Capsicum pubescens Ruiz & Pav., Fl. Peruv. [Ruiz & Pavon] 2: 30. 1799.

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

36. Capsicum pubescens Ruiz & Pav., Fl. Peruv. [Ruiz & Pavon] 2: 30. 1799.

Figs 103 , 104

Capsicum violaceum Kunth, Nov. Gen. Sp. [H.B.K.] 3: 49. 1818. Type. Ecuador. Pichincha: Quito, [no date], F.W.H.A. von Humboldt & A.J.A. Bonpland 3027 (holotype: P [P00670654]).

Capsicum quitense Willd. ex Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 809. 1819. Type. Ecuador. Pichincha: In Quito (holotype: B [B-W04433-01-0]).

Brachistus Brachistus ? Brachistus lanceifolius Miers, Ann. Mag. Nat. Hist. ser. 2, 3(16): 267. 1849, as " lanceaefolius ". Type. Ecuador. Loja: Loja, Aug 1847, B.C. Seemann 879 (lectotype, designated here: K [K000585923]; isolectotype: BM [BM000992131].

Capsicum maximowiczii Regel & Rach, Index Seminum [St. Petersburg (Petropolitanus)]: 40. 1859. Type. Cultivated in St. Petersburg, Russia [protologue "Cultum in hortis circa Valparaiso sub nomine "Agi dulce". Semina misit Maximowicz. (Rch.)"] "Ex horto bot. Petropolitano", 27 May 1858, L.T. Rach (no specimens cited; lectotype, designated here: LE).

Capsicum pubescens Ruiz & Pav. var. oviforme Hassk., Bonplandia (Hanover) 8(6): 95. 1860. Type. "Ab incolis Peruviae" (no specimens cited; no original material found).

Capsicum lanceifolium (Miers) Kuntze, Revis. Gen. Pl. 2: 449. 1891, as " lanceaefolium ". Type. Based on Brachistus ? Brachistus lanceifolius Miers.

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

Type.

Peru. Pasco: "Ex Pozuzo" [protologue - "Habitat affatim in Peruviae cultis, praesertim ad Panatahuarum Provinciam et in Andium nemoribus"], H. Ruiz & J. Pavón s.n. (lectotype, designated here: MA [MA-815154]; isolectotypes: CORD [CORD 00101751, fragment from G], G, MA [MA-815153, MA-815155]) .

Description.

