Based on Parascopolia acapulcensis Baill.

Figure 6. 

Image of herbarium specimen of L. acapulcensis, Dean 209 (DAV). Image used with permission from the UC Davis Center for Plant Diversity.

Perennial herb from moniliform storage roots, decumbent to erect, 0.1–0.5 (1) m tall, dying back each season. Indument of white, uniseriate, multicellular, simple or dendritically branched, eglandular, spreading to appressed-retrorse trichomes 0.1–1 (2) mm long. Stems green to green-purple, sparsely to moderately pubescent, much compressed upon drying in a plant press, woody with age, especially near the base; first stem (1.5) 5–30 (70) cm long to first inflorescence, the internodes 2–10 (14); first two sympodial branching points dichasial, followed by monochasial branching, this sometimes very extensive (in some Costa Rican and Nicaraguan populations the stems spreading along the ground and rooting at the nodes). Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 3–18 × 1–8 cm, the smaller ones with blades 1/4 to 3/4 the size of the larger, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, chartaceous to thick chartaceous, glabrous to moderately pubescent, the primary veins 4–7 on each side of midvein, the base cuneate (rarely truncate), short attenuate or decurrent onto the petiole, slightly oblique on smaller leaves, the margin entire, usually irregularly undulate, the apex acute to short-acuminate (rarely long-acuminate), the petioles 0.5–1.5 (2.5) cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels (10) 30–60 (85) mm and erect in flower, 20–70 (90) mm long and deflexed in fruit, sparsely to moderately pubescent; calyx (2) 2.5–5.5 (6.25) mm long, 3.5–5 (6) mm in diameter, obconic, campanulate, or urceolate, glabrous to moderately pubescent, the margin truncate, with (5) 10 linear, spreading to reflexed appendages 1–6.5 (9) mm long emerging 0.5–1 mm below the calyx rim; fruiting calyx enlarged, (1.5) 2–4 (6) mm long, 5–12.5 (14) mm in diameter, the appendages to 10 mm long, usually reflexed (sometimes appressed to fruit), often broken; corolla 1.1–2.7 cm long (2–5 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white, sometimes with darker maroon to purple stripes along the major veins adaxially, green near the major veins abaxially, glabrous; stamens unequal, straight, the filaments of three lengths, the two shortest filaments 1–2.5 (3.5) mm long, the two medium filaments 1–3.5 (4.5) mm long, the one long filament 4–9 mm long, the length of the long filament nearly always 2–4 times that of medium filaments, glabrous, the anthers 4.5–7.5 mm long, lanceolate to oblong (rarely ovate), free of one another, yellow, glabrous, poricidal at the tips, the pores linear to ovate, dehiscing distally or away from the style, not opening into longitudinal slits; pollen grains dicolporate; pistil with glabrous ovary, the style 9–14 mm, linear, straight to slightly curved, glabrous, the stigma usually strongly bilobed (sometimes weakly bilobed or capitate). Fruit a berry, remaining attached to calyx at maturity, pendent, (10) 15–50 (70) mm long, (4.5) 9–20 mm in diameter, short-ovoid to elongate fusiform, the tip apiculate to long-attenuate, the exocarp glossy blue-black, grey-blue, bright blue, or dull purple, glabrous, the mesocarp ranging from dark purple and juicy to light purple and powdery, lacking sclerotic granules, the placental area light purple and powdery. Seeds (11) 20–80 (90) per fruit, 2.5–3.5 × 3–4.2 mm, not compressed, irregularly depressed obovate to depressed rhombic, ridged and blistered along one side, black, the surface reticulum with a rough, loose serpentine pattern with deep luminae.

2n = 24 from Dean 313, 314, 329 (Dean 2004).

Mexico (Chiapas, Colima, Guerrero, Jalisco, México, Michoacán, Morelos, Oaxaca), Guatemala (Huehuetenango, Retalhuleu, Suchitepéquez), El Salvador, Nicaragua, and Costa Rica in clearings and disturbed areas in oak or coniferous forest, shrublands, tropical moist forest, and tropical dry forest, generally on volcanic soils (rarely on limestone, granite, or shale) at 450–2600 m in elevation (Fig. 7).

Figure 7. 

Map of geographic distribution of L. acapulcensis based on herbarium specimen data.

Mexico. Fruit edible; maravilla, huevo de cuervo, chimpin, tsibu (Dean 2004).

Flowering specimens have been collected from May to September; specimens with mature fruits have been collected between September and November. In the field, the first author has observed that the corollas open in the very early morning and close by late morning. The pollen in this species has a lemony fragrance.

Lycianthes acapulcensis is a widespread species in Mexico and Central America, represented by 116 collections and occurring in seven protected areas. This species was given a preliminary conservation assessment by Anguiano-Constante et al. (2018) of Least Concern (LC).

Lycianthes acapulcensis is a very variable species, and it may be that some of the local forms deserve varietal status. It is variable in habit, indument (both trichome type and density), leaf shape, presence or absence of purple stripes on the corolla, fruit shape, and fruit coloration. However, the variation extremes are connected by intermediate populations (Dean 2004).

Lycianthes acapulcensis may be confused with L. ciliolata Bitter and L. rzedowskii. It is separated from those species by its combination of white corollas that may or may not have maroon to purple stripes and its pattern of filament lengths (the longest filament nearly always more than twice as long as the adjacent filaments). The anthers have a lemony fragrance, which is unlike that of any other anther (pollen) fragrance in similar Mexican and Guatemalan species of Lycianthes. The root shape (moniliform rather than fusiform segments) is helpful if underground parts are available for examination. On dried specimens, the length of the pedicels of the youngest mature flowers relative to their subtending leaves is often a useful character for separating L. acapulcensis from L. ciliolata. In the former, the length of those pedicels is usually less than that of the subtending leaves, while in L. ciliolata the length of the pedicels generally exceeds that of the leaves. Lycianthes acapulcensis appears to hybridize with L. moziniana and L. rzedowskii where the species co-occur (Dean 2004).

When Baillon (1888) published the name Parascopolia acapulcensis, he did not specify a specimen or herbarium. Similarly, when D’Arcy (1986b) transferred this species to Lycianthes, he did not cite a type specimen. There is only one specimen of this species seen by Baillon, and it is at P [P00070403]. Therefore, we are here designating specimen P00070403 as the lectotype of P. acapulcensis.

Guatemala. Huehuetenango: Mpio. Jacaltenango, 15.6744, -91.7353, 1627 m, 11 Jul 2006, M. Véliz 17055 (BIGU). Retalhuleu: S. Sebastian, [14.55, -91.65], Sep 1874, C. Bernoulli 2404 (GOET). Suchitepéquez: Patutlul, Finca Los Tarrales, [14.5364, -91.17], 300 m, 30 Jul 2004, S. Montiel s.n. (BIGU). Mexico. Chiapas: El Ranchito, sobre la carretera de los miradores, Parque Nacional Cañon del Sumidero, 16.8192, -93.0736, 1301 m, 24 Aug 2007, J.A. Espinosa-Jiménez 306 (MO). Colima: Rancho El Jabalí, 22 air km NNW of Colima in the SW foothills of the Volcán de Colima, on border of Colima and Jalisco, [19.45, -103.7], 1300 m, 15 Jul 1991, L. Vázquez-Villagran 887 (MEXU, DAV). Guerrero: Arroyo Cumiapa, a 1.44 km en línea recta al noroeste de la Comisaría de Arroyo Cumpiapa, sobre el camino que va a Cerro Zapote, en el terreno del Sr. Lauro Cortez, 16.8826, -98.6266, 531 m, 2 Aug 2017, K. Velazco-G 40590 (DAV). Jalisco: Sierra del Halo, ca. 2 rd mi along rd to San Isidro (or Jilotlan) that leaves old Colima-Tecalitlán rd c. 7 rd mi S of Tecalitlán, [19.3171, -103.2696], 1340 m, 23 Nov 1991, E. Dean 329 (DAV, MEXU). México: Cruz de los Pozitos, 18.9025, -99.7428, 2340 m, 20 Aug 2011, F. D. Dorantes-Hernández 408 (MEXU). Michoacán: 4 km al sur de Doctor Miguel Silva, sobre la carretera a la Huacana, [19.1368, -101.7215], 500 m, 22 Jul 2001, J. Rzedowski 53805 (MEXU). Morelos: noroeste de La Barranca de Atzingo, [18.9455, -99.2754], 1800 m, 12 Aug 1987, E. Estrada-Loera 1708 (MEXU). Oaxaca: San José del Chilar, 17.7007, -96.9321, 683 m, 10 Nov 2009, O. Vargas-Ponce 2084 (IBUG).

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

Figure 8. 

Image of herbarium specimen of L. amatitlanensis, Breedlove 55371 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Perennial herb to shrub, 0.4–4 m tall. Indument of off-white, tan or purplish (reddish), uniseriate, multicellular, simple, acute, straight to curved, eglandular, spreading or ascending trichomes 0.5–3 mm long. Stems green when young, moderately to densely pubescent, somewhat compressed upon drying in a plant press, light brown and woody with age; upper sympodial branching points usually monochasial with a few dichasial branching points. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 6.5–26 × 1.5–8.5 cm, ovate, elliptic, or obovate, the smaller ones with blades 0.3–4 (7) × 0.2–2 (3.5) cm, usually ovate, the leaf pairs similar in texture, chartaceous, sparsely to densely pubescent, the trichomes along the midvein of the abaxial side spreading (at a 90 degree angle) to ascending (at a 45 degree angle), the base cuneate (sometimes rounded in the smaller leaves), usually oblique, the margin entire, usually delicately undulate, the apex acute to acuminate, the petiole 0.1–1.5 cm long, sometimes absent, the large leaf blades with (7) 10–22 primary veins on each side of the midvein. Flowers solitary, axillary, oriented horizontally to nodding; peduncles usually absent, sometimes present as a 1–3 mm long peg with overlapping pedicel scars; pedicels 4–12 mm and arching in flower, 6–16 mm long and erect to arching in fruit, moderately to densely pubescent; calyx 1–2 mm long, 2–3 mm in diameter, obconic to narrowly campanulate, moderately pubescent, the margin truncate to undulate, with 5–10 narrow, linear, spreading to reflexed appendages 0.8–4 mm long emerging 0.25–0.5 mm below the calyx rim; fruiting calyx slightly enlarged, widely bowl-shaped to plate-shaped, 1–2.5 mm long, 3.5–6 mm in diameter, the appendages very narrow and weak, to 5 mm long, sometimes withering in age; corolla 0.5–0.8 cm long, campanulate to reflexed in orientation, stellate in outline, divided 1/2 to 2/3 of the way to the base, interpetalar tissue present, white to pale yellow, adaxial markings unknown, moderately pubescent on abaxial surface with tuft of trichomes on distal end of lobe; stamens equal, straight, the filaments 1–2 mm long, glabrous, the anthers 2.5–3 mm long, lanceolate, free of one another, yellowish, glabrous, attenuate and abruptly narrowed at the tip, the narrowed portion ca. 0.5 mm long, poricidal at the tip, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 4–6 mm long, linear, straight, glabrous, widened distally into the stigma, the stigma capitate, decurrent down two sides. Fruit a berry, 5–8 mm long, 5–8 mm in diameter, globose, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 25–80 per fruit, 0.9–1.1 × 0.75–1 mm, compressed but not flat, sometimes with one shallow ridge, semi-circular, depressed ovate, triangular, or rhombic in outline, orange, the surface reticulum with tight, shallow serpentine pattern with shallow luminae.

Unknown.

Southern Mexico (Chiapas, Tabasco, Veracruz), Guatemala (Alta Verapaz, Escuintla, Guatemala, Huehuetenango, Petén), south to Belize, Honduras, Nicaragua, Costa Rica, Panama, and possibly South America. Tall forest, tropical rain forest, tropical moist forest, wet premontane forest, and cloud forest, in shady canyons, slopes, drainages (often near rivers or streams), sometimes in disturbed areas or coffee plantations, sometimes on limestone, usually 100–1000 m in elevation, rarely up to 1800 m (Fig. 9).

Figure 9. 

Map of geographic distribution of L. amatitlanensis from Mexico to Honduras based on herbarium specimen data.

Guatemala: Alta Verapaz: kaki saki maï (I. Kunkel 186, 398); same location: maï (I. Kunkel 211).

Flowering specimens have been collected March through November; fruiting specimens have been collected May through February. De Nevers (1986) documented the pollination of L. amatitlanensis in eastern Panama. He described the flowers as being pendulous, positioned below the leaves. Pollinators (halictid bees) were observed visiting the flowers early morning to late afternoon and at 1 a.m. at night. No floral scents, nectar, or diurnal movements were noted. The flowers on most herbarium specimens are open, indicating that if there are diurnal movements, the flowers close for a very short time or only at night.

Lycianthes amatitlanensis is a widespread species ranging from eastern Mexico to Panama, represented by 48 collections and occurring in nine protected areas. The EOO is 687,839.069 km², and the AOO is 192 km². Following the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Lycianthes amatitlanensis is a wide ranging species with small white to pale yellow flowers (pubescent on the abaxial side of the corolla lobes and with tufted trichomes at the lobe tips) and long, coarse trichomes that spread away from the midvein on the abaxial side of the leaf (usually with some trichomes at an angle close to ninety degrees). Lycianthes amatitlanensis is morphologically similar and perhaps closely related to three other species occurring in Mexico and/or Central America: L. glabripetala (endemic to Mexico); L. inconspicua (Central America); and L. inaequilatera (Rusby) Bitter (Costa Rica, Panama and South America). Lycianthes inconspicua differs from L. amatitlanensis in having longer pedicels (15–30 mm in flower and 30–35 mm in fruit), appressed trichomes along the midvein of the abaxial side of the leaf, and ovate anthers with a shorter attenuate portion at the tip (ca. 0.25 mm long). Lycianthes inaequilatera has pedicels of similar length to those of L. amatitlanensis, but it has short, soft, appressed trichomes along the midvein on the abaxial side of the leaf. The Mexican species Lycianthes glabripetala has larger, nearly glabrous corollas and appressed, wavy trichomes along the midvein on the abaxial side of the leaf blade; L. glabripetala does not overlap in distribution with L. amatitlanensis. Where the distribution of L. amatitlanensis overlaps with L. inconspicua and L. inaequilatera, L. amatitlanensis tends to occur at lower elevations than the other two species. Intermediates between L. inaequilatera and L. amatitlanensis occur in Costa Rica and Panama, and these two species may eventually be treated as a single entity. Lycianthes inaequilatera is a South American species, originally described from Bolivia that has not been reported further north than Costa Rica. Lycianthes amatitlanensis is a Mexican and Central American species, originally described from Guatemala but reported in South America. Further study is needed to understand the ranges and variation of the two species. If united, L. inaequilatera is the earlier and correct name.

