Capscium baccatum L. var. pendulum (Willd.) Eshbaugh, Taxon 17: 51. 1968.

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

2b. Capscium baccatum L. var. pendulum (Willd.) Eshbaugh, Taxon 17: 51. 1968.

Fig. 28

Capsicum pendulum Willd., Enum. Pl. [Willdenow]: 242. 1809. Type. Cultivated in the Berlin Botanic Garden, Germany "Habitat ... [Country unknown]. Cult. in Hort. Bot. Berol"., C.L. Willdenow s.n. (lectotype, designated here: B [B-W04431-01-0]).

Capsicum frutescens L. var. pendulum (Willd.) Besser, Cat. Jard. Bot. Krzemieniec: 29. 1816. Type. Based on Capsicum pendulum Willd.

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

Capsicum pendulum Willd. var. majus Dunal, Prodr. [A. P. de Candolle] 13(1): 425. 1852. Type. No locality cited (no specimens cited; no original material located; Dunal may have considered this the typical variety).

Type.

Based on Capsicum pendulum Willd.

Description.

Erect shrubs or perennial herbs (0.60-) 1-2.5 m tall, with the main stem 1.5-2.5 cm in diameter at base, much branched from near the base and above, the branches spreading in a typical “zig-zag” appearance. Young stems 3-4-angled, fragile, dark green or green, mostly glabrous to sparsely or moderately pubescent with appressed-antrorse, short to long, simple, uniseriate, 4-9-celled, eglandular trichomes 0.5-1.3 mm long; nodes green; bark of older stems green with light brown fissures, glabrous; lenticels absent or few. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, equal or subequal in shape. Leaves membranous, discolorous, dark green above, light green beneath, glabrous to glabrescent with short eglandular trichomes in margins and long, spreading, 5-9-celled, eglandular trichomes along the primary veins and in the vein axils beneath; blades of major leaves 5-12 (-14.5) cm long, 3-5 (-7) cm wide, ovate, the major veins 5-7 on each side of mid-vein, the base asymmetric and attenuate or symmetric and rounded, the margins entire, the apex acute or acuminate; petioles 2.5-7.5 cm long, sparsely pubescent; blades of minor leaves 3-5.8 cm long, 1.3-2.5 cm wide, ovate or elliptic, the major veins 3-4 on each side of mid-vein, the base rounded, the margins entire, the apex acute; petioles 1.7-2 cm long, sparsely to moderately pubescent. Inflorescences axillary, 2-3 flowers per axil or flowers solitary; flowering pedicels 20-50 mm long, terete or angled, erect, sometimes curved, geniculate at anthesis, glabrous, glabrescent to moderately pubescent, the trichomes short, antrorse; pedicels scars inconspicuous. Buds globose, white with greenish-yellow spots, occasionally purple. Flowers 5-8-merous. Calyx 2-3 mm long, 3-4.2 mm wide, cup-shaped, thick, strongly 10-nerved, green, glabrous or glabrescent, the calyx appendages 5-6, rarely up to 8, 0.9-2.5 mm long, 0.2 mm wide, subequal, thick, erect, cylindrical, inserted close to the margin, with the same pubescence as calyx tube. Corolla 8.5-15 mm long, 12-16 (-20) mm in diameter, thick, white with greenish-yellow to tan spots and a white centre outside and within, rotate to rotate-stellate, with interpetalar membrane, lobed less than 1/3 of the way to the base, pubescent adaxially with short glandular trichomes (stalk 1-3-celled; head globose, peltate, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 4-5 mm long, the lobes 3-3.3 (-5) mm long, 3.5-6 mm wide, triangular to broadly triangular, spreading, the margins finely ciliate, the tips acute, papillate. Stamens 5-8, equal; filaments (2.5-) 3-4 mm long, white, inserted on the corolla 1.2-1.5 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 2-2.6 mm long, ellipsoid or ovoid, yellow, brownish post-dehiscent, not connivent (rarely connivent) at anthesis. Gynoecium with ovary 2.5-3 mm long, 1.6-2.5 mm in diameter, 2-5-carpelar, light green, ovoid; ovules more than two per locule; nectary ca. 1.2 mm tall, yellowish-green; styles dimorphic, short style 1.3-2 mm long, not exceeding the anthers length, long style 2.5-3.5 mm long, at the same level or slightly exserted beyond the anthers, yellowish-white, cylindrical; stigma ca. 0.2 mm long, 0.7-0.8 mm wide, discoid or bilobed, pale green. Berry 20-180 mm long, (10-) 20-40 (-50) mm in diameter, usually elongate or elongate-curved, triangular or campanulate, rarely subglobose, the base truncate or obtuse, the apex rounded, blunt or pointed, dark green or green when immature, green, bright yellow, orange, brown or red at maturity, persistent, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels (35-) 50-95 mm long, pendent, sometimes strongly curved, terete or slightly angled, widened distally, green; fruiting calyx 9-18 (-20) mm in diameter, persistent, slightly accrescent, campanulate, thick, somewhat corrugated or not, green, the appendages 0.5-2 mm long, appressed to the berry. Seeds 30-80 per fruit, (3-) 4-5.2 mm long, 3-3.8 (-4) mm wide, reniform or C-shaped, pale yellow to yellow, the seed coat smooth or slightly reticulate (SM), minutely reticulate (SEM), the cells polygonal to irregular in shape, the lateral walls straight to wavy; embryo imbricate.

