Capsicum baccatum L. var. baccatum

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

2a. Capsicum baccatum L. var. baccatum

Figs 25 , 26

Capsicum pulchellum Salisb., Prodr. Stirp. Chap. Allerton: 134. 1796, nom. illeg. superfl. Type. Based on Capsicum baccatum L. (cited in synonymy).

Capsicum microcarpum Cav., Descr. Pl. (Cavanilles): 371. 1802. Type. Cultivated in the Royal Botanical Garden in Madrid, Spain "H.R.M. [Hortus Regis Matritensis]. Se cría en la Havana... y se cultiva en el Jardín botánico” (lectotype, designated here: MA [MA-307276]).

Capsicum ciliare Willd., Enum. Pl. [Willdenow] 1: 243. 1809. Type. Cultivated in Berlin, Germany, of unknown origin "Cult. in Hort. Bot. Berol.", C.L. Willdenow s.n. (lectotype, designated here: B [B-W04430-01-0]).

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

Capsicum comarim Vell., Fl. Flumin.: 60. 1829 ( “1825”); Fl. Flumin. Icon. 2: t. 2. 1831 ( “1827”). Type. Brazil. [Rio de Janeiro]: "Colitur hortis, et sponte undequaque crescit" (lectotype, designated by Knapp et al. 2015, pag. 284: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_005] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 2. 1831).

Capsicum cumanense Fingerh., Monogr. Capsic.: 17. 1832, nom. illeg. superfl. Type. Based on (renaming of) " Capsicum baccatum Kunth" [= C. baccatum L.] (cited in synonymy).

Capsicum microcarpum DC. forma fruticosum Sendtn., Fl. Bras. (Martius) 10(6): 146. 1846. Type. Brazil "In Brasilia", Pohl s.n. (lectotype, designated here: M [M-0171544]).

Capsicum microcarpum DC. forma herbaceum Sendtn., Fl. Bras. (Martius) 10(6): 146. 1846. Type. Brazil. "Martius Mss. in Itinerario n. 132", Prope Polafoco, Sept., C.F.P. Martius 132 (lectotype, designated here: M [M-0171543]; isolectotype, CORD [CORD00101765]).

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

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

Capsicum annuum L. var. microcarpum (Cav.) Voss, in Vilm. Blumengärtn., ed. 3. 1: 723. 1894. Type. Based on Capsicum microcarpum Cav.

Capsicum frutescens L. var. baccatum (L.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 99. 1898. Type. Based on Capsicum baccatum L.

Capsicum microcarpum Cav. var. glabrescens Hassl., Repert. Spec. Nov. Regni Veg. 15: 244. 1918. Type. Paraguay. Canindeyú: "Iter ad Yerbales montium Sierra de Maracayu, in regione cursus superioris fluminis Jejui guazú”, Dec. 1898-99, É Hassler 5703 (lectotype, designated by Barboza 2011, pg. 28, second step designated here: G [G00390268]; isolectotypes: BM [BM000074084, acc. # 5447772; BM000074084a, acc. # 4575837; G [G00390266 two sheets with same barcode, G00390267], GH [00936720], K [K000585896], MO [MO-503802, acc. # 1574551], NY [00138600], P [P00410160, P00410161, P00482076], UC [UC944854], W [acc. # 1902-0002869]).

Capsicum annuum L. subsp. baccatum (L.) Terpó, Feddes Repert. 72: 173. 1966. Type. Based on Capsicum baccatum L.

Description.

