Capsicum frutescens L., Sp. Pl. 1: 189. 1753.

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

19. Capsicum frutescens L., Sp. Pl. 1: 189. 1753.

Figs 67 , 68

Capsicum conoide Mill., Gard. Dict. ed. 8, Capsicum no. 8. 1768. Type. Cultivated at the Chelsea Physic Garden (England), seeds from Antigua (Antilles) (no specimens cited).

Capsicum conicum G.Mey., Prim. Fl. Esseq.: 112. 1818, nom. illeg., non Capsicum conicum Lam. (1794). Type. "In plantationibus" (no specimens cited) [Guyana]. Rio Essequebo, Sept, E.K. Rodschied 30 (lectotype, designated here: GOET [GOET003418]).

Capsicum minimum Roxb., Fl. Ind., ed. Carey & Wall. 2: 261. 1824, nom. illeg., not C. minimum Mill. (1768). Type. India. "East India" (no specimens cited), 18 Dec 1814, W. Roxburgh s.n. [Wallich Cat. No. 2641A) (lectotype, designated here: K [K001116724]).

Capsicum fastigiatum Blume, Bijdr. Fl. Ned. Ind. 13: 705. 1826. Type. " Crescit : in hortis et locis incultis … Nomen: Tjabe rawiet" [Indonesia]. Java, C.L. Blume s.n. (lectotype, designated here: L [L 0003564, acc. # 202560]).

Capsicum indicum Dierb. var. conoide (Mill.) Dierb., Arch. Apotheker-Vereins Nördl. Teutschl. 30(1): 28. 1829. Type. Based on Capsicum conoide Mill.

Capsicum indicum Dierb. var. berberideum Dierb., Arch. Apotheker-Vereins Nördl. Teutschl. 30(1): 29. 1829. Type. Based on Capsicum frutescens Mill. (= C. frutescens L.)

Capsicum baccatum Vell., Fl. Flumin.: 60. 1829 ( “1825”); Fl. Flumin. Icon. 2: t. 3. 1831 ( “1827”), nom. illeg., non Capsicum baccatum L. (1753). Type. Brazil. [Rio de Janeiro]: "Sponte crescit, et 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_006] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 3. 1831).

Capsicum conoide Mill. var. sulcatum Fingerh., Monogr. Capsic.: 15. 1832. Type. No locality cited (no specimens cited; lectotype, designated here: [illustration] Fingerhuth, Monogr. Capsic. Tab. III c. 1832]).

Capsicum conoide Mill. var. chordale Fingerh., Monogr. Capsic.: 15. 1832. Type. " Crescit in America et Indiis"; no specimens cited; lectotype, designated here: [illustration] Fingerhuth, Monogr. Capsic. Tab. III d. 1832]).

Capsicum flexuosum Sendtn. var. perrottetti Dunal, Prodr. [A. P. de Candolle] 13(1): 413. 1852. Type. French Guiana. Sin. loc., 1820, S. Perrottet 218 (holotype: G-DC [G00131855]).

Capsicum frutescens L. var. multilobatum Dunal, Prodr. [A. P. de Candolle] 13(1): 413. 1852. Type. No locality indicated, possibly from plants in cultivation (holotype: G-DC [G00131856]).

Capsicum conicum G.Mey. var. orientale Dunal, Prodr. [A. P. de Candolle] 13(1): 415. 1852. Type. Sudan. [Kurdufan] "ad pagum Cordofan in Milbes", 4 Dec 1839, G.C.T. Kotschy 292 (lectotype, designated here: G-DC [G00131905]; isolectotypes: BM [BM001016438, BM001016449], E [E00687052], G [G00442768], LE).).

Capsicum conicum G.Mey. [unranked “variat”] Capsicum latifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 415. 1852. Type. Sudan. “Senaar”, 1831, G. Acerbi s.n. (lectotype, designated here: G-DC [G00131885]).

Capsicum conicum G.Mey. [unranked “variat”] Capsicum angustifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 415. 1852. Type. Oman. Muscat "in horto Mascate", P.M.R. Aucher-Eloy 5039 (lectotype, designated here: G [G00390279]; isolectotype: G [G00390280]).

