Lycianthes arrazolensis (J.M.Coult. & Donn.Sm.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 388. 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

5 Lycianthes arrazolensis (J.M.Coult. & Donn.Sm.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 388. 1919 Fig. 14

Solanum arrazolense J.M.Coult. & Donn.Sm., Bot. Gazette 37: 421. 1904. Type: Guatemala. Guatemala: Arrazola, 5500 ft [1,600 m], Apr 1893 [protologue says Apr, 1893 but writing on label looks like Mar or Mai, 1893], E.T. Heyde et E. Lux 4736 (holotype: F [0073068F, acc. # 264950, photo negative 49338]; isotypes: B [not seen, cited by Bitter (1919), probably destroyed], G [G00379120], GH [01652206], K [K000585744], US [01014241, 00027461], probably elsewhere).

Type.

Based on Solanum arrazolense J.M.Coult. & Donn.Sm.

Description.

Shrub, 0.9-5 m tall, sometimes vining or arching through neighboring vegetation. Indument of light yellow (sometimes appearing tan or off-white), uniseriate, multicellular, simple, eglandular, spreading or appressed-ascending trichomes 0.1-1.5 (1.75) mm long, sometimes the stems with very small appressed trichomes between longer spreading trichomes. Stems green to violet when young (drying tan) with maroon/purple lenticular vertical striations (drying blackish), 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 or dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 7.5-22 × (1.8) 3-9 (11) cm, the smaller ones with blades (1) 3-9 (12.5) × (0.4) 1.2-5.7 (7) cm, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate (rarely lanceolate), chartaceous, glabrous to densely pubescent, the trichomes usually densely spreading outward (towards the margins) along the abaxial veins, especially at the base of the main vein, the base cuneate to attenuate, sometimes oblique, the margin entire, usually irregularly undulate, the apex acuminate, the petiole 0.2-3 cm long, sometimes absent, the larger leaf blades with 5-7 primary veins on each side of the midvein. Flowers solitary or in groups of 2-10 (19), axillary, oriented horizontally; peduncles absent; pedicels 4-15 mm long and erect in flower, 6-16 (21) mm long and erect in fruit, sparsely to densely pubescent; calyx (1) 1.5-2.5 (3) mm long, 2-3 (3.5) mm in diameter, obconic to campanulate, sparsely to densely pubescent (sometimes nearly glabrous in Oaxaca), the margin truncate, with 10 spreading, linear appendages 0.5-2.5 mm long (atypically to 5 mm in low elevation Guerrero populations) emerging 0.5-1 mm below the calyx rim; fruiting calyx enlarged, bowl-shaped to rotate, 1-2 mm long, 4-7 mm in diameter, the appendages to 2.5 (4) mm long (probably longer in Guerrero); corolla 0.6-1.2 (1.6) cm long, rotate to campanulate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white to pale violet, adaxially sometimes with purple stripes along the major veins of the lobes or with three green spots located between the short stamens, glabrous, the abaxial side of the lobes green, glabrous to sparsely puberulent near the veins; stamens unequal, straight, the four short filaments 0.5-2 mm long, the one long filament (1.25) 3-4 (5) mm long, glabrous, the anthers 2.5-4 (4.5) mm long, lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, the pores of the longest stamen dehiscing toward the style, the pores of the shorter stamens usually dehiscing away from the style (sometimes dehiscing distally, rarely inward), not opening into longitudinal slits; pistil with glabrous ovary, the style (5) 6.5-9 (10) mm long, linear, straight to curved upward at the tip, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 5-10 (11) mm long, 5-10 (11) mm in diameter, globose, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds (3) 10-108 per fruit, 1.2-2.5 (3) × 1-2.5 mm, flattened, elliptic, irregularly triangular, or oval in outline, not obviously notched (if slightly indented, indentation is usually less than 0.3 mm), yellow-orange, surface reticulum rough with indistinct serpentine pattern with shallow luminae.

Chromosome number.

2n = 24 ( Gentry and Pearce 1977).

Distribution and habitat.

Mexico (Chiapas, Guerrero, Jalisco, México, Michoacán, Morelos, Oaxaca), Guatemala (Alta Verapaz, Baja Verapaz, Chimaltenango, El Progreso, Escuintla, Guatemala, Quiché, Sacatepéquez, Sololá, Suchitepéquez), Belize, El Salvador, Honduras, and Nicaragua in wet canyons and drainages, often in riparian forest or disturbed forest, in oak, oak/pine, and tropical dry forest (higher elevation populations are often in hardwood cloud forest; south of Guatemala, it has been collected in high-elevation, dwarf cloud forest and Cupressus forest), 500-3000 m in elevation (Fig. 15 ).

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. The corollas of this species are open in the morning and closed by late morning.

Preliminary conservation status.

