Capsicum mirabile Mart., Fl. Bras. (Martius) 10(6): 144. 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

29. Capsicum mirabile Mart., Fl. Bras. (Martius) 10(6): 144. 1846.

Figs 88 , 89

Capsicum mirabile Mart. var. grandiflorum Sendtn., Fl. Bras. (Martius) 10(6): 144. 1846. Type. [Brazil]. Brasilia, F. Sellow 209 (possible original material B, destroyed [F neg. 2871], no additional material found).

Capsicum buforum Hunz., Kurtziana 5: 394. 1969. Type. Brazil. São Paulo: Eugênio Lefévre, a unos 26 km de Campos do Jordão, ca. 1400 m elev., 5 Dec 1967, A.T. Hunziker 1956 2 (lectotype, designated here: CORD [CORD00006616]; isolectotypes: CORD [CORD00006617, CORD00006618]).

Type.

Brazil. "Habitat in sylvis fere ubique per Provinciam Sebastianopol. et Paulinam", Dec., C.F.P. von Martius s.n. (lectotype, designated by Barboza 2011, pg. 30: M [M-0171538]) .

Description.

Erect shrubs or subshrubs (0.70-) 1.2-2.5 (-3) m tall or rarely small trees, with the main stem thick 2.5-4 cm in diameter at base, much branched above, the branches dichotomously spreading in a typical “zig-zag” appearance. Young stems angled, fragile, green, glabrous or glabrescent, with antrorse, curved, simple, uniseriate, 3-4-celled, eglandular trichomes 0.1-0.3 mm long, new growth with sparse whitish pubescence; nodes purple, slightly purple or green; bark of older stems dark brown, angled, glabrous; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous to coriaceous, slightly discolorous to discolorous, green above, light green beneath, glabrous adaxially, glabrescent abaxially and margins, especially on the veins, with sparse, 2-3-(-6)-celled, eglandular trichomes 0.2-0.5 mm long; blades of major leaves 5-13.5 cm long, (1-) 1.3-3 (-5.5) cm wide, elliptic or narrowly elliptic to ovate, the major veins 4-6 (-7) on each side of mid-vein, the base asymmetric and attenuate, the margins entire, the apex acuminate to long-acuminate; petioles 0.7-2 (-2.5) cm long, glabrous; the blades of minor leaves 2-3.3 (-4) cm long, 0.9-1.7 cm wide, elliptic or ovate, the major veins 3-4 on each side of mid-vein, the base attenuate, the margins entire, the apex acute; petioles 0.2-0.4 cm long, glabrous. Inflorescences axillary, 2-5 (-7) flowers per axil, rarely flowers solitary; flowering pedicels (13-) 16-25 mm long, terete or slightly angled, erect or slightly spreading, geniculate at anthesis, green or greenish-purple, glabrous or glabrescent, the eglandular trichomes short, antrorse; pedicels scars inconspicuous. Buds globose to ovoid, inflated, greenish-purple or dark purple or clearer near anthesis. Flowers 5-merous. Calyx 1.5-2.2 mm long, ca. 2-2.5 mm wide, cup-shaped, thin, yellowish-green or green, glabrous or glabrescent, the calyx appendages five, (0.4-) 0.5-1.5 (-3) mm long, subequal, thick, green, erect or spreading, cylindrical, inserted very close to the margin. Corolla (6-) 7.5-12 mm long, (9-) 10-13 mm in diameter, thick, purple with white margin and yellowish-green at the base outside, similar colour within, but sometimes the purple or reddish-brown spots diffuse up to absent, stellate with interpetalar membrane, lobed halfway or less of the way to the base, the tube 3.5-5 mm long, with a continuous ring of glandular trichomes (stalk long, 1-3-celled; head globose, peltate, unicellular) adaxially, glabrous abaxially, the lobes 3-5 mm long, 3.2-4 mm wide, broadly triangular, spreading, glabrous abaxially and adaxially, the margins papillate, the tips cucullate, papillate. Stamens five, equal; filaments (2.8-) 3-3.8 (-4) mm long, equal, light green or greenish-white, inserted on the corolla 1-1.8 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5-2 mm long, ellipsoid, yellow when young and reddish-brown or grey-purple post-dehiscent, not connivent at anthesis. Gynoecium with ovary 1.1-1.5 mm long, 1-1.2 mm in diameter, green, globose to ovoid; ovules more than two per locule; nectary ca. 0.3 mm tall; styles homomorphic, 3.5-4.7 mm long, at the same level or barely exserted beyond the anthers, yellowish-white or light green, clavate; stigma 0.2 mm long, 0.6-0.8 mm wide, discoid or globose, pale green. Berry 7-9 mm in diameter, globose, green when immature, greenish-golden yellow, deciduous, pungent, the pericarp thin, translucent, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 20-32 mm long, pendent and curved, angled, widened and with a slight constriction distally, green; fruiting calyx 3-3.5 mm in diameter, persistent, not accrescent, green, discoid, the appendages 0.7-3.5 mm long, 0.3-0.4 mm wide, spreading. Seeds (2-) 3-8 (-10) per fruit, 2.5-3.2 mm long, 2-2.5 mm wide, C-shaped, brownish-black to black, the seed coat reticulate and tuberculate at margins (SM), reticulate with pillar-like outgrowths at margins (SEM), the cells polygonal in shape, the lateral walls straight; embryo coiled.

