Lycianthes tricolor (Dunal) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 385. 1919

Dean, Ellen, Poore, Jennifer, Anguiano-Constante, Marco Antonio, Nee, Michael H., Kang, Hannah, Starbuck, Thomas, Rodrigues, Annamarie & Conner, Matthew, 2020, The genus Lycianthes (Solanaceae, Capsiceae) in Mexico and Guatemala, PhytoKeys 168, pp. 1-333 : 1

48 Lycianthes tricolor (Dunal) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 385. 1919 Fig. 108

Solanum tricolor Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 756. 1814 [1813] (official date of publication of volume 3 is 1813, but pages 369-780 were published in 1814 [ Stafleu and Cowan 1979]). Type: Painting made during the Royal Expedition to New Spain (1787-1803) under the direction of Martin de Sessé y Lacasta (lectotype designated by Dean et al. 2017a, pg. 205: Hunt Institute for Botanical Documentation HI Art acc. # 6331.04610).

Solanum monodynamum Visiani, Sem. Hort. Patav. (1841): 3. 1841. Type: cultivated plant in the botanical garden at Padua (Padova), Italy, seed source from Mexico (neotype designated by Dean et al. 2017a, pg. 205: La Mexique, Jardin de Padoue, 2 Oct 1844, collector unknown s.n. (G-DC [G00145628]).

Solanum pilosissimum M.Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12(1): 139. 1845. Type: Mexico, Oaxaca, Yavezia, 7000 ft, Nov-Apr, 1840, H. Galeotti 1228 (lectotype designated by Dean et al. 2017a pg. 205: BR [00000552342]; isolectotypes: BR [00000552309], G [G00359732], P [P00368470], W [acc. # 3080]).

Solanum quadriflorum M.Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12(1): 139. 1845. Type: Mexico, Oaxaca, Cordillera, 7000 ft, Nov-Apr 1840, H. Galeotti 1231 (lectotype designated by Dean et al. 2017a pg. 206: BR [00000552205]; isolectotypes: BR [00000552238], P [P00369232, two sheets]).

Solanum nyctaginoides Dunal, Prodr. [A. P. de Candolle]13(1): 172. 1852. Type: Hort. Mexican (also on another duplicate Jardín Bot. D. Mexico), 12/7 (12 Jul) 1827, J. Berlandier 553 (lectotype designated by Dean et al. 2017a, pg. 205: G-DC [G00145660]; isolectotypes: G [G00359731, G00301639], P [P00385102, P00385101], W [acc. # 0003084].

Lycianthes pilosissima (M.Martens & Galeotti) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 378. 1919. Type: Based on Solanum pilosissimum M.Martens & Galeotti.

Lycianthes tricolor (Dunal) Bitter var. flavidipila Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 387. 1919. Type: Guatemala, Vulcán de Agua, 2,500 m, 31 May 1882, F. C. Lehmann 1486 (lectotype designated by Dean et al. 2017a, pg. 205: G [G00414712, 2 sheets]; isolectotypes: US [00027898], K [K000585741], B [cited by Bitter (1919), not found, likely destroyed]).

Lycianthes tricolor (Dunal) Bitter var. primoaurata Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 387. 1919. Type: Guatemala, Volcán de Fuego, ridges above calderas, 8300 ft, Sep 1873, O. Salvin s.n. (holotype: W [acc. # 1886-8331]).

Lycianthes tricolor (Dunal) Bitter var. hirsutior Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 388. 1919. Type: Mexico, Castresana [locality likely Cuatresana near San Pedro Nolasco, Oaxaca],1842, F. Liebmann 1438 (lectotype designated by Dean et al. 2017a, pg. 205: C [10021746]; isolectotypes: C [10021745], US [00027899]).

Lycianthes arrazolensis (J.M.Coult. & Donn.Sm.) Bitter var. patentipila Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 391. 1919. Type: Mexico. Oaxaca, 1750 m, Conzatti & González 1071 (holotype: GH [00936247]).

Type.

Based on Solanum tricolor Dunal.

Description.

