Lycianthes heteroclita (Sendtn.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 494. 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

21 Lycianthes heteroclita (Sendtn.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 494. 1919 Fig. 48

Solanum heteroclitum Sendtn. Flora 29 (13): 193 [as 177]. 1846. Type: Guatemala. No exact location, Friedrichsthal s.n. (lectotype designated here: W [acc. # 0003646]).

Brachistus escuintlensis J.M.Coult., Bot. Gaz. 16: 144. 1891. Type: Guatemala. Escuintla: Escuintla, 1100 ft, Mar 1890, J. Donnell Smith 2267 (lectotype designated by D’Arcy 1973b, pg. 116: US [00624003, acc. # 1335155]; isolectotypes: F [F0072758F, acc. # 267053], G [G00379121], GH [00076928], K [K000585751], M [M-0171852]).

Bassovia purpusii Brandeg. Univ. Calif. Publ. Bot. 6: 372. 1917. Type: Mexico. Veracruz: Zacuapan [ca. 19°13'N, 96°53'W, 1025 m], Jul 1915, C. Purpus 7502 (holotype: UC [178570]).

Solanum mitratum Greenm., Bot. Gaz. 37: 211. 1904. Type: Costa Rica. Cartago: Atirro, 600 m, Mar [April on US specimen] 1896, J. Donnell Smith 6673 (lectotype designated by D’Arcy 1973a, pg. 636: GH [00077517]; isolectotype US [00624007]).

Lycianthes mitrata (Greenm.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 500. 1919. Type: Based on Solanum mitratum Greenm.

Lycianthes heteroclita (Sendtn.) Bitter ssp. coalescens Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 496. 1919. Type: Guatemala. Alta Verapaz: Cubilqüitz, [Cubilhuitz], [15.6675, -90.4293], 350 m, Aug 1907, H. von Tuerkheim II 813 (lectotype designated by D’Arcy (1973a, pg. 636): US [00027880]; isolectotypes: G, NY, W [acc. # 1908-3590, acc. # 1908-3589]).

Lycianthes heteroclita (Sendtn.) Bitter var. gracilis Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 496. 1919. Type: Panama. Canal Zone: Railroad relocation between Gorgona and Gatun, 10-50 m, Pittier 2281 (lectotype designated by Dean and Reyes 2018a, pg. 42: US [acc. # 676535]; isolectotype US [acc. # 676536]).

Bassovia escuintlensis (J.M.Coult.) Standl., Contrib. U. S. Natl. Herb. 23: 1304. 1924. Type: Based on Brachistus escuintlensis J.M.Coult.

Capsicum escuintlense (J.M.Coult.) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 12: 347 1936. Type: Based on Brachistus escuintlensis J.M.Coult.

Solanum escuintlense (J.M.Coult.) Hunz., Kurtziana 5: 166. 1969. Type: Based on Brachistus escuintlensis J.M.Coult.

Lycianthes escuintlensis (J.M.Coult.) D’Arcy, Phytologia 25: 116. 1973. Type: Based on Brachistus escuintlensis J.M.Coult.

Type.

Based on Solanum heteroclitum Sendtn.

Description.

Herb, shrub, to treelet, sometimes epiphytic, erect, 0.5-5 m tall. Indument of very small, white to tan, uniseriate, multicellular, simple, eglandular, appressed-ascending trichomes <0.1 (0.2) mm long. Stems green when young, glabrous to sparsely pubescent, compressed (hollowed at the nodes) upon drying in a plant press, 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 7-32 × 2.5-15 cm, the smaller ones with blades 3-12 × 1.5-6 cm, the leaf pairs usually similar in shape, the blades ovate to elliptic (rarely obovate), thin-membranaceous, glabrous to very sparsely pubescent, the base cuneate to attenuate, often oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 0.3-5 cm long, the larger leaf blades with 5-8 primary veins on each side of the midvein. Flowers solitary or in groups of 2-12, axillary, erect; peduncles absent; pedicels 8-20 mm and erect in flower, to 25 mm long and erect in fruit, usually glabrous to sparsely pubescent; calyx 2-3 (4) mm long, 3-6 mm in diameter, campanulate, sparsely puberulent, the margin truncate, undulate, the appendages lacking (but ribs sometimes prominent and dark green); fruiting calyx enlarged, widely bowl-shaped to plate-like and slightly reflexed, 1-3.5 mm long, 4.5-10 mm in diameter; corolla 1-1.6 cm long, campanulate to reflexed in orientation, stellate in outline, usually divided 3/4 of the way to all of the way to the base, the lobes usually with scarce interpetalar tissue, white to lilac and glabrous adaxially, white to green abaxially, puberulent abaxially; stamens equal, straight, the filaments 0.5-1.5 mm long, glabrous, the anthers 4-7 mm long, lanceolate, usually partially connivent or connate to the adjacent anther (at least near the middle or base of the anther, not at the tips), white to yellow, glabrous, poricidal at the tips, the pores round, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 8-11 mm long, linear, straight or curved at tip, glabrous, the stigma oblong. Fruit a berry, 6-13 mm long, 8-13 (15) mm in diameter, globose to depressed globose, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 50-300 per fruit, 1-2.5 × 1-1.5 mm, flattened, circular, ovate, depressed ovate or somewhat triangular in outline, yellow-orange to orange-brown, often lighter in color near the margin, the surface reticulum with minute, serpentine pattern and shallow luminae.

