Capsicum flexuosum Sendtn., Fl. Bras. (Martius) 10(6): 143. 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

17. Capsicum flexuosum Sendtn., Fl. Bras. (Martius) 10(6): 143. 1846.

Figs 62 , 63

Capsicum schottianum Sendtn. var. leptophyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 416. 1852. Type. Brazil. "In Brasiliâ australiore. (Sellow e Sendtn. l.c.)" (no herbaria cited; no original material found).

Capsicum parvifolium Sendtn. var. sellowianum Dunal, Prodr. [A. P. de Candolle] 13(1): 419. 1852. Type. Brazil. "In Brasilia australiore. (Sellow.)" (no herbaria cited; no original material found).

Capsicum campylopodium Sendtn. forma magis-puberula Chodat, Bull. Herb. Boissier ser. 2, 2: 815. 1902. Type. Paraguay. [ Canindeyú]: Sierra de Maracayú, in silva Ipé hú, Oct 1898-1899, É. Hassler 5134 (lectotype, designated by Barboza 2011, pg. 26 [G], second step designated here: G [G00390263]; isolectotypes: A [00936719], BM [BM000074110], CORD [CORD00087947, fragment ex G], G [G00390264], G [G00390265], K [K000648540], NY [04206101], P [P00410080, P 00410081], S [S16-28249], UC [UC-944853]).

Capsicum campylopodium Sendtn. forma laurifolium Chodat, Bull. Herb. Boissier ser. 2, 2: 815. 1902. Type. Paraguay. [ Itapuá]: in silva pr. fl. Capibary, 5 Dec 1898-1899, É. Hassler 5893 (lectotype, designated here: G [G00390270]; isolectotypes: BM [BM000074083], G [G00390269], K [K000648539], P[P00410127], S [S16-27826], UC [UC950167]).

Capsicum mositicum Toledo, Arq. Bot. Estado São Paulo 3(2): 64, f. 2. 1953. Type. Brazil. São Paulo: Mun. de Amparo, Monte Alegre, margem da estrada para Socorro, 740 m elev., 17 Dec 1942, M. Kuhlmann 144 (holotype: SP [001631, acc. # 47939]; isotypes: CORD [CORD00006628, CORD00006629]).

Capsicum ramosissimum Witasek, Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 79(2): 320. 1910. Type. Brazil. "Prov. São Paulo: Prope "Fazenda bella vista", in districtu urbis Santa Cruz ad flumen Rio Pardo, ca. 500 m elev., VII, leg. v. Wettstein et Schiffner ", Jul 1901, R. v. Wettstein & V. Schiffner [338] (lectotype, designated here: WU [acc. # 0037945]; isolectotypes: CORD [CORD00006633], F [v0093723F, acc. # 871103, fragment], W [1922-0001510], WU [acc. # 0037946], Z [Z-000038683]).

Capsicum schottianum Sendtn. var. flexuosum (Sendtn.) Hunz., Huitième Congr. Int. Bot. Paris. Comptes Rend. Séances Rapp. & Commun. 1954, sect.4: 73. 1956. Type. Based on Capsicum flexuosum Sendtn.

Type.

Brazil. "In Brasilia australiore: Sellow" (no original material located; Brazil. Paraná: Mun. Curitiba, Parque Iguacu , 27 Dec 1979, R. Kummrow 1307; neotype, designated here: MBM [MBM064252]; isoneotype, CORD [CORD00101762]) .

Description.