Erect shrubs or perennial herbs 0.50-3 (-3.5) m tall, rarely small trees, the main stem 2-2.5 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, green, sometimes the ridges purple, mostly glabrous to sparsely or moderately pubescent with appressed-antrorse, simple, uniseriate, 4-7-celled, eglandular trichomes 0.2-1.2 mm long; nodes usually purple; bark of older stems fissured, dark brown, glabrous; lenticels abundant. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous, slightly discolorous, dark green above, light green beneath, glabrescent to moderately pubescent with appressed-antrorse trichomes like those of the stems on both surfaces and margins; blades of major leaves 4.5-10 cm long, 2.5-6 cm wide, ovate, the major veins 5-8 on each side of mid-vein, the base somewhat asymmetric and attenuate, the margins entire, the apex acute; petioles (1.5-) 2-4 cm long, moderately to densely pubescent; blades of minor leaves 3.5-4.5 cm long, 2-3 cm wide, ovate, the major veins 4-5 on each side of mid-vein, the base rounded or truncate, the apex acute; petioles 0.5-1 cm long, moderately to densely pubescent. Inflorescences axillary, 2-3 flowers per axil, rarely flowers solitary; flowering pedicels (17-) 20-35 mm long, angled, erect or slightly spreading, geniculate at anthesis, glabrescent to moderately pubescent, the eglandular trichomes short, spreading or antrorse; pedicel scars inconspicuous. Buds globose, white with greenish-yellow spots, occasionally purple. Flowers 5-merous. Calyx 1.5-2 (-2.5) mm long, ca. 2-2.5 mm wide, cup-shaped, thick, green, pubescent with the same trichomes as pedicels and some glandular trichomes, the calyx appendages 5, (0.3-) 0.5-2 mm long, 0.2 mm wide, subequal, thick, erect, cylindrical, inserted close to the margin, pubescent with the same trichomes as calyx tube. Corolla 4.5-7.5 mm long, 10-13 mm in diameter, thick, white with greenish-yellow spots and white centre outside and within, rotate or rotate-stellate, with interpetalar membrane, lobed 1/3 or less 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 2.5-2.7 mm long, 3.4-3.5 mm wide, broadly triangular, spreading, the margins with very short eglandular trichomes, the tips acute, papillate. Stamens five, equal; filaments 2.5-3.5 mm long, white, inserted on the corolla 1-1.1 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5-1.8 mm long, ellipsoid, white or pale yellow, more rarely greyish, not connivent at anthesis. Gynoecium with ovary 2.5-2.7 mm long, ca. 2 mm in diameter, ovoid, green; nectary 0.3-0.5 mm tall; styles dimorphic, short style 2-2.5 mm long, not exceeding the anthers length, long style ca. 3.5 mm long, exserted 1.4-1.7 mm beyond the anthers, cylindrical, white; stigma 0.3 mm in diameter, globose or discoid, pale green. Berry 6-8 (-10) mm in diameter, globose or subglobose, less frequently ellipsoid with truncate or flattened apex, 10-20 mm long, 4-7 mm in diameter, green when immature turning to greenish-black and bright red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 20-35 mm long, erect, strongly angled, widened distally, green; fruiting calyx 3-4.5 mm in diameter, persistent, not accrescent, cup-shaped or discoid, green, the appendages 1.5-2.3 mm long, appressed to the berry or spreading. Seeds 12-15 per fruit, 2.5-4 mm long, 2.3-3 mm wide, ovoid, subglobose or C-shaped, pale yellow to yellow, the seed coat smooth or slightly reticulate (SM), cerebelloid (SEM), the cells irregular in shape, the lateral walls sinuate strongly sinuate; embryo imbricate.

Distribution.

Capsicum baccatum var. baccatum is widely distributed in South America from northern Venezuela and Colombia through Peru, Bolivia, Paraguay, northern and north-eastern Argentina to south and eastern Brazil (Fig. 27 ).

Ecology.

Capsicum baccatum var. baccatum occurs in dry or humid subtropical or tropical forests with semi-deciduous or deciduous vegetation, between 150 and 1,900 m elevation; it is quite common in Chaco scrub forests, in gallery forests and in the margins or interior of secondary forests. It is frequently a ruderal in disturbed areas.

Phenology.

Flowering from October to May. Fruiting from late November to September.

Chromosome number.

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

Common names.

Argentina: Coincho (Jujuy, Fabris 3454), Cumbarí (Misiones, Montes 15164), Quitucho (Salta, Hunziker 1985), Puta-parió (Corrientes, Martínez Crovetto 11125), Ají quitucho (Salta, Hunziker 1579), Ají del campo, Ajitucho (Salta, West 8389), Ají del monte (Salta, Rial Alberti s.n.), Pimenta del monte (Misiones, Montes 15164), Pimentón del monte, ají cumbarí (Misiones, Montes 15202); Bolivia: Arabibi (Santa Cruz, Zenteno-R 12798), Aribibe (Santa Cruz, Hurtado 296), Aribibi (Chuquisaca, Debouck 3019; Santa Cruz, de Michel 159), Arivivi (Chuquisaca, Serrano 1903; Santa Cruz, Cárdenas 4702), Cobincho (Tarija, Krapovickas & Schinini 39010), Ají aribibi (Cochabamba, Thomas 705), Aribibi silvestre (Beni, Rivero 218), Ají del campo (Tarija, Krapovickas & Schinini 39010), Arivivi grande o cumbarito ( Fundación PROINPA 2007), Arivivi last’a o miska ( Fundación PROINPA 2007), Arivivi morado ( Fundación PROINPA 2007), Arivivi tuna árbol grande ( Fundación PROINPA 2007), Arivivi tuna con flor blanca ( Fundación PROINPA 2007); Brazil: Cumari ( Espírito Santo, Crepaldi 59), Pimenta-cumari ( Goiás, Mendonça et al. 5971), Pimenta de passarinho (Bahia, Mori 11603); Colombia: Ají (Vichada, Rodríguez 164), Ají de babilla (Amazonas, Cárdenas 9423), Ají de la capitania (Amazonas, Cárdenas 9403); Peru: Aji Ayucllo ( Junín, Bosland and Votava 2000).