Guatemala. Alta Verapaz: Cubilquitz [Cubilhuitz], [15.6675, -90.4293], 350 m, July 1907, H. von Tuerkheim 153 (US). Escuintla: Río Guacalate, 600 m, 16 Dec 1938, P.C. Standley 60200 (US). Guatemala: Barranca de Eminencia, 1400 ft, Feb 1892, J. Donnell Smith 1457 (US). Huehuetenango: between Ixcan and Finca San Rafael, Sierra des los Cuchumatanes, 200–800 m, 24 Jul 1942, J.A. Steyermark 49396, (NY). Izabal: Cerro San Gil, 15.6333, -88.8167, 803 m, 8 Feb 2012, M. Véliz 23523 (BIGU). Petén: La Cumbre, in zapotal, on parcela de José León, 3 km east, 14 Aug 1976, C.L. Lundell 20140 (LL). Mexico. Chiapas: Ejido Tres Picos, 16.2272, -93.5808, 1780 m, 19 Apr 2002, A. Reyes-García 4437 (MEXU). Tabasco: vicinity of Teapa, along road between Teapan and Tacotalpa, 3.1 m. E of Teapa along stream and limestone cliffs ca 1/4 mi S of Hwy, 17.55, -92.9833, 150 m, 19 Feb 1987, T.B. Croat 65349 (MO). Veracruz: Mpio. Jesús Carranza, lomas al S de Pob. 2 (ca. 3 km al S de entronque de terracería La Laguna-Sarabia con camino al N a Pob. 2), 17.2, -94.65, 150 m, 8 Jul 1988, T. Wendt 6064 (MO).

Mexico. Veracruz: Río Blanco, Orizaba, Bourgeau 2536 (lectotype designated by Dean and Reyes 2018a, pg. 40: BR [000000552905]; isolectotypes: G [G00415142], GH [00077065], K [K000063121], MPU [MPU310734], P [P00385091, P00385092], S [cited by Bitter (1919), but not seen]).

Figure 10. 

Image of herbarium specimen of L. anomala, Diggs 2731 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Herb, shrub, to tree, sometimes epiphytic, semi-epiphytic, or vine-like, erect, 2–10 (15) m tall. Indument of tan to brownish, uniseriate, multicellular, simple or dendritically branched, eglandular, spreading trichomes 0.25–0.5 mm long. Stems green when young, glabrous to sparsely pubescent, not compressed upon drying in a plant press, quickly becoming woody (glossy pale grey with longitudinal wrinkles upon drying); upper sympodial branching points mostly monochasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 10–25 × 5–15 cm, ovate, oblong, or elliptic (rarely obovate), the smaller ones with blades 2–10 × 1.5–5 cm, orbicular to ovate, the leaf pairs similar in texture, coriaceous, usually glabrous adaxially, abaxially with tufts of trichomes in the axils of the major veins, the base rounded to cuneate, usually oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 1–4 cm long, the larger leaf blades with 6–9 primary veins on each side of the midvein. Flowers solitary or in groups of 2–6, axillary, erect; peduncles absent or present as a short stub with many pedicel scars, 5–10 mm long; pedicels 8–20 mm and erect in flower, to 35 mm long and erect in fruit, glabrous; calyx 4–5 mm long, 6–8 mm in diameter, widely campanulate, glabrous, coriaceous in texture, the margin truncate, very well developed, with 5–10 reflexed appendages, 0.25–1 mm long (sometimes just a bulge), connate at their bases, emerging 1–2 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 3–4 mm long, 8–10 mm in diameter, the appendages 0.5–2 mm long, reflexed as a connate unit; corolla 1–1.5 cm long, reflexed in orientation, stellate in outline, divided 1/3–1/2 of the way to the base, with scant interpetalar tissue, the lobes purple adaxially, nearly glabrous; stamens equal, straight, the filaments ca. 1 mm long, glabrous, the anthers 4–6 mm long, elliptic, connate at edges to adjacent anther, forming a cone, yellow, glabrous, poricidal at the tips, the pores dehiscing distally and opening into longitudinal slits that extend ca. 1/3 of the way from apex to base, the slit forming between the thecae of adjacent anthers; pistil with glabrous ovary, the style ca. 8 mm long, linear, straight to curved, glabrous, the stigma truncate to capitate. Fruit a berry, 7–10 mm long, 7–10 mm in diameter, globose to depressed globose, green to white when immature, purple at maturity, glabrous, lacking sclerotic granules. Seeds ca. 100 per fruit, 1–1.5 × 1–1.25 mm, flattened, slightly curved, triangular to depressed-ovate in outline, yellow, the surface reticulum with tight, minute serpentine pattern with shallow luminae.

Unknown.

Mexico (Oaxaca, Veracruz), in tropical moist forest and cloud forest, sometimes in sandy soil or on limestone, often in disturbed areas, such as secondary forest or coffee plantations, 450–1300 m in elevation (Fig. 11).

Figure 11. 

Map of geographic distribution of L. anomala based on herbarium specimen data.

None known.

Flowering specimens have been collected June through November. Specimens with immature fruits have been collected June through March. Specimens with mature fruits have been collected June through February. Possibly flowering and fruiting throughout the year in some locations. The diurnal movements of the corolla of this species are unknown, but it has been noted that the flowers are sometimes open at midday (Nee 1986).

Lycianthes anomala is a Mexican endemic, represented by 20 collections, none of which are in protected areas. The EOO is 7,552.355 km², and the AOO is 80 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

This species is similar to the Central American species L. synanthera in having pale woody stems with petioles that darken upon drying, tufts of trichomes in the vein axils of the abaxial side of the leaf, and fused anthers that dehisce by slits formed between adjacent thecae. It differs from that species in having connate, reflexed appendages on the calyx (versus no appendages in L. synanthera) and purple fruits (versus yellow/orange fruits in L. synanthera). Reyes-Cornejo (2015) indicated that this species is also found in Central America, and we have studied specimens from Costa Rica and Panama that resemble L. synanthera but have calyx appendages. These specimens need further study.

Mexico. Oaxaca: Santa María Tlalixtac, orillas del Río Cóndor, brecha entre Santa María Tlalixtac y Chiquihuitlán de Benito Juárez, [17.9541, -96.7179], 675 m, 25 Nov 2004, G. Juárez-García 877 (MEXU). Veracruz: Ladera de cerro al E de Coetzala, 18.7806, -96.9144, 650 m, 11 Nov 2001, A. Rincón G. 2811 (IEB, MEXU, XAL).

Mexico. Quintana Roo: Cobá, east of the ruins, in advanced deciduous forest, [20.4800, -87.7300], 1 Jun 1938, C.L. & A.A. Lundell 7800 (holotype: US [00027868]; isotypes: A [00936248], F [0072898F, acc. # 1307280], US [01014254, 01014255]).

Figure 12. 

Image of herbarium specimen of L. armentalis, Cabrera 5160 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Clambering shrub to vine, 0.5–4 (7) m tall. Indument of pale yellow to orange-brown, uniseriate, multicellular, sessile to stalked, multangulate-stellate, eglandular, spreading trichomes 0.1–0.75 mm long, 0.2–0.5 mm in diameter, the rays (3) 5–8 (10) per whorl, straight, rarely rebranched, often with an enlarged sphere where the rays join, rarely some dendritically branched trichomes also present. Stems green to light brown when young, sparsely to densely pubescent (appearing like dense felt), not compressed when dried in a plant press, becoming woody early; upper sympodial branching points a mixture of monochasial and dichasial branching, the branching divaricate (diverging at wide angles). Leaves simple, the leaves of the upper sympodia paired or not, the leaves often appearing like they terminate short shoots with the pairs arranged at 90 degree angles, the pairs unequal in size, the larger ones with blades 3–9 (13) × 2–4.5 (7) cm, the smaller ones (often not developing) with blades 0.9–2.5 × 0.8–2 cm, the leaf pairs similar in shape, the blades ovate, elliptic, obovate, or suborbicular, thick chartaceous, sparsely to moderately pubescent (denser on the abaxial side, especially along the veins, the adaxial side sometimes nearly glabrous), the base cuneate to rounded, sometimes oblique, the margin entire, usually irregularly undulate, the apex rounded, obtuse, acute or acuminate, the petiole 0.3–1.2 cm long, the larger leaf blades with 3–5 primary veins on each side of the midvein. Flowers solitary or in groups of 2–6, axillary, erect; peduncles absent; pedicels 6–13 (20) mm long and erect in flower, 9–25 mm long and erect in fruit, moderately pubescent; calyx 2.5–3 mm long, 3–4 mm in diameter, campanulate, moderately pubescent, the margin truncate, with 10 spreading linear appendages 0.5–3.5 mm long emerging ca. 0.3 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 2–4 mm long, 6–10 mm in diameter, the appendages to 5 mm long; corolla 0.8–1.2 cm long, campanulate to rotate in orientation, stellate in outline (divided ca. 1/4–1/2of the way to the base), with abundant interpetalar tissue, white, with a few scattered trichomes on the adaxial side of the lobes near the major veins, sparsely to moderately puberulent on the lobes near the major veins abaxially; stamens slightly to very unequal, straight, the four short filaments 0.5–1 (1.5) mm long, the one long filament 1–2 (3) mm long, glabrous, the anthers 3–4 mm long, elliptic to lanceolate, free of one another, yellow, sparsely pubescent on the inner face, poricidal at the tips, the pores ovate, terminal, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–8 mm long, linear, straight to curved, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 5–12 mm long, 5–12 mm in diameter, globose, red-orange when mature, glabrous, lacking sclerotic granules. Seeds 20–30 per fruit, 2.5–3 × 2–2.5 mm, flattened, circular to depressed ovate in outline, thickened on the edges, thin and semi-transparent in the center, yellow to dark orange, the surface reticulum in the center nearly smooth, the edges with minute serpentine pattern with shallow luminae.

Unknown.

Mexico (the Yucatán Peninsula, including Campeche, Quintana Roo, Yucatán), Guatemala (El Progreso, Petén), and Belize, in forest (often secondary), usually in tropical rain forest, tropical moist forest, or tropical dry forest, sometimes on limestone, 0–500 m in elevation (Fig. 13).

Figure 13. 

Map of geographic distribution of L. armentalis based on herbarium specimen data.

None known.

Flowering specimens have been collected from May to October; specimens with mature fruits have been collected June to March. The timing of the diurnal corolla movements for this species are not known, but many specimens have been collected with closed flowers indicating that the flowers are open for a limited period during the day, probably in the early morning.

Lycianthes armentalis is a widespread species ranging from southeastern Mexico to Belize represented by 116 collections and occurring in seven protected areas; unfortunately, the habitat of this species is vulnerable. The EOO is 122,903.444 km² (LC) and AOO is 404 km² (EN). Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Lycianthes armentalis is often confused with L. sideroxyloides, L. rafatorresii, and L. scandens var. scandens (previously known as L. lenta (Cav.) Bitter). Lycianthes armentalis differs from those species in its combination of multangulate-stellate (not geminate-stellate) trichomes, widely divaricate branching, calyx appendages that are not enlarged at the tip, white, shallowly stellate corollas, and unequal stamens. Lycianthes sideroxyloides has geminate-stellate trichomes, appendages that are enlarged at the tip, deeply stellate corollas, and equal stamens. Lycianthes scandens var. scandens has purple, mostly entire corollas and lacks widely divaricate branching. Lycianthes rafatorresii has similar flowers and trichomes to L. armentalis but lacks widely divaricate branching and has calyx appendages that are enlarged at the tip. Lycianthes armentalis occurs at relatively low elevations on the Yucatán Peninsula, a distribution that overlaps with L. scandens var. scandens, but not the other two species.

Guatemala. El Progreso: Tulumaje, [14.9256, -90.0469], 346 m, 23 Nov 2003, R. Ávila 71 (BIGU). Petén: Mpio. Melchor de Mencos, sitio arqueológico El Naranjo, 17.1319, -95.2606, 297 m, 18 Jun 2009, L. Velásquez 413 (BIGU). Mexico. Campeche: a 2 km al E de X-Mejía, 19.2347, -89.3592, 150 m, 24 Jun 2005, E. Martínez- Salas 38011 (MEXU). Quintana Roo: camino a Zafarrancho, 0.73 km al N de Zafarrancho, 19.52, -88.8864, 91 m, 22 Aug 2005, E. Martínez-Salas 38092 (MEXU). Yucatán: carretera Noholal-Sudzal-Chico, 19.75, -89.25, 18 Nov 1992, F. May 766 (MEXU).

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

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.

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.

2n = 24 (Gentry and Pearce 1977).

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).

Figure 15. 

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

None known.

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.

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).

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).

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).

Guatemala. Suchitepéquez: Volcán Santa Clara, between Finca El Naranjo and upper slopes, 1250–2650 m, 23 May 1942, J.A. Steyermark 46653 (holotype: F [0072901F, acc.# 1148517]; isotypes: NY [00138703], US [00624009]).

Figure 16. 

Image of herbarium specimen of L. barbatula. Pascual 2155 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Shrub, erect to scandent (sometimes described as a vine), 3–5 m tall. Indument of small white, uniseriate, multicellular, simple, curved, eglandular, appressed-ascending trichomes 0.1–1 mm long. Stems green when young, glabrous to sparsely pubescent, ribbed to angled 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 4–15 × 1.5–6.5 cm, the smaller ones with blades 1–9 × 0.7–4 cm, the leaf pairs usually similar in shape, the blades narrowly ovate to elliptic or obovate, thick chartaceous, glabrous except for tufts of trichomes located in the axils along the midvein of the abaxial side, the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 0.2–2 cm long, sometimes absent, the larger leaf blades with 5–10 primary veins on each side of the midvein. Flowers solitary or in groups of 2–8, axillary, oriented horizontally to nodding; peduncles absent; pedicels 20–30 mm long and arching in flower, to 40 mm long and spreading to deflexed in fruit, glabrous to sparsely pubescent; calyx 1.5–2.5 mm long, 3–4 mm in diameter, obconic to campanulate, sparsely pubescent, the margin truncate, with 10 linear, erect to spreading appendages 0.5–2 mm long, emerging 0.5 mm below the calyx rim; fruiting calyx usually enlarged, widely campanulate to bowl-shaped, sometimes splitting, 3–4 mm long, 5–8 mm in diameter, the appendages 2.5–4 mm long, spreading; corolla 0.7–2 cm long, rotate to campanulate in orientation, entire to slightly stellate in outline, divided 0–1/5 of the way to the base, with abundant interpetalar tissue, adaxially white with lavender ring near stamen insertion, abaxial color unknown, glabrous; stamens equal or nearly so, the filaments 2–2.5 mm long, glabrous, the anthers 2.5–3 mm long, elliptic, free of one another, brownish-yellow, glabrous, poricidal at the tips, the pores round, large, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–7 mm long, linear, straight, glabrous; stigma capitate to oblong, decurrent down two sides. Fruit a berry, 10–12 mm long, 10–12 mm in diameter, globose to ovoid, changing from green to white as it matures, possibly remaining white or changing to blue-grey or purple at maturity, glabrous, lacking sclerotic granules. Seeds ca. 20 per fruit, 3–3.5 × 2–2.5 mm, flattened, triangular to depressed ovate in outline, tan to orange-brown, the surface reticulum with minute serpentine pattern with shallow luminae.

