Capsicum tovarii Eshbaugh, P.G.Sm. & Nickrent, Brittonia 35(1): 55. 1983.

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

42. Capsicum tovarii Eshbaugh, P.G.Sm. & Nickrent, Brittonia 35(1): 55. 1983.

Figs 117 , 118

Type.

Peru. Huancavelica: Prov. Tayacaja: Andaimarca, entre Colcabamba y Surcubamba, valle del Mantaro , 2000 m elev., 14 Apr 1954, O. Tovar 1867 (holotype: USM [USM000895]; isotypes: MU (n.v.), US [00027402, acc. # 3048334], USM [USM248309]).

Description.

Erect subshrubs or shrubs, 0.5-1.5 (-2.5) m tall, with the main stem thick, 1.5-3.5 cm in diameter at base, much branched from the base, the branches fragile and scandent. Young stems 2-3-angled, green, rigid, glabrescent to moderately pubescent with appressed-antrorse or spreading, simple, uniseriate, 2-6-celled, eglandular trichomes 0.08-0.9 (-1.2) mm long and sparse small, simple, glandular trichomes (stalk short, unicellular; head dark, oblong, multicellular); nodes green, bark of older stems greenish-brown, fissured, glabrescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair subequal in size and shape. Leaves membranous, discolorous, dark green above, greenish-grey below, moderately to densely pubescent, with simple eglandular trichomes and glandular trichomes similar to those of the stems, these latter especially abundant on the veins abaxially; blades of all leaves (2.5-) 4-15.5 cm long, (1-) 1.5-5.7 cm wide, ovate or lanceolate, the major veins 3-5 on each side of mid-vein, the base short-attenuate and asymmetric, the margins entire, the apex acuminate; petioles 1-3 cm long, densely pubescent. Inflorescences axillary, (2-) 4-8 flowers; flowering pedicels 3-10 mm long, short, angled or terete, spreading to pendent, non-geniculate at anthesis, green, moderately pubescent, the eglandular trichomes long, spreading; pedicel scars inconspicuous. Buds globose to ovoid, yellowish-green. Flowers 5-merous. Calyx 1.5-2 mm long, 1.8-2.5 mm wide, cup-shaped, pentagonal in outline, thin, light green with dark green veins, moderately pubescent with the same eglandular trichomes as stems and leaves, sometimes sparse glandular trichomes, the calyx appendages absent or five, equal, 0.5 mm long, inserted very close to the margin, sparsely pubescent. Corolla (4.5-) 6-8 mm long, 8-10 mm in diameter, yellowish-green with lilac interpetalar membrane outside, purple with greenish-yellow spots in the lobes and greenish-yellow centre within or lobes mostly yellowish-green and diffuse lilac pigmentation within, campanulate or campanulate-stellate, lobed less than 1/3 of the way to the base, the tube 3.8-4.62 mm long, with sparse small glandular trichomes (stalk unicellular; head globose, unicellular) adaxially, glabrous abaxially, the lobes 2.5-3.2 mm long, 2.5-3.5 mm wide, ovate, erect, glabrous adaxially and abaxially, the margins papillate, the tips acute, slightly papillate. Stamens five, equal; filaments 1.6-2.5 mm long, lilac, inserted on the corolla 0.8-1.3 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5-1.8 (-2) mm long, ellipsoid, purple or lilac, somewhat connivent or not connivent at anthesis. Gynoecium with ovary 1.2-2 mm long, 1-1.5 mm in diameter, 2 (-4)-carpellate, light green, ovoid or subglobose; ovules more than two per locule; nectary ca. 0.4 mm tall; styles dimorphic, short style 1-1.6 mm, not exceeding the anthers, long style 4.2-7 mm long, exserted ca. 1 mm beyond the anthers, cream with lilac pigmentation, clavate; stigma 0.2-0.37 mm long, ca. 0.75 mm wide, capitate or lobed, pale yellow. Berry 5-7.2 mm in diameter, globose, green turning nearly black when immature, bright red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 7-12 (-15) mm long, erect, strongly angled, widened distally, green; fruiting calyx 4-6 mm in diameter, persistent, not accrescent, discoid, the appendages 0.0-0.6 mm long, appressed to the berry. Seeds 4-12 per fruit, 3.7-4.8 mm long, 2-3 mm wide, C-shaped or subglobose, yellow to brownish-yellow, the seed coat reticulate (SM), reticulate-cerebelloid (SEM), the cells irregular in shape, the lateral walls mostly sinuate, wavy at margins; embryo imbricate or coiled.

Distribution.

Capsicum tovarii is an Andean endemic species from central Peru ( Junín and Huancavelica Departments) (Fig. 119 ).

Ecology.