Distribution.

This species is endemic to east and south-eastern Brazil (Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro and São Paulo States) (Fig. 86 ).

Ecology.

Capsicum mirabile is found in the understorey, edges and interior of the wet Atlantic Forest (Mata Atlântica), primarily in the Floresta Ombrófila Densa Montana, at 800-1,900 m elevation.

Phenology.

Flowering from November to April and fruiting from December to May.

Chromosome number.

n = 13 ( Tong and Bosland 2003; Pozzobon and Schifino-Wittmann 2006, as Capsicum sp 6 and C. buforum ); 2 n = 2x = 26 ( Moscone et al. 2007).

Common name.

Brazil: Pimenta do sapo ( São Paulo, Hunziker 1956 2).

Uses.

None recorded.

Preliminary conservation assessment.

EOO (190,248.624 km2); AOO (224 km2). Based on the large extent of occurrence, as well the many collections made in several protected areas (Parque Nacional Serra das Lontras, PN do Caparaó, Parque Estadual Serra do Brigadeiro, PN Itatiaia, Parque Estadual Pico do Itacolomi, Reserva Florestal Uaimi, Parque Estadual Três Picos, PN da Serra dos Órgãos, Parque Estadual Maciço da Pedra Branca, Estação Biológica de Boracéia and PN Serra do Bocaina), we assign Capsicum mirabile the Least Concern (LC) category. Subpopulations are not rare in the interior of forests, but this species could be at risk due to the fragmentation and loss of its primary forest habitat outside the legal conservation units.

Discussion.

Capsicum mirabile is an enigmatic and morphologically variable species of the Brazilian Atlantic Forest clade ( Carrizo García et al. 2016) and its interspecific relationships are still not well understood. Smith and Downs (1966) confused plants of C. recurvatum (e.g. Smith 7974, Reitz & Klein 5766, 5943 and Klein 2232) and Athenaea spp. (e.g. Reitz 4417, Ule 699, Klein 2417 and others) from Santa Catarina, Brazil, with C. mirabile , a region where this latter species does not occur. Furthermore, Hunziker’s use of the epithet Capsicum mirabile in herbaria has been erratic, making it difficult to understand its real circumscription. Initially, Hunziker (in litt.) had a broad and inclusive concept for this species, which motivated him to annotate specimens he considered different, but related to C. mirabile with the imprecise name " C. mirabile var.". Later, Hunziker (2001) accepted six species for eastern Brazil ( C. baccatum var. praetermissum , C. campylopodium , C. dusenii (here a synonym of C. cornutum ), C. mirabile , C. schottianum and C. villosum ), but he did not mention C. buforum which he described in 1969 and here is treated as a synonym of C. mirabile . Both names, C. mirabile and C. buforum have been used in literature for accessions used in cytogenetic studies ( Tong and Bosland 2003; Pozzobon and Schifino-Wittmann 2006; Pozzobon et al. 2006; Moscone et al. 2007). Barboza and Bianchetti (2005) and Barboza et al. (2011, 2020a) provided identification keys for the Brazilian species, finally resolving the ambiguities surrounding C. mirabile .