Indigenous names.

Bolivia: Pochetii (Trinitario, Cochabamba, Thomas 705), Winno, Sachimi (Yuracare, Cochabamba, Thomas 705); Colombia: Azi (Piapocos, Vichada, Rodríguez 164), Gugsobia (Amazonas, Cárdenas 9423), Kulana (And, Amazonas, Castro & Matapí 564), Kulana (Yucuna, Amazonas, Cárdenas 9403), Kuraraka (Letuama, Amazonas, Cárdenas 9401); Paraguay: Hõmpita (Ayoreo, Boquerón, Gragson 124), Nuuhá (Alto Paraguay, Schmeda 1584).

Uses.

As fruits are generally extremely pungent, they are collected and stored for use as a food condiment by native populations. Some accessions of this wild pepper ( “arivivi”) in Bolivia have been considered promising for their interesting agro-morphological and biochemical characteristics with potential for the development of high value products for different markets ( Fundación PROINPA 2007; van Zonneveld et al. 2015). However, commercialisation is still marginal and fruits are cultivated in home gardens or gathered from nature for self-consumption or to be distributed locally ( Jäger et al. 2013).

Preliminary conservation assessment.

EOO (11,809,545.422 km2); AOO (1,212 km2). Capsicum baccatum var. baccatum is considered Least Concern (LC) for the time being.

Discussion.

Capsicum baccatum var. baccatum is a member of the Baccatum clade and is related to C. rabenii and C. chacoense ( Carrizo García et al. 2016). This entity is considered to be the wild progenitor of the cultivated C. baccatum var. pendulum and is widespread in South America. Its main centre of domestication is thought to be in the Bolivian Amazonia and inter-Andean valleys ( Scaldaferro et al. 2018).

In an effort to clarify the taxonomy of C. baccatum , Eshbaugh and collaborators ( Eshbaugh 1968, 1970, 1976; D’Arcy and Eshbaugh 1974; Jensen et al. 1979; Pickersgill et al. 1979; McLeod et al. 1983a, 1983b; Mitchell et al. 1989) and others ( Kuriachan 1981; Egawa and Tanaka 1984; Jarret 2007; Scaldaferro et al. 2018) have studied the wild and domesticated forms of this species with a multitude of techniques (morphological, breeding, cytogenetic, biochemical, molecular, phylogeographical); the results support the wild progenitor-domesticate association proposed by Eshbaugh (1968).

Capsicum baccatum var. baccatum typically exhibits 2-3 flowers per node, rarely solitary flowers, geniculate pedicels that are erect or declining at anthesis, 5-merous flowers with white rotate or rotate-stellate corollas with greenish-yellow spots, dimorphic styles and small, globose, subglobose or ellipsoid, erect, deciduous, red fruits (Fig. 26 ). The domesticated genetic lines mainly differ in having larger, 5-8-merous flowers, an ovary with 2-5 locules, quite diverse fruits that vary in size, colour (green, yellow, brown, orange, red) and shape (pendent, usually elongate or of different forms) and larger seeds.

Fruiting specimens of C. baccatum var. baccatum are very similar to C. rabenii and it is sometimes impossible to distinguish the two, especially if there are no annotations about the corolla colour (in C. rabenii , corolla lobe margins are purple). However, C. baccatum var. baccatum usually has glabrescent to moderately pubescent leaves in contrast to the densely lanose pubescence found abaxially along the main veins in C. rabenii (Fig. 106B ).

Capsicum baccatum var. baccatum differs from C. chacoense , with which it is sympatric in some localities of Argentina, Bolivia and Paraguay, by usually having five calyx appendages, a larger and rotate or rotate-stellate corolla with greenish-yellow pigmentation within, staminal plaques with auricles fused to the corolla and long and short styles. In contrast, C. chacoense has a calyx with 5-10 unequal appendages, entirely white and smaller corollas (4-6 mm long), staminal plaques with auricles not fused to the corolla and homomorphic styles (Fig. 46 ). Fruiting specimens of C. baccatum var. baccatum sometimes are difficult to distinguish from C. chacoense . The fruiting pedicels of C. baccatum var. baccatum are usually 2-3 per node and the fruiting calyces have five subequal appendages, without any evidence of a mucro below the calyx margin, which differs from the solitary fruiting pedicels and 5-10 unequal calyx appendages in C. chacoense .