Erect and scandent shrubs or perennial herbs, 1-3 (-4) m tall, with the main stem ca. 3 cm in diameter at base, much branched above, the branches in a typical “zig-zag” appearance. Young stems angled, fragile, green or green with purple spots and dark brown ridges, glabrescent to densely pubescent with a soft whitish pubescence of long, spreading, simple, uniseriate, 4-11-celled, eglandular trichomes (some forked) 0.5-1.5 mm long; nodes frequently dark purple; bark of older stems dark brown, smooth or striate, glabrescent to densely pubescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair markedly unequal in size, similar or dissimilar in shape. Leaves membranous, concolorous or discolorous, green above, light green beneath, rugose (the youngest leaves) or somewhat smooth with the mid-vein and primary veins raised abaxially, glabrescent to densely pubescent on both surfaces and margins, with similar trichomes like those on stems; blades of major leaves 6-12 (-16) cm long, 2.4-4 (-7) cm wide, ovate or more rarely elliptic, the major veins 4-6 on each side of mid-vein, the base asymmetric and attenuate or cuneate, the margins entire, the apex acuminate; petioles 1.5-2 cm long, densely pubescent to glabrescent; blades of minor leaves (2.5-) 3.5-5 (-6) cm long, 1.8-2.4 cm wide, ovate, the major veins 3-4 on each side of mid-vein, the base rounded, the margins entire, the apex acute or acuminate; petioles 0.5-0.7 cm long, densely pubescent to glabrescent. Inflorescences axillary, 1-2 flowers per axil, rarely up to four flowers; flowering pedicels 15-25 mm long, angled, erect, geniculate at anthesis, green or purple-ribbed, moderately to densely pubescent, the eglandular trichomes long, spreading; pedicels scars conspicuous. Buds globose or ovoid, dark purple. Flowers 4-8-merous. Calyx 2-3 mm long, ca. 4-4.3 mm wide, cup-shaped, thick, green, moderately to densely pubescent with the same trichomes as pedicels, sometimes sparse forked trichomes, the calyx appendages 4-8, (0.3-) 0.5-1.5 (-1.7) mm long, subequal or somewhat unequal, thin, erect, cylindrical, green, inserted close to the margin, pubescent with the same trichomes as calyx tube. Corolla 10-15 mm long, 15-22 (-25) mm in diameter, thick, dark purple or violet with a white centre outside and within (sometimes with a weak yellowish-green centre within), rotate to stellate, with thin interpetalar membrane, lobed 1/3 or a little more of the way to the base, pubescent adaxially with short glandular trichomes (stalk 1-2-celled; head globose, unicellular) in the throat and base of the lobes, the tube 5-8 mm long, glabrous abaxially, the lobes 3.5-6.5 (-7.2) mm long, 4.7-7.4 (-8.5) mm wide, broadly triangular, spreading, with eglandular trichomes abaxially especially on the veins, the margins pubescent with very short purple eglandular trichomes, the tips acute or obtuse, cucullate or not, sometimes papillate. Stamens 4-8, equal; filaments 2-3.3 (-4.25) mm long, purple or lilac, inserted on the corolla 1.4-1.6 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 2-2.8 mm long, ellipsoid or ovoid, purple with a wide cream connective, not connivent at anthesis. Gynoecium with ovary 2-3-carpelar, 2-2.5 mm long, ca. 2.5 mm in diameter, light green, ovoid or pear-shaped; ovules more than two per locule; nectary ca. 1.2 mm tall; styles heteromorphic, short, 3-3.5 mm, not exceeding the anthers, medium near the same length as the anthers or long 4.5-5.2 mm, exserted 1-1.5 mm beyond the anthers, lilac or purple, clavate; stigma 0.3 mm long, 0.7 mm wide, discoid or slightly globose, light green. Berry 20-40 mm long, 17-25 mm in diameter (semi-domesticated plants) or larger up to 50 mm long, 55 mm in diameter (cultivated plants), round, blocky or elongate-curved or not, the base obtuse, truncate or lobate, sometimes narrowed forming a neck-like, the apex blunt or sunken, rarely pointed, green when immature, brightly coloured at maturity (from red to light yellow or blackish), persistent, very pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 35-50 (-55) mm long, pendent, stout, curved or not, strongly angled, widened distally, usually green; fruiting calyx 8-13 mm in diameter, persistent, slightly accrescent, discoid, green, the appendages 0.8-2.1 mm long, ca. 0.3-0.4 mm wide, spreading. Seeds 15-45 per fruit, 5.5-7 mm long, 4.8-6 mm wide, C-shaped, subglobose or irregular, brownish-black to black, the seed coat reticulate (SM), reticulate-cerebelloid (SEM), the cells polygonal or irregular in shape, the lateral walls wavy to sinuate in the seed body, straight at margins; embryo imbricate.

Distribution.

Capsicum pubescens is native to Bolivia and Peru (Fig. 105 ), probably originating in mid-elevation southern Andes ( Eshbaugh 1976, 1993; Pickersgill 1984), but confirmed wild populations have not been recorded. Currently, this species is confined to the Americas being moderately cultivated in North America (mainly in Mexico) and Central America and more intensively along the Andean Region of South America, especially from Colombia to Bolivia and less in northern Argentina (Salta and Jujuy), Chile (Arica and Parinacota, cf. Tapia and Campos 2016) and Venezuela. Cultivation of C. pubescens outside the Americas is very restricted. A few populations have been reported for Asia (north-western China and Tibetan mountains, Djian-Caporalino et al. 2007) and an introduction was confirmed in Java, Indonesia ( Yamamoto et al. 2013). Recent introductions as cash crops were recorded in Nagano, Japan ( Matsushima et al. 2010) and a mention as a potential crop in the United Kingdom ( Samuels 2014).