Erect subshrubs or shrubs, (0.3-) 0.9-2 m tall, more rarely small trees 2-2.5 m tall, with the main stem thick 2-2.5 cm in diameter at base, much branched above, the branches dichotomously spreading in a typical “zig-zag” appearance. Young stems terete or slightly ridged, fragile, green, glabrous to moderately or densely pubescent, with antrorse simple, uniseriate, 2-6-celled, eglandular trichomes 0.2-0.9 mm long; nodes solid, green; bark of older stems dark brown, glabrous, rarely pubescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair unequal or more or less equal in size, similar in shape. Leaves membranous, discolorous, dark green above, light green or greenish-grey beneath, moderately pubescent on both sides or, if glabrescent, with an evident tuft of trichomes in the basal vein axils abaxially, the trichomes appressed-antrorse similar to those of the stems; blades of major leaves 5.2-14 cm long, 1.5-4.5 (7.5-) cm wide, ovate to elliptic, the major veins 4-7 on each side of mid-vein, the base attenuate or short-attenuate, the margins entire, the apex acuminate; petioles 0.5-1.5 cm, glabrous to moderately pubescent; blades of minor leaves 1.7-3.5 (-4.5) cm long, 0.5-2 cm wide, ovate to elliptic, the major veins 3-4 on each side of mid-vein, the base attenuate, the margins entire, the apex acute to acuminate; petioles 0-0.5 cm, with same pubescence as the major leaves. Inflorescences axillary, 2-3 (-6)-flowers per axil or flowers solitary; flowering pedicels 10-21 mm long, terete, pendent, more rarely spreading, non-geniculate at anthesis, glabrous to moderately pubescent, the eglandular trichomes short, antrorse; pedicels scars inconspicuous. Buds globose, inflated, white with light green spots. Flowers 5-merous. Calyx 1.3-2 mm long, 3-3.5 mm wide, cup-shaped, green or greenish-yellow, hyaline at the margin, glabrous to moderately pubescent, the calyx appendages five, inconspicuous, umbo-like and the calyx pentagonal in outline with a tuft of short uniseriate eglandular trichomes on each angle, if the calyx appendages absent, the calyx circular in outline. Corolla 6-11 mm long, ca. 13 mm in diameter, white with yellowish-green pigmentation outside, white with variously yellowish-green spots in the base of lobes and throat within, exceptionally also with purple spots, stellate or rotate-stellate, with thin interpetalar membrane, lobed less than or up to nearly halfway to the base, the tube 3-6 mm long, pubescent adaxially with a continuous ring of glandular trichomes (stalk long 1-3-celled; head globose, peltate, unicellular), glabrous abaxially, the lobes 3-5 mm long, 2-4.5 (-5) mm wide, broadly triangular, spreading, glabrous adaxially and abaxially, the margins involute, papillate, the tips cucullate, densely papillate. Stamens five, equal, filaments 2.5-3.5 mm long, cream, inserted on the corolla 1-1.3 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.1-1.9 mm, ellipsoid or ovoid, yellow, not connivent at anthesis. Gynoecium with ovary 1.3-2 mm long, 1.2-1.4 mm in diameter, light green, ovoid; ovules more than two per locule; nectary ca. 0.4 mm tall; styles homomorphic, (3.8-) 4-5.7 mm, exserted 0.8-1 mm beyond the anthers, white, clavate; stigma 0.2 mm long, 0.8 mm wide, discoid, pale green. Berry 5-8 mm in diameter, subglobose or globose, brilliant green when immature, reddish-orange or red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 15-26 mm long, pendent, terete, widened distally, green; fruiting calyx 4-5 mm in diameter, persistent, not accrescent, discoid, light green, sometimes the margin ripped. Seeds (4-) 5-25 per fruit, 2.8-3.4 mm long, 2.2-3 mm wide, C-shaped, subglobose or teardrop-shaped, brownish-black, the seed coat faintly reticulate (SM), reticulate (SEM), the cells polygonal to irregular in shape, the lateral walls sinuate in the seed body and straight to wavy at margins; embryo imbricate or coiled.

Distribution.

Capsicum flexuosum is a widespread species occupying a more or less continuous range from north-eastern Argentina (Corrientes and Misiones Provinces) and southern Paraguay (Alto Paraná, Amambay, Caaguazú, Caazapá, Canindeyú, Central, Guairá, Itapuá, Paraguarí and San Pedro Departments) to south and south-eastern Brazil (Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina and São Paulo States) (Fig. 61 ).

Ecology.

Capsicum flexuosum is very common in the interior and margins of primary and secondary forests, in remnants of forests or disturbed areas. In Brazil, it occurs in a wide range of vegetation types, primarily in the Atlantic Rainforest (Floresta Estacional Decidual, Floresta Estacional Semidecidual Montana and Submontana, Floresta Ombrófila Densa Submontana and Montana, Floresta Ombrófila Mista Montana, Floresta Ombrófila Densa das Terras Baixas), at low and medium altitudes, between 30 and 1,300 m.

Phenology.

Flowers nearly all year around; fruiting occurs from January to July and from October to December.