Some earlier botanists submerged the epithet Capsicum baccatum under C. annuum ( Kuntze 1891) or C. frutescens ( Irish 1898) to name specimens belonging to C. annuum var. glabriusculum or wild forms of C. baccatum , an interpretation followed by other researchers (Smith and Heiser 1957; Emboden 1961; Terpó 1966).

Although the pungency of C. baccatum is regarded as low-mild ( Libreros et al. 2014; Tanaka et al. 2017), some accessions of C. baccatum var. baccatum from Bolivia have been reported as non-pungent (e.g. Chuquisaca: Manchego CBNP 04, 05 & 06; Santa Cruz: Manchego CBNP 01, 02 & 03; also see Tewksbury et al. 2006).

Cavanilles (1802) described C. microcarpum , based on material cultivated at the Real Jardín Botánico de Madrid from seeds sent from Cuba. We found two Cavanilles specimens at MA gathered in the Real Jardín Botánico de Madrid in 1802: MA-307276, separated as type material in the Cavanillesii Typi collection ( Garilleti 1993; https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.ma307276?loggedin=true) and MA-307278, which is filed in the general collection. Both specimens are labelled as Capsicum microcarpum . According to Garilleti (pers. comm.), the original writing on the sheet label MA-307278 (the left label) is not from Cavanilles, but from Demetrio Rodríguez; since this plant was grown in the Real Jardín Botánico, Rodríguez could have collected it later and Cavanilles might not have seen it. Therefore, the only unequivocal original material is that with the label in Cavanilles’ handwriting (MA-307276), which is here selected as the lectotype; this sheet consists of three branches with flowers and fruit.

Willdenow (1809) based the description of C. ciliare on a specimen of unknown origin cultivated in the Botanical Garden of Berlin. At B, there is one sheet in Willdenow’s Herbarium with the script " C. ciliare , Hort. Bot. Berol. W."; this sheet is selected as the lectotype.

Sendtner (1846) described C. microcarpum forma fruticosum , based on two collections, one from Corego de Jaragua, São Paulo ("leg. Pohl") and the other from "prov. Sebastianopolitana, ad Lagoa de Freitas" [Rio de Janeiro], but he cited no herbaria. We have found only the specimen collected by Pohl (M-0171544), which is selected as the lectotype.

For C. microcarpum forma herbaceum , Sendtner (1846) mentioned the specimen "Mart. Mss. in Itinerario n. 132", but cited no herbaria. We found a collection at M (M-0171543) with a handwritten description that is in agreement with Sendtner’s diagnosis and has the number 132. Therefore, this sheet is designated the lectotype.

When Kuntze (1891) proposed the combination C. annuum var. baccatum (L.) Kuntze, he was referring to specimens that actually correspond to the spontaneous forms of C. annuum (cfr. Kuntze s.n., NY barcode 01008231), but he incorrectly used the epithet Capsicum baccatum L. which corresponds to a different species ( C. baccatum L.).

When Terpó (1966) proposed the combination C. annuum subsp. baccatum , based on C. baccatum L., he clearly stated "Mexikanischer Wildpaprika" which actually belongs to the spontaneous forms of C. annuum .

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 25. Capsicum baccatum var. baccatum A flowering branch B calyx C glandular trichome of the calyx D eglandular trichome of the calyx E flower, upper view F sector of opened corolla G gynoecium H fruit I seed J seed, in cross section K embryo A-G from Hunziker 7350 H-K from Hunziker 1579. Drawn by N. de Flury. Published in Barboza (2013), courtesy of the Board of the Instituto Darwinion (San Isidro, Buenos Aires, Argentina), reproduced with permission.

Figure 26. Capsicum baccatum var. baccatum A plant B flower bud C purple flower bud D, E, F flowers at anthesis, different views G young fruiting branch showing ovary with long and short styles H immature fruits I mature fruit A, C from Barboza 4913 B, D-F, H from Barboza 5038 G from Barboza 2431 bis, I from Barboza et al. 3419. Photos by G. E. Barboza.

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

Figure 46. Capsicum chacoense A plant B flower bud C flower, in pre-anthesis D flower, in front view E immature fruit F mature fruit A-F from Barboza et al. 4910. Photos by G. E. Barboza.

Figure 106. Capsicum rabenii A plant B leaf, abaxial surface showing pubescence on the veins C reproductive nodes D bud on geniculate pedicel E flower, lateral view F rotate corolla G stellate corolla H flower showing the exserted style I immature fruits J globose mature fruits K ellipsoid mature fruit and fruiting calyx A-I, K from Barboza & Deanna 5003 J from Barboza et al. 1646. Photos by G. E. Barboza.