (American name). He distinguished this species from C. annuum primarily by its peculiarly clustered leaves and fruits at the summit of the plant. We found no herbarium material corresponding to this name, but Sturtevant stated in the protologue that C. fasciculatum is "well figured under the name Tenjikumamori in a Japanese botanical work …” referring to Tanaka and Ono (1874). The figure cited by Sturtevant (1888b) is very accurate and represents very well one of the many variations of the domesticated C. annuum and is here selected as the lectotype.

Greenman (1903) based C. frutescens var. lanicaule on four syntypes and characterised it as a more pubescent variant of C. frutescens . We studied all the syntypes (Palmer 639, 640, 642 and González 975). Palmer’s collections are flowering and fruiting branches, while González’s specimen has flowers and very young fruits. The fruiting calyx is critical in assigning the correct placement of this name and, amongst the fruiting Palmer specimens, the most informative one is Palmer 639 which has mature fruits typical of C. annuum var. annuum and is designated here as the lectotype.

Capsicum velutinum ( De Wildeman 1922) was described based on five syntypes from different localities and collectors in what is now the Democratic Republic of the Congo, all of them well preserved at BR (Blommaert s.n., Jespersen s.n., Lamboray 22 and Lescrauwaet 315). All the specimens consist of strikingly pubescent flowering or fruiting branches or both. We designate Lamboray 22 (BR 000000649909) as the lectotype for this name, since it matches the protologue most closely.

Figures

Figure 20. Capsicum annuum var. annum. Lectotype (BM). Copyright The Trustees of the Natural History Museum, London. Reproduced with permission.

Figure 21. Capsicum annuum var. annuum A flower bud on pendent pedicel B flower with connivent anthers C flower with hexamerous corolla (note nectar droplets on the limb) and style near the same length as the anthers D flower with heptamerous purple corolla E flower with pentamerous corolla and style exceeding the anthers F, H mature fruits on pendent pedicels G mature fruits on upright pedicels A-H no specimen vouchers, photos by G. E. Barboza and C. Carrizo Garcia taken at different greenhouses.