Distribution.

Capsicum baccatum var. pendulum is found from low to mid-Andean elevations, mainly in Bolivia and Peru, extending to Ecuador and Colombia in the north and reaching Argentina, Paraguay and south-western Brazil in the south (Fig. 27 ). Introductions to North and Central America ( D’Arcy and Eshbaugh 1974; Thampi 2003), northern Brazil (Roraima, Barbosa et al. 2006), Europe ( Rodríguez-Burruezo et al. 2009), Japan ( D’Arcy and Eshbaugh 1974) and India ( Thampi 2003; Rudrapal and Sarwa 2020) are recent.

Ecology.

Capsicum baccatum var. pendulum is a cultivated plant adapted to many different ecological conditions between 150 and 3,400 m elevation.

Phenology.

Flowering and fruiting all year.

Chromosome number.

2 n = 2x = 24 ( Pickersgill 1977; Moscone et al. 2003, 2007).

Common names.

Argentina: Ají (San Juan, Ariza Espinar 3214; Corrientes, Benítez 76); Varita (Salta, Krapovickas & Schinini 28134), Ají picante (Salta, Hunziker 25496); Ají vainilla (Salta, Hilgert 1363), Puta parió (Corrientes, Martínez Crovetto 11125), Varita larga (Salta, Krapovickas & Schinini 28132), Ají huevo de gallo (Salta, Hilgert 1374); Bolivia: Ají (Santa Cruz, Williams 696; Tarija, Krapovickas & Schinini 39321), Aribibe (Santa Cruz, Hurtado 296), Aribibi (Santa Cruz, Heiser C281a, La Paz, Debouck et al. 3016), Ulupica (Tarija, Krapovickas & Schinini 31056), Aji acabeche (La Paz, Hinojosa & Wásra 1133), Ají amarillo (Cochabamba, Moscone 205), Aji Picante (Santa Cruz, Krapovickas & Schinini 32133), Locato largo (Santa Cruz, Heiser C290), Ají churcu, Ají rojo, Ají redondo, Pimentón colorado, Ají colorado gigantón (Santa Cruz, Libreros et al. 2014), Asta de buey amarillo, Huacareteño Duraznal, Huacareteño naranjo, Pa púca, Plomadita amarillo, Plomadita rojo, Chicotillo grueso, Huacarateño ancho amarillo, Cola de ratón amarillo, Cola de ratón naranjo (Chuquisaca, Libreros et al. 2014), Chicotillo, Huacareteño, Huacareteño amarillo, Ají amarillo, Ají colorado, Cumbaru rojo, Asta de buey, Asta de toro, Astay toro amarillo, Astay toro rojo, Astay toro anaranjado, Puntay lanza rojo, Puntay lanza anaranjado, Astay buey rojo (Chuquisaca, Jäger et al. 2013), Lata y Toro (Cochabamba, Moscone 209); Brazil: Pimenta (Rio de Janeiro, Krapovickas et al. 23428), Pimenta ardida ( São Paulo, Alves de Paiva 01), Chifre de Veado ( São Paulo, Heiser BGH 952), Dedo de moça ( Paraná, Freire de Carvalho s.n.; Roraima, Barbosa et al. 2006), Pimenta de Passarinho ( Espírito Santo, Mori et al. 11603), Pimenta dedo-de-moça ( Amapá, Pereira et al. 1819; Minas Gerais, Vianna s.n.; Paraná, Leitão s.n.), Pimenta de cheiro amarela ( Amapá, Pereira et al. 1830), Pimenta de cheiro vermelha ( Amapá, P ereira & Severino 1851), Pimenta-unha-de velha ( Río de Janeiro, Borges 66); Chile: Chota cabra (Santiago, Krapovickas 22232); Paraguay: Ají ( Paraguarí, Galander 1877), Locotito ( Paraguarí, Williams et al. 127), Locotito chico ( Guairá, Williams et al. 121); Peru: Ají (San Martín, Belshaw 3210), Ají escabeche (Ancash, Francia 42); Amarillo Moquegua (Lima, Velarde Nuñez 6), Amarillo Tacna (Lima, Velarde Nuñez 23), Amarillo Trujillo (Lima, Velarde Nuñez 10), Cilindro amarillo (Lima, Velarde Nuñez 13), Cilindro colorado (Lima, Velarde Nuñez 15), Colorado Tacna (Lima, Velarde Nuñez 24), Escabeche amarillo (Lima, Velarde Nuñez 11), Escabeche intermedio (Lima, Velarde Nuñez 27), Escabeche Lurín (Lima, Velarde Nuñez 2), Escabeche Moquegua (Lima, Velarde Nuñez 3), Extra Lurín (Lima, Velarde Nuñez 21), Escabeche colorado ENA (Lima, Velarde Nuñez 1), Ají Amarillo, Ají escabeche ( Bosland and Votava 2000), Ayuyo, Challuaruro (San Martín, Libreros et al. 2013), Ají uña de gallina (Lambayeque, Libreros et al. 2013).

