Corresponding author: Ellen Dean (
Academic editor: Sandy Knapp
Dean E, Poore J, Anguiano-Constante MA, Nee MH, Kang H, Starbuck T, Rodrígues A, Conner M (2020) The genus
The only taxonomic monograph of
Georg
The Swiss botanist Emil Hassler raised
Two names for the concept of
Since Bitter’s monograph, many additional New World
Morphological revisions have been completed on five of Bitter’s
To date, a phylogeny of the entire genus has not yet been published, although work on this project using molecular markers is ongoing at the laboratory of Lynn Bohs at the University of Utah, and preliminary results are completed. Research into the evolution of
Herbarium collections of many of the species of
Over the past four years, the first author has been investigating the
Geographic distribution of the native
Species | Distribution |
---|---|
Mexico to Costa Rica | |
Mexico to South America | |
Mexico | |
Mexico, Guatemala, Belize | |
Mexico to Nicaragua | |
Mexico, Guatemala | |
Mexico | |
Mexico | |
Mexico, Guatemala | |
|
Mexico, Guatemala |
Mexico to Nicaragua | |
Mexico, Guatemala | |
Mexico, Guatemala | |
Guatemala | |
Mexico | |
Guatemala | |
Mexico | |
Mexico | |
Guatemala | |
Mexico, Guatemala, Belize | |
Mexico | |
Mexico to South America | |
Mexico | |
Mexico to Honduras | |
Guatemala to Panama | |
Mexico | |
Mexico, Guatemala, Belize | |
Mexico to El Salvador | |
Mexico to Nicaragua | |
Mexico | |
Mexico | |
|
Mexico |
Mexico | |
Mexico to Panama | |
Mexico and Guatemala | |
Mexico and Guatemala | |
Mexico | |
Mexico | |
Mexico | |
Mexico to Honduras | |
Mexico and Guatemala | |
Mexico | |
Mexico | |
Mexico to Costa Rica | |
|
Mexico to South America and the Caribbean |
Mexico to Nicargua | |
Mexico | |
Mexico to Honduras | |
Mexico | |
Mexico | |
Mexico to El Salvador | |
Mexico |
The circumscriptions of the species treated here are based on examination of herbarium specimens, cultivated plants, and field observations and are supported by morphological evidence. We examined specimens from the following herbaria either in person or as images: A, ANSM, ARIZ, ASU, BIGU, BRIT, BM, BR, BREM, C, CAS, CIIDIR,
Throughout this work, type specimens with a known barcode number are cited with the herbarium code followed by the number (for example: holotype: P [P00070402]). In cases where the specimen has no barcode number (or the herbarium wishes the accession number to be used instead, for example F), the accession number is provided (preceded by “acc. #”). If no number is cited for a type specimen, none was provided on the specimens or was unavailable to the authors. Nearly all type specimens cited were examined as either a digital photo or in person by one of the authors; therefore, herbarium codes are not followed with an exclamation mark. If a specimen was not seen by us, we indicate this. When lectotypes are designated in the nomenclature section of a species treatment, remarks justifying the choice of the lectotype are included in the commentary section of that same treatment.
Specimens examined are listed in Appendix
In order to create maps of the Mexican species, specimens were georeferenced by using either Geolocate, an online software-mapping package (
The georeferenced location data were analyzed in order to provide conservation assessments for each species using GeoCAT (
Our species concept is a morphological one (
The sympodial branching pattern of
Trichome type is important in the identification of many
Furcate (forked) trichomes (Fig.
In some species with simple trichomes, the cells collapse and flatten upon drying, appearing like a flattened ribbon (Fig.
Only four species included here have glandular trichomes (Fig.