Unknown.

Mexico (Oaxaca, probably also Chiapas) and Guatemala (Chimaltenango, Quezaltenango, Suchitepéquez), in tropical moist forest, cloud forest, and near coffee plantations, 920–1600 m in elevation. [Note: type specimen has a high elevational range of 2650 m, but the specimen may not have been collected that high.] Our knowledge of the distribution and ecology of this species is incomplete due to the paucity of specimens in herbaria (Fig. 17).

Figure 17. 

Map of geographic distribution of L. barbatula based on herbarium specimen data.

None known.

Flowering specimens have been collected in May and June. Specimens with mature fruits have been collected in September and January. Immature fruits have been collected in January. The phenological record is incomplete due to a paucity of specimens. The corollas on the specimens of this species are often open; this indicates that the corollas are open for a substantial amount of time each day.

Lycianthes barbatula is a rarely collected species of Mexico and Guatemala, represented by only 11 collections, only one of which is from a protected area (Cuenca del Lago Atitlán, Guatemala). The EOO is 20,500.215 km², and the AOO is 28 km². Following the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Lycianthes barbatula is morphologically similar to L. manantlanensis and L. orogenes with which it shares relatively thick leaves (thick chartaceous to coriaceous), relatively glabrous foliage, long, delicate, arching pedicels, white corollas, and equal to nearly equal stamens with yellowish anthers, sometimes with a brownish connective. Lycianthes barbatula differs from the other two species in having longer calyx appendages and tufts of trichomes in the vein axils along the midvein of the abaxial side of the leaf blade. The fruit color of L. barbatula has been recorded on several specimen labels as being white, and this is also the color given in Gentry and Standley (1974), but there is uncertainty as to whether this is the final color at maturity or a transitional color, as other specimens mention blue-grey or blue-purple fruits.

Guatemala. Chimaltenango: Volcán Acatenanago, Aldea Quisache, 14.51806, -90.2844, 1500 m, 19 May 2004, M. Véliz 15261 (BIGU, MEXU). Quetzaltenango: above Finca Montevideo, along Barranco Espinazo and tributary of Río Pantaleón, lower and middle southwestern slopes of Volcán Fuego, 1200–1600 m, 20 Sep 1942, J.A. Steyermark 52055 (US). Suchitépequez: Volcán Santa Clara, between Finca El Naranjo and upper slopes. 1250–2650 m, 23 May 1942, J.A. Steyermark 46653 (US). Mexico. Oaxaca: Dto. Pochutla, El Vigía, 16.0125, -96.1108, 1519 m, 24 Jun 2008, J. Pascual 2155 (DAV).

Mexico. Chiapas: Mpio. La Independencia, third ridge along logging road from Las Margaritas to Campo Alegre, [16.4756, -91.8234], 2300 m, 6 May 1973, D. Breedlove 34793 (holotype: CAS [480622]; isotypes: LL [00226970], MO [acc. # 2602916]).

Figure 18. 

Image of isotype of L. breedlovei, Breedlove 34793 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Vine to scandent shrub, 2–3.5 (5) m tall (perhaps taller, if a vine). Indument of orange to pale yellow (yellow-grey), uniseriate, multicellular, stalked, multangulate-stellate, eglandular, spreading trichomes 0.25–1.5 (2) mm long, ca. 0.75 in diameter, the rays 3–5 per whorl, straight, often rebranched, sometimes several times. Stems pale green (drying tan) when young, moderately to densely pubescent, not compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points a mixture of monochasial and dichasial, the branching divaricate, the branches diverging at wide angles. Leaves simple, the leaves of the upper sympodia usually unpaired, if paired, then unequal in size, the larger ones with blades 3.5–10 × 1.5–4.5 cm, the smaller ones (usually not developing) with blades 1–3.5 × 0.5–2 cm, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, chartaceous, sparsely to densely pubescent (denser on the abaxial side, especially along the veins), the base cuneate to rounded, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate, the petiole 0.3–1 cm long, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–5, axillary, oriented horizontally; peduncles absent; pedicels 9–16 mm long and erect in flower, 10–25 mm long and erect in fruit, densely pubescent; calyx 2.5–3.5 mm long, 3.5–4.5 mm in diameter, campanulate, pale green (sometimes nearly translucent) with dark ribs, sparsely to densely pubescent, the margin truncate, with 10 spreading linear appendages 1–3 mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 2–3 mm long, 6–8 mm in diameter, the appendages to 5 mm long; corolla 0.9–1.5 cm long, rotate in orientation, shallowly stellate in outline, divided ca. 1/2 of the way to the base, with abundant interpetalar tissue, white to lilac, adaxially with darker purple stripes on the lobes, sparsely pubescent with few scattered trichomes, abaxially usually densely puberulent on the lobes (best seen in bud); stamens slightly unequal, straight, the four short filaments 0.5–1 mm long, the one long filament 1–2 mm long, glabrous, the anthers 3–4 mm long, elliptic, free of one another, yellow, sometimes pubescent on the inner face along the connective, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–8 mm long, linear, straight to curved, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 4–10 mm long, 5–11 mm in diameter, depressed globose, orange when mature, glabrous, lacking sclerotic granules. Seeds 5–30 per fruit, 3–3.5 × 2–2.5 mm, flattened, thickened on the edges, reniform to depressed ovate in outline, usually with small notch on one side, orange, the surface reticulum with tight serpentine pattern and shallow luminae, the margin thickened and rougher in texture than the center.

Unknown.

Mexico (Chiapas), in cloud forest, often in oak forest, sometimes associated with Pinus, Abies, Magnolia, or Podocarpus, sometimes near disturbed areas, such as milpas, 2000–3000 m in elevation (Fig. 19).

Figure 19. 

Map of geographic distribution of L. breedlovei based on herbarium specimen data.

Mexico. Chiapas: chichol mut (Tzeltal) (C. Santíz R. 854); penko antivo (Tzotzil) (C. Santíz R. 904); tunatzak (Tzeltal) (A. Médez Ton 5054).

Flowering specimens have been collected from April through July; specimens with mature fruits have been collected from August to November. The timing of the diurnal movements of the corolla of this species is not known, but many specimens have been collected with open flowers indicating that the flowers are open for an extended period during the day.

Lycianthes breedlovei is restricted to cloud forest habitat in the state of Chiapas, represented by 18 collections, only one of which is from a protected area. This species was previously given a preliminary assessment by Dean et al. (2019a) of Endangered.

Lycianthes breedlovei is a shrub to vine with zigzag branching due to widely divaricate branching angles, yellow to orange branched pubescence, white flowers with violet markings, and unequal stamens. It is closely related to L. hortulana Standley & L.O.Williams, described from Honduras. The two species are geographically isolated from one another, with no populations of either species known to occur in Guatemala. They have diverged from one another in pedicel length (L. hortulana flowers have pedicels 3–9 mm long vs 9–16 mm long), flower size (L. hortulana has corollas that are 0.6–1 cm long vs 0.9–1.5 cm), corolla pubescence (L. hortulana has very sparse pubescence on the abaxial side of the corolla lobes vs dense), and stamen length (L. hortulana has equal stamens vs usually unequal) (Dean et al. 2019a).

Mexico. Chiapas: Mpio. Tenejapa, along the road to the town of Matzam, ca. 1.5 km from the eastern outskirts of the town of Las Ollas where the road forks, about 2.6 km from the intersection with the San Cristóbal de las Casas-Tenejapa road, just west and upslope of the settlement of Paraje Cruz Tzibaltic, on ridge where there is an intersection with an undeveloped road, 16.7832, -92.5275, 2484 m, 13 Sep 2017, E. Dean 9531 (DAV226596).

Mexico. Veracruz: District Cordoba, Cerro de Chocomán, 12 May 1907, C. Seler & E. Seler 5168 (holotype: B [not seen, cited by Bitter (1919), probably destroyed]; isotype: GH [00936203]).

Figure 20. 

Image of herbarium specimen of L. caeciliae, Dean 10030 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Shrub to small tree, 1.5–3 m tall. Indument of off-white to light brown, uniseriate, multicellular, simple (very rarely furcate), acute, eglandular, appressed to spreading trichomes 0.25–1 mm long, these usually remaining cylindrical and acute upon drying. Stems green to purple-green when young, glabrous to moderately pubescent, partly to fully compressed upon drying in a plant press, brown and woody with age; upper sympodial branching points mostly monochasial with a few dichasial branching points. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 4–16 × 1–5 cm, elliptic to obovate, the smaller ones with blades 1–6 × 0.5–3 cm, ovate, lanceolate, or obovate, the blades of both the large and small leaves chartaceous to thick chartaceous, glabrous to moderately pubescent, denser on the veins, the base cuneate, sometimes oblique, the margin entire, undulate, the apex acute to acuminate, the petiole to 0.8 cm long, sometimes absent, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–3, axillary, oriented horizontally to nodding; peduncles absent; pedicels 15–30 mm long, arching to deflexed in flower, to 42 mm long, arching to deflexed in fruit, glabrous to moderately pubescent; calyx 3–3.5 mm long, 4–4.5 mm in diameter, campanulate, green, sometimes with a purple hue, glabrous to sparsely pubescent, the margin truncate, with 10 spreading, linear-subulate appendages 2–5 mm long emerging 0.5–1 mm below the prominent, undulating calyx rim; fruiting calyx enlarged, widely bowl-shaped, 2–2.5 mm long, 6–8 mm in diameter, the appendages widening but not significantly elongating, to 7 mm long; corolla 0.7–1.6 cm long, campanulate to reflexed in orientation, stellate in outline, divided 1/3–2/3 of the way to the base, with interpetalar tissue, purple adaxially with green at the base of each lobe near the stamen insertion, purple abaxially, sometimes with a single white line down the middle, nearly glabrous; stamens equal, straight, the filaments 0.5–2 mm long glabrous, the anthers 4–4.5 mm long, ovate to lanceolate, free of one another, purple, glabrous, poricidal at the tips, the pores ovate, dehiscing toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 5–7.5 mm long, linear, glabrous; stigma capitate to oblong. Fruit a berry, 7–13 mm long, 6–13 mm in diameter, globose to ovoid, dark purple at maturity, glabrous, lacking sclerotic granules. Seeds 10–50 per fruit, 2.5–3 × 3–4 mm, compressed but not flat, depressed ovate or reniform (with small notch) in outline, brown, the surface reticulum with a tight, serpentine pattern with deep luminae, with fibrils protruding from the cell walls.

Unknown.

Mexico (Veracruz), on the eastern slopes of two volcanos, Cofre de Perote and Citlaltépetl [Orizaba], in cloud forest and oak forest, 1750–2250 m in elevation (Fig. 21).

Figure 21. 

Map of geographic distribution of L. caeciliae based on herbarium specimen data.

None known.

Specimens have been collected with both flowers and mature fruits April to September. In the field, the first author observed that most of the corollas of this species were closed by 1 pm, but some of the corollas still remained open at that time. Corollas opening for the first time (the smallest on the plant) are a deep purple, while older, larger flowers are a pale purple. The green ring at the base of the corolla is more prominent in older flowers.

Lycianthes caeciliae is a Mexican species of vulnerable cloud forest habitat in the state of Veracruz that is known from only four locations, with only one in a protected area (La Cortadura Ecological Reserve). The EOO is 149.767 km², and the AOO is 12 km². Following the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

As detailed elsewhere (Dean et al. 2019b), Lycianthes caeciliae is a species that was poorly collected until the 21^st century. Other than the type collection, it was just collected one other time in the 20^th century by Matuda. It was then collected numerous times at La Cortadura on the slopes of Cofre de Perote by Gonzalo Castillo-Campos between 2005 and 2007. The species is most abundant in original oak cloud forest where it is in flower and fruit; although observed by the first author in secondary forest of Alnus or Liquidambar at lower elevations in the area of La Cortadura, the plants are small and often sterile. Lycianthes caeciliae is closely related to L. pilifera, a Oaxacan cloud forest endemic, with which it shares simple, straight, spreading trichomes that do not collapse upon drying, corollas with purple and green coloration, purple anthers, purple coloration in the calyx, and dark purple fruit with large brown seeds. Lycianthes caeciliae could be confused with L. stephanocalyx, a rhizomatous herb to subshrub which also occurs in Veracruz, but at lower elevations. Lycianthes stephanocalyx differs from L. caeciliae in having small incurved trichomes, yellow connivent anthers, red fruits, and tan seeds (Dean et al. 2019b).

Mexico. Veracruz: Mpio. Coatepec, La Cortadura, falda este del Cofre de Perote, 19.4900, -97.0427, 1800 m, 8 May 2019, E. Dean 10030 (DAV230605).

Based on Brachistus ceratocalycius Donn.Sm.

Figure 22. 

Image of herbarium specimen of L. ceratocalycia, Dean 9510 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Herb to shrub, erect, 0.5–2 m tall, sometimes epiphytic. Indument of off-white to tan, uniseriate, multicellular, simple, eglandular, appressed-ascending trichomes 0.1–0.2 mm long. Stems green when young, the surface with brownish scurfy horizontal lines (perpendicular to stem axis), glabrous to moderately pubescent, compressed 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 4–14 × 2–4 cm, the smaller ones with blades 1.5–6 × 1–2.5 cm, the leaf pairs usually similar in shape, the blades ovate to elliptic (sometimes narrowly), membranaceous to chartaceous, sometimes purple abaxially, glabrous to sparsely pubescent, the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 0.2–0.7 cm long, sometimes absent, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers in groups of 2–5 (15), axillary, erect; peduncles absent; pedicels 7–17 mm and erect in flower, to 25 mm long and erect in fruit, glabrous to moderately pubescent; calyx 2.5–4 mm long, 4–5 mm in diameter, campanulate, puberulent with very small trichomes, the margin truncate, undulate, very well developed, with 10 very short, reflexed appendages 0.25–1 mm long emerging 1–1.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 2–2.5 mm long, 6–8 mm in diameter, the appendages not changing in length; corolla 0.5–1.2 cm long, campanulate to reflexed in orientation, stellate in outline, divided at least 3/4 of the way to the base, with scarce interpetalar tissue, lilac to purple adaxially and abaxially, with deeper purple markings adaxially near the stamens, pubescent abaxially with very short trichomes; stamens equal, straight, the filaments ca. 1 mm long, glabrous, the anthers 3–4.5 mm long, lanceolate, free of one another, pale yellow (sometimes with brown shiny pigment on the outer side), glabrous, poricidal at the tips, the pores round, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–7 mm long, linear, straight to curved at the tip, glabrous; stigma capitate, decurrent down two sides. Fruit a berry, 5–9 mm long, 5–9 mm in diameter, globose to ovoid, red at maturity, glabrous, lacking sclerotic granules. Seeds 10–60 per fruit, 1.5–2.5 × 1.5–2 mm, flattened with slightly raised and thickened edges, depressed ovate to circular in outline, tan to orange, with the surface reticulum with minute serpentine pattern and shallow luminae.