Chromosome number.

n = 12 ( Moscone 1992; Pozzobon and Schifino-Wittmann 2006); 2 n = 2x = 24 ( Pozzobon et al. 2006; Moscone et al. 2007).

Common names.

Argentina: Cumbarí (Misiones, Montes 2139), Pimentina (Misiones, Montes 2375), Pimentiña (Misiones, Montes 2375), Ai Jesu (Misiones, Buchinger & Rodríguez 3222), Ají Cumbarí (Corrientes, Bonpland s.n.), Pimenta silvestre (Misiones, Montes 2140), Pimienta del monte (Misiones, Schwindt 4310), Pimiento del monte (Misiones, Buchinger & Rodríguez 3222), Pimentón del monte (Misiones, Schwindt 1871); Brazil: Pimenteira (Rio Grande do Sul, Santos et al. 2869), Pimenta-braba (Rio Grande do Sul, Abruzzi 576), Pimenta de passarinho ( Paraná, Lleras Pérez et al. 1947), Pimenta do bugni (Santa Catarina, Neubert 216), Pimento-do-mato, pimenta-braba (Santa Catarina, Schwirkowski 1519); Paraguay: Locote ( Canindeyú, Montes 3280), Pimiento silvestre ( Canindeyú, Montes 3280).

Indigenous names.

Argentina: Guachu ky’i ( Guabyrá poty, Misiones, Keller & Ferreira 1349), Guach ky’yi (Takuapi, Misiones, Keller 2957), ‘Ka’a ete’y’ ( Guabyrá poty, Misiones, Keller & Ferreira 288); Paraguay: Ke-jiú ( Canindeyú, Montes 3280).

Uses.

Herbarium labels record that fruits are used as condiments in Argentina and Paraguay due to their high pungency. There are no records of uses from Brazil.

Species conservation assessment.

EOO (778,640.645 km2); AOO (1,348 km2). Capsicum flexuosum is quite abundant throughout its distribution. Based on the EOO, the AOO and its presence in many conservation units, we consider C. flexuosum is not under risk and assign the category of Least Concern (LC).

Discussion.

Capsicum flexuosum is the single member of the Flexuosum clade ( Carrizo García et al. 2016). It is a highly variable species in its pubescence density that varies from absent (plants glabrous) to moderate or dense on young branches, leaves and calyx; on leaves, the simple eglandular trichomes are located mainly on the abaxial surface of the lamina. The name C. mositicum was applied to glabrous or glabrescent specimens with a tuft of trichomes in the abaxial leaf vein axils. Another striking aspect of C. flexuosum is the profuse, dichotomous-divaricate branching at the apex of young branches that give the plant a messy aspect, with crowded leaves, flowers and fruits; C. ramosissimum was described, based on this peculiar trait. It is possible that this growth pattern is caused by a plant/pathogen (bacterium, fungus, virus or insect).

Another variable trait is the greenish-yellow pigmentation of the white corolla; this colouration can be present as many small spots interrupted by white lines covering the throat and the base of the lobes or it can be present as five large spots covering the surface (Fig. 63C ) or the greenish-yellow pigmentation can extend over the lobes, leaving a thin white edge.

A striking population from around Monteiro Lobato ( São Paulo State, Brazil) is morphologically very similar to C. flexuosum in its habit, pendent, non-geniculate flowering pedicels, pentagonal calyx, orange-red globose pungent fruits and black seeds. However, unlike other C. flexuosum populations in which the corolla is clearly stellate and white with greenish-yellow spots (Fig. 63E-H ), the population in Monteiro Lobato has a rotate-stellate white corolla with greenish-yellow pigmentation in the throat and purple spots in the lobes (similar to C. schottianum ). An accession of this population (as Capsicum aff. flexuosum ) was included in the molecular phylogeny ( Carrizo García et al. 2016) and was recovered within the Flexuosum clade. In addition, its chromosome number is 2n = 24 (see Table 2 ). Since this population occurs within other typical populations of C. flexuosum ( Pozzobon et al. 2006; Moscone et al. 2007), is treated here as a local variant of C. flexuosum .