Capsicum crispum Dunal, Prodr. [A. P. de Candolle] 13(1): 415. 1852. Type. Republic of Mauritius. "Cult. au Jardin des Pamplemousses", 1839, L. Bouton s.n. (holotype: G-DC [G00131904]; isotype: MPU [MPU023045]).

Capsicum crispum Dunal var. piper-rabiosum Dunal, Prodr. [A. P. de Candolle] 13(1): 416. 1852. Type. France. Île de Bourbon [= Réunion Island], 1821, Anonymous s.n. (holotype: G-DC [G00131903]; isotype: MPU [MPU023044]).

Capsicum conoide Mill. var. oblongoconicum Dunal, Prodr. [A. P. de Candolle] 13(1): 415. 1852. Type. No locality cited (no specimens cited; lectotype designated here: [illustration] Fingerhuth, Monogr. Capsic. Tab. III, b. 1832).

Capsicum curvipes Dunal, Prodr. [A. P. de Candolle] 13(1): 423. 1852. Type: French Guiana. "In Guianâ (h. Moric.)", Anonymous s.n. (holotype: G [G00390275]; isotype: MPU [MPU023048].

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

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

Capsicum annuum L. var. subconicum Alef., Landw. Fl.: 133. 1866. Type: Based on Capsicum conoide Mill.

Capsicum annuum L. var. conoide (Mill.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 65. 1898, as " Capsicum conoides ". Type: Based on Capsicum conoide Mill.

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

Capsicum annuum L. var. parvo-acuminatum Makino, J. Jap. Bot. 3(8): 30. 1926. Type. Cultivated in Japan "Hab. JAPAN, cultivated" (no specimens cited, no original material located).

Capsicum frutescens L. var. queenslandicum Domin, Biblioth. Bot. 89: 572. 1928. Type: Australia. Queensland: "Nordost-Queensland: Harweys Creek", Jan 1910, K. Domin 8247 (holotype: PR [acc. # 530853]).

Capsicum annuum L. var. oblongo-conicum (Dunal) Cufod., Bull. Jard. Bot. État Bruxelles 33 (Suppl.): 860. 1963. Type: Based on Capsicum conoide Mill. var. oblongo-conicum Dunal.

Type.

" Habitat in India " Herb. A. van Royen n. 908.244-150 (lectotype, designated by Heiser and Pickersgill 1969, pg. 280: L [L 0053043, acc. # 902560, branch in the lower half of the sheet]) .

Description.