Unknown.

Mexico (Chiapas) and Guatemala (Alta Verapaz, Huehuetenango, and Quiché), in cloud forest, including oak forest, sometimes in disturbed or open areas, often on calcareous soil, 1300–2100 m in elevation (Fig. 23).

Figure 23. 

Map of geographic distribution of L. ceratocalycia based on herbarium specimen data.

None known.

Flowering specimens have been collected April to September. Specimens with mature fruits have been collected in August. In the field in Mexico and Guatemala, the first author observed that the corollas are open in the morning and closed in the afternoon.

Lycianthes ceratocalycia is a species restricted to Guatemala and lands immediately adjacent in Chiapas that is known from only 10 locations, only one of which is from a protected area (Lagunas de Montebello, Mexico). The EOO is 2,169.823 km², and the AOO is 40 km². Following the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Lycianthes ceratocalycia was rarely collected until recently. It can be distinguished from similar species with stellate corollas and equal stamens by the scurfy horizontal lines on the young stems.

Guatemala. Alta Verapaz: San Cristóbal, Finca Pamac II, 15.3986, -90.5883, 2146 m, 16 Aug 2015, A. Borrayo 5 (BIGU). Huehuetenango: northwestern region near the border with Mexico. Mpio. San Mateo Ixtatán, E of the town of Aguacate, along road to town of Yalanhuitz, 16.0377, -91.4662, 1567 m, 15 Aug 2017, E. Dean 9510 (DAV). Quiché: camino a Amachel, 15.6878, -90.9928, 1553 m, 25 Jun 2006, R. Ávila 3038 (MEXU, MEXU). Mexico. Chiapas: Mpio. La Trinitaria, Parque Nacional Lagunas de Montebello, at an outlook (mirador) above and to the E of Dos Lagunas, on the west side of Hwy 307 to the E of Lago Tziscao, ca. 7 km E of the intersection of Hwy 307 and the road to Cinco Lagos, 16.0939, -91.6361, 1461 m, 14 Sep 2017, E. Dean 9532 (DAV).

Based on Solanum chiapense Brandegee.

Guatemala. Suchitepéquez: Las Nubes, Nov 1875, K. Bernoulli & O. Cario 2397 (holotype: GOET [GOET003443]).

Figure 24. 

Image of herbarium specimen of L. chiapensis var. chiapensis, Standley 84958 (F). Specimen used with permission from the Field Museum of Natural History.

Woody vine to 10 m tall, probably taller. Indument of tan, pale yellow to brown, uniseriate, multicellular, stalked, multangulate-stellate (also some simple and dendritic), eglandular, spreading trichomes 0.1–0.5 (1 mm) long, 0.75–1 mm in diameter, the rays of the multangulate trichomes 3–6 per whorl, straight, rarely rebranched. Stems greenish-tan when young, sparsely to moderately pubescent, not compressed when dried in a plant press, becoming woody with age; upper sympodial branching usually monochasial, sometimes dichasial. Leaves simple, the leaves of the upper sympodia sometimes paired and unequal in size, the larger ones with blades 4–14 × 2–4.5 cm, the smaller ones with blades 2–7 × 1.5–3.5 cm, the leaf pairs similar in shape, the blades ovate, elliptic or obovate (sometimes the small geminate leaf nearly orbicular), thick chartaceous, sparsely to moderately pubescent especially along the veins (sometimes nearly glabrous), the base cuneate to rounded, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate (rarely rounded on smaller leaves), the petiole 0.2–2.5 cm long, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–3, axillary, oriented horizontally to ascending; peduncles absent; pedicels 10–25 mm and erect to arching in flower, to 40 mm long and erect to arching in fruit, glabrous to sparsely pubescent; calyx 4–5 mm long, 4–5 mm in diameter, campanulate, glabrous to sparsely pubescent, the margin truncate, slightly membranous, wavy or shallowly lobed, with 10 erect to spreading, linear appendages 0.25–3 mm long emerging 0.1–1.5 mm below the calyx rim; fruiting calyx enlarged, campanulate, remaining close to the fruit, 6–8 mm long, 10–13 mm in diameter, the appendages to 5 mm long; corolla 0.7–1.5 cm long, open corolla orientation unknown, stellate in outline, divided ca. 1/3 of the way to the base, with abundant interpetalar tissue, white, additional markings unknown, nearly glabrous to moderately pubescent with short trichomes abaxially near the veins; stamens subequal to unequal, straight, the four short filaments ca. 1 mm long, the long filament 1–2 mm long, glabrous, the anthers 5–5.5 mm long, lanceolate, free of one another, yellow, sometimes sparsely pubescent on inner surface along connective, poricidal at the tips, the pores obovate, dehiscing distally or toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 8–10 mm long, linear, straight, glabrous, the stigma ovoid. Fruit a berry, 10–20 mm long, 7–15 mm in diameter, ovoid, orange at maturity, glabrous, lacking sclerotic granules. Seeds 20–30 per fruit, 3.5–4 × 2.5–3 mm, flattened, with slightly thickened rim, depressed ovate in outline, yellow-orange to brown, the surface reticulum with minute, serpentine pattern with shallow luminae.

Unknown.

Mexico (Chiapas) and Guatemala (Quetzaltenango, San Marcos, Suchitepéquez), in tropical moist forest or cloud forest, sometimes on sand formations, 1500–2400 m in elevation (Fig. 25).

Figure 25. 

Map of geographic distribution of L. chiapensis var. chiapensis based on herbarium specimen data.

None known.

Flowering specimens have been collected in June. Specimens with mature fruits have been collected in December and January. The timing of the diurnal movements of the corollas of this species is not known, but the corollas are usually closed on herbarium specimens, indicating that they are probably open for a short time, most likely in the morning.

Lycianthes chiapensis var. chiapensis is a rarely collected variety of southwestern Guatemala and adjacent Mexico, represented by only 12 collections, three of which are from protected areas. The EOO is 3,020.311 km², and the AOO is 44 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Lycianthes chiapensis var. chiapensis is an upper elevation wet forest taxon that is localized in the southern tip of Chiapas and the western region of Guatemala along the Pacific slope. Unlike the more common variety, L. chiapensis var. sparsistellata Standl. & Steyerm., this variety is under-collected, and little is known about its appearance and growth form; it is likely a large liana like var. sparsistellata. The lower sympodial units merge into sinuate woody branches as the plant ages. The mature wood is dark brown and lustrous. This variety differs from var. sparsistellata in having a larger flowering calyx that remains campanulate in fruit and adheres to the fruit as it ages; var. sparsistellata has a much smaller calyx that becomes plate-like in fruit. The fruit of var. chiapensis is also larger, ovoid, and has more seeds. This variety was the first variety to be described, and the type specimens mainly have buds on them, but the larger size of the buds are obvious and different from those of var. sparsistellata. In addition, the leaves of var. chiapensis are somewhat thicker in texture and often more glabrous than those of var. sparsistellata. Although most authors of floras have synonymized the two varieties (for example: Gentry and Standley 1974; Nee 1986), they appear to be very different with regard to calyx morphology and fruit size, and so we are keeping them separate in this treatment. Standley and Steyermark (1940) introduced confusion into the description of the two varieties when describing var. sparsistellata for the first time. The type of var. sparsistellata is clearly that of the small calyx form, but then one of the paratypes cited (Purpus 7166 from Cerro del Boquerón, Chiapas) is clearly the large calyx form. In describing var. sparsistellata, the authors chose to emphasize pubescence density and ignored the differences in calyx and fruit size.

Guatemala. Quetzaltenango: Above Mujuliá, between San Martín Chile Verde and Colomba, 1800 m, 1 Feb 1941, P.C. Standley 85684 (F). San Marcos: near Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo, west facing slope of the Sierra Madre Mountains, 1800–2400 m, 10–18 Dec 1963, L.O. Williams 26281 (NY). Suchitepéquez: barranca by Loma Grande, above Finca El Naranjo, on Volcán Santa Clara, 1950–2100 m, 2 Jun 1942, J.A. Steyermark 46833 (NY). Mexico. Chiapas: Reserva de la Biosfera El Triunfo, Poligono III, Cerro Quetzal, 50 km al Sur de la Colonia Independencia, 15.7067, -92.9378, 1856 m, 1 Apr 2001, G. López-Hernández (MO).

Guatemala. Chiquimula: Tixixí (Tishishí), 3–5 miles north of Jocotán, 500–1500 m, 10 Nov 1939, J.A. Steyermark 31555 (holotype: F [0072902F, acc. # 1039909]; isotype WIS).

Figure 26. 

Image of herbarium specimen of L. chiapensis var. sparsistellata, Téllez 7946 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Shrub to woody vine, beginning as a shrub, becoming a large liana climbing into tree crowns up to 25 m tall and spreading in the crown up to 10 m wide, the lower stem to 15 cm in diameter. Indument of tannish-yellow to brown, uniseriate, multicellular, stalked, multangulate-stellate, eglandular, spreading trichomes 0.25–1 mm long, 0.75–1.2 mm in diameter, the rays 3–6 per whorl, straight, rarely rebranched. Stems light green when young (drying tan), moderately to densely pubescent, not compressed when dried in a plant press, becoming dark brown and woody with age, the stems sinuous, sometimes glabrate; upper sympodial branching usually monochasial, sometimes dichasial. Leaves simple, the leaves of the upper sympodia sometimes paired and unequal in size, the larger ones with blades 2.5–10 × 1–4 cm, the smaller ones with blades 0.8–4 × 0.5–2 cm, the leaf pairs similar in shape, the blades ovate, elliptic or obovate (sometimes the small geminate leaf nearly orbicular), thin chartaceous to chartaceous, sparsely to moderately pubescent especially along the veins (sometimes nearly glabrous), the base cuneate to rounded, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate (rarely rounded on smaller leaves), the petiole 0.2–1 cm long, the larger leaf blades with 4–5 primary veins on each side of the midvein. Flowers solitary or in groups of 2–3 (5), axillary, oriented horizontally to ascending; peduncles absent; pedicels (5) 7–24 mm and erect to arching in flower, to 30 mm long and erect to arching in fruit, sparsely to moderately pubescent; calyx (2) 2.5–3.5 mm long, 3–4.5 mm in diameter, campanulate, sparsely to moderately pubescent, the margin truncate, slightly membranous, truncate to wavy or shallowly lobed, with 10 erect to spreading, linear appendages 0.5–2.5 mm long emerging 0.25–0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl- to plate-shaped, 2–3(4) mm long, 5–8(10) mm in diameter, the appendages not lengthening and often breaking off; corolla 0.6–1.5 cm long, rotate to slightly reflexed in orientation, nearly entire to stellate in outline, divided ca. 1/3–2/3 of the way to the base, with abundant interpetalar tissue, white, the adaxial lobes sometimes with a green spot at the base near the insertion of the shorter stamens, sparsely to moderately pubescent with short trichomes abaxially near the veins; stamens unequal, the four short filaments 0.5–1 mm long, the fifth filament 2.5–4 mm long, glabrous, the anthers 4–5 mm long, lanceolate, free of one another, yellow, pubescent on the inner face, poricidal at the tips, the pores obovate, those of the shorter stamens dehiscing distally or toward the style, those of the long stamen dehiscing toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–8 mm long, linear, straight to slightly curved, glabrous, the stigma capitate. Fruit a berry, 5–10 (13) mm long, 5–10 (13) mm in diameter, globose, green to white when immature, orange to red at maturity, glabrous to sparsely pubescent, lacking sclerotic granules. Seeds 5–20 per fruit, 3.5–4.5 × 2.5–3.5 mm, flattened, with slightly thickened rim, depressed ovate in outline, yellow-orange to brown, the surface reticulum with minute serpentine pattern and shallow luminae.

Unknown.

Mexico (Chiapas, Oaxaca, and Veracruz), Guatemala (Baja Verapaz, Chiquimula, El Progresso, probably elsewhere), Honduras (Copán, Cortés, Ocotepeque), Nicaragua (Jinotega, Matagalpa), in primary or secondary cloud forest (including oak forest), montane rain forest, and tropical dry forest, (500) 900–2000 m in elevation (Fig. 27).

Figure 27. 

Map of geographic distribution of L. chiapensis var. sparsistellata from Mexico to Honduras based on herbarium specimen data.

None known.

In most parts of the range, flowering specimens have been collected July through November (January to March in Nicaragua). Specimens with immature and mature fruits have been collected throughout the year. In the field in Guatemala, the first author observed that the corollas of this species were still open at noon and closed later in the day.

Lycianthes chiapensis var. sparsistellata is a widespread variety of cloud forest habitat ranging from Mexico to Nicaragua, represented by 23 collections and occurring in five protected areas. The EOO is 156,415.154 km², and the AOO is 84 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Lycianthes chiapensis var. sparsistellata is an upper elevation cloud forest taxon that ranges from Veracruz to Honduras (possibly Nicaragua) mostly along the Caribbean slope. It can grow into a very tall liana that can cover the tree canopy, supported by a very large twining woody stem. The lower sympodial units merge into sinuate woody branches as the plant ages. The mature wood is dark brown and lustrous. This variety is the more common of the two varieties of L. chiapensis and differs from var. chiapensis in having a smaller flowering calyx that becomes plate-like as the plant fruits. The other variety has a larger flowering calyx that adheres to the fruit as it ages and a larger fruit with more seeds. See further discussion of the two varieties under var. chiapensis.

Guatemala. Alta Verapaz: at Orchigonia orchid nursery/preserve outside of the city of Cobán along Guatemala Highway 14, 15.4373, -90.4120, 1487 m, 10 Aug 2017, E. Dean 9507 (DAV). Chiquimula: Cerro Tixixí (Tishishí), 3–5 m north of Jocotán, 500–1500 m, 10 Nov 1939, J.A. Steyermark 31555 (F, WIS). El Progreso: Cerro Pinalón, Sierra de las Minas, San Acasaguastlán, 15.0656, 89.9833, 2230 m, 1 Mar 2007, M. Flores 3548 (MO). Mexico. Chiapas: cima del Cerro Salomón, al NO de Benito Juárez, ca. 44 km en línea recta al N de San Pedro Tapantepec, 16.7708, -94.1953, 1770 m, 7 Apr 1986, M. Ishiki 1451 (NY). Oaxaca: Cerro Sabinal, ca. 2 km al SO de Cerro Guayabitos, ca. 3 km en línea recta al NNO de Díaz Ordaz, ca. 40 km en línea recta al N de San Pedro Tapanatepec, al O de la cima del cerro, 16.7333, -94.1917, 1500 m, 21 Dec 1984, T. Wendt 4678 (NY). Veracruz: along trails to base of Volcán Santa Marta, 0–3 km E village of Santa Marta, [18.35, -95.8667], 1100–1200 m, 29 Jun 1982, M. Nee 24700 (F, NY, XAL).

Based on Solanum ciliolatum M.Martens & Galeotti.

Figure 28. 