Indigenous names.

Argentina: Keuí (= picante) (Corrientes, Hunziker 7339); Bolivia: Kîî ( Guaraní, Santa Cruz, Roca 0689); Paraguay: Ky y’ ( Guaraní, Cordillera, Williams et al. 135), Pimenta í ( Guaraní, Guairá, Williams et al. 121).

Uses.

This domesticated variety, mostly known as ‘Ají’, ‘Ají amarillo’ or ‘Ají escabeche’, is an important component of the diet of the Bolivian and Peruvian native population, less so in Argentina, Brazil, Ecuador and Colombia. In Bolivia and Peru, many different pod types occur that differ both in morphological (shape, colour, size) and biochemical attributes (e.g. capsaicinoids, vitamin E, flavonoids and quercetin content and antioxidant capacity). These forms are consumed in regional cuisines as spices and vegetables, fresh or dehydrated and ground ( Libreros et al. 2013, 2014).

Preliminary conservation assessment.

EOO (11,296,813 km2); AOO (356 km2). Capsicum baccatum var. pendulum is a very widespread cultivated plant and can be assigned a category of Least Concern (LC).

Discussion.

Capsicum baccatum var. pendulum is a member of the Baccatum clade ( Carrizo García et al. 2016). Morphologically, the corollas are similar in shape and colour to those of its wild progenitor, but they are larger, reaching 20 mm in diameter at anthesis; more consistent differences are the position of the fruiting pedicels and fruits (pendent), fruits that vary in size, shape and colour, the number of seeds (up to 80 per fruit) and the larger seeds (3-5.2 mm long, 3-4 mm wide). The presence of heterostylous flowers is more frequent in var. Capsicum baccatum pendulum than in var. Capsicum baccatum baccatum (Barboza, pers. obs.). This is an uncommon character within Capsicum that has been observed in both domesticated ( Perera and Poulos 1993; Peña-Yam et al. 2019) and wild species ( C. benoistii and C. tovarii ).

Many studies have been carried out that demonstrate the potential of this domesticated form in crop improvement. Capsicum baccatum var. pendulum encompasses a wide range of fruit morphology (e.g. fruit weight, size, shape and ripe colour), health-related compounds (flavonoids, polyphenols, quercetins, vitamin E, ascorbic acid, fat, amongst others) and capsaicinoids content (low to mild) ( Rodríguez-Burruezo et al. 2009; Kollmannsberger et al. 2011; Libreros et al. 2013, 2014; van Zonneveld et al. 2015). Furthermore, a particularly strong vegetable-like aroma has been detected in the fruits, due to diverse volatiles in accessions from Ecuador and Peru, with additional fruity/exotic notes in some genotypes ( Kollmannsberger et al. 2011).

Willdenow (1809) provided the description of C. pendulum , based on cultivated material of unknown provenance. In Willdenow’s Herbarium housed at B, we found a sheet with original material (B-W04431-01-0) consisting of a fruiting specimen with the inscription " C. pendulum , Hort. bot. Berol. W."; this sheet is designated the lectotype.

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 27. Distribution of C. baccatum var. baccatum, C. baccatum var. pendulum and C. baccatum var. umbilicatum.

Figure 28. Capsicum baccatum var. pendulum A plant B flower bud C flower, in pre-anthesis D flower short-styled, longitudinal section E flower long-styled, lateral view F-H flowers hexamerous showing connivent anthers and not connivent anthers I immature fruit J-M mature fruit A, B, E from Barboza 4886 C, D, I, K, L, M no specimen vouchers (cult. Pairumani, Cochabamba-Bolivia) F, G, H from Palombo 3 J from Barboza et al. 4824. Photos by G. E. Barboza.