Low subshrubs or shrubs, herbaceous or woody at the base, 0.3-1.5 (-2.5) m tall, much branched from near the base. Young stems slightly angled, fragile, green or purple or green with purple ridges, glabrous to sparsely pubescent with simple, uniseriate, 4-9 (-11)-celled, eglandular trichomes 0.3-1.4 mm long; nodes solid, green or purple; bark of older stems brown; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair more or less similar in size and shape. Leaves membranous, concolorous or slightly discolorous, glabrescent or glabrous on both surfaces, sometimes with eglandular trichomes on the main veins or a tuft of trichomes in the vein axils abaxially; blades of all leaves (4-) 4.4-8.2 (-12.5) cm long, 2-4.5 (-6) cm wide, ovate or narrowly elliptic, the major veins 5-7 on each side of mid-vein, the base asymmetric, cuneate or attenuate, the margins entire, the apex acuminate to long-acuminate; petioles 0.5-3 cm, glabrescent or glabrous. Inflorescences axillary, 2-4 (-5) flowers per axil, rarely flowers solitary; flowering pedicels 9-30 (-40) mm long, erect, geniculate at anthesis, glabrous or sparsely pubescent; the eglandular trichomes minute, antrorse; pedices scars inconspicuous. Buds ovoid, cream or greenish-white. Flowers 5-7-merous, spreading or pendent. Calyx 1.2-2.5 mm long, cup-shaped, thick, green, glabrous or sparsely pubescent, the calyx appendages absent or five, ca. 0.5 mm long. Corolla 3.75-6.5 mm long, 5-15 mm in diameter, usually dull white or greenish-white outside and within, stellate with interpetalar membrane, lobed nearly the halfway to the base, pubescent adaxially with small glandular trichomes (stalk 1-2-celled; head unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 3-4 mm long, the lobes 3-5 mm long, 2-3.5 mm wide, the margins finely ciliate, the tips acute, papillate. Stamens five (-seven), equal; filaments 1-1.5 mm long, cream or purple, inserted on the corolla 1-1.6 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5-2.85 mm long, ellipsoid, bluish-grey or purplish or (rarely) dark green or yellow, connivent at anthesis. Gynoecium with ovary 2.5-4 mm long, 1.3-1.8 mm in diameter, oblong-ovoid, pale green; ovules more than two per locule; nectary ca. 0.3 mm tall; styles homomorphic, 3-4 mm, exserted 1.5-2 mm beyond the anthers, cream or purple, cylindrical; stigma 0.1 mm long, 0.25 mm wide, discoid, light green. Berry usually 9-30 (-60) mm long, 4-12 (-15) mm in diameter, usually elongate and narrowly triangular, with the apex pointed or blunt and the base narrowed, usually green and yellowish-green when immature with transitions to yellow, orange-red, orange-yellow, orange to red at maturity, deciduous or persistent, pungent, rarely non-pungent, pericarp thick and opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 16-35 (-50) mm long, usually erect, sometimes pendent, terete to strongly angled, widened distally, green; fruiting calyx (3-) 4-6 (-7) mm in diameter, persistent, somewhat accrescent, deeply cup-shaped, without an evident constriction in its base and the junction with the pedicel, sometimes the margin ripped, green. Seeds 10-52 per fruit, 2-4 mm long, 2.2-3.3 (-4) mm wide, C-shaped to subglobose, pale yellow, the seed coat smooth to obscurely reticulate (SM), reticulate-cerebelloid (SEM), the cells irregular in shape, the lateral walls slightly to strongly sinuate; embryo imbricate.

Distribution.

Capsicum frutescens has a broad distribution in lowland tropical America (Fig. 69 ), ranging from Brazil to Central America and the West Indies. It is not found in Chile, Argentina and Uruguay in the field, but it is sold in local markets ( Peralta and Liverotti 2018). The species has increased its worldwide distribution more as a weed than as a cultivated plant. In the Old World, it has been introduced and established as a weedy component or it has escaped from cultivation into the flora of Papua New Guinea ( Pickersgill 1989), Nepal ( Baral and Bosland 2002), Philippines ( Yamamoto and Nawata 2009) and Africa. It is cultivated in many countries around the world ( Bosland and Votava 2000; Yamamoto and Nawata 2005, 2009), especially in the United States of America (Louisiana), where the cultivar Tabasco is grown commercially ( Pickersgill 1971; DeWitt and Bosland 1997) and in Brazil, where the cultivar Malagueta is very popular ( Bosland and Votava 2000).

Ecology.

Capsicum frutescens grows in low semi-deciduous forests or in disturbed areas and agricultural clearings; it is commonly cultivated near homesites and in the chakras of many local communities, between 10 and 1,500 (2,000) m elevation.

Phenology.

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

Chromosome number.

2 n = 2x = 24 ( Pickersgill 1977, Moscone et al. 1996, 2007).

Common names.