Image of herbarium specimen of L. ciliolata, Dean 227 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Perennial herb from fusiform storage roots, usually erect (0.1) 0.2–0.6 (1 m) tall, dying back each season. Indument of white, uniseriate, multicellular, simple or dendritically branched, eglandular, spreading to appressed trichomes 0.1–1 (1.5) mm long. Stems greenish purple, sparsely to moderately pubescent, usually not much compressed upon drying in a plant press, woody with age, especially at the base of the plant; first stem 5–60 cm long to the first inflorescence, the internodes 4–10 (19); first two sympodial branching points dichasial, followed by monochasial branching, this usually extensive. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades 2–14 × 1–7 cm, the smaller ones with blades 1/4 to 3/4 (to equal) the size of the larger, the leaf pairs similar in shape, the blades ovate, lanceolate, or elliptic, (rarely obovate), chartaceous, sparsely to moderately pubescent, the primary veins 4–7 on each side of midvein, the base truncate, obtuse, or cuneate, attenuate onto the petiole, often slightly oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate (rarely long-acuminate), the petiole of larger leaves winged and poorly defined, 0.3–2.5 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 30–90 mm and erect in flower, (30) 40.5–80 (110) mm long and deflexed in fruit, sparsely to moderately pubescent with spreading to appressed-retrorse trichomes; calyx 2.5–4.5 mm long, (2.5) 3.5–5.5 (6.5) mm in diameter, obconic to campanulate glabrous to moderately pubescent, the margin truncate, with 10 linear, reflexed appendages 3–9 (11) mm long emerging ca. 0.5–1 mm below calyx rim; fruiting calyx enlarged, (1.5) 2–4 (6) mm long, 5–12.5 (14) mm in diameter, the appendages 3–11 mm long, reflexed, often broken; corolla 1.1–2.7 cm long (2.1–5.3 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, lilac, with maroon to purple stripes along the major veins adaxially, green near the major veins abaxially, usually glabrous; stamens unequal, straight, the filaments of three lengths, the two shortest filaments 1(0.5) 1–3.5 (4.25) mm long, the two medium filaments 1–4.5 (5.5) mm long, the one long filament (2) 3–7 (8) mm long, the length of the long filament nearly always 1.8–3 times that of medium filaments (rarely 1.5–1.8 times), glabrous, the anthers (3) 4–6.25 (8) mm long, lanceolate to oblong, free of one another, yellow, glabrous, poricidal at the tips, the pores linear to ovate, usually dehiscing distally or toward the style, not opening into longitudinal slits; pollen grains dicolporate with remnant third pore; pistil with glabrous ovary, the style 9–12 (13) mm long, linear, straight to slightly curved, glabrous, the stigma round to weakly bilobed (in Oaxaca) to very bilobed (in Guatemala). Fruit a berry, remaining attached to calyx at maturity, pendent, 14–39 mm long, (7) 9–22 mm in diameter, ovoid to conic, the exocarp dark purple to rose-colored, glabrous, the mesocarp rose-colored or dark purple, soft and juicy, lacking sclerotic granules, the placental area light purple and powdery. Seeds (8) 30–80 (102) per fruit, 2.9–4.1 × 2.5–3.9 mm, not compressed, depressed obovate, ridged and blistered along one side, black, the surface reticulum rough in texture with loose serpentine pattern and deep luminae.

2n = 24 from Dean 271, 295 (Dean 2004)

Mexico (Chiapas, Guanajuato, Oaxaca, Puebla, Querétaro, San Luis Potosí) and Guatemala (Baja Verapaz, Huehuetenango, Totonicapán), in oak or oak/pine forest (that may be intermixed with palms, Juniperus or Yucca) or in xerophilous scrub in southern Mexico, on slopes, in drainages, in canyons, along paths, and in agricultural fields, on limestone soils, 755–3000 m in elevation (Fig. 29).

Figure 29. 

Map of geographic distribution of L. ciliolata based on herbarium specimen data.

Flowering specimens have been collected June to October, depending on region. Specimens with mature fruits have been collected between September and November. The first author observed in the field that the corollas are open in the very early morning and closed by late morning. The pollen has a sweet scent. Solitary bees in the genera Thygator and Pseudoaugochloropsis visit this species (Dean 2001).

Mexico. Hoh, yich hoh, yichjoj, tintolón, ma’ ‘u’ dsea nuu jgiaa, u dsea niquia, yich balam, guì in-dèm, ngûd-dèm, guizh-dam, kuan xille bekue, campanilla, chichi de vaca, chichi de perro, chichi de venado, binduchi, binduchito, rshtisti katya, tronchichi, tonchicho, manzanillo del campo, shashasto, la pera, chile de ratón. People eat the fruits of this species in the states of Puebla, Oaxaca, and Chiapas. The edible fruits are gathered from wild plants, often from plants growing as volunteers in agricultural fields (Dean 2004).

Lycianthes ciliolata is a widespread species ranging from central Mexico to Guatemala represented by 117 collections and occurring in six protected areas. Anguiano-Constante et al. (2018) provided a preliminary conservation assessment of Least Concern (LC).

Lycianthes ciliolata is similar to L. rzedowskii and L. acapulcensis. It differs from those species in having lilac rather than white corollas (or the very pale lilac sometimes found in L. rzedowskii). The pattern of filament lengths can also be useful in separating it from L. rzedowskii. In L. ciliolata the longest stamen filament is often more than twice as long as the medium-short filaments, while in L. rzedowskii the longest stamen filament is less than twice as long as the medium-short filaments. The lengths of the pedicels of the youngest mature flowers relative to their subtending leaves is often a useful character for separating L. ciliolata from L. acapulcensis. In the latter, the length of those pedicels is usually less than that of the subtending leaves, while in L. ciliolata the length of the pedicels generally exceeds that of the leaves (Dean 2004).

Guatemala. Baja Verapaz: Patal, 1600 m, H. von Tuerkheim 2317 (W, BR, E, F , G, MO, NY, US). Huehuetenango: Cerro Pixpix, above San Ildefonso, Ixtahuacan, [15.4675, -91.8116], 1600–200 m, 15 Aug 1942, J.A. Steyermark 50597 (NY no #). Totonicapán: Cerro María Tecum, Sierra Madre Mountains, 10–20 km east of Totonicapán, [14.8959, -91.2636], 3100–3400 m, 16 Dec 1962, L.O. Williams 23156 (MO). Mexico. Chiapas: Col. Carrizal, 700 m al oriente de la Escuela, 16.6575, -92.6975, 2250 m, 6 Jun 1995, H. Mejía 429 (XAL). Guanajuato: Rancho Beltrán, 10 km al oeste de Xichu, [21.3164, -100.0987], 2000 m, 9 Dec 1990, E. Ventura 6473 (XAL). Oaxaca: Santa María Jaltianguis, [17.361, -96.5282], 7200 ft, 20 Oct 1991, E. Dean 295 (DAV, IEB, MEXU). Puebla: hills to the SW of the city of Tehuacán, up dirt road near El Riego, [18.4433, -97.4235], 1677 m, 27 Sep 1991, E. Dean 272 (DAV). San Luis Potosí: Los Aguajitos, 11 km al NE de Guadalcázar, hacia Pozo de Acuña, 22.625, -100.3167, 2000 m, 17 Nov 1996, R. Torres C. 14874 (MEXU).

Guatemala. Huehuetenango: Cruz de Limón, between San Mateo Ixtatán and Mucá, Sierra de los Cuchumatanes, 2600–3000 m, 31 Jul 1942, J.A. Steyermark 49828 (holotype: F [0072906F, acc. # 1199398]).

Figure 30. 

Image of herbarium specimen of L. connata, Dean 9530 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Shrub, erect, 0.5–5 (7) m tall. Indument of white, off-white, or brownish, uniseriate, multicellular, simple, curved, eglandular, appressed-ascending trichomes 0.25–1 mm long. Stems green to purplish when young, glabrous to sparsely pubescent, compressed 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 4.5–16.5 (20) × 1.5–8.5 cm, the smaller ones with blades 1.5–8.5 × 0.5–4 cm, the leaf pairs usually similar in shape, the blades ovate, elliptic, or obovate, membranaceous, glabrous to sparsely pubescent, sometimes with purple veins, the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 0.1–3 cm long, sometimes absent, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–5, axillary, erect or oriented horizontally; peduncles absent; pedicels 15–35 mm and erect to arching in flower, to 40 mm long, and erect to arching in fruit, usually glabrous; calyx 2–4 mm long, 4–5 mm in diameter, campanulate to widely bowl shaped, sometimes appearing flat-bottomed, usually nearly glabrous, the margin truncate, very well developed, with 10 linear, ascending to reflexed appendages 0.5–4 mm long, connate to each other at the base, emerging 1–3 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 2–2.5 mm long, 5–7 mm in diameter, the appendages 1.5–3.5 mm long, spreading to reflexed; corolla 0.5–1.7 cm long, rotate to reflexed in orientation, entire to slightly stellate in outline, divided ca. 1/4 of the way to the base, with well-developed interpetalar tissue, white, sometimes with a purple ring in the center adaxially, mostly glabrous; stamens nearly equal to unequal, the four short filaments 1–1.5 mm long, the one long filament 1.5–2.5 mm long, glabrous, the anthers 3.5–5 mm long, lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores oval, dehiscing distally, sometimes enlarging by slitting laterally down the side of the anther; pistil with glabrous ovary, the style 6–7 mm long, linear, straight to slightly curved, glabrous, the stigma capitate. Fruit a berry, 5–9 mm long, 5–9 mm in diameter, globose to ovoid, orange at maturity, glabrous, lacking sclerotic granules. Seeds 20–60 per fruit, 1.5–2 × 1.25–1.5 mm, flattened, depressed ovate to reniform in outline, with notch on one side, orange-yellow to orange-brown, the surface reticulum with minute serpentine pattern and shallow luminae.

Unknown.

Mexico (Chiapas, Oaxaca) and Guatemala (El Progreso, Huehuetenango, Zacapa) in cloud forest and montane rain forest, often with Quercus, Pinus, Podocarpus, Magnolia, sometimes on slopes, 1600–3000 m in elevation (Fig. 31).

Figure 31. 

Map of geographic distribution of L. connata based on herbarium specimen data.

None known.

Flowering specimens have been collected from March through December. Specimens with mature fruits have been collected October through March. The diurnal movements of the corolla are not known, but as some specimens have open corollas, the corollas must stay open at least until late morning.

Lycianthes connata is a cloud forest species of Oaxaca, Chiapas and Guatemala, represented by 27 collections, four of which are from two protected areas. The EOO is 71,681.613 km² and the AOO is 92 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Lycianthes connata is a very distinctive species due to the structure of its calyx which has a very long, sleeve-like margin and appendages that are connate at their bases that are reflexed in fruit. The only species that can be confused with Lycianthes connata are L. ceratocalycia and L. gongylodes, both of which have stems that compress upon drying and short, bulging appendages that can give a somewhat similar appearance to the calyx. Lycianthes ceratocalycia differs in having stellate, purple corollas, equal stamens, and young stems with scurfy, horizontal lines. L. gongylodes differs in having equal stamens and stems that are moderately pubescent with curling trichomes. Lycianthes connata was originally described from Guatemala and has been collected often in Chiapas. There are fewer collections from high elevation Oaxaca in the Sierra de Juárez. The Oaxacan populations are very like the Chiapas populations except that the calyx appendages are notably shorter, making the calyx look more like that of L. gongylodes or L. ceratocalycia.

Guatemala. El Progreso: On top of Montaña Piamonte, along Joya Pacayal, 3000 m, 7 Feb 1942, J.A. Steyermark 43677 (NY). Huehuetenango: Cruz de Limón, between San Mateo Ixtatán and Mucá [Nucá], Sierra de los Cuchumatanes, [15.8306, -91.4447], 2600–3000 m, 31 Jul 1942, J.A. Steyermark 49828 (F). Zacapa: Mpio. Río Hondo, 1.5 horas N Finca Alejandra, 30 minutes S Cerro Paloma (P26-Proyecto Deslaves), 15.1569, -89.6178, 2512 m, 17 Mar 2012, C. Cifuentes 435 (BIGU). Mexico. Chiapas: Tzontehuitz. Mpio. Chamula, 16.6856, -92.5714, 2897 m, 28 Aug 1999, L.Y. Domínguez-Torres 105 (MEXU). Oaxaca: Dto. Mixe, Kets tekum, tonun Kux, [17.2523, -96.0292], 17 Jul 1994, Rivera-Reyes 3156 (IEB, MEXU).

Guatemala: Huehuetenango: between Xoxlac and Nucapuxlac, Sierra de los Cuchumatanes, 1600–2500 m, 17 Jul 1942, J.A. Steyermark 48925 (holotype: F [0072907F, acc. # 1188543]; isotype: A [00934886]).

Figure 32. 

Image of holotype of L. cuchumatanensis, Steyermark 48925 (F). Specimen used with permission from the Field Museum of Natural History.

Scandent shrub to vine, height unknown. Indument of pale yellow to reddish-brown, uniseriate, multicellular, sessile to stalked, multangulate stellate to geminate stellate (multistoried), eglandular, spreading trichomes, 0.25–1 mm long, 0.5–1 mm in diameter, the rays 5–8 per whorl, straight, not rebranched. Stems pale green (drying tan) when young, sparsely to densely pubescent (the surface often obscured), not compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points monochasial and dichasial, the branching divaricate (diverging at wide angles). Leaves simple, the leaves of the upper sympodia rarely paired and unequal in size, the blades of the larger ones 5–10 × 2–3.5 cm, the blades of the smaller ones 2–3 × 0.5–1.5 cm, ovate, obovate, lanceolate, or elliptic, subcoriaceous to coriaceous, adaxially sparsely to moderately pubescent (with trichomes concentrated along the veins), abaxially moderately to densely pubescent (with leaf surface sometimes obscured by pubescence), the base cuneate, the margin entire, usually irregularly undulate, the apex acute to acuminate, the petiole 0.5–1.2 cm long, the larger leaf blades with 3–5 primary veins on each side of the midvein. Flowers usually in groups of 2–4, axillary, erect; peduncles absent; pedicels 10–14 mm long and erect in flower, to 15 mm long and erect in fruit, densely pubescent (the surface often obscured); calyx 2.5–3.5 mm long, 2.5–3.5 mm in diameter, campanulate, densely pubescent (the surface usually obscured), the margin truncate, prominent, undulate, scarious (sometimes torn), with 10 obovate, spreading appendages 1–1.5 mm long emerging 1–2 mm below the calyx rim; fruiting calyx slightly enlarged, bowl-shaped, sometimes splitting, 2.5–3 mm long, 4–5 mm in diameter, the appendages to 2 mm long, spreading to reflexing; corolla 0.7–1.1 cm long, open orientation unknown, stellate in outline, divided 1/2 of the way to the base, with scant interpetalar tissue, adaxially purple, glabrous, abaxially densely pubescent on the lobes; stamens equal, straight, the filaments ca. 1 mm long, glabrous, the anthers 3–3.5 mm long, elliptic, free of one another, color uncertain, sparsely pubescent with scattered trichomes, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–7 mm long, linear, straight, glabrous, the stigma capitate. Fruit a berry, ca. 7 mm long, 7 mm in diameter, color unknown, glabrous, lacking sclerotic granules. Seeds number per fruit and seed details unknown, ca. 2.5–3 mm long.