Ecology.

Capsicum pubescens is frequently found in the Andean mid-elevations to highlands from (800-) 1,200-3,500 m and rarely below 500 m elevation. In Indonesia, the cultivation of C. pubescens occurs also in highlands, over 1,400 m elevation ( Yamamoto et al. 2013).

Phenology.

Probably flowering and fruiting all year, depending on the cultivation area.

Chromosome number.

2 n = 2x = 24 ( Pickersgill 1977, 1991; Moscone et al. 1993, 1995, 1996a, 2007).

Common names.

Bolivia: Locato (La Paz, Heiser C272), Locoto (La Paz, Lewis 88629; Santa Cruz, Saldías P. 563), Locote (La Paz, Duke & Winters 17330), Locotito (Santa Cruz, Vargas C. 932); Colombia: Ají ( Nariño, López Jurado & Riascos 613; Putumayo, Bristol 1115), Ají rocoto (Huila, Romero Castañeda 6674); Ecuador: Ají (Chimborazo, Moina Z 23; Loja, Ellemann 66689; Tungurahua, Cascante 6), Ají rocoto (Azuay, Steyermark 52690; Pichincha, Mejía 002); Guatemala: Siete caldos, Caballo ( Meckelmann et al. 2015), Chile cuadrocaldo (San Marcos, Steyermark 36930), Chile caballo (San Marcos, Steyermark 36930); Honduras: Chile garrapata (Alta Verapaz, Standley 91227), Chile petenero ( Morazán, Valerio R. 3237); Mexico: Chile (Morelos, Aguilar P. s.n.), Jalapeño, Perón, Manzano, Ciruelo (Laborde et al. 1982), Chile cera (Veracruz, Ventura A. 9739), Chile manzano ( México, Monsalvo J. 12; Michoacán, Soto Núñez et al. 6369), Chile pimiento ( México, Bonilla Beas 346), Chile canario (Oaxaca, García R. & Montaño M. 348; Veracruz, Castillo C. et al. 1757), Chile de cepa (Veracruz, Chazaro B. 2558), Chile de cera (Veracruz, Calzada 10856), Chile gordo (Veracruz, Castillo C. et al. 214), Morrón (Veracruz, Castillo C. et al. 1757), Moro Ich (Chiapas, Sánchez León 1139); Peru: Alú, ahi (Loreto, Killip & Smith 28864), Ají (Loreto, Williams 3405), Locoto ( Junín, Ochoa 602), Rocobo (Pasco, Ruiz 1940), Rocoto (Ancash, Gamarra 439; Cajamarca, Campos & Nuñez 4266; Cuzco, Cook & Gilbert 1017; Lima, Cerrate de Ferreyra 7644; San Martín, Quipuscoa & Bardales 979; Pasco, Chuck 137), Roccoto ( Ruiz and Pavón 1799), Ají rocoto (Andahuaylas, West 3739); Venezuela: Ají ( Mérida, Pittier 12707), Ají vocato (Humboldt & Bonpland & 3027)

Indigenous names.

Bolivia: Uchu rocoto (= ají globoso) (Rentzell s.n.); Colombia: Totsha ( Kamsá, Putumayo, Bristol 1115); Ecuador: Huchu (Quechua, Loja, Ellemann 66689).

Uses.