Argentina: Tabasco (Salta, Hunziker 25489); Antilles: Bird pepper ( Pickersgill 1984); Belize: Bird pepper (Cayo, Atha et al. 1022), Chile (Cayo, Balick et al. 2272), Chili del monte (Cayo, Arvigo et al. 148); Bolivia: Ají (Beni, Williams 933), Arivivi (Santa Cruz: Krapovickas & Schinini 36348), Aribibi chico (La Paz, Williams 764), Aribibi silvestre (Beni, Scolnik & Luti 672); Brazil: Malagueta (Amazonas, Hill et al. 12996; Bahia, Thomas et al. 9080; Goiás, Vieira 709; Mato Grosso do Sul, Pott et al. 5562; Piauí, Pickersgill RU72-140; Rio Grande do Norte, Pickersgill RU72-357; Rondonia, Walter et al. 581), Malaguetinha, malaguetão (Roraima, Barbosa et al. 2006), Olho-de-caranguejo ( Paraná, Leitão s.n.), Pimenta camari (Minas Gerais, Grandi 261), Pimenta malaguêta (Amazonas, Hill 12996; Bahia, André et al. 6863; Espírito Santo, Folli 2995; Mato Grosso do Sul, Bortolotto & Rodrigues B-615; Minas Gerais, Pereira 3219; Paraíba, Agra 625; Tocantins, Eiten & Eiten 10140), Pimenta malaguetinha ( Amapá, Pereira & Severino 1855), Pimenta malaguetão ( Amapá, Pereira et al. 1836), Pimenta malagueta doce (Rondonia, Walter et al. 575); Colombia: Ají (Amazonas, Henao 167; Bolívar, Bro. Heriberto 231; Guainía, Triana 18; Magdalena, de Romero 72; Vaupes, Zarucchi 2173), Chivatillo ( Nariño, de Benavides 4692), Ají Ajuja ( Guainía, Marín & Rodríguez 508), Ají clavito (Amazonas, Torres & Rodríguez 2020), Ají chivatillo ( Nariño, Caballero 40), Ají churere ( Guainía, Marín & Rodríguez 509), Ají malagueta (Amazonas, Cárdenas et al. 9441), Aji pequeñito ( Caquetá, Cárdenas 9307; Guainía, Rodriguez 2), Ají largo ( Bolívar, Espina 586), Ají Quiñapira (Guaviare, Rodríguez & Coy s.n.), Ají de ajoja ( Vaupés, Rodríguez 112), Ají de blancos (Amazonas, La Rotta 273), Ají diente, chiche (Amazonas, Cárdenas et al. 9413), Ají pico de pájaro ( Bolívar, Espina 571), Diente de chucha (Amazonas, Cárdenas et al. 9412; Caquetá, Cárdenas et al. 9325); Costa Rica: Chili (Puntarenas, Tonduz 7255); Ecuador: Ají (Esmeralda, Yañez et al. 1407; Pichincha, Kvist 40201), Ají chivatillo (Esmeralda, Acosta Solías 13937; Pichincha, Acosta Solís 13927), Ají clavo, ají gallinazo (El Oro, van den Eynden & Cueva 654; Esmeraldas, Cerón 14561), Ají patate (Chimborazo, Angulo 007), Ají silvestre (Loja, Vivar C. 4003), Ají de ratón ( Manabí, Deanna 53); El Salvador: Chiltepe (El Salvador, Velasco 8925), Chilpepe, Chile de zope (Sonsonate, Standley 22237), Chile chocolate (San Salvador, Calderón 1198), Guatemala: Chili (Izabal, Standley 25055; Peten, Lundell 16480), Chile largo (Izabal, Standley 24297), Chile chiltepe (Izabal, Standley 23755); Haiti: Piment-chien (Ile La Tortue, Ekman s.n.); Honduras: Chile bravo ( Atlántida, Standley 53392), Pico de Pájaro ( Morazán, Molina R. 1177); Mexico: Chilpaya o Chilayate (Veracruz, Martínez A. 346), Chile chocolate (Chiapas, Matuda 17642), Chile picapalo (Tabasco, Becerril Pérez & Ortiz Cornejo 7), Chile tabaquero (Oaxaca, Alcocer & Morales s.n.), Pico Paloma (Tabasco, Ortega et al. 876), Chile del monte ( Yucatán, Gaumer 864), Chile pico paloma (Tabasco, Alegría O. 47), Diente de perro (Chiquimula, Kufer 100); Panamá: Ají (Balboa, Standley 25504); Peru: Arnaucho, Aji arnaucho (Peru, Dunal 1852), Malageta (Loreto, Martin & Lau-Cam 1242), Malaguete (San Martín, Libreros et al. 2013), Ají chuncho ( Junín, Smith s.n.), Ají mono, Ají pinguita de mono (Lambayeque, Llatas Quiroz 3452), Mala guctí ( Huánuco, Schunke V. 1397), Pipí de mono (Loreto, Libreros et al. 2013); Trinidad and Tobago: Bird pepper (Tobago, Broadway 4498; Trinidad, Heiser C 259); United States of America: Tabasco (Louisiana, Meyer & Mazzeo 11988); Panamá: Ají (Balboa, Standley 26493); Venezuela: Chirel (Margarita Island, Smith-Davis Ac 1495), Chirere (Carabobo, Pittier 7909), Aji pico pájaro (Margarita Island, Smith-Davis Ac 1498).

