Capsicum rabenii Sendtn., Fl. Bras. (Martius) 10(6): 145. 1846.

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

37. Capsicum rabenii Sendtn., Fl. Bras. (Martius) 10(6): 145. 1846.

Fig. 106

Capsicum microcarpum Cav. var. tomentosum Chodat & Hassl., Bull. Herb. Boissier 2,4: 80. 1903. Type. Paraguay. Paraguarí: In regione collium, Cerros de Paraguay, Dec 1900, É. Hassler 6498 (lectotype, designated by Barboza 2011, pg. 27, second step designated here: G [G00390271]; isolectotypes: BM [BM000087632, acc. # 5447784; BM000087632a, acc. # 4575838], CORD [CORD00006947 fragment from G], G [G00390272, G00390273], K [K000585894], P [P00410166], MO [MO-2530203, acc. # 1575002], MPU [MPU023043], S [S16-28128], UC [UC944377]).

Capsicum praetermissum Heiser & P.G.Sm., Brittonia 10(4): 198. 1958. Type. Brazil. Minas Gerais: Viçosa, road to São Miguel, near km 4, 700 m elev., 3 Jan 1930, Y. Mexia 4205 (holotype: F [F-0093991F]; isotypes: BM [BM000992134], CAS [CAS0002376], CORD [CORD00086136], GH [GH00077006], K [K001073038], NY [00138603], MICH [1109872], MO [MO-503526, acc. # 1164250], U [U.1736360], UC [UC509516], S [S-04-2815], US [US00027398], Z [Z000028484]).

Capsicum baccatum L. var. praetermissum (Heiser & P.G.Sm.) Hunz., Kurtziana 6: 242. 1971. Type. Based on Capsicum praetermissum Heiser & P.G.Smith

Type.

Brazil. Rio de Janeiro: " In prov. Sebastianopolitana: Com. de [F. C.] Raben (n. 301)" (lectotype, designated here: BR [BR 0000005529261].

Description.

Erect shrubs or scandent subshrubs (0.50-) 0.70-2.5 (-3) m tall, with the main stem 2-2.5 cm in diameter at base, rarely perennial herbs, much branched from near the base and above. Young stems angled, somewhat rigid, green, pubescent with appressed-antrorse or spreading, simple, uniseriate, 4-8-celled, eglandular trichomes 0.3-1.5 mm long, the new growth with a dense whitish pubescence; nodes green or purple; bark of older stems striate, dark brown, glabrescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair subequal in size and similar or dissimilar in shape. Leaves membranous, slightly discolorous, green above, light green and with the primary veins raised beneath, glabrescent to moderately pubescent, with appressed-antrorse trichomes like those on stems on adaxial surface and margins, sparsely pubescent on the lamina, but with tufts of eglandular trichomes in the vein axils and with long spreading trichomes along the veins giving a woolly appearance abaxially; blades of major leaves 5.3-10 cm long, 2.4-6.5 cm wide, ovate, the major veins 4-6 (-7) on each side of mid-vein, the base asymmetric and attenuate, the margins entire, the apex acuminate or long-acuminate; petioles (1.5-) 2.5-3.5 cm long, winged distally for the decurrent base of the lamina, moderately to densely pubescent; blades of minor leaves 3-4 cm long, 1.4-2 cm wide, ovate or elliptic, the major veins 2-3 on each side of mid-vein, the base rounded or truncate, the margins entire, the apex acute or acuminate; petioles 0.8-1.2 cm long, moderately to densely pubescent. Inflorescences axillary, 2-4 flowers per axil; flowering pedicels (8-) 12-18 (-25) mm long, angled, erect, geniculate at anthesis, moderately to densely pubescent, the eglandular trichomes spreading or antrorse; pedicels scars inconspicuous. Buds globose, whitish-green or with green spots at the base. Flowers 5-merous, rarely 4-merous. Calyx 1.5-2 mm long, ca. 2-2.5 mm wide, cup-shaped, thick, green, pubescent with the same trichomes as pedicels, the calyx appendages 5, 0.8-1.1 mm long, 0.3 mm wide, subequal, thick, erect, cylindrical, inserted close to the margin, moderately pubescent with the same trichomes as calyx tube. Corolla 6-7 mm long, (9-) 12-15 mm in diameter, thick, white or lilac and greenish-yellow outside, mostly purple or lilac with greenish-yellow or green spots and cream centre within, rotate to stellate with interpetalar membrane, lobed less than 1/3 to nearly halfway to the base, pubescent adaxially with short glandular trichomes (stalk 1-3-celled; head globose, peltate, unicellular), glabrous abaxially, the tube 4-4.5 mm long, the lobes 2-2.5 mm long, 2.2-4 mm wide, broadly triangular, spreading, the margins with very short eglandular trichomes, the tips acute, papillate. Stamens five, rarely four, equal; filaments (0.8-) 1.2-1.5 (-2) mm long, purple or lilac, inserted on the corolla 0.8 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.2-1.5 (-1.8) mm long, ellipsoid, white, cream or yellow, not connivent at anthesis. Gynoecium with ovary 1.3-1.5 mm long, 1.4-1.5 mm in diameter, green, ovoid; carpels two, ovules more than two per locule; nectary ca. 0.3 mm tall; styles homomorphic, 2.6-2.8 mm long, exserted 0.8-1.2 mm beyond the anthers, yellowish-white, cylindrical; stigma 0.5 mm long, 0.7-0.75 mm wide, discoid, pale green. Berry globose or subglobose, 5-7.5 mm in diameter or ellipsoid, 7-11 mm long, (4-) 6-8 mm in diameter, green when immature, orange-coloured or bright red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels (14-) 20-28 mm long, erect, strongly angled, widened distally, green; fruiting calyx 4-4.5 mm in diameter, persistent, not accrescent, cup-shaped or discoid, green, the appendages 1-2 mm long, ca. 0.3-0.4 mm wide, erect or spreading. Seeds (5-) 7-15 per fruit, 3-4.2 mm long, 2.5-2.8 mm wide, C-shaped or subglobose, pale yellow or yellow, the seed coat smooth (SM), reticulate-cerebelloid (SEM), the cells irregular in shape, the lateral walls sinuate in seed body, straight at margins; embryo imbricate.