Shrub, 1.2-4.5 m tall, sometimes vining or arching through neighboring vegetation. Indument of light yellow, uniseriate, multicellular, simple, often curly, eglandular, spreading to appressed trichomes 0.2-1.75 mm long, sometimes with very small glistening bumps between longer trichomes. Stems green to purple (drying tan) with dark lenticular vertical striations when young, sparsely to densely pubescent, not much compressed when dried in a plant press, becoming light brown and woody with age; upper sympodial branching points monochasial and dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades (4.6) 6.3-11.8 (15) × (1.9) 2.5-6.5 (8.8) cm, the smaller ones with blades 1.7-5.2 (8.5) × 0.9-3.9 (6.3) cm, the leaf pairs similar in shape, the blades ovate to elliptic, chartaceous, moderately pubescent, the trichomes usually densely spreading outward (towards the margins) along the abaxial veins, especially at the base of the main vein, the base usually cuneate (rarely attenuate), sometimes oblique, the margin entire, usually irregularly undulate (rarely coarsely dentate), the apex acuminate, the petiole 0.5-3.3 cm long, the larger leaf blades with 4-6 primary veins on each side of the midvein. Flowers solitary or in groups of 2-7 (10), axillary, oriented horizontally; peduncles absent; pedicels (8) 13-33 mm long and erect in flower, (15) 18-33 (40) mm long and erect in fruit, moderately pubescent; calyx 2.3-3.5 (4.1) mm long, 2.5-3.5 (4) mm in diameter, obconic to campanulate, moderately to densely pubescent, the margin truncate, with 10 erect, linear appendages (0.5) 1-4.3 mm long emerging 0.25-0.5 mm below the calyx rim; fruiting calyx enlarged, bowl-shaped to rotate, 1.5-3.2 mm long, 5-8.5 mm in diameter, the appendages to 6 mm long; corolla (0.7) 0.9-1.8 cm long, rotate in orientation, entire in outline, with abundant interpetalar tissue, adaxially white to pale violet with purple stripes along the major veins, with three green spots located between the short stamens, glabrous, abaxially green on the lobes, moderately puberulent near the veins; stamens unequal, straight, the four short filaments 1-2 (3) mm long, the one long filament (2.75) 3.5-4.2 (5) mm long, glabrous, the anthers 2.5-4.5 mm long, lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing toward the style or distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6-9 mm long, linear (sometimes curved upward at the tip), glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 6-11 mm long, 6-10 (15) mm in diameter, globose, red at maturity, glabrous, lacking sclerotic granules. Seeds 10-30 per fruit, (2) 2.5-4 × 2-3 mm, flattened, reniform in outline, with deep notch on side (usually more than 0.5 mm deep), yellow-orange, the surface reticulum with minute serpentine pattern and shallow luminae.

Chromosome number.

Unknown.

Distribution and habitat.

Mexico (Chiapas, Guerrero, Jalisco, Michoacán, Oaxaca), Guatemala (Chimaltenango, Escuintla, Guatemala, Huehuetenango, Quetzaltenango, Quiché, Sacatepéquez, San Marocis, Sololá), and El Salvador on steep slopes, in ravines, and in semi-disturbed areas such as roadside thickets, path edges, and disturbed agricultural areas in mixed broadleaved forest, oak forest, oak/pine forest, fir forest, and cloud forest, 2100-3000 m in elevation (Fig. 109 ).

Common names and uses.

None known.

Phenology.

Flowering specimens have been collected from February through October; specimens with mature fruits have been collected January through December. Field observation of the corollas indicates that the corollas are open in the early morning and closed by late morning ( Dean et al. 2017a).

Preliminary conservation status.

Lycianthes tricolor is a widespread species ranging from southwestern Mexico to El Salvador, represented by 82 collections and occurring in seven protected areas. The EOO is 22,404.475 km2, and the AOO is 308 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion.