Capsicum mirabile is characterised by shrubby habit with dichotomous branching, glabrous or glabrescent pubescence, short petioles, elliptic to ovate leaves, few flowers (2-7) per node, five short to long calyx appendages, purple corolla with white margin and yellowish-green or yellowish-white centre and greenish-golden yellow fruits (Table 4 , Fig. 89 ).

The distinctive corolla spots of this species were not mentioned in the protologue (Martius 1846), although the other diagnostic characters were clearly stated; this may have introduced confusion into the understanding of this species. Variations in some of the morphological features have been observed in the field, such as the length of the calyx appendages, which can range from very short (<1 mm long, Rio de Janeiro, Espírito Santo and Minas Gerais States) to longer (up to 3 mm, São Paulo and Minas Gerais States) and the shape of the leaves (usually elliptic, but also ovate). The most variable trait is the extent and intensity of tones of the reddish-brown spots in the corolla (Fig. 89C-H ), covering partially or almost completely the lobes within ( Carrizo García et al. 2013 and see photographs in http://www.pepperfriends.org/dbpf/capsicum-mirabile_001.asp and http://www.pepperfriends.org/dbpf/capsicum-mirabile_003.asp). Variation in the corolla colour has also been observed in specimens in cultivation at the Federal University of Viçosa (seeds from Espírito Santo). Each corolla lobe had a unique purplish spot in some specimens (Hunziker 25235) or two intense purple spots (Hunziker 25243) or the spots were scarcely purple or nearly lacking all together (Hunziker 25236).

Capsicum mirabile is sympatric with C. friburgense near the top of Cerro Calêdonia (Nova Friburgo, Rio de Janeiro) where the latter is endemic. Both species share geniculate pedicels, a calyx with five appendages, greenish-golden yellow fruits and blackish-brown or black seeds. They are easily told apart by the shape and colour of the corolla (stellate and multi-coloured in C. mirabile vs. campanulate-urceolate and entirely lilac or pink in C. friburgense ) and the leaf shape (elliptic to ovate in C. mirabile vs. ovate in C. friburgense ).

The two species similar to C. mirabile are C. villosum and C. muticum . The main difference amongst the three species is the pubescence, since C. mirabile is almost completely glabrous (or sparsely pubescent) and the other two species are densely pubescent on stems, leaves, pedicels and calyx. Capsicum villosum has five calyx appendages (as C. mirabile ), but the corolla is smaller and the lobes are more white than purple and the greenish-yellow centre is more extensive in comparison with C. mirabile . Capsicum muticum lacks calyx appendages and the purple pigmentation is very scarce, as lines or small spots bordering the greenish-yellow centre of the corolla.

When Hunziker (1969b) described C. buforum , he stated that Hunziker 1956 2 was "Typus speciei (CORD)". Three sheets of this collection are held in CORD, but none of them has an annotation indicating intent by Hunziker; we select here the best of these (CORD00006616) as the lectotype.

Specimens examined.

See Suppl. material 4: Appendix 4.

Figures

Figure 86. Distribution of C. lycianthoides, C. minutiflorum and C. mirabile.

Figure 88. Capsicum mirabile A flowering branch B, C major and minor leaf D flower bud E flower, in lateral view F, G flowers, upper views H sector of opened corolla I gynoecium J fruit K fruit, in cross section L seed. From Hunziker 1956 2. Drawn by L. Sanchez.

Figure 89. Capsicum mirabile A plant B flower buds on geniculate pedicels C flower, in pre-anthesis D flower, seen from behind E, H flowers, in front view, showing corollas with different colouration F flower, in full anthesis G flower, seen from behind I immature fruits J mature fruit A, B, D-J from Barboza & Deanna 5024 C from Bianchetti & Bustamante 1518 A, B, D-H photos by G. E. Barboza C photo by L. Bianchetti I, J photos by R. Deanna.