Distribution.

Capsicum rabenii is native to eastern and central Brazil, occupying a large range in Brazil from Bahia to Rio Grande do Sul States and into central Paraguay (Fig. 105 ).

Ecology.

Capsicum rabenii occurs in a wide variety of vegetation types, from the Atlantic Forest (Semi-deciduous Seasonal Forest, Ombrophilous Forest) to xerophytic forests in the Cerrado and Caatinga; it can be found in disturbed gallery forest, calcareous outcrops vegetation, creek and river margins and as ruderal around anthropogenic and cultivated areas, between (35-) 100 and 1,300 m elevation. It is also frequent as an escape from cultivation or in cultivation around houses and growing commercially in small farms.

Phenology.

Flowering from November to May. Fruiting from December to May and June extending to September when in cultivation.

Chromosome number.

n = 12 (Heiser and Smith 1958, as C. praetermissum ; Pozzobon and Schifino-Wittmann 2006 as C. baccatum var. praetermissum ); 2 n = 2x = 24 ( Pickersgill 1977, as C. praetermissum ; Pozzobon et al. 2006; Moscone et al. 2007; Scaldaferro et al. 2016, counts as C. baccatum var. praetermissum ).

Common names.

Brazil: Comarim (Minas Gerais, Krieger 5212), Pimenta (Mina Gerais, Mexia 4205; São Paulo, Briske s.n.), Pimentinha (Santa Catarina, Reitz 4749), Pimenta brava (Minas Gerais, Anonymous s.n.), Pimenta comary ( São Paulo, Handro 860), Pimenta camari ( São Paulo, Coelho s.n.), Pimenta cumarí ( Goiás, Vieira 700; Minas Gerais, Krieger 5212; São Paulo, Hoehne & Gehrt s.n.), Pimenta cumarina (Minas Gerais, Tavares et al. 17), Pimenta-cumari-verdadeira (Santa Catarina, Schwirkowski 885), Pimenta Jorobinha (Rio de Janeiro, Glaziou 8841), Pimenta Passarinho (Southeast Brazil, Ferreira Pinto et al. 2016), Pimenta de combari (Minas Gerais, Lindberg 177), Pimenta do mato (Rio Grande do Sul, Costa Sacco 1332; Espírito Santo, Sucre & Soderstrom 8987).

Uses.

In Brazil, fresh fruits are very appreciated in markets ( Ferreira Pinto et al. 2016), used as a condiment and consumed as pickles and in hot sauces.

Preliminary conservation assessment.

EOO (2,746,764 km2); AOO (404 km2). Capsicum rabenii is well-adapted to different environments including highly disturbed anthropogenic areas and is easily cultivated in farms, which suggest that it is not under threat. This species is proposed as Least Concern (LC).

Discussion.