Unknown.

Guatemala (Alta Verapaz, Huehuetenango), 1500–2600 m in elevation. Nothing is known about the habitat where this species grows, but it may be cloud forest (Fig. 33).

Figure 33. 

Map of geographic distribution of L. cuchumatanensis based on herbarium specimen data.

None known.

Flowering specimens have been collected in July and August, and fruiting specimens have been collected in July. Very little is known about this species. The corollas are closed on the few specimens that exist of this species, indicating that the corollas have diurnal movements, but the timing is unknown.

Lycianthes cuchumatanensis is a rarely collected species of Guatemala, represented by only three collections, all outside of protected areas. The EOO is 773.642 km², and the AOO is 12 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Critically Endangered (CR).

Although Gentry (1973) thought Lycianthes cuchumatanensis to be a close relative of L. limitanea, L. cuchumatanensis does not resemble L. limitanea. It is very similar to L. sideroxyloides in its pubescence, branching pattern, solitary leaves, stellate corollas, and equal stamens. It differs from that species in having leaf blades that are usually chartaceous to thick chartaceous, rather than coriaceous, have more rounded bases (rather than cuneate), and have less dense pubescence on the abaxial side. Lycianthes sideroxyloides also has a smaller seed size (1.5–2 mm long) than that cited in the protologue for L. cuchumatanensis (Gentry 1973). The paratype cited by Gentry in the protologue (Steyermark 48625) differs from the holotype in having less dense pubescence and leaf blades that are thinner in texture. Further field work is necessary to locate extant populations at the type locality of L. cuchumatanensis to determine if it is conspecific with L. sideroxyloides. The name Lycianthes cuchumatanensis has been misapplied to L. breedlovei and L. fredyclaudiae (Dean et al. 2019a).

Guatemala. Alta Verapaz: Mpio. San Juan Chamelco, Chicacnab, La Laguna, 15.3844, -90.1639, 2300 m, 4 Aug 1998, M. Robles 124 (MSB). Huehuetenango: Cerro Huitz between Mimanhuitz and Yulhuitz, Sierra de los Cuchumatanes, [15.8550, -91.3244] 1500–2600 m, 14 Jul 1942, J.A. Steyermark 48617 (G).

Based on Solanum dejectum Fernald.

Figure 34. 

Image of herbarium specimen of L. dejecta, Dean 224 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Perennial herb from very large fusiform storage roots, decumbent to erect, decumbent forms to 0.5 m in diameter, tall forms often scrambling, supported by nearby shrubs, to 0.7 m tall, dying back each season. Indument of white, uniseriate, multicellular, simple, dendritically branched, and multangulate-stellate, eglandular, spreading trichomes 0.1–0.5 (0.75) mm long, 0.5–0.75 mm in diameter, the rays of the multangulate trichomes 3–4 per whorl, straight, often rebranched. Stems green with darker green and purple striations, moderately to densely pubescent, much compressed when pressed and dried, becoming woody with age only near the base; first stem 1–35 cm long to the first inflorescence, the internodes 2–7 (13); first sympodial branching point dichasial, followed by a mixture of monochasial and dichasial branching, this branching extensive. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades to 3.5–14.5 × 2–8.5 cm, the smaller ones with blades ca. 1/4–3/4 the size of the larger, the leaf pairs similar in shape, the blades ovate, deltate, or reniform, chartaceous, moderately to densely pubescent, the primary veins 3–5 on each side of the midvein, the base truncate, cuneate, attenuate, decurrent onto the petiole, slightly oblique, the margin entire, usually slightly undulate, the apex acute, the petioles winged and poorly defined, 2–5.5 cm long. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 29–80 (115) mm and erect in flower, (35) 50–110 (125) mm long and deflexed in fruit, moderately to densely pubescent with spreading trichomes; calyx 2.5–4 mm long, 2.5–6 mm in diameter, obconic, the margin truncate, with 10 linear, somewhat reflexed appendages (1) 2–7 (8) mm long emerging ca. 0.5 mm below the calyx rim (usually obscured by trichomes); fruiting calyx enlarged, (5) 6–10 (11.5) mm long, (9) 11–20 (23) mm in diameter, the appendages reflexed to curved, often broken, (3.5) 4–15 (19) mm long; corolla 1–2.2 cm long (1.9–4.1 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white to lilac, with maroon to purple stripes along the major veins adaxially, green, white, or purple and densely pubescent near the major veins abaxially; stamens unequal, straight to curved, the filaments of three lengths, the two shortest (1.25) 1.5–4 mm long, the two medium filaments (1.5) 2–5 mm long, the one long filament (2) 3–7.5 mm long, the length of the long filament 1.1–1.8 (2) times that of the medium filaments, glabrous; anthers 3–6 mm, ovate to lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pollen grains tricolporate; pistil with glabrous ovary, the style (6) 7–11.5 mm, linear, straight to curved downward, the stigma lobed. Fruit a berry, remaining attached to calyx at maturity, pendent, sometimes near the ground, 17–39 mm long, 12–24 mm in diameter, ovoid to conic, the exocarp light to dark green with purple or black lines (becoming yellowish or brown in age), the mesocarp white to green and juicy, lacking sclerotic granules, the placental area narrow, greenish-white, juicy. Seeds (40) 50–170 (185) per fruit, 2.2–2.8 × 1.5–2.5 mm, rounded, slightly compressed, reniform to depressed-obovate, dark brown to black, surface reticulum with loose serpentine pattern with deep luminae and microscopic fibrils protruding from the cell walls.

2n = 24, Dean 276, 309 (Dean 2004)

Mexico (Baja California Sur, Distrito Federal, Durango, Guanajuato, Hidalgo, Jalisco, México, Michoacán, Nuevo León, Oaxaca, Puebla, Querétaro) on limestone on either side of the transvolcanic belt, as well as in eroded, ancient agricultural areas within the transvolcanic belt (rarely on rhyolite), usually in xerophilous shrub; it has also been found in disturbed relictual tropical dry forest or oak forest. It has been suggested that eroded volcanic areas within the Valley of Mexico are often home to calciphiles, because erosion has exposed a lower soil layer that is calcium rich (Rzedowski 1986). Habitats include pastures, paths, the sides of agricultural fields, and within abandoned fields at 1800–2900 m in elevation (Fig. 35).

Figure 35. 

Map of geographic distribution of L. dejecta based on herbarium specimen data.

Mexico. Trompeta, chichi de perra (Dean 2004); used to treat stomachache in Guanajuato (Ocampo 47).

Flowering specimens have been collected June to August; specimens with mature fruits have been collected September to October. The first author observed in the field that the corollas open after sunrise and close by early afternoon. The pollen in this species has a sweet scent. Solitary bees in the genus Colletes visit this species (Dean 2001).

Lycianthes dejecta is a widespread Mexican endemic, represented by 55 collections and occurring in three protected areas (Sierra la Laguna, Sierra Gorda and Tehuacán-Cuicatlan Valley). Anguiano-Constante et al. (2018) provided a preliminary conservation assessment of Least Concern (LC).

Lycianthes dejecta is a perennial herb recognized by its dense, dendritic to multangulate-stellate trichomes, which cover all parts of the plant, and the truncate bases of its leaf lamina. Its fruits and seed type are similar to those of L. moziniana. It differs from that species in having maroon to black lines on its fruits, reflexed to curled calyx teeth, and microscopic fibrils on its seeds. All parts of this plant, including the fruits, have a bitter taste (Dean 2004).

Mexico. Baja California Sur: Mpio. La Paz. El Paraje de Cano, Sierra de la Victoria, 23.5833, -109.9167, 1670 m, 30 Sep 1994, M. Domínquez L. 800 (HCIB). Distrito Federal: Sierra de Guadalupe, [19.5908, -99.1203], 7000 ft, Balls 5073 (BM, K, UC). Durango: [24.0237, -104.6580], Apr to Nov 1896, E. Palmer 347 (BM, C, CAS, G, UC, F, US, MO, NY). Guanajuato: cañada a 5 km de Santa Anita, cerro La Meza, [20.9667, -100.3], 2300 m, 22 Sep 2002, Castillejos-Cruz 1229 (MEXU). Hidalgo: Cañon de las Ajuntas, Santa María Macua, 20.1125, -99.4625, 2150 m, 15 Jun 2003, L. Romero 75 (MEXU). Jalisco: carretera Lagos de Moreno-Leon, km 31, [21.1739, -101.7238], 2020 m, 15 Jul 1991, H. Arreola-Navas 1270a (MEXU). México: N of Huehuetoca along the road to Apaxco, c. 4.2 road mi from building “los arcos” (in dowtown Huehuetoca), W side of rd, [19.8894, -99.2141], 7100 ft, 3 Aug 1991, E. Dean 243 (DAV). Michoacán: Vic. Morelia, Punguato, [19.6954, -101.1381], 2100 m, 20 June 1912, Arsène 8300 (F, GH, MO, NY). Nuevo León: Cerro El Gallo, [24.92, -99.78], 2085 m, 15 Jun 1991, G. Hinton 21019 (GH, IEB, TEX). Oaxaca: a las afueras de Guadalupe Membrillos, 18.0228, -97.5508, 2276 m, 12 Aug 2004, O. Téllez-V. 17009 (MEXU). Puebla: Mpio. Caltepec, between Cerro Pochote and Cerro Gavilán Chico in hills SE of town of Caltepec, along road to Atolotitlan, near small valley called La Laguna, [18.1784, -97.4698], 6800–6900 ft, E. Dean 228 (DAV). Querétaro: 2.09 km al NO de Bernal, Ezequiel Montes, 20.7508, -99.9572, 2240 m, 21 Sep 2012, O. Rubio-García 263 (IEB).

Guatemala. Baja Verapaz: Niño Perdido, Cerro Verde, east of km 150 of Cobán Road, in high forest, elevation not recorded, 3 Dec 1976, C.L Lundell 20419 (holotype: LL 00490012; isotypes: F [acc. # 1912542], MO [acc. # 3342033], LL [00490006]).

Figure 36. 

Image of herbarium specimen of L. fredyclaudiae, Lundell 21085 (LL). Specimen used with permission from the Lundell Herbarium, University of Texas at Austin.

Scandent shrub to weak vine, sometimes epiphytic, 2–3 m tall. Indument of tan, pale yellow, or orange, uniseriate, multicellular, stalked, multangulate-stellate, eglandular, spreading trichomes 0.25–1 mm long, 0.75–1.2 mm in diameter, the rays 3–5 per whorl, straight, often rebranched. Stems tan-green to purple-green when young, moderately to densely pubescent, not compressed when dried in a plant press, becoming dark brown and woody with age, the surface of the stems shiny and somewhat longitudinally wrinkled upon drying; upper sympodial branching points mostly monochasial, with some dichasial branching point, the branching not widely divaricate, the adjoining branches often forming straight continuous axes. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 3–8.5 × 1.5–4 cm, the smaller ones with blades 1–3.5 × 0.5–2.5 cm, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, the smaller leaf sometimes nearly round, coriaceous, sparsely to densely pubescent, sometimes nearly glabrous adaxially, the trichomes usually dense on the abaxial side, especially along the veins, the base cuneate to rounded, sometimes oblique, the margin entire, usually irregularly undulate, the apex obtuse, acute, or short-acuminate, the petiole 0.2–1.4 cm long, sometimes absent, the larger leaf blades with 3–5 primary veins on each side of the midvein. Flowers in groups of 2–8, axillary, oriented horizontally; peduncles absent; pedicels 8–25 mm long and erect in flower, to 31 mm and erect in fruit, moderately to densely pubescent; calyx 2–3.5 mm long, 3–4.5 mm in diameter, campanulate, moderately to densely pubescent, the margin truncate, with 10 spreading linear appendages 0.5–2 mm long emerging 0.25–0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 2–3 mm long, 5–8 mm in diameter, the appendages not elongating; corolla 0.7–1.7 cm long, rotate in orientation, entire in outline, with abundant interpetalar tissue, adaxially white to pale lilac, very sparsely puberulent, abaxially white to pale lilac, the lobes sometimes greenish, moderately to densely puberulent; stamens unequal, straight, the four short filaments 0.5–2 mm long, the one long filament 2–4 mm long, glabrous, the anthers 4–5 mm long, lanceolate, free of one another, yellow to purple, usually with small, white trichomes scattered on either the face of the anther or on the two lobes at the very bottom of the anther, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–8 mm long, linear, straight, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 5–13 mm long, 4–12 mm diameter, globose to depressed globose, green to white when immature, yellow to orange when mature, sometimes with a few scattered trichomes, lacking sclerotic granules. Seeds 10–40 per fruit, 2.5–4 × 2–3.5 mm, flattened, thickened on the edges, circular to depressed ovate in outline, sometimes reniform with small notch on one side, yellow-orange to orange-brown, the surface reticulum with loose serpentine pattern and deep luminae, the margin rougher in texture than the center

Unknown.

Guatemala (Baja Verapaz), in cloud forest, “tall forest,” wet forest thickets, and forest floors, sometimes along drainages or on slopes, prefers undisturbed forest, 1500–1800 m in elevation (Fig. 37).

Figure 37. 

Map of geographic distribution of L. fredyclaudiae based on herbarium specimen data.

None known.

Flowering specimens have been collected all months of the year except September to November and January to February; specimens with mature fruits have been collected all months of the year except April and May. In the field, the first author observed that the corollas were open in the morning and closed in the afternoon.

Lycianthes fredyclaudiae is a Guatemalan cloud forest endemic, represented by 13 collections from a relatively small area, with six collections from protected areas (Mario Dary Rivera and Sierra de Minas). Dean et al. (2019a) provided a preliminary conservation assessment of Endangered (EN) for this species.

Lycianthes fredyclaudiae is a species of cloud forest that is known only from the area south of Cobán in the Department of Baja Verapaz. In the 1970s, this species was often collected in primary forest in the vicinity of Union Barrios (collections include those made by the fourth author of this paper), but when this area was revisited in 2017, most of the forest had been cleared, and it was difficult to find the species in the remnants along the highway. One plant was located growing in a roadside thicket, after two days of searching the area, including two reserves. Therefore, it is assumed that the species prefers more undisturbed, lower light habitats, and the species may now be a rare plant (Dean et al. 2019a). Lycianthes fredyclaudiae has been misidentified as L. cuchumatanensis, a species related to L. sideroxyloides. It differs from that species in having rotate, mostly entire corollas (vs stellate corollas), and unequal stamens (vs equal). It is most similar to L. chiapensis, L. breedlovei, and L. hortulana. Lycianthes chiapensis differs in being a large vine climbing high into the canopy, having thinner leaves, and having trichomes with rays that are mostly unbranched. Lycianthes breedlovei and L. hortulana differ from L. fredyclaudiae in having stellate corollas and divaricate branching (Dean et al. 2019a).