The fruits of Capsicum pubescens are one of the most appreciated in the Andean cuisine for their unique aroma, flavour, meatiness, juiciness and pungency ( Meckelmann et al. 2015). Fruits are used fresh, cooked or in powder (as condiment) in different ways in traditional and popular meals ("rocoto relleno", “ceviche”, "picante de gallina", "pique macho", cfr. Meckelmann et al. 2015; Tapia and Campos 2016; Barboza, pers. obs.). In Mexico, fruits are also consumed as a spice and in a great variety of industrial products ( Montes Hernández 2010). In South America, indigenous communities (Saraguro people, Ecuador) have attributed medicinal properties to leaves and fruits; in Peru, fruits are used in veterinary practice (see Table 3 for details). In Indonesia, immature and mature fruits are consumed as vegetables and spices ( Yamamoto et al. 2013).

Preliminary conservation assessment.

EOO (7,150,643 km2); AOO (536 km2). Capsicum pubescens is a widespread cultivated species across the Americas and can be assigned the Least Concern (LC) status.

Discussion.

Capsicum pubescens belongs to the Pubescens clade ( Carrizo García et al. 2016). The origin and affinities of C. pubescens are being analysed in depth and preliminary results (using genome-wide SNP data obtained through RAD-sequencing) show C. pubescens is sister to a clade formed by C. eximium , C. eshbaughii and C. carde -nasii ( Carrizo García et al. 2019; CCG, pers. obs.). Therefore, the circumscription of the Pubescens and Purple corolla clades (after Carrizo García et al. 2016) is being re-assessed (Carrizo García and Palombo 2019; CCG, pers. obs.). Capsicum pubescens was domesticated and has been highly appreciated by early Peruvian peoples for 4,000 years before the present ( Perry et al. 2007), while its introduction in Central America and Mexico has occurred in the twentieth century (Laborde et al. 1982; McLeod et al. 1982). Capsicum pubescens is very distinctive in the combination of the following characters, with minor differences in cultivars: habit, general pubescence, shape, size and colour of flowers and seeds and heteromorphic style (Figs 103 , 104 ). Capsicum pubescens is an erect to scandent shrub up to 4 m high, with a dense white, soft pubescence covering stems, leaves, pedicels and calyx (sometimes plants are glabrescent), with rugose young leaves, large rotate or rotate-stellate 4-8-merous corollas that are usually deep purple, heteromorphic styles with three different lengths and the largest brownish-black to black seeds (5.5-7 mm long, 4.8-6 mm wide) in the genus. Variations in corolla colour have been observed throughout its distribution, from dark purple to lighter tonalities (near rose colour) or completely lacking purple pigmentation ( Eshbaugh 1979). Pure white corollas (filaments and style white) have also been observed as a rare mutant in cultivated C. pubescens plants in Indonesia ( Yamamoto et al. 2013).

The fruit is the most variable character in this species and its different common names refer to this (Heiser and Smith 1948; Rick 1950; Eshbaugh 1979). In the Andean highlands, the most popular names (and some deviations of these words), are “locoto”, an Aymara word (luqutu), meaning ‘piquant’, used in Bolivia (also in Argentina and Chile) and “rocoto”, a Quechua word (rukutu) meaning 'pepper very piquant’ as is mostly known in Peru and Ecuador. In Central America and Mexico, the common names refer to fruit shapes and colours, to the sensation caused by capsaicinoids in humans or to a particular use in the cuisine ( DeWitt and Bosland 2009; Meckelmann et al. 2015). Fruits are large as in the other cultivated species and very attractive at maturity because of their shape and bright colours (red, orange-red, orange, orange-yellow, yellow, light yellow, nearly black). They can be more or less spherical, blocky or elongate (Fig. 104L-P ); in the first case, they are depressed with the apex truncate or rounded, such as in the “canario” (bird canary, fruit roundish and yellow), “manzano” (apple-shaped and red) and “peron” (large pear-shaped and yellow fruit) varietals ( DeWitt and Bosland 2009); the elongate fruits can be ovoid or elliptic, curved or not, with the apex truncate, rounded or acute and the base sometimes narrowed forming a conspicuous neck-like shape. In Mexico, C. pubescens fruits are also called "chile de cera" (cera = wax) in allusion to the soft and brilliant pericarp.