Lycianthes arrazolensis is a widespread species ranging from southern Mexico to Nicaragua, represented by 193 collections and occurring in eight protected areas. The EOO is 380,617.675 km2, and the AOO is 676 km2. Following the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion.

This species is very similar to Lycianthes tricolor . It can be easily distinguished from L. tricolor 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 usually less than 0.25 mm and never sharply notched ( Dean et al. 2017a). Non-fruiting specimens can be challenging to identify. If a specimen was collected below 1700 m in elevation, it is most likely to be Lycianthes arrazolensis . In addition, 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 pedicels of the oldest, largest flowers and the pedicels of the most mature fruits of L. arrazolensis are usually shorter than those of L. tricolor , although there is overlap in this characteristic; and the leaves of typical L. arrazolensis tend to have obvious geminate leaf pairs with elliptic to obovate laminas 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).

There is a wide range of morphological variation within Lycianthes arrazolensis as circumscribed here, especially in leaf size, leaf shape, and density of pubescence. This is particularly true of populations in the state of Oaxaca and neighboring Guerrero where small-leaved forms with very dense pubescence as well as large-leaved forms with sparse pubescence are found. The populations with larger leaves and sparser pubescence are usually found below 1,500 m, while those with denser pubescence are usually found above 2,000 m. The degree of variation found in this species is worthy of more study ( Dean et al. 2017a).

Representative specimens examined.

Guatemala. Alta Verapaz: 3 kms de Villa Hermosa, [14.87, -91.57], 1400 m, 16 May 1963, A. Molina R. 12362 (NY): Baja Verapaz: Mpio. San Jerónimo, Santa Elena la Cumbre, 15.0292, -90.2167, 2263 m, 12 Nov 2009, A. Cóbar 1980 (BIGU). Chimaltenango: Volcán Acatenango, Aldea Quisache, 14.5181, -90.2844, 500 m, 19 May 2004, M. Velíz 15260 (MEXU). El Progreso: 15 km N of Morazo, [14.975, -90.2067], 17 Jul 1970, W.E. Harmon 3207 (MO). Escuintla: 2 km W of San Vicente Pacaya, [14.4225, -90.6464], 1500 m, 31 May 1970, W.E. Harmon 2438 (MO). Guatemala: Villa Canales, Fea. San Agustín Las Minas, 14.5264, -90.495, 1839 m, 12 Aug 2010, L. Velásquez 1460 (BIGU). Quiché: Nebaj, Batzchocolá, 15.5714, -91.1035, 1300 m, 1 Aug 2017, E. Tribouillier 3 (DAV). Sacatepéquez: Mpio. Alotenango, astillero municipal, 14.4619, -90.8147, 1332 m, 20 Apr 2011, M. Véliz 23680 (BIGU). Sololá: Patanatic, 6 km to Panajachel [14.7634, -91.1354], 1700 m, 20 Sep 1971, A. Molina-R. 26664 (MEXU). Suchitepéquez: Volcán Santa Clara, 1.5, 2 miles W of Finca El Naranjo [14.6077, -91.3364], 1250 m, 1 Jun 1942, J.A. Steyermark 46787 (NY). Mexico. Chiapas: Ejido Sierra Morena, 16.1522, -93.5902, 1550 m, 30 May 2002, A. Reyes-García 4788 (MEXU). Guerrero: 10 km al suroeste de Puerto del Gallo, sobre el camino a Atoyac, 17.4717, -100.1969 2100 m, 13 Mar 2007, Y. Ramírez-Amezcua 964 (DAV, IEB, NY). Jalisco: Mpio. Tecalitlán, 46 km Carr. Cd. Guzmán-Pihuamo, por brecha Llanitos-Canutillo, a 24 km, [19.4692, -103.3064], 1750 m, 14 May 1988, V. Pichardo A. 42 (NY). México: El Potrero, Cañada de agua fría, 3 km al sur de Tlatlaya, 18.6106, -100.2139, 1650 m, 6 Aug 2004, I. Martínez de la Cruz 212 (MEXU). Michoacán: Mpio. Coalcomán, 4.5 km (en línea recta) al oeste de Las Joyas sobre una brecha maderera, 18.5014, -103.0939, 1970 m, 29 Aug 2008, V.W. Steinmann 6347 (DAV). Morelos: Barranca Tepecapa, 18.9683, -99.0156, 1849 m, 17 Jul 2010, R. Hernández-Cardenás 445.2 (IEB). Oaxaca: Dto. Tlaxiaco. Santiago Yosondua. Paraje El Limón, a 100 m del Río Yutama, 16.8, -97.5833, 1470 m, 17 Jul 2013, D. Sandoval-Gutiérrez 967 (MEXU).

Figures

Figure 14. Image of herbarium specimen of L. arrazolensis, Thomas 3721 (NY). Specimen used with permission from the William and Lynda Steere Herbarium, New York Botanical Garden.

Figure 15. Map of geographic distribution of L. arrazolensis based on herbarium specimen data.