Guatemala. Baja Verapaz: along Guatemala Highway 14 just south of highway marker 158 and La Ram Tzul nature reserve, W side of the road, 15.2064, -90.2065, 1610 m, 11 Aug 2017, E. Dean 9508 (DAV226622).

Based on Solanum geminiflorum M.Martens & Galeotti.

Figure 38. 

Image of herbarium specimen of L. geminiflora, Dean 9523 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Herb, shrub, to treelet, erect, 0.5–5 m tall. Indument of very small, off-white to tan, uniseriate, multicellular, simple, eglandular, appressed-ascending trichomes < 0.1 (0.2) mm long. Stems green when young, sparsely to moderately pubescent, compressed 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 5–15 × 1.5–6 cm, the smaller ones with blades 2–6 × 0.5–3 cm, the leaf pairs usually similar in shape, the blades ovate, elliptic, or 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 0.2–0.5 (0.7) cm long, the larger leaf blades with 5–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–8, axillary, erect; peduncles absent; pedicels 8–20 mm and erect in flower, to 25 mm long and erect in fruit, glabrous to sparsely pubescent; calyx 1–1.5 mm long, 2.5–3 mm in diameter, widely campanulate, puberulent with very small trichomes, the margin truncate, undulate, the appendages lacking; fruiting calyx not very enlarged, widely bowl-shaped, 0.5–1.5 mm long, 3–4 mm in diameter; corolla 0.6–1.2 cm long, campanulate to reflexed in orientation, stellate in outline, divided nearly to the base, interpetalar tissue mostly lacking, white and glabrous adaxially, yellow-green to green and puberulent abaxially; stamens equal, straight, the filaments ca. 1 mm long, glabrous, the anthers 3–3.5 mm long, elliptic, usually partially connivent to the adjacent anther (at least near the middle or base of the anther, not at the tips), yellow, glabrous, poricidal at the tips, the pores round, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–8 mm long, linear, straight to curved, glabrous; stigma capitate, decurrent down two sides. Fruit a berry, 4–7 mm long, 4–10 mm in diameter, globose to depressed globose, orange-red at maturity, glabrous, lacking sclerotic granules. Seeds 20–70 per fruit, 1.25–1.75 × 1–1.25 mm, flattened, depressed ovate in outline, tan to orange, the surface reticulum with pitted serpentine pattern with deep luminae.

Unknown.

Mexico (Hidalgo, Oaxaca, Puebla, Veracruz) in cloud forest, tropical dry forest, oak and oak-pine forest, sometimes in disturbed habitats, secondary forest, slopes, coffee plantations, 800–2200 (3000) m in elevation (Fig. 39).

Figure 39. 

Map of geographic distribution of L. geminiflora based on herbarium specimen data.

Mexico. Oaxaca: leaves used as an edible green (Boyle 2631); ndia-sku-ya (Mazatec) (Giovannini 236a). Puebla: plant used as an edible green;, ndazkjuyoo (Mazateco) and cuajquilitl (Nahuatl) (C. Mota Cruz 652). Veracruz: hierba mora (Dorantes 100).

Flowering and fruits specimens have been collected most months of the year. Based on field observations by the first author, the corollas of this species are open for much of the day in some locations.

Lycianthes geminiflora is a species of threatened cloud forests of central and southern Mexico, represented by 55 collections, all collected outside of protected areas. The EOO is 39,209.944 km², and the AOO is 200 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Lycianthes geminiflora is a large herb to treelet, usually found in cloud forest on slopes. It is very similar to L. heteroclita (Sendtn.) Bitter, with which it can overlap in distribution in Mexico. The two species differ in the size of the flowering calyx, with that of L. geminiflora 1.5 mm long or less, and that of L. heteroclita usually 2 mm long or more.

Mexico. Hidalgo: Mpio. Tianguistengo, 10 km al E de Tianguistengo, [20.7265, -98.5279], 6 Jul 1995, M. Sousa Peña 594 (IEB, MEXU). Oaxaca: Sierra de Juárez, Mpio. Comaltepec, along Hwy 175 to the NE of the turnoff to Comaltepec, and NE of the cabins and restaurant of Mirador, trail on the southeast side of the road called Sendero Relampago, 17.5918, -96.3999, 2080 m, 10 Sep 2017, E. Dean 9521 (DAV). Puebla: La Guacamaya, [18.3375, -96.8230], 1100 m, Oct 2006, C. Mota-Cruz 652 (XAL). Veracruz: Rancho El Riscal, 19.4544, -96.9964, 2180 m, 26 Sep 2007, J. Hernández-M. 24 (XAL).

Mexico. Querétaro: Mpio. Landa, 10 km al noreste de Agua Zarca, sobre camino a Neblinas, 1100 m, 23 Jun 1988, J. Rzedowski 46837 (holotype: DAV [acc. # 217731]; isotypes: IEB [acc. # 193504], TEX [00449100]).

Figure 40. 

Image of holotype of L. glabripetala, Rzedowski 46837 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Perennial herb to small shrub, 0.5–2 m tall. Indument of off-white to tan, uniseriate, multicellular, simple, acute, curved to crisped, eglandular, appressed-ascending (rarely spreading) trichomes, 0.25–1.25 mm long. Stems green when young, moderately to densely pubescent, somewhat compressed upon drying in a plant press, light brown and woody with age; upper sympodial branching points usually monochasial with a few dichasial branching points. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades (4.5) 8.5–14 × (1.8) 2.5–5 cm, ovate to elliptic, the smaller ones with blades 1.3–4.5 × 0.8–2.1 cm, usually ovate, the blades of both the large and small leaves chartaceous, moderately to densely pubescent, the pubescence densest along the veins of the abaxial side, the trichomes along the midvein of the abaxial side appressed and appearing woolly, the base cuneate, usually oblique (sometimes rounded in the smaller leaves), the margin entire, usually delicately undulate, the apex acute to acuminate, the petiole 0.1–1.5 cm long, sometimes absent, the large leaf blades with (6) 8–11 primary veins on each side of the midvein. Flowers often solitary, sometimes in groups of 2–3, axillary, oriented horizontally to nodding; peduncles absent; pedicels 9–15 mm long and arching in flower, 12–20 mm long and arching in fruit, moderately to densely pubescent; calyx 2–2.5 mm long, 2.5–3 mm in diameter, obconic to narrowly campanulate, moderately pubescent, the margin truncate to undulate, with 5–10 narrow, linear, spreading appendages 0.5–2 mm long emerging 0.25–0.5 mm below the calyx rim; fruiting calyx slightly enlarged, widely bowl-shaped to plate-shaped, 1–2 mm long, 4–6 mm in diameter, the appendages withering in age; corolla 1–1.2 cm long, campanulate to reflexed in orientation, stellate in outline, divided nearly to the base, interpetalar tissue present near base, white, adaxial markings unknown, sparsely pubescent on abaxial surface along the midvein; stamens equal, straight, the filaments 0.75–1 mm long, glabrous, the anthers ca. 3 mm long, lanceolate, somewhat narrowed at the tip (the narrowed portion ca. 0.25 mm long), free of one another, color unknown, glabrous, poricidal at the tip, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style ca. 8 mm long, linear, glabrous, widened distally into the stigma; stigma capitate, decurrent down two sides. Fruit a berry, 3.2–8 mm long, 3.1–7 mm in diameter, globose, orange at maturity, glabrous, lacking sclerotic granules. Seeds 30–60 per fruit, 1–1.2 × 0.5–1 mm, compressed but not flat, sometimes with one shallow ridge, semi-circular, depressed ovate, triangular, or rhombic in outline, orange, the surface reticulum with serpentine pattern and shallow luminae.

Unknown.

Mexico (Querétaro, Veracruz) in tropical dry forest and cloud forest, including Quercus, Carpinus, and Liquidambar forest, in shady canyons and on slopes, sometimes on limestone, 1040–1450 m in elevation (Fig. 41).

Figure 41. 

Map of geographic distribution of L. glabripetala based on herbarium specimen data.

None known.

Flowering specimens have been collected in June; specimens with mature fruits have been collected in January, July, and October. The timing of the corolla movements is not known, but since the corollas on the specimens of this species are open, the flowers are probably open during the day, as in the morphologically similar Lycianthes amatitlanensis (Coult. & Donn.Sm.) Bitter.

Lycianthes glabripetala is a rarely collected species of endangered cloud forest habitat of central Mexico, represented by only six collections, two of which are from a protected area (Sierra Gorda). The EOO is 8,363.379 km², and the AOO is 24 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Vulnerable (VU).

Lycianthes glabripetala is an endemic Mexican species morphologically similar to L. amatitlanensis (of Mexico and Central America), L. inconspicua (of Central America), and L. inaequilatera (of Central and South America). Lycianthes glabripetala differs from those species in combining woolly curved trichomes on the abaxial side of the leaves, a relatively large corolla (to 1.2 cm long), nearly glabrous surfaces on the abaxial side of the corolla lobes, and a pedicel length of 9–15 mm in flower and 12–20 mm in fruit. Lycianthes amatitlanensis usually has straight trichomes that project at a 90-degree angle from the midvein of the abaxial leaf surface, corollas 0.5–0.8 cm long, and very evident long trichomes on the abaxial side of the corolla lobes with these trichomes usually tufted at the tip of the lobe. Lycianthes inconspicua Bitter can have flowers as long as L. glabripetala, and has variable pubescence on the abaxial side of the corolla lobes, but it has longer pedicels (15–30 mm in flower and 30–35 mm in fruit) and delicate straight trichomes that are tightly appressed to the midvein of the abaxial leaf surface; it also differs in having oval anthers. Lycianthes inaequilatera has smaller corollas and has pubescence much like L. inconspicua, and it occurs far south of the range of L. glabripetala (Dean et al. 2018b).

Lycianthes glabripetala is known at this time from the highlands of central Mexico in the states of Querétaro and Veracruz in cloud forest vegetation above 1000 m in elevation; this habitat is similar to that of L. inconspicua but differs from the most common habitat of L. amatitlanensis, a species that is usually found below 1000 m in elevation, often below 500 m, in humid tropical forest.

Mexico. Querétaro: 1 km al sureste de El Naranjo, [21.2421, -99.1014], 1050 m, 24 Jul 1989 H. Rubio 909 (IEB, DAV). Veracruz: Mpio. Zontecomatlán, along Huayacocotla-Zontecomatlán road, 1 km NE of San Antonio Ixtatetla, 20.7, -98.3833, 1300 m, 27 Apr 1983, M. Nee 26820 (NY, XAL).

Guatemala. Huehuetenango: Mpio. San Mateo Ixtatán, 4 miles east of San Mateo Ixtatán on road to Barillas, 8500 ft, 7 Feb 1965, D. Breedlove 8771 (holotype: F [0072910F, acc. # 1624724]; isotype: CAS [0003289]).

Figure 42. 

Image of herbarium specimen of L. gongylodes, Proctor 25432 (LL). Specimen used with permission from the Lundell Herbarium, University of Texas at Austin.

Herb to shrub, erect, 1.5–3.5 m tall. Indument of pale yellow to light brown, uniseriate, multicellular, usually simple (sometimes dendritic), curling to crisped, eglandular, spreading to appressed trichomes 0.1–0.75 (1) mm long. Stems green when young, sparsely to moderately pubescent, compressed 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 7–17.5 × 3.5–6.5 cm, the smaller ones with blades 3.5–10 × 2–4 cm, the leaf pairs usually similar in shape, the blades ovate to elliptic, membranaceous, sparsely pubescent, especially along the veins, the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 0.5–2.5 cm long, sometimes absent, the larger leaf blades with 5–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–5, axillary, erect or oriented horizontally; peduncles absent; pedicels 10–15 mm and erect to arching in flower, to 30 mm long and erect to arching in fruit; calyx 2–3 mm long, 3.5–4.5 mm in diameter, widely bowl shaped, glabrous to sparsely pubescent, the margin truncate, undulate, very well developed, with 10 very short, reflexed appendages 0.25–0.5 mm long emerging 1–1.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, sometimes appearing flat-bottomed, 1.5–3 mm long, 6–9 mm in diameter, the appendages not changing in length; corolla 0.6–1 cm long, rotate to reflexed in orientation, shallowly to deeply stellate in outline, sometimes divided to below the middle, interpetalar tissue present, white adaxially, glabrous, white abaxially, sparsely pubescent with very short trichomes; stamens equal, the filaments ca. 1 mm long, glabrous, the anthers 4–5 mm long, lanceolate, free of one another, pale yellow, glabrous, poricidal at the tips, the pores round, dehiscing distally; pistil with glabrous ovary, the style ca. 7 mm, linear, straight, glabrous, the stigma capitate, decurrent down two sides. Fruit a berry, 7–10 mm long, 7–9 mm in diameter, globose, orange at maturity, glabrous, lacking sclerotic granules. Seeds 20–60 per fruit, 2–2.5 mm × 1.5–2 mm, flattened, depressed ovate to oval in outline, with shallow notch on one side, yellow-orange, the surface reticulum with minute serpentine pattern with shallow luminae.

Unknown.

Guatemala (Huehuetenango, Quiché), in cloud forest, 2400–3000 m in elevation (Fig. 43).

Figure 43. 

Map of geographic distribution of L. gongylodes based on herbarium specimen data.

None known.

Flowering specimens and specimens with mature fruits have been collected in February and from June to August. All flowering specimens of this species have open flowers indicating that the flowers are probably open for most of the day.

Lycianthes gongylodes is a rarely collected species of Guatemala, represented by only four collections, all made before 1970, and none from protected areas. The EOO is 43.078 km², and the AOO is 12 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Critically Endangered (CR).

Lycianthes gongylodes is known from only four collections made by Breedlove, Proctor, and Steyermark. The species is most likely allied to L. heteroclita, with which it shares green herbaceous stems that collapse upon drying. The form of the calyx in the two species is similar, although L. gongylodes differs in having small calyx appendages which make the calyx appear thicker and more bowl-shaped in flower and flat-bottomed in fruit (a feature it shares with L. connata). Lycianthes heteroclita usually lacks appendages on the calyx, which looks campanulate in flower and bowl-shaped to plate-like in fruit and, unlike L. gongylodes, it usually lacks obvious interpetalar tissue connecting the lobes of the corolla. The curly trichomes that are present on the stems and leaves of L. gongylodes are quite distinctive and different than the very small trichomes present in L. heteroclita; in addition, L. gongylodes lacks the tiny groups of white trichomes that appear like small granules on the calyx of L. heteroclita. Lycianthes gongylodes could be confused with L. ceratocalycia, another allied species that occurs in the same region of Guatemala and adjacent Mexico. Lycianthes ceratocalycia differs in having purple, stellate corollas with sparse interpetalar tissue and scurfy horizontal lines on the young branches. A sterile specimen of L. gongylodes could be misidentified as L. tricolor (Dunal) Bitter, something that was done in Dean et al. (2017a), where Steyermark 49839 (a paratype of L. gongylodes) is cited under L. tricolor.