Chromosome number.

Unknown.

Distribution and habitat.

Mexico (Chiapas, Guerrero, Jalisco, Oaxaca, Tabasco, Veracruz), Guatemala (Alta Verapaz, Escuintla, Huehuetenango, Izabal, Petén, Quetzaltenango, Quiché, Santa Rosa, Zacapan), Belize, El Salvador, Honduras, Nicaragua, Costa Rica, and Panama, in tropical rainforest, tropical moist forest, tropical dry forest, cloud forest, rarely in shrublands, sometimes along streams or on limestone, common in secondary forest, agricultural areas (such as coffee plantations), and along roadsides, 0-1000 (2000) m in elevation, perhaps cultivated in some areas (Fig. 49 ).

Common names and uses.

Mexico. Chiapas: fruits and leaves eaten after cooking in water, yerbabuena (Spanish), ashinte (Tzeltal) (L. Bohs 3962); ch’ayok (Maya Lacandon) (M. González-Espinosa 755); tumat tez (Tzeltal) (A. Méndez Ton 5132); ajchkintez (Tzeltal) (Alush Méndez Ton 6741); leaves eaten (S. Levy T. 43); quilete (Matuda 17477); zajchkintez (Tzeltal) (A. Méndez T. 5215); xote nuk (Mayan) (B. Paniagua 57); the leaves are eaten boiled, used medicinally for bones ( utz’ak abaker), ubojo ch’ayok’ (B. Paniagua 264). Guerrero: eaten boiled as an edible green (Mixtec) (K. Velazco-G. 40341). Veracruz: hierba mora, hierba mora cimarrona (W. Marquez R. 256, 275).

Phenology.

Flowering specimens and specimens with mature fruits have been collected throughout the year in most locations. Corollas usually open in very early morning, closing in late morning.

Preliminary conservation status.

Lycianthes heteroclita is a widespread species ranging from Mexico to Panama, represented by many collections. The EOO is 1,336,839.05 km2, and the AOO is 1,328 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion.

Lycianthes heteroclita is one of the most common and wide-ranging Lycianthes species in Central America. Its lifeform ranges from a large herb growing epiphytically or on the ground to a treelet, becoming woody at the base. When pressed, its herbaceous stems compress at the nodes. The variability in the flowers of this species deserves more study. From Mexico to Nicaragua, the corolla is usually white on the adaxial side and deeply stellate with very little interpetalar tissue. However, populations with corollas with more interpetalar tissue at the base of the lobes have been observed and collected in Mexico (Chiapas), Guatemala, El Salvador, and Nicaragua; this form of the corolla may be stellate only half-way to the base. In Costa Rica and Panama, the corollas are usually deeply stellate and purple on the adaxial side; this form was first described as Solanum mitratum (equal to L. mitrata ) and later by Georg Bitter as L. heteroclita var. gracilis (both listed as synonyms above). Further study may show that these flower variants are geographically distinct and deserve recognition at the species level.

In Mexico, Lycianthes heteroclita is sometimes confused with the Mexican endemic L. geminiflora . In general, the two species can be separated by flowering calyx size, with that of L. heteroclita usually equal to or greater than 2 mm in length and that of L. geminiflora usually up to 1.5 mm in length. Also, L. heteroclita is usually found at lower elevations than L. geminiflora , which is a high-elevation cloud forest species, but the two species do co-occur in Oaxaca and Veracruz. In the areas of overlap, L. heteroclita is usually found below 1000 m, while L. geminiflora is usually found above 800 m. In addition, L. heteroclita specimens have been collected in Guatemala at elevations of 2000 m or more, which is unusual. In Chiapas, Honduras and Costa Rica, there are many specimens collected from elevations between 1200 and 1700 m, as well as elevations below 1000 m.

