Capsicum cardenasii Heiser & P.G.Sm., Brittonia 10(4): 195. 1958.

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

8. Capsicum cardenasii Heiser & P.G.Sm., Brittonia 10(4): 195. 1958.

Figs 42 , 43

Type.

Cultivated at Indiana University greenhouse from seeds sent by M. Cárdenas from market in La Paz, Bolivia, 15 Aug 1956, C.B. Heiser Jr. 4196 (Paul Smith Ac. -1793) (lectotype, designated here: IND [IND1000063, acc. # 139347]; isolectotype: IND [IND1000064, acc. # 139348]) .

Description.

Erect shrubs or subshrubs, 0.8-2 (-2.5) m tall, with the main stem 1-1.5 cm in diameter at base, much branched from near the base, the fragile branches in a typical “zig-zag” appearance above. Young stems strongly angled, green, glabrescent with sparse appressed-antrorse, simple, uniseriate, 4-6 (-7)-celled eglandular trichomes 0.08-0.6 mm long and minute, simple, glandular trichomes (stalk short; head dark); nodes green; bark of older stems greyish-white or with light brown-green fissures, glabrescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair subequal in size and shape. Leaves membranous, slightly discolorous, glabrescent, with sparse eglandular trichomes similar to those on stems and many small glandular trichomes (stalk unicellular; head dark, multicellular) on both surfaces, the glandular trichomes more abundant along mid-vein abaxially; blades of major leaves 3-5 (-6.5) cm long, 1.4-2.5 cm wide, narrowly ovate or ovate-lanceolate, the major veins 3-4 on each side of mid-vein, the base attenuate, the margins entire, the apex acute; petioles 1.2-2 cm long, glabrous or glabrescent; the blades of minor leaves 2-3.5 cm long, 0.9-1.2 cm wide, narrowly ovate or ovate-lanceolate, the major veins 2-3 on each side of mid-vein, the base attenuate, the margins entire, the apex acute or obtuse; petioles 0.5-0.8 cm long. Inflorescences axillary, 2-3 flowers per axil or flowers solitary; flowering pedicels 8-18 (-22) mm long, angled, erect to slightly spreading, geniculate at anthesis, entirely green, or purple distally, with moderate small glandular trichomes (stalk transparent, uni-bicellular; head dark, multicellular) and sparse short, antrorse eglandular trichomes; pedicels scars inconspicuous. Buds ellipsoid or ovoid, lilac or violet. Flowers 5-merous. Calyx 1-3 mm long, 2-3 mm wide, cup-shaped, thick, green or green with violet spots, moderately pubescent with the same glandular and eglandular trichomes as the pedicels, the calyx appendages five, 1-2 mm long, 0.3 mm wide, subequal, thick, erect or spreading, cylindrical, inserted close to the margin, sparsely pubescent with the same trichomes as the calyx tube. Corolla (6-) 6.5-12 mm long, 8-11 (-13) mm in diameter, thick, almost completely violet or lilac, but white at the base and along the main veins outside and within, sometimes greenish-yellow spots near the base within, campanulate with interpetalar membrane, lobed 1/3 or less of the way to the base, the tube 7-9 mm long, pubescent adaxially with short glandular trichomes (stalk 1-2-celled; head globose, unicellular) up to near its base, glabrous abaxially, the lobes (1.5-) 3-3.2 mm long, 2-2.4 mm wide, triangular, erect or spreading, alternating with five minute interlobes, glabrous adaxially and abaxially, the margins papillate, the tips acute, papillate. Stamens five, equal; filaments (4-) 6-7 mm long, whitish or lilac, inserted on the corolla 1.5-2 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5-1.8 mm long, ellipsoid, lilac or bluish, not connivent at anthesis. Gynoecium with ovary 1.5-1.85 mm long, 1.2-1.6 mm in diameter, green, ovoid or pear-shaped; ovules more than two per locule; nectary 0.4-0.6 mm tall, light green; styles homomorphic, 4.5-5.7 mm long, exserted ca. 1 mm beyond the anthers, lilac or purple, clavate; stigma ca. 0.2 mm long, 0.8 mm wide, discoid or globose, pale green. Berry 6-10 mm in diameter, globose or subglobose, green when immature, orange-red to bright red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 10-24 mm long, pendent, strongly angled, slightly widened distally, usually green; fruiting calyx 2-4 mm in diameter, persistent, not accrescent, discoid, green, the appendages 1-2.5 mm long, ca. 0.3 mm wide, appressed to the berry, spreading or reflexed. Seeds (4-) 5-13 per fruit, (2.5-) 3-4.2 mm long, (2.2-) 2.5-2.8 mm wide, C-shaped or subglobose, pale yellow to brownish-yellow, the seed coat reticulate to obscurely reticulate (SM), mostly cerebelloid (SEM), the cells irregular in shape, the lateral walls strongly sinuate in the central zone, rectangular to subpolygonal at margins; embryo imbricate.