In the protologue of C. pubescens , Ruiz and Pavón (1799) stated "Habitat affatim in Peruviae cultis, praesertim ad Panatahuarum Provinciam et in Andium nemoribus". The three specimens of C. pubescens in the Ruiz and Pavón Herbarium at MA (MA-815153, MA-815154, MA-815155) are labelled as being collected in Pozuzo ("Ex Pozuzu") and are not in exact agreement with the protologue of C. pubescens . In his posthumously published journals, however, Ruiz (1940) clearly states the record of the travels of the expedition "Among the plants that I described while we remained in Puzuzo are the following [p. 175] … Capsicum frutescens L., arnaucho and Capsicum pubescens , rocobo; both species very abundant in Peru" [p. 176]. We, therefore, assume that all of these specimens labelled as coming from “Pozuzu” belong to the type gathering of C. pubescens . From amongst these three sheets, we have selected the best fertile specimen as the lectotype (MA-815154). A sheet at G from "herb. Pavon" appears to be a duplicate of MA-815154 and a fragment taken from the sheet at G is housed at CORD.

Miers (1849) cited two collections in the protologue of Brachistus? lanceifolius : Seemann 879 & McLean s.n. that he saw in "herb. Hook.", now held at Kew. The Seemann collection (K000585923) consists of two flowering branches and has Seemann’s original label with the collecting locality; the McLean collection (K000585921) has mounted on it another branch collected near Lima by W. Nation (K000585922) that also fits within the circumscription of C. pubescens , but is not part of the type material. We select the Seemann collection (K000585923) as the lectotype of B. lanceifolius as it is the most complete and best-preserved of the specimens cited by Miers.

The protologue of C. maximowiczii ( Kuester et al. 1859) provides a complete description for this species, based on a specimen grown in St. Petersburg from seeds sent to Karl Maximowicz (at the time curator of the Herbarium in St. Petersburg) of a plant cultivated in Valparaiso (Chile). No specimens are indicated and it is likely the description was based on living plants. The corollas are said to be hexamerous and violet and the seeds black, which matches with C. pubescens . However, the common name "agi dulce" referring to a sweet pepper is not usually used in reference to C. pubescens and could be an error. We found a sterile branch with the script "v.v. Rach" [seen alive by Rach] dated 27 May 1858 at LE; this is certainly original material and we designate it the lectotype.

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 103. Capsicum pubescens A reproductive branch B trichome of the leaf C calyx D section of the calyx showing the venation E flower, upper view F sector of opened corolla G, H anthers, dorsal and ventral views, respectively I, J, K gynoecium with short, medium and long style, respectively L anatomical detail of the pericarp (note the giant cell in the mesocarp) M seed N seed, in cross section O structure of seed coat at the seed margin P structure of seed coat at the seed body Q embryo. From Hunziker 25484. Drawn by J. de Ugarte.

Figure 104. Capsicum pubescens A plant B reproductive nodes C young leaves, adaxial surface D flower bud on geniculate pedicel E-H flowers, in front view, showing 4 - 7 - merous corollas I, J flowers, seen from behind K flower, in lateral view L-O mature fruits from different provenance (L, N from Bolivia, M from Peru, O from Mexico) P fruit, in longitudinal section, showing the black seeds A-D, G Barboza et al. 4889 E, P from Barboza et al. 4890 F from Palombo 21 H from Barboza et al. 1847 I from Palombo 22 J from Palombo 23 K, L no specimen vouchers (cult. Cordoba, Argentina) M no specimen voucher (cult. Huancayo-Peru) N from Carrizo Garcia et al. 35 O no voucher specimen (bought in Mexico) A-D, G, H, L, M, O, P photos by G. E. Barboza E, F, I-K, N photos by N. Palombo.

Figure 105. Distribution of C. pubescens and C. rabenii.