Indigenous names.

Belize: Mash-ík (Mayal, Cayo, Balick et al. 2272), Smash-ík (Cayo, Balick et al. 2391); Bolivia: Bido (Tacana, La Paz, Williams 764), Naris (Chiquitano, Santa Cruz, Del Aguila et al. 662); Colombia: Bee-a’ (Tucano, Amazonas, Schultes & Cabrera 13046), Biaá (Yucana, Amazonas, Cárdenas 9405), Cog (Puinave, Guainía, Triana 18), Eviviaa (Tucano, Guainía, Marín & Rodríguez 508), Fekorai (Huitoto-Mɨnɨka, Amazonas, Henao 167), Ferocoi (Huitoto, Caquetá, Cárdenas et al. 9300), Ichiriay (Matapi, Amazonas, Cárdenas et al. 9413), Jibirai ( Caquetá, Cárdenas 9345), Jichiri (Yucuna, Amazonas, Cárdenas et al. 9441), Lehirihay (Yucuma, Caquetá, Cárdenas et al. 9325), Meniray (Huitoto, Caquetá, Cárdenas 9307), Mèe (Colona, Amazonas, Torres & Rodríguez 2020), Pidootú (And, Amazonas, Castro & Andoke 608), Pipitatu (And, Amazonas, Castro & Andoke 606), Pxrxtú (And, Amazonas, Castro & Andoke 597), Tsirrerreji (Sikuare guahibo, Guainía, Rodriguez 2), Viaa (Kuboes, Guainía, Marín & Rodríguez 509); Ecuador: Aatyu (Chapalaachi, Esmeraldas, Yañez et al. 1407), Ampy (Shuar, Zamora-Chinchipe, Ortega et al. 51), Chimidu dio (Cayapa, Esmeralda, Kvist & E. Asanza 40456), Ma pipi pia (Siona and Secoya Indians, Napo, Vickers 208), Ocoma ( Cofán, Napo, Cerón 194), Panduchu (Achuar Jívaro, Pastaza, Lewis et al. 14011), Pía (Secoya, Sucumbíos, Miranda-Moyano 225), Pipetio (Cayapa, Esmeraldas, Kvist 40565), Pucuitape (Cayapa, Esmeraldas, Kvist & Asanza 40356), Sun’nyo pipi pia (Siona & Secoya Indians, Napo, Vickers 226), Tun (Colorado, Pichincha, Kvist 40201), Uchu (Pastaza, Lewis et al. 14011); Guatemala: Ich ( Ch’orti, Chiquimula, Kufer 100); Paraguay: Ki’ï (Central, Schinini & Bordas 24511); Peru: Shunaru’ca (Chayahuita, Loreto, Odonne 641), Tsukagka (Amazonas, Salaün 177), Yancuru’ca (Chayahuita, Loreto, Odonne 664). Mexico: Guiiña xcuuchu (Zapoteco, Oaxaca, Sánchez L. et al. 953).

Uses.

Capsicum frutescens has medicinal, ornamental, ritual and food applications. The fresh or cooked fruits are an important component of daily meals in India (soups, sauces, jams, pepper powder). The famous hot Tabasco sauce is made from tabasco peppers, a varietal of C. frutescens . In the Brazilian Amazon, herbarium labels report plants with sweet fruits (Walter et al. 575) that are used for decorating dishes and salads and in smoking rituals ( Barbosa et al. 2002, 2006). The leaves and fruits have many reported medicinal uses (see Table 3 ).

Preliminary conservation assessment.

EOO (19,757,841 km2); AOO (1,336 km2). Capsicum frutescens is a widespread cultivated species across the Americas and can be assigned the Least Concern (LC) status.