Distribution.

Capsicum cardenasii is a narrow endemic species restricted mainly to the highlands of La Paz Department (Bolivia, Fig. 32 ). Only one collection from Tarija, probably introduced.

Ecology.

Capsicum cardenasii is a typical component of the warm and dry hillsides and remnants of forests in the inter-Andean valleys, growing preferentially in open places between cactus and Cassia , at 2,400-3,000 m elevation. It is cultivated by local people on small farms for local or family use of the fruits (Barboza, pers. obs.).

Phenology.

Flowering from December to March; fruiting from February to April.

Chromosome number.

n = 12 (Heiser and Smith 1958); 2 n = 2x = 24 ( Pickersgill 1977; Moscone et al. 2007; Scaldaferro et al. 2013, 2016).

Common name.

Bolivia. Ulupica (La Paz, Heiser & Smith 4196).

Indigenous name.

Bolivia. Uaika ( Aymará, La Paz, Barboza 4881).

Uses.

The fruits are harvested directly from wild plants and marketed locally on a small scale ( Jäger et al. 2013), mainly in La Paz, Bolivia. People consume dehydrated or fresh fruits in the preparation of a hot sauce called 'Jallpa huayka’ (in aymara), 'uchu llajfua’ (in quechua) (Heiser and Smith 1958; Cárdenas 1969), ‘llajwa’ (in quechua) or ‘llaswa’ (most popular), a mix of ‘tomato’, ‘onion’, ‘ulupica’ or other ‘chiles’ and aromatic herbs. Fruits are also preserved in vinegar or in oil and used as pickle ( ‘escabeche’) or they are cooked in boiling water before being sold ( National Research Council 1989; Barboza, pers. obs.).

Preliminary conservation assessment.

EOO (1,032.864 km2); AOO (32 km2). Capsicum cardenasii is a geographically isolated species from the dry valleys of Luribay (Prov. Loayza), not far from La Paz; based on its extent of occurrence and the number of localities (6), it is assigned a status of Endangered (EN; B1ab(iii,iv)). It is harvested by local people; its area of distribution is poorly known.

Discussion.

Capsicum cardenasii is resolved within the Purple corolla clade ( Carrizo García et al. 2016). More recent preliminary phylogenetic evidence showed that C. pubescens is sister to this clade ( Carrizo García et al. 2019; CCG, pers. obs.), thus circumscription of these taxa is under revision (CCG, pers. obs.; see under C. pubescens description). The fruits of C. cardenasii are very similar to those of C. eximium and C. eshbaughii , both also known as “ulupica”. The three species can be differentiated by their general pubescence and corollas. Capsicum eshbaughii (Fig. 58 ) has a dense pubescence of long furcate glandular trichomes and stellate white corollas with greenish-yellow spots around the throat (rarely purple lines in the lobes). In contrast, C. cardenasii and C. eximium have sparse to moderate pubescence of eglandular simple trichomes and minute simple glandular trichomes. In addition, C. cardenasii has lilac to purple, broadly campanulate corollas (Fig. 43C-F ), while C. eximium has mostly purple, lilac or magenta, stellate corollas with greenish-yellow pigmentation within (Fig. 60H-J ).