Guatemala. Huehuetenango: Sierra de los Cuchumatanes, near the place called Kurus Lemun, 4 miles E of San Mateo Ixtatán along road to Barillas, [15.82, -91.4264], 8500 ft, 7 Aug 1965, D. Breedlove 11628 (TEX). Quiché: El Boquerón, 8000–8200 ft, 6 Aug 1964, G.R. Proctor 25432 (BRIT, MO, TEX).

Guatemala. Sacluc, Aug 1877, K. Bernoulli & O. Cario 2357 (holotype: GOET [GOET003446]).

Figure 44. 

Image of herbarium specimen of L. gorgonea, Dean 9528 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Shrub to woody vine, 1–4 m tall. Indument of white to tan, uniseriate, multicellular, simple, eglandular and/or glandular, spreading trichomes 0.5–3 mm long. Stems greenish-tan when young, moderately to densely pubescent, not compressed when dried in a plant press, becoming woody with age; upper sympodial branching points a mixture of dichasial and monochasial, the branching divaricate and zigzagging. Leaves simple, the leaves of the upper sympodia usually paired, the leaf pairs often conspicuously different in size and shape, the larger ones with blades 4–11 × 2–4 cm, ovate to lanceolate, the smaller ones with blades 0.5–4 × 0.5–2 cm, orbicular to ovate, the leaf pairs similar in texture, membranaceous, moderately pubescent (densely pubescent along the midvein of the abaxial side), the base cuneate to rounded, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole to 0.5 (1) cm long, sometimes absent, the larger leaf blades with 5–7 primary veins on each side of the midvein. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 15–25 mm and erect to arching in flower, to 40 mm long and spreading in fruit, densely pubescent; calyx 2–2.5 mm long, 3–3.5 mm in diameter, obconic to campanulate, densely pubescent (sometimes nearly obscured), the margin truncate, with 10 very long spreading linear appendages 7–15 mm long emerging 0.5 mm below the calyx rim; fruiting calyx usually enlarged, widely campanulate to bowl-shaped, 2.5–5 mm long, 6–9 mm in diameter, the appendages to 20 mm long, spreading; corolla 1–1.5 cm long, rotate in orientation, entire in outline, with abundant interpetalar tissue, white to pale blue-violet, glabrous adaxially, with long trichomes near the veins abaxially; stamens equal or nearly so, straight, the filaments 1–2 mm long, glabrous, the anthers 4–4.5 mm long, elliptic, connate to one another at their edges, yellow, glabrous, poricidal at the tips, the pores round, large, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–9 mm long, linear, straight to curved, glabrous; stigma capitate, decurrent down two sides. Fruit a berry, 5–8 mm long, 5–9 mm in diameter, globose, red at maturity, glabrous, lacking sclerotic granules. Seeds 15–25 per fruit, 2.25–2.5 × 2.5–3 mm, flattened to unevenly thickened and curved, triangular to depressed ovate in outline, yellow-orange, the surface reticulum with minute serpentine pattern and shallow luminae.

Unknown.

Mexico (Chiapas, Oaxaca, Tabasco, Veracruz), Guatemala (Alta Verapaz, Petén), and Belize (El Cayo), in high forest, lower montane rain forest, and tropical moist forest, often on limestone ridges or in canyons, sometimes near streams, 200–1000 m in elevation (Fig. 45).

Figure 45. 

Map of geographic distribution of L. gorgonea based on herbarium specimen data.

None known.

Flowering specimens have been collected November through August. Specimens with mature fruits have been collected April and July through November. Many specimens have been collected with immature fruits, and these have been collected throughout the year. In Belize, the corollas of this species open at sunrise and close at sunset (Smith and Knapp 2002).

Lycianthes gorgonea is a species of lowlands of southern Mexico, Guatemala, and Belize, represented by 31 collections and occurring in six protected areas. The EOO is 79,973.926 km², and the AOO is 108 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Lycianthes gorgonea is a distinctive species of lower elevation tropical rain forest, often found on limestone. The divaricate, zigzag branching of this species, in combination with a distinctive size/shape difference of the paired geminate leaves, soft long, pale trichomes, and very long calyx appendages, is quite different from any other species of Lycianthes in Mexico and Central America. The pollination of this species was studied in Belize by Smith and Knapp (2002), and they found that the flowers are visited by the bee genus Paratetrapedia.

Guatemala. Alta Verapaz: between Limón and Chisec, 200–230 m, 19 Mar 1942, J.A. Steyermark 45116 (NY, LL). Petén: Uaxactun, on Dos Lagunas road, in zapotal/ramonal, 3 km W, 22 Jan 1977, C. L. Lundell 20534 (MO, LL). Mexico. Chiapas: Mpio. Barriozábal, along road from Berriozábal to Las Maravillas, ca. 1.4 km S of the town of Efraín A. Gutiérrez, in remnant of tall forest called La Mata Café, 16.8711, -93.2956, 1005 m, 12 Sep 2017, E. Dean 9528 (DAV). Oaxaca: Dto. Tuxtepec, predio La Joya del Obispo, [17.8599, -96.2060], 12 Aug 1990, C. H. Ramos 435 (IEB, MEXU, XAL). Tabasco: ladera W del Cerro del Madrigal, cerca de la base, Puyacatengo, [17.5213, -92.9225], 25 Mar 1992, M. A. Guadarrama-O. 1244 (TEX). Veracruz: Mpio. Minatitlán, 6.6 km al Norte de la terracería La Laguna-Río Grande, sobre el camino nuevo (no completo) a Ejido Belisario Domínguez, el cual sale de la terracería 14.7 km al E. de La Laguna, 17.3333, -94.3667, 130 m, 13 Jul 1980, T. Wendt 2557 (MO).

Mexico. Chiapas: [Mpio. Siltepec?] Letrero, near Siltepec [15.5564, -92.3233], 2000 m, 6 Jul 1941, E. Matuda 4350 (holotype: MEXU [acc. # 82114]; isotypes: A [00934887], [LL [00226934, 00227071], MO [acc. # 1244040]).

Figure 46. 

Image of herbarium specimen of L. grandifolia, Heath 1012 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Shrub, 2–3 m tall, erect. Indument of light yellow (sometimes appearing tan or off-white), uniseriate, multicellular, simple, eglandular, ascending-appressed to spreading trichomes 0.1–1.5 mm long. Stems green, angled when young, sparsely to moderately pubescent, somewhat compressed and ribbed when dried in a plant press, becoming 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 8–22 × 5–12 cm, the smaller ones with blades 3–9 × 2–4.5 cm, the leaf pairs similar in shape, the blades ovate (often widely so), thin chartaceous, sparsely pubescent, ciliate along the margin, the base usually long attenuate (cuneate), sometimes oblique, the margin entire, usually irregularly undulate, the apex acuminate, the petiole 0.2–5 cm long, those of the longer leaves 2 cm long or longer, the larger leaf blades with 5–6 primary veins on each side of the midvein. Flowers in groups of 2–8, axillary, oriented horizontally; peduncles absent; pedicels 10–25 mm long and erect in flower, sparsely to moderately pubescent, mature fruiting pedicels not yet seen; calyx 2.5–3.5 mm long, 3–4 mm in diameter, obconic to campanulate, sparsely to moderately pubescent, the margin truncate, with 10 spreading linear appendages 2–7.5 mm long (at least some appendages on a calyx 4 mm long or longer), emerging 0.25–0.5 mm below the calyx rim; mature fruiting calyx not yet seen; corolla 0.8–1.6 cm long, rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white, glabrous adaxially, the abaxial side of the lobes moderately puberulent near the major veins; stamens unequal, straight, the four short filaments 1–2 mm long, the one long filament 3–4.5 mm long, glabrous, the anthers 3–4 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 dehiscing distally or away from the style, not opening into longitudinal slits; pistil with glabrous ovary, the style ca. 7 mm long, linear, glabrous, the stigma capitate, decurrent down two sides, slightly lobed. Fruit a berry, 4–5 mm long, 4–5 mm in diameter, globose to depressed-globose, green when immature (mature fruit not yet seen), glabrous, lacking sclerotic granules. Seeds 10–30 per fruit, only seen when immature, mature size and details not yet known, not notched.

Unknown.

Mexico (Chiapas), in cloud forest, 1700–2000 m in elevation (Fig. 47).

Figure 47. 

Map of geographic distribution of L. grandifolia based on herbarium specimen data.

None known.

Flowering specimens have been collected in June and July, presumably fruiting after that. The corollas on the specimens of this species are usually closed, indicating that the corollas may just open very early in the morning and then close by late morning.

Lycianthes grandifolia is a rarely collected species of southern Mexico, represented by only five collections, two from the protected area El Triunfo. The EOO is 828.005.215 km², and the AOO is 16 km². Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

This species belongs to series Tricolores and is distinguished from most of the other species in the series by long and broad leaves and occurring in cloud forest at elevations close to 2000 m. Every specimen of this species that we have examined has at least one leaf with a length greater than or equal to 18 cm. Other species within series Tricolores with leaves that can reach this length (but not typically) are L. arrazolensis, L. jalicensis, and L. michaelneei. Lycianthes grandifolia differs from L. michaelneei in having white corollas (rather than purple) and having less dense pubescence. It differs from L. arrazolensis by usually having longer calyx appendages that are inserted < 0.5 mm from the calyx rim (versus > 0.5 mm). It differs from L. jalicensis in having pubescent calyces and corollas and occurring at elevations near 2000 m (versus < 1400 m) (Dean et al. 2018c).

Lycianthes grandifolia occurs in habitats similar to those of L. tricolor (high elevation forest types), and it has pedicels of similar length. It differs from L. tricolor by longer leaves and angled and ribbed young stems that compress when dried in a plant press. We have not seen the mature fruits or seeds of L. grandifolia, however the immature seeds resemble those of L. arrazolensis, which are unnotched, rather than those of L. tricolor, which are notched (Dean et al. 2018c).

This species is poorly known and has been rarely collected. One collection from southeastern Chiapas that may belong to L. grandifolia: Mt. Pasitar (Paxtar), 3–4 Aug 1937, Matuda 1642 (MO-1807983; US-1807982) differs in having the calyx appendages densely pubescent with long, spreading trichomes (2 mm long). Therefore, we did not include it in the species description. The elevation and exact location of this collection is unknown. If from low elevations, it may represent a new species belonging to series Tricolores. More fieldwork is necessary to fill out the morphological details of L. grandifolia (Dean et al. 2018c)

Mexico. Chiapas: Mpio. Mapastepec, Reserva El Triunfo, Deslave-Cipresal, 15.65, -92.8, 1750 m, 14 Jun 1990, M. Heath 1012 (MO3801177).

Based on Solanum heteroclitum Sendtn.

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.

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.

Unknown.

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).

Figure 49. 

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

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).

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.

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).

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.

Guatemala. Alta Verapaz: city of Cobán, garden of residence along 1^st 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).

Mexico. Nuevo León: Mpio. Aramberri, Cerro El Viejo, 1200 m, 28 Jul1993, Hinton et al. 22882 (holotype: DAV [DAV155244]; isotypes: CIIDIR [CIIDIR022490]; GBH [GBH022882]; TEX [00208091]).

Figure 50. 

Image of herbarium specimen of L. hintonii, Hinton 23263 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Perennial herb from storage roots of unknown shape, usually erect, to ca. 0.5 m tall, dying back each season. Indument of white, uniseriate, multicellular, simple (rarely forked or dendritically branched), eglandular, spreading to appressed-retrorse trichomes 0.1–1 mm long. Stems green with darker green vertical markings, sparsely to moderately pubescent, somewhat compressed upon drying in a plant press, somewhat woody with age, especially at the base of the plant; first stem ca. 25 cm long to the first inflorescence, the internodes ca. 13; first two sympodial branching points dichasial, followed by monochasial branching. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades 8–15 × 5.5–6.5 cm, the smaller ones with blades 1/2 to 2/3 the size of the larger, the leaf pairs similar in shape, the blades deltoid, ovate, or elliptic, thin chartaceous, sparsely pubescent, the primary veins 4–5 on either side of the midvein, the base attenuate or decurrent onto the petiole, slightly oblique on smaller leaves, the margin entire, and usually irregularly undulate, the apex rounded, acute, or short-acuminate, the petioles poorly defined, 1–3 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 40–70 mm and erect in flower, ca. 110 mm long (or longer) in fruit (material with mature fruits and fruiting pedicels not yet seen), sparsely pubescent with spreading to appressed trichomes; calyx 3–4 mm long, 4–5 mm in diameter, campanulate, sparsely pubescent, the margin truncate, with 10 linear, spreading to reflexed appendages 4–11 mm long emerging ca. 1 mm below the calyx rim; fruiting calyx not yet seen; corolla 1–2.5 cm long (ca. 2–4 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white, green near the major veins abaxially, glabrous; stamens unequal, straight, the filaments of three lengths, the two shortest filaments 1.5–3.5 mm long, the two medium filaments 2.5–4 mm long, the one long filament 3–6 mm long, the length of the long filament nearly always 1.2–1.5 times that of the medium filament, glabrous, the anthers 4.5–6 mm long, ovate-lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pollen grains tricolporate; pistil with glabrous ovary, the style 8.5–11 mm long, linear, slightly curved, glabrous, the stigma capitate (sometimes weakly bilobed). Fruits and seeds not yet seen.

Unknown.

Mexico (Nuevo León), in oak forests on limestone soils in the mountains in the vicinity of Cerro El Viejo, 1200–1500 m in elevation (Fig. 51).

Figure 51. 

Map of geographic distribution of L. hintonii based on herbarium specimen data.

None known.

Flowering specimens have been collected July to August. Fruit not yet seen. The diurnal movements of the corolla have not been observed in the field; the corollas are probably open in the early morning and closed in the late morning. Unlike the other herbs of series Meizonodonatae, the scent of the pollen of this species is unknown.

Lycianthes hintonii is a rarely collected species of northern Mexico, represented by only two collections made before 1993, both from the same location (Cerro El Viejo, Nuevo León), which is not a protected area. Anguiano-Constante et al. (2018) provided a preliminary assessment of Critically Endangered (CR).

Although this species has not been observed in the field, it is obviously related to the species of series Meizonodontae and it is assumed that it has the characteristic tuberous roots. The fruit type is unknown and could be either green, like Lycianthes moziniana, or dark purple like L. ciliolata. This species is similar to L. rzedowskii in its white flowers, but it differs from that species in having fewer, larger leaves on the first stem to emerge from the ground, triangular, tricolporate pollen, and in growing in basic limestone soils. Its distribution is quite disjunct from the other populations of L. rzedowskii (Dean 2004).

Mexico. Nuevo León: Mpio. Aramberri, Cerro El Viejo, [23.9885, -99.7612], 1495 m, 3 Aug 1993, Hinton 23263 (DAV, GBH, TEX, UC).

Belize. Orange Walk: 10 Oct 1926, H. W. Winzerling V-14 (holotype: US [00027883]; isotypes: F [0072913F, acc. # 573777]), G [G00379122], WIS [00000961MAD]).