The upper sympodial units of the
The sympodial units of
The pedicels of the flowers can be erect, ascending, spreading, or reflexed. It is common for the pedicel to be recurved at the tip as the flower is developing (this is the case in the herbs of series
In all
The corolla in
The corollas of
The color of the corollas of
Stamens in
Anthers can be connate, connivent, or free. They can be glabrous or pubescent; most of the
The pollen of most
The gynoecium in the
The fruits of the
Seed shape terminology used in this paper is taken from
The color of the seeds of the species included here can be tan, yellow, orange, brown, or black. The seed reticulum (surface) has a serpentine pattern formed by the periclinal walls of the surface cells; the luminae of the cells can be shallow to deep. Often, lenticular seeds have a smooth center with an indistinct serpentine pattern and cells with shallow luminae; however, the thickened margin of these seeds have a reticulum with much deeper luminae. Seeds that are more three-dimensional often have a reticulum with a more pronounced serpentine pattern and cells with much deeper luminae. As discussed in previous papers (
Twenty-two taxa of
The preliminary conservation assessment for each species is discussed under each species treatment. The conservation assessments range from Least Concern for common and widespread species to Critically Endangered for species that are only known from one location (Table
Preliminary conservation assessments for the native
Species | Assessment |
---|---|
|
Least Concern |
|
Least Concern |
|
Endangered |
|
Least Concern |
|
Least Concern |
|
Endangered |
|
Endangered |
|
Endangered |
|
Endangered |
|
Endangered |
|
Least Concern |
|
Least Concern |
|
Least Concern |
|
Critically Endangered |
|
Least Concern |
|
Endangered |
|
Near Threatened |
|
Vulnerable |
|
Critically Endangered |
|
Least Concern |
|
Endangered |
|
Least Concern |
|
Critically Endangered |
|
Least Concern |
|
Endangered |
|
Vulnerable |
|
Least Concern |
|
Least Concern |
|
Least Concern |
|
Endangered |
|
Near Threatened |
|
Least Concern |
|
Vulnerable |
|
Least Concern |
|
Endangered |
|
Endangered |
|
Least Concern |
|
Endangered |
|
Endangered |
|
Least Concern |
|
Endangered |
|
Least Concern |
|
Endangered |
|
Least Concern |
|
Least Concern |
|
Least Concern |
|
Endangered |
|
Least Concern |
|
Least Concern |
|
Critically Endangered |
|
Least Concern |
|
Endangered |
Based on
Perennial herbs (from stolons, rhizomes, or tuberous roots), shrubs, or vines, sometimes epiphytic. Pubescence of glandular or eglandular, simple, dendritic, or stellate trichomes. Stems with sympodial growth. Leaves alternate, geminate, or solitary, simple, usually entire, usually petiolate, the base often unequal, the leaf pairs sometimes anysophyllous. Inflorescences axillary, the peduncles very short or absent, with one to many pedicelled flowers; calyx with truncate rim, often enlarging in fruit, 10-nerved, often with five to ten (25) appendages protruding from the calyx below the margin; corolla with five lobes, the lobes often connected by interpetalar tissue, the shape entire to stellate, opening and closing daily for several days in a row (sometimes opening only in the very early morning), campanulate, rotate or reflexed when open, white to purple or blue; stamens inserted near the base of the corolla, the filaments equal or not, the anthers free, connivent, or connate, dehiscing by pores (rarely lengthwise); pistil 2-carpellate, ovary spherical, ovoid, or conical, style straight or curved, stigma capitate to oblong, entire to lobed, ovules usually numerous; fruit a berry, round to ovoid, the exocarp purple, red, orange, yellow, or green, some species with sclerotic granules in the outer part of the mesocarp; seeds usually numerous, lenticular compressed to round or angular in outline, tan, yellow, orange, brown or black.
Genus name based on the type species
Key to the Groups
1 | Herbaceous perennial from tuberous roots, dying back to the ground each season; inflorescence one flowered; corollas rotate to reflexed in orientation, mostly entire (not stellate) in outline; stamens unequal |
|
– | Herbaceous perennial from rhizomes, subshrubs, shrubs, epiphytes, or vines, generally persisting above ground from one season to the next; inflorescence of one to many flowers; corollas campanulate, rotate, or reflexed in orientation, entire to stellate in outline; stamens equal or unequal |
|
2 | Plant with obvious glandular trichomes (to 1 mm long or more), at least on the pedicels and/or calyx, sometimes also on the leaves and stem |
|
– | Plant lacking obvious glandular trichomes, sometimes with small obscure glandular trichomes inside the calyx or elsewhere |
|
3 | Simple, furcate and/or dendritic trichomes forming a majority of the indument on stems and leaves, or plant completely glabrous |
|
– | Multangulate- or geminate-stellate trichomes (with more than two rays at a node, the rays sometimes rebranched) forming a majority of the indument on stems and leaves (simple or dendritic trichomes may also be present) |
|
Herbaceous perennial from tuberous roots, dying back to the ground each season; inflorescence one flowered; corollas rotate to reflexed in orientation, mostly entire (not stellate) in outline; stamens unequal.