This species is very similar to Lycianthes arrazolensis from which it can be distinguished by seed shape. The seeds of L. tricolor have a definite sharp notch that is usually deeper than 0.5 mm, whereas the seeds of L. arrazolensis lack this notch. In some Mexican L. arrazolensis populations in the states of Morelos and México, the seeds are shallowly indented on one side, but this indentation is less than 0.5 mm and usually less than 0.25 mm, and never sharply notched ( Dean et al. 2017a). Non-fruiting specimens can be challenging to identify. The following non-seed characters can be helpful: the calyx rim of L. arrazolensis tends to be more prominent, often protruding beyond the appendage insertion by over 0.5 mm, while the calyx rim of L. tricolor is usually less than 0.5 mm; the appendages of L. arrazolensis tend to bend away from the rim, exposing the rim, while the appendages of L. tricolor are oriented closer to the rim and corolla, hiding the rim; the pores of the short stamens in L. arrazolensis usually face away from the style, while those of L. tricolor usually face toward the style; the flowering pedicels of L. tricolor are usually 13-33 mm long, becoming much longer in fruit, while those of L. arrazolensis are usually less than 15 mm in flower and 21 mm in fruit (however shorter pedicels in L. tricolor are found in Guatemala and Chiapas); and the leaves of typical L. arrazolensis tend to have obvious geminate leaf pairs with elliptic to obovate blades and leaf bases often attenuate into the petiole, while in L. tricolor the small geminate leaf often abscises early, and the laminas are more ovate with a less-attenuate leaf base ( Dean et al. 2017a).

Representative specimens examined.

Guatemala. Chimaltenango: Reserva Natural Privada El Encanto de Tecpán, Caserlo Panimachavac, Aldea Chajaljya, Tecpán Chimaltenango, 14.8142, -90.9872, 2317 m, 8 Apr 2014, B. Escobar 87 (BIGU). Escuintla: Volcán de Agua of Finca Rosario de La Vista Hermosa, [14.4411, -90.7506], 6000-8000 ft, 22 May 1971, R.L. Wilbur 14748 (DUKE). Guatemala: along old road to San Lucas, vicinity of San Rafael, 1800 m, 27 Sep 1972, A. Molina 27614 (MEXU, TEX). Huehuetenango: Chanximil, Aldea San Martín, todos Santos Cuchumatán, 15.5531, -91.6986, 2185 m, 16 Sep 2006, J. Morales 3889 (MO). Quetzaltenango: Mpio. Zunil, road to Fuentes Georgiñas, 4 km S of Zunil, northwest slopes of Volcán Zunil, [14.7542, -91.4828], 2442 m, 4 Jan 2009, T. Sultan Quedensley 7013 (BIGU). Quiché: Nebaj, about 4 km W, 6700 ft, 11 Jun 1964, E. Contreras 4953 (MO, LL). Sacatepéquez: Volcán de Agua, [14.4708, -90.7392], 2500 m, 15 Aug 2003, P. García 26 (BIGU, MEXU, TEX). San Marcos: 20 miles S of San Marcos along road from San Raphael, 2100 m, 13 Jul 1977, T. B. Croat 41006 (MO). Sololá: Mpio. San Pedro La Laguna, Volcán San Pedro, [14.6564, -91.2672], 2500 m, 27 Jul 2005, P. Pardo 665 (BIGU). Mexico. Chiapas: Reserva de la Biosfera El Triunfo, sendero Finca Prusia-Campamento El Triunfo, 22.15, -99.5667, 1900 m, 12 Nov 2004, N. Martínez- Meléndrez 587 (MEXU). Guerrero: Omiltemi, Barranca Potrerillos, [17.5595, -99.6759], 2170 m, 28 Nov 1993, C. González 241 (MEXU). Jalisco: parte alta del ejido de Pabelo, cerca del predio Las Iglesias, 4.4 km al sur-suroeste de Santa Monica (Mpio. Ayutla) y 4 km al este-noreste de Plaza de Gallos, 19.8922, -104.5464, 2120-2160 m, 12 Aug 2012, P. Carrillo-Reyes 6743 (IBUG, IEB, MEXU). Michoacán: Mpio. Coalcomán, 14 km al SE de Varaloso, sobre el camino a Barranca Seca, 18.6536, -102.9769, 2000 m, 22 Dec 2007, V. Steinmann 6121 (DAV). Oaxaca: Dto. Sol de Vega, Pena Ahumada, 16.7256, -97.0781, 2 Sep 2006, A. Zárate Marcos 407 (IEB).

Figures

Figure 108. Image of herbarium specimen of L. tricolor, Breedlove 25699 (MO). Specimen used with permission from the Missouri Botanical Garden (http: // www. tropicos. org).

Figure 109. Map of geographic distribution of L. tricolor based on herbarium specimen data.