Discussion.

Capsicum frutescens belongs to the Annuum clade ( Carrizo García et al. 2016); it is a shrubby plant with 2-5 flowers per node, dull white or greenish-white corollas, bluish-grey or purplish anthers, upright elongate and narrowly triangular pungent fruits, a deeply cup-shaped fruiting calyx enclosing the narrowed fruit base and smooth, pale yellow seeds. It does not have as many cultivars as C. annuum and C. chinense ( Bosland and Votava 2000).

Morphoagronomic and molecular characterisation studies have been carried out on many accessions of C. frutescens from different germplasm banks ( Carvalho et al. 2017; Tripodi and Greco 2018; Brilhante et al. 2021), as well as on commercial crops ( Peñuela et al. 2020), with differing results as to the most useful morphological diagnostic descriptors for the species. Carvalho et al. (2017) identified three morphological patterns of fruits: malagueta type (the most common type with detailed characterisation), Tabasco type and malaquetinha type, based on qualitative and quantitative morphological descriptors and SSR markers in 103 accessions of C. frutescens from Brazil. Molecular results in the samae study recovered six groups with genetic variability between and within each group.

Many authors have recognised close affinities between C. annuum , C. frutescens and C. chinense (e.g. Jensen et al. 1979; McLeod et al. 1983b; Prince et al. 1995; Baral and Bosland 2004; Ibiza et al. 2012; Carrizo García et al. 2016; D’Agostino et al. 2018). Together, these species constitute the Capsicum annuum primary gene pool ( van Zonneveld et al. 2015).

Flowering specimens of C. frutescens and C. chinense are sometimes difficult to distinguish in herbaria when fruits are missing. Venation of flowering calyx is a good distinguishing feature between both species since, in C. frutescens , the main nerves are often completely immersed in the calyx surface (Fig. 68C, G ), while in C. chinense , they protrude conspicuously from the calyx (Fig. 47C, J, L ) or may protrude distally. In fruit, the distinction between these two taxa is remarkable; C. frutescens has deeply cup-shaped calyx enclosing the narrowed base of the elongate fruit (Fig. 68G, H ), the pedicels are usually erect and a constriction at the junction calyx-pedicel is totally absent. Capsicum chinense has a flat discoid calyx appressed or reflexed to the fruit base, the pedicels are usually pendent and a noticeable circular constriction at the junction with the fruiting calyx is present.

Capsicum conoide was coined by Miller (1768) for shrubby plants with conical, erect red fruits "about one half inch" long that he grew from seeds received from Antigua under the name "Hen Pepper". The identity of C. conoide has been controversial in literature. Dierbach (1829) considered C. conoide an infraspecific category of C. indicum and it was submerged under C. annuum var. glabriusculum by D’Arcy and Eshbaugh (1974), due to the size of the fruit. Fingerhuth (1832) accepted C. conoide , added two varietal names ( C. conoide var. sulcatum and var. chordale ) and provided illustrations for all of them. Irish (1898) recognised C. conoide as a variety of C. annuum , assuming a wide concept for this latter taxon with the inclusion of garden cultivars currently accepted in different species (Heiser and Pickersgill 1975). We found no original material for C. conoide , but we interpret this name (and Fingerhuth’s varieties) as synonyms of C. frutescens , based on the descriptions.

For C. conicum , Georg Meyer (1818) cited no specimens in his "Florae Essequeboensis" (Guyana). At GOET, we found original material in Herbarium Meyer (Rodschied 30) with ' Capsicum frutescens ' written in Rodschied’s hand. This matches the text stated in the protologue of C. conicum . We select this sheet (GOET003418) as the lectotype.