1 | Indument of multangulate stellate trichomes mixed with dendritic and simple trichomes, the dendritic trichomes usually up to 0.5 mm long; species of arid habitats |
|
– | Indument of simple and/or dendritic trichomes (not stellate), if present, the dendritic trichomes often greater than 0.5 mm long; species of both arid and non-arid habitats |
|
2 | Plant prostrate (rarely decumbent or ascending); berry with yellow sclerotic granules in the mesocarp; style often strongly curved downward; ovary rounded to ovoid, less than 2 mm long; indument of simple, antrorsely appressed trichomes |
|
– | Plants prostrate or not; berry without sclerotic granules; style straight to slightly curved, never strongly curved downward; ovary conical, usually longer than 2 mm (sometimes shorter in |
|
3 | Corolla white, with or without maroon to purple nectar guides; corolla lobes usually glabrous abaxially; plant body usually erect, with first sympodial unit well developed above the ground; first two branching points on the plant body usually dichasial |
|
– | Corolla lilac, violet or light purple, with maroon to purple nectar guides; corolla lobes glabrous or pubescent abaxially; plant body erect, decumbent or prostrate with first sympodial unit sometimes not well developed above ground; first branching point on the plant body dichasial, second sympodial branching point sometimes monochasial |
|
4 | Filament of the longest stamen usually more than twice as long as those of the lateral stamens; pores of the anther of the longest stamen lateral, dehiscing toward the style, usually narrow and linear; stigmas usually deeply bilobed (rarely just capitate); widespread species of the transvolcanic belt, southern Mexico and Central America |
|
– | Filament of the longest stamen usually less than twice as long as those of the lateral stamens; pores of the anther of the longest stamen nearly terminal, oval; stigmas not deeply bilobed; species of various regions of Mexico and Guatemala |
|
5 | First sympodial unit usually very well developed (to 90 cm long) with numerous internodes (usually 10–21) and leaves; lateral branching from the nodes of the first sympodial unit usually not present at the time of flowering; subsequent sympodial growth poorly developed; pollen dicolporate; states of Morelos, Michoacán and México, on volcanic soils |
|
– | First sympodial unit |
|
6 | Berry green to tan at maturity, sometimes with purple blotches; seeds less than 3 mm long, smooth and shiny to the naked eye; second sympodial branching point usually monochasial |
|
– | Berry purple to black-purple at maturity; seeds greater than 3 mm long, rough-textured and dull to the naked eye; second sympodial branching point usually dichasial |
|
7 | Calyx teeth in flower lax, laying against the corolla; calyx teeth in fruit appressed to the berry, not spreading; abaxial side of corolla lobes densely hairy; leaves of first sympodial unit cuneate at base, not attenuate; leaves sessile or petiole less than 5 mm long; agricultural areas of the transvolcanic belt |
|
– | Calyx teeth in flower slightly spreading; calyx teeth in fruit spreading, not appressed to the berry; abaxial side of the corolla lobes slightly hairy to nearly glabrous (glabrous in Nuevo Leon); leaves of first sympodial unit attenuate at base; petiole to 1.5 cm long; not of the region of the transvolcanic belt (states of San Luis Potosi, Nuevo Leon, Oaxaca) often growing on limestone soil |
|
8 | Filaments glabrous; exocarp of berry green; placental area green and juicy; disturbed clearings and agricultural fields of mountains of state of Oaxaca |
|
– | Filaments glabrous or pubescent; exocarp of berry green or tan, often with purple blotches; placental area often purplish and powdery; forested areas or more disturbed situations on limestone in northern Mexico (states of San Luis Potosí and Nuevo León) |
|
9 | Calyx teeth at anthesis lax or slightly spreading, not widely spreading or reflexed; corolla lobes usually noticeably pubescent abaxially; first sympodial unit poorly developed above ground, the plant body often prostrate; rare species of the Sierra de Nanchititla (state of México) |
|
– | Calyx teeth at anthesis widely spreading to reflexed; corolla lobes generally glabrous abaxially (rarely slightly pubescent in Oaxacan plants); first sympodial unit well-developed above ground, the plant body usually erect; widespread in SE Mexico and Guatemala |