Confusion about what name to use for this species exists in the literature. In his treatment of the Solanaceae for Flora of Panama, D’Arcy (1973a) used the name Lycianthes esquintlensis for this species, and under that name, he synonymized the names L. heteroclita ssp. coalescens and Solanum mitratum . D’Arcy also included the species L. synanthera in his treatment. But he then annotated many specimens of L. synanthera as L. esquintlensis . In addition, in their treatment of Lycianthes for the Flora of Guatemala, Gentry and Standley (1974) only recognized L. synanthera and synonymized L. heteroclita , L. escuintlensis , and S. mitratum under that name, and their annotations reflected this. This created confusion in the identification of this species for several decades. Nee (1986) in his treatment for The Flora of Veracruz and Bohs (2015) in her recent treatment for the Flora of Costa Rica have corrected these errors.

Where they co-occur, Lycianthes heteroclita may be confused with L. synanthera and L. nitida Bitter, because all three species have calyces lacking appendages and may be epiphytic. Both L. synanthera and L. nitida have woody upper stems that do not collapse at the nodes upon drying. In addition, L. synanthera usually has hairs in the axils of the primary veins on the abaxial leaf surface, while L. nitida has coriaceous leaves with geminate pairs in which the minor leaf is much smaller in size and more rounded than the larger leaf.

The name Solanum heteroclitum Sendtn. is lectotypified here. In the protologue, Sendtner (1846) cites only one collection: Guatemala. No exact location, Friedrichsthal s.n., but does not specify a particular specimen or herbarium. We were only able to locate one specimen seen by Sendtner at W [acc. # 0003646], and we are choosing that specimen as the lectotype.

Representative specimens examined.

Guatemala. Alta Verapaz: city of Cobán, garden of residence along 1st St belonging to the family of Fredy Archila, seed of this plant originally collected at Finca Siguanha in forest at 1400 m in elevation, 15.4749, -90.3722, 1335 m, 9 Aug 2017, E. Dean 9500 (DAV). Escuintla: Palín Finca Comunal, El Chilar, 14.3536, -90.7283, 959 m, 10 Nov 2010, M. Véliz 22280 (BIGU). Huehuetenango: Between Ixcan and Finca San Rafael, Sierra de los Cuchumatanes 200-800 m, 24 Jul 1942, J.A. Steyermark 49399 (NY). Izabal: W of Santo Tomás de Castilla, near Guatel antennas on one of the summits of Cerro San Gil, 15.6703, -88.6997, 800-900 m, 21 Sep 1997, Nee 47320 (MO, NY). Petén: La Cumbre. Pusila road, 5 km, 17 Aug 1976, C.L. Lundell 20194 (MO, LL). Quetzaltenango: Colomba, Fca. San Francisco Pie de la Cuesta, 14.7281, -91.7178, 1113 m, 15 Feb 2011, L. Velásquez 1728 (BIGU). Quiché: Nebaj, about 12 km west, [15.4058, -91.1461], 8000 ft, 4 Jul 1964, E. Contreras 5196 (MEXU, NY, LL, TEX). Santa Rosa: Region of Platanares, between Taxisco and Guazacapán, 220 m, 3 Dec 1940, P.C. Standley 79062 (MO). Zacapan: Gualan, 122 m, 28 Dec 1905, W.A. Kellerman 5678 (LL). Mexico. Chiapas: road from Ocosingo to Palenque in pueblo de Bahtaj, about 26 km NE of Ocosingo, roadside, 17.1447, -92.1283, 858 m, 5 Dec 2012, L. Bohs 3962 (DAV, MEXU). Guerrero: Yoloxóchitl, 4.37 km en línea recta al norte de la comunidad, en el terreno del Sr. Enrique Rómulo (tierras de uso común), sobre el arroyo que va al paraje Salto de la Mona, 16.8531, -98.6723, 545 m, 22 Apr 2017, K. Velazco-G. 40341 (DAV). Jalisco: Mpio. Casimiro Castillo, parte poniente del puente "La Calera" por la carretera Guadalajara-Barra de Navidad, [19.6730, -104.4287], 550 m, 17 Aug 1990, R. López -V. 213 (WIS). Oaxaca: Sierra de Juárez, Mpio. Comaltepec, along Hwy 175 between km 66 and 67 just south of the town of Metates, 17.6860, -96.3289, 870 m, 11 Sep 2017, E. Dean 9524 (DAV). Tabasco: Sierra El Madrigal, al E del edificio principal del centro regional Tropical Puyacatengo, [17.5172, -92.9028], 600 m, 6 Jun 1991, A.M.H. Alipi 440 (MEXU). Veracruz: Cerro del Marinero, poblado Adolfo López Mateos, 18 km este de Catemaco, 18.4444, -94.9633, 500 m, 8 Jun 1991, M. Torres 495 (MEXU).

Figures

Figure 48. Image of herbarium specimen of L. heteroclita, Cate s. n. (NY). Specimen used with permission from the William and Lynda Steere Herbarium, New York Botanical Garden.

Figure 49. Map of geographic distribution of L. heteroclita based on herbarium specimen data.