Capsicum tovarii grows in xerophytic to mesophytic habitats with abundant columnar cacti, Bombax ruizii K.Schum. ( Eshbaugh et al. 1983) and Prosopis spp. across the Rio Mantaro Basin. It is common in full sun or protected under trees, between 850 and 2,500 m elevation.

Phenology.

The collections seen are from February to May, all in flower and fruit.

Chromosome number.

n = 12 ( Eshbaugh et al. 1983); 2 n = 2x = 24 ( Moscone et al. 2007; Scaldaferro et al. 2016).

Common names.

Peru. Ají silvestre (Huancavelica, Tovar 1867), Mucuru (Huancavelica, Tovar 1867; Junín, Barboza 5044), Mucuru-uchu (Huancavelica, Tovar 1867).

Uses.

The fruits are very pungent and eaten by local people (Tovar 1867, Barboza 5044), used as a spice ( Eshbaugh et al. 1983).

Preliminary conservation assessment.

EOO (1,422.442 km2); AOO (28 km2). Capsicum tovarii occupies a very narrow geographic range in Central Peru. Based on its EOO, AOO and the few nearby locations where it was found, none of them in protected areas and considering that the quality of the habitat will decline by the increased agricultural development, we assign the Endangered (EN; B1+B2ab(iii)) status to this species.

Discussion.

Capsicum tovarii appears as an isolated branch in the phylogeny of the genus ( Carrizo García et al. 2016, see Fig. 1 ) and it is recognised as the distinct Tovarii clade. The closest affinities of C. tovarii have been the subject of debate (see Carrizo García et al. 2016 for detailed information); it has been suggested to be close to either the purple-flowered group ( C. eximium , C. cardenasii or C. pubescens ) or the white-flowered species ( C. baccatum complex). An isolated position as sister group to these two groups (the Baccatum and Annuum clades which encompass the most important domesticated species) has recently been proposed for this species ( Carrizo García et al. 2016).

This species has been rarely collected (<10 collections) in Central Peru and the majority of the collections have been made by the late Peruvian botanist Oscar Tovar (USM). The species deserves more field observations to better understand the morphological variation in trichome type, corolla colour, heteromorphism of styles, number of carpels, breeding system and area of distribution. The pubescence varies from glabrescent with a predominance of small glandular trichomes along leaf veins, petioles and pedicels to moderately pubescent with predominance of antrorse eglandular trichomes hiding the glandular ones (but these were somewhat visible by virtue of their dark head). The corolla colour has been annotated as purple or purple-blue in Tovar´s labels (e.g. Tovar 1867, 5012 & 5363), but corollas that are entirely cream or cream with two green spots within are also reported in the protologue (Eshbaugh et al. 2013). Similar cream or pale yellow corollas have been observed in plants obtained from seeds from Pariahuanca-Huancayo (sent by O. Tovar in 1999 to A. T. Hunziker, CORD), in which corollas were pale yellow with two green spots at the base of each lobes and throat, while others had purple tones in the interpetalar membrane (Hunziker 25654). In a different plant from the same seed accession (Hunziker 25655), corollas displayed an evident purple pigmentation in the lobes and adjacent zone of the throat, as was seen by Barboza in another population from Potrero, Huancayo (Barboza 5044, Fig. 118F, G ). The style is dimorphic, flowers either have short (1-1.6 mm long) or long styles (4.2-7 mm long - only a few styles measured) and the ovary is 2-4-carpellar (also needs more observations). Eshbaugh et al. (1983) mentioned that the flowers were functionally unisexual or bisexual, but more information is needed to associate this with any morphological trait.

Figures

Figure 1. Capsicum phylogeny. Cladogram summarising findings from Carrizo Garcia et al. (2016) and Barboza et al. (2019, 2020 b). Only one sample per species has been included; species names have been updated. Clades distinguished are labelled. Asterisks indicate branches with moderate / low support, letters ' d' clades with domesticated taxa.

Figure 117. Capsicum tovarii A flowering branch B, C eglandular trichomes of the leaf D, E glandular trichomes of the leaf F flower G calyx showing the venation H sector of opened corolla I glandular trichome of the abaxial surface of the corolla J gynoecium with long style K gynoecium with short style L fruit M seed N seed, cross section O structure of seed coat at the seed margin P structure of seed coat at the seed body Q embryo. From Eshbaugh E 1137. Drawn by N. de Flury.

Figure 118. Capsicum tovarii A plant B leaf, adaxial surface C leaf, abaxial surface D node with flower buds on pendent pedicels E flower, in lateral view F, G flower in front view, showing different pattern of purple colouration H immature fruits I mature fruit. From Barboza 5044. Photos by G. E. Barboza.

Figure 119. Distribution of C. tovarii and C. villosum.