|
Plants with obvious glandular trichomes (to 1 mm long or more), at least on the pedicels and/or calyx, sometimes also on the leaves and stem
1 | Calyx appendages less than 2 mm long, or reduced to small protuberances, the calyx sometimes tearing, appearing lobed or two-lipped; berry ovoid, orange |
|
– | Calyx appendages usually greater than 2 mm long, the calyx margin truncate, never appearing lobed or two-lipped; berry turbinate, globose, or depressed globose, orange to red |
|
2 | Corolla purple; berry turbinate, the tip pointed, orange; seed surface with widely spaced serpentine cell pattern, noticeably pitted; state of Oaxaca |
|
– | Corolla white to lilac, sometimes with purple or green markings; berry globose to depressed globose, the tip round, orange to red; seed surface finely marked with shallow serpentine cell pattern, not noticeably pitted; widely distributed |
|
3 | Leaf pairs of similar shape; inflorescence of 1–5 flowers; calyx appendages in flower 2–10 mm long, to 11 mm long in fruit; stamens unequal with one filament noticeably longer than the other four, the anthers free of one another; from state of Sinaloa to Oaxaca, 670–2200 m in elevation |
|
– | Leaf pairs usually very different in shape, the smaller leaf blade orbicular to ovate, the larger leaf blade narrowly ovate to lanceolate; inflorescence of a single flower; calyx appendages in flower 7–15 mm long, to 20 mm long in fruit; stamens equal or nearly so, the filaments of approximately the same length, the anthers connate to one another at their edges; Mexico (states of Chiapas, Oaxaca, Tabasco, and Veracruz) to Belize and Guatemala, 200–1000 m in elevation |
|
Plants with simple, furcate and/or dendritic trichomes forming a majority of the indument on stems and leaves, or plant completely glabrous
1 | Shrub; stems angular or with prominent striations; corolla dark purple; berries often not developing, the exocarp yellow to light orange when mature, the mesocarp with many sclerotic granules; cultivated plants |
|
– | Shrub or other life form; stems angular and striated or not; corolla white to light purple, not usually dark purple; berries often developing, the exocarp yellow, orange, red, or dark purple when mature, the mesocarp lacking sclerotic granules; non-cultivated plants |
|
2 | Leaf blades glabrous except for trichomes restricted to the axils where the primary veins meet the midvein on the abaxial side; stamens usually equal in length |
|
– | Leaf blades glabrous to pubescent, but trichomes not restricted to the axils where the primary veins meet the midvein on the abaxial side; stamens equal or unequal in length |
|
3 | Shrub or vine, often epiphytic; calyx coriaceous and fleshy, the appendages 0.25–1 mm long, connate at their bases, reflexed as a unit; corolla stellate in outline, blue to purple adaxially; anthers connate at their edges |
|
– | Shrub, terrestrial; calyx membranaceous, not fleshy, the appendages 0.5 to 2 mm long, free at their bases and not reflexed; corolla entire to very slightly stellate in outline, white with lavender ring at the base adaxially; anthers free at their edges |
|
4 | Stamens equal in length or nearly so |
|
– | Stamens unequal in length, one filament noticeably longer than the other four |
|
5 | Calyx lacking appendages; corolla shallowly to deeply stellate in outline, sometimes without interpetalar tissue |
|
– | Calyx with appendages 0.25–20 mm long; corolla entire to stellate in outline, with interpetalar tissue present at least at the base of the corolla lobes |
|
6 | Calyx margin often irregularly notched or torn in flower and/or fruit, sometimes appearing lobed; corolla stellate, divided 1/2 to nearly all the way to the base, with abundant interpetalar tissue; anthers often with dark connective |
|
– | Calyx margin entire, truncate, in flower and fruit; corolla stellate, deeply divided to the base, lacking interpetalar tissue; anthers without dark connective |
|
7 | Upper stem nodes remaining round in cross-section upon drying, quickly becoming woody; leaves completely glabrous, coriaceous, the geminate leaf pairs very different in size and shape, the smaller leaf usually 1/4 or less the length of the larger and round to ovate, the larger leaf narrowly ovate to lanceolate; anthers 4 mm long or more and connivent to one another at their edges; woody shrub to vine, usually epiphytic |