Phytogeographically, Eshbaugh (1979) suspected that C. cardenasii and C. eximium could be sympatric on the eastern margin of the range of C. cardenasii . He had observed some intermediate plants between both taxa in the Luribay Valley ( Eshbaugh 1976, 1979, but specimens not cited) and, at that time, no collections of the typical C. eximium were known from that area ( Eshbaugh 1979). We recently collected C. eximium (e.g. Barboza 4885) in Luribay, very near to the sites where C. cardenasii grows abundantly. Luribay Valley deserves to be explored intensively to document the presence of natural hybrids in the area and to find out if their level of fertility is as high as in experimental crosses (Heiser and Smith 1958; Eshbaugh 1976). Evidence of hybridisation was found while attempting to identify and cytogenetically characterise these Capsicum species, using a molecular cytogenetic approach (seeds from Gene Bank, Nijmegen University, The Netherlands); however, the fertility of the hybrid has not been possible to ascertain ( Scaldaferro 2019).

Eshbaugh and Smith (1971) also obtained successful crosses between C. cardenasii and C. eshbaughii (e.g. IND 139349), though F1 and F2 hybrids were less fertile than with C. eximium . The narrow distributions of Capsicum cardenasii and C. eshbaughii are allopatric, with the first species in north-western Bolivia (La Paz: Luribay) and the second concentrated in central-eastern Bolivia (mainly Santa Cruz: Samaipata). Their distributions are separated by nearly 700 km of distance, making hybridisation in the wild highly unlikely.

Although the number of collections of C. cardenasii obtained in the field are scarce (8), the ease with which the seeds of this species germinate and produce fertile plants explains the large numbers (> 20) of specimens (and duplicates) gathered from plants in cultivation that are housed in many herbaria (see Specimens Examined).

The type collection of C. cardenasii consists of flowering specimens (two sheets dated 15 Aug 1956 at IND) obtained from seeds bought at the La Paz (Bolivia) marketplace; it is supposed that fruits came from warm, dry places along the Río Abajo, near La Paz, at 2400 m altitude (Heiser and Smith 1958). Of the two specimens in IND, that with barcode 1000063 is the most complete, is labelled type and is here designated as the lectotype. There are specimens distributed in other herbaria (e.g. CORD, IND, LIL, US) with the same collection number as the lectotype (Heiser 4196, Paul Smith Acc. 1793), but these have different dates of collection and should not be considered as duplicates of the lectotype.

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 32. Distribution of C. benoistii, C. caatingae, C. caballeroi, C. campylopodium and C. cardenasii

Figure 42. Capsicum cardenasii A flowering branch B eglandular trichome of the calyx C glandular trichome of the corolla D glandular trichome of the adaxial surface of the calyx E flower F section of the calyx showing the venation G sector of opened corolla H gynoecium I fruit J anatomical detail of the pericarp (note the giant cell in the mesocarp) K seed L seed, in cross section M structure of seed coat at the seed margin N structure of seed coat at the seed body O embryo A-H from Eshbaugh 1527 I-O from Eshbaugh 2046 J. Drawn by N. de Flury.

Figure 43. Capsicum cardenasii A plant B flower bud C, D flower, in lateral view E, F flower, in front view (note the different colouration inside the corolla) G immature fruits H mature fruit A, B, D, E, G from Barboza 4882, C, F, H from Barboza 4881. Photos by G. E. Barboza.

Figure 58. Capsicum eshbaughii A plant B flower buds C flower, in pre-anthesis D-G stellate corollas with different colouration patterns H flower buds and flower in anthesis, seen from behind I immature fruit J mature fruits A, E, H-J from Carrizo Garcia 67, photos by G. E. Barboza and C. Carrizo Garcia B, C, D, F, G from Palombo 19, photos by N. Palombo.

Figure 60. Capsicum eximium A plant (wet habitat) B plant (dry habitat) C main fissured stem D young inflorescence E-G flower buds with different colouration H-K flowers, in front view, with different corolla colouration within L flower, in lateral view M, N flowers seen from behind O mature fruits A, G, N from Barboza et al. 4914 B, E, M from Barboza et al. 4885 C, K from Barboza et al. 4903 D from Wageningen Netherlands University germplasm collection F, I from Barboza et al. 4895 H, O from Barboza 1919 (cult.) J from Barboza et al. 4896 L from Barboza et al. 3543 A-C, E-O photos by G. E. Barboza, D photo by P. Bosland.