Capsicum schottianum and C. campylopodium are the most morphologically similar species to C. flexuosum . All three of these species lack calyx appendages and are extremely difficult to distinguish from one another in herbarium material, when the corolla colour is not stated on the labels or only fruit is present. Capsicum flexuosum differs from C. campylopodium and C. schottianum in its non-geniculate (vs. geniculate) pedicels, yellowish-green spots on the adaxial surface of the corolla (absent in C. campylopodium and in combination with purple spots in C. schottianum ) and edible reddish-orange or red fruit (vs. greenish-golden yellow fruits).

For over 100 years, C. flexuosum has been misinterpreted in both literature and herbaria. Chodat (1902) and Chodat and Hassler (1903) cited specimens of C. flexuosum from Paraguay under C. campylopodium , a Brazilian endemic. In addition, these authors described infraspecific names, such as Capsicum campylopodium forma magis-puberula and Capsicum campylopodium forma laurifolium . Hunziker (1950) confused C. flexuosum with C. schottianum , but later amended the error ( Hunziker 1998) and restricted the distribution of C. schottianum to Brazil ( Hunziker 2001). Finally, in floristic Brazilian works, some specimens of C. recurvatum have been referred to as C. flexuosum ( Freire de Carvalho 1985).

Sendtner (1846) described C. flexuosum and cited a single un-numbered collection made by Friedrich Sellow from "Brasilia australiore", but as was usual at the time, he did not cite a herbarium. Despite searching in the many herbaria (e.g., BM, F, G, K, M, W) where Sellow’s collections are held today, the only Sellow collection we have seen that corresponds to C. flexuosum was held in Berlin (F neg. 2868) and was destroyed in World War II. In the interests of fixing the application of this name, we are designating a neotype using a modern collection that shows all the diagnostic characters of the species (R. Kummrow 1307: MBM064252; CORD00101762).

Dunal (1852) proposed C. schottianum var. leptophyllum for the unnamed var. β of Sellow’s C. schottianum (Sellow 1846: 144). This name refers to plants with narrower leaves and fruiting pedicels that are not curved, but arched throughout their length in contrast to the typical fruiting pedicels that are initially rigid, geniculate (curved) and erect to spreading in C. schottianum (pedicels pendent in mature fruits). As no original material was found, we propose the varietal name Capsicum leptophyllum under C. flexuosum , based on the pedicel features that fit exactly with that species (i.e. pedicels never curved in flower and fruit).

Dunal (1852) based the name C. parvifolium var. sellowianum on a previous Sellow’s description (Sellow 1846: 146) given for a fruitful specimen with "foliis glabrioribus, ... calyci deplanato 5-angulari ...". As no original material was found, we propose this name as a synonym of C. flexusoum due to glabrous leaves and pentagonal calyx (calyx appendages minute) described in the protologue which fit best with C. flexuosum . Capsicum parvifolium has pubescent leaves and five well-developed calyx appendages.

Chodat (1902) based the description of C. campylopodium forma laurifolium on the collection Hassler 5893, without citing a herbarium. We found two sheets of this gathering at G, both consisting of very well-preserved reproductive branches. We select the better of these, G00390270, as the lectotype.

Witasek (1910) described C. ramosissimum on the basis of two Wettstein & Schiffner specimens, both from "Fazenda bella vista" ( São Paulo, Brazil). We found the two syntypes at WU where the collections she used are held; the specimen numbered Wettstein & Schiffner 338 (WU 0037945) is selected as the lectotype, because it is well-preserved and has duplicates housed at CORD, F, W, WU and Z.

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 61. Distribution of C. eximium and C. flexuosum.

Figure 62. Capsicum flexuosum A flowering branch B flower C calyx D glandular trichome of the abaxial surface of the calyx E eglandular trichome of the adaxial surface of the calyx F sector of opened corolla G glandular trichome of the abaxial surface of the corolla H gynoecium I fruit J seed K seed, in cross section L embryo A from Hatschbach 18030 B-H from Subils & Moscone 4273 I-L from Hunziker et al. 24993. 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 63. Capsicum flexuosum A fruiting branch B flowering node C stellate corolla with greenish spots D, H mature fruits E plant F flower bud G rotate-stellate corolla with purple and greenish spots A-D from Barboza et al. 1034 E Barboza et al. 3631 F-H from Carrizo Garcia 84. Photos by C. Carrizo Garcia.