|
– | Upper stem nodes compressed upon drying, lower stems sometimes woody; leaves glabrous to short pubescent, membranaceous, the geminate leaf pairs unequal in size but similar in shape; anthers less than 4 mm long and free of one another; large herb to shrub, sometimes epiphytic |
|
8 | Calyx up to 1.5 mm long; corolla 0.6–1.2 cm long; anthers 3–3.5 mm long; endemic to Mexico (states of Hidalgo, Oaxaca, Puebla, and Veracruz), usually over 800 m in elevation |
|
– | Calyx 2 mm long or more; corolla 1–1.6 cm long; anthers 4–7 mm long; widespread in Mexico and Central America, usually up to 1000 m in elevation |
|
9 | Upper stem epidermis with rough texture formed by scurfy horizontal lines; corolla deeply stellate in outline, divided 3/4 of the way to the base, lilac to purple adaxially, the interpetalar tissue only present below the middle of the lobes; Mexico (state of Chiapas) and Guatemala, in cloud forest, usually above 1300 m in elevation |
|
– | Upper stem epidermis smooth, angular or roughened by vertical lenticels, but without scurfy horizontal lines; corolla deeply stellate to rotate in outline, white, pale yellow, or purple adaxially, the sometimes abundant interpetalar tissue not restricted to the lower part of the lobes; widely distributed in Mexico and Central America in various habitats above or below 1300 m in elevation |
|
10 | Number of veins on either side of the leaf blade midvein of largest leaves usually 8 or more, the blades moderately to densely pubescent, the blade base very oblique; calyx teeth in flower up to 4 mm long, very narrow (less than 0.25 mm wide), often withering in fruit; corollas white to pale yellow, stellate in outline, divided 1/2 to nearly all the way to the base; anthers free of one another, up to 3 mm long, abruptly attenuate at the tip; berry globose, orange-red at maturity |
|
– | Number of veins on either side of the leaf blade midvein of largest leaves usually less than 8 (if 8, the blades glabrous), the blades glabrous to pubescent, the blade base not oblique to somewhat oblique; calyx teeth in flower up to 15 mm long, not very narrow (greater than 0.25 mm wide), not withering in fruit; corollas white to purple, stellate to rotate in outline; anthers free of one another, connivent, or connate, usually greater than 3 mm long, not truncate, rounded, or acute at the tip, not abruptly attenuate; berry globose to ovoid, orange-red or dark purple at maturity |
|
11 | Number of veins on either side of the larger leaf blades usually 10–22, the blade trichomes usually spreading along the midvein on the abaxial side; corolla 0.5–0.8 cm long, the lobes moderately pubescent abaxially, with a tuft of trichomes at the tip; Mexico (southern Veracruz) to Central America, often below 1000 m in elevation |
|
– | Number of veins on either side of the larger leaf blades usually 8–12, the blade trichomes appressed along the midvein on the abaxial side; corolla 0.8–1.2 cm long, the lobes glabrous to sparsely pubescent abaxially, lacking tuft of trichomes at the tip; Mexico (states of Querétaro and Veracruz), Guatemala, and Central America usually above 1000 m in elevation |
|
12 | Trichomes along midvein on abaxial leaf blade surface bent to wavy, appearing woolly; pedicels in flower 9–15 mm long, in fruit 12–20 mm long; corolla 1–1.2 cm long, nearly glabrous abaxially except for sparse hairs near the lobe tip; endemic to Mexico (northern state of Veracruz to Querétaro) |
|
– | Trichomes along midvein on abaxial leaf blade surface mostly straight and appressed, not appearing woolly; pedicels in flower (11) 15–30 mm long, in fruit 30–36 mm long; corolla 0.5–1 cm long, sparsely pubescent abaxially, densest near the lobe tip; Guatemala to Panama |
|
13 | Perennial herb to woody vine or shrub; mature berry red; anthers connate to connivent; usually occurring at or below 1000 m in elevation |
|
– | Herb, shrub or treelet; mature berry orange, red or dark purple; anthers free from one another; usually occurring above 1000 m in elevation |
|
14 | Rhizomatous herb to climbing shrub; plant glabrous to sparsely pubescent, the appressed to ascending trichomes to 0.6 mm long, eglandular; calyx appendages 1.5–5 mm long in flower, to 8 mm long in fruit |
|
– | Weak shrub to vine; plant moderately to densely pubescent, the spreading trichomes to 3 mm long, sometimes glandular; calyx appendages 7–15 mm