When Roxburgh used the name Capsicum minimum for the first time ( Roxburgh 1814: 17), he provided no descriptions, but he stated that the plant was from India. Later, the name was validly published ( Roxburgh 1824), accompanied by a short description and the vernacular names "East Indian Bird chilly or Cayenne-pepper capsicum". In the Wallich Herbarium at Kew, there are two sheets, one from Francis Buchanan-Hamilton’s Herbarium (K001116724) labelled with the Wallich catalogue number 2641A and the other from "hort. Bot. Calcutta" with the Wallich catalogue number 2641B (K001116725). These are not duplicates (see discussion of Wallich’s catalogue numbering system at http://wallich.rbge.info/). The sheet from Buchanan-Hamilton’s Herbarium (K001116724) has several labels, the upper left of which is in Roxburgh’s handwriting (see Forman 1997), stating " Capsicum minimum /Botanical garden 18 Decr 1814". The other sheet in the Wallich Herbarium (K001116725) has no specific label with either the scientific name or place of origin; this sheet is not likely to be a duplicate and should not be considered original material. A sheet in G-DC (G00131883) is a good match for the centre stem on Wallich Cat. No. 2461B and is potentially a duplicate. We consider the sheet from the Buchanan-Hamilton herbarium (K00116724) to be the only authentic original material that we have seen and we here designate it the lectotype for C. minimum .

The description of C. fastigiatum ( Blume 1826) cited no herbarium material. A specimen at L collected by Blume in Java with the handwritten inscription Capsicum fastigiatum is here considered original material, since many of Blume’s collections are housed at L ( Stafleu and Mennega 1993). Heiser and Pickersgill (1969) commented that they examined the lectotype of C. fastigiatum , probably alluding to the specimen at L, but they were not specific. We designate here the sheet in Leiden (L 0003564) (presumably the sheet referred to by others) as the lectotype for C. fastigiatum .

Capsicum conoide var. sulcatum and var. chordale were both coined by Fingerhuth (1832), with no specific herbarium material cited. However, illustrations were provided for each variety that depict small fruiting branches with the elongate fruit typical of C. frutescens . We have selected these illustrations as the lectotypes for both of these varietal names.

In describing C. frutescens var. multilobatum , Dunal (1852) based its description on a specimen "v. s. in h. DC", with no additional information. A sheet in G-DC (G00131856) has no locality label, but is annotated " Capsicum frutescens β Capsicum multilobatum " in Dunal’s hand; we interpret this as the holotype.

Dunal (1852) described an infraspecific category for C. conicum G. Mey. named β [var.] Capsicum orientale that he distinguished by its woody branches and smaller leaves. Within it, he recognised two unranked categories ( “Variat”, that is, it varies) Capsicum latifolium and Capsicum angustifolium , for which he cited specimens. The two specimens, cited for the unranked Capsicum latifolium (Kotschy 292 and Acerbi s.n.), are mounted on a same sheet and they unequivocally belong to C. frutescens , based on the shape of the fruit and morphology of the fruiting calyx. We select Kotschy 292 (G00131905) as the lectotype for var. Capsicum frutescens orientale , because it has duplicates in many herbaria. In order to avoid homotypy for [unranked “Variat”] Capsicum latifolium , we select the other collection Dunal used for this name (Acerbi s.n. G00131885), mounted on the same sheet as the Kotschy collection, as the lectotype. For [unranked “Variat”] Capsicum angustifolium , Dunal cited a collection made in Muscat by P.M.R. Aucher-Eloy in "h. Boiss."; two sheets of Aucher-Eloy 5039 are held in the general herbarium at G and we select the better of these (G00390279) as the lectotype.

In describing C. conoide var. oblong-conicum , Dunal cited no herbarium material, but he referred to the Fingerhuth’s illustration "Tab 3, f. b" as the element he has seen. Therefore, he is clearly differentiating this illustration from C. conoides and it is the original material for the varietal name. We select this illustration as the lectotype.

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 67. Capsicum frutescens A reproductive branch B eglandular trichome of the leaf C flower D section of the calyx showing the venation E glandular trichome of the abaxial surface of the calyx F opened corolla G glandular trichome of the abaxial surface of the corolla H, I anthers, dorsal and ventral views, respectively J gynoecium K anatomical detail of the pericarp (note the giant cells in the mesocarp) L seed M seed, in cross section N structure of seed coat at the seed margin O structure of seed coat at the seed body P embryo. From Hunziker 25489. Drawn by N. de Flury.

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.

Figure 69. Distribution of C. frutescens.