Capsicum rabenii is a member of the Baccatum clade ( Carrizo García et al. 2016, as C. praetermissum ). The species has a wide distribution in Brazil where it is mostly known as 'pimenta comari’ or “Cumari” (as are also C. baccatum , C. chacoense and C. flexuosum ) in allusion to its pungent red fruits. It grows spontaneously in many areas but can also be found semi-domesticated and under cultivation in small gardens or in larger scale in farms for commercial use ( Pickersgill 1969a; Eshbaugh 1993; Ferraz et al. 2016). In Paraguay, this species is less common and the fruits appear to be not so appreciated and consumed as in Brazil.

Flowering and fruiting specimens of C. rabenii are very similar to C. baccatum var. baccatum on herbarium specimens, it sometimes being impossible to distinguish one from another if there are no annotations of corolla colour or if specimens are only in fruit. Both species have long petiolate leaves, rotate or rotate-stellate corollas, orange to red globose or elliptic upright fruits and pale yellow seeds. Capsicum rabenii is distinguished, however, not only for its purple-margined corollas (Fig. 106E-H ), but also for the dense woolly pubescence of the leaves abaxially with spreading trichomes along the main veins and a conspicuous tuft of trichomes in the vein axils (Fig. 106B ); these features distinguish C. rabenii from C. baccatum var. baccatum that lacks anthocyanin pigmentation in the corollas and has usually glabrescent leaves ( Hunziker 1998; Barboza 2013). Only one individual of C. rabenii has been found with completely white corollas in São Paulo State (Brazil, Hunziker 1956 4) growing in the same environmental conditions with other specimens with purple-margined corollas ( Hunziker 1956 3); this lack of pigmentation may be due to a recessive gene. Similarly, some specimens from Goias State (Brazil) lack dense pubescence along the veins but the tuft of trichomes on the secondary vein axils and the typical corollas with the purple band at the margins are present in these populations.

Capsicum rabenii differs from C. chacoense , with which is sympatric in Paraguay, in its rotate, coloured corollas and the characteristic pubescence of the leaves beneath, while C. chacoense has stellate, entirely white corollas and a variable pubescence, but is never woolly (Fig. 46 ).

It is possible that C. rabenii hybridises naturally with C. baccatum and C. frutescens in Brazil as has already been shown through reciprocal crosses that produced some fruits with viable seeds (Heiser and Smith 1958), but intermediates amongst these species have not been observed in field. Our observations indicate that the three species are usually distinguishable using the corolla colour, the pubescence (see above) and calyx features.

When Heiser and Smith (1958) described C. praetermissum , they stated that none of the species of "Flora Brasiliensis" ( Sendtner 1846) matched with their new species; they did not realise that the description of C. rabenii fitted very well with C. praetermissum in the pubescence of leaves and shape of the corolla, characters clearly observable in the type collections of both species. Hunziker (1971) proposed C. praetermissum as a variety of C. baccatum and he recognised C. rabenii as synonym of his new varietal name. Hunziker preferred to use the epithet Capsicum praetermissum because of the valuable and unequivocal information that Heiser and Smith (1958) provided on this plant. According to the ICN (Art. 11.2, Turland et al. 2018), Hunziker could use the name Capsicum praetermissum because Capsicum rabenii was in a different rank and, consequently, the epithet Capsicum rabenii passed into oblivion. Since then, many authors have spread both names C. praetermissum and C. baccatum var. praetermissum in literature and on herbarium specimens. Enzymatic ( McLeod et al. 1983b), cytogenetic ( Moscone et al. 2007) and molecular studies ( Kochieva et al. 2004; Ibiza et al. 2012; Carvalho et al. 2014; Carrizo García et al. 2016), plus the clear morphological characters discussed above, provide more than enough evidence that C. baccatum and C. praetermissum are distinct species. As C. rabenii has priority over C. praetermissum , the first becomes the correct name for this species, despite the previous more common usage of the latter in both literature and on herbarium sheets.

Sendtner (1846) cited a single collection in the description of C. rabenii , "Com. de Raben (n.301)" (Danish Count Frederik Christian Raben). A specimen at BR labelled "Com. Raben" with the number 301 written on it in another hand is the only original material we have found and we designate it the lectotype. A specimen in C (C10019144) labelled as "Herbarium Fr. Chr. Comitis Raben" and "Lago de Rodriguez. R. de Janeiro. 1835", has been suggested as original material by Olof Ryding on an additional annotation slip on this sheet stating “… It matches the holotype of Capsicum rabenii Sendtn. at BR. The two [BR and C] specimens apparently belonged to the same collection … O. Ryding 1999" (see also Ryding 2002 for further information on Raben collections). We compared these specimens and they are clearly not the same. The lectotype at BR has leaves that are densely pubescent abaxially, matching Sendtner’s (1846) description, while the specimen at C has leaves that are glabrous or glabrescent on both surfaces, belonging to C. baccatum var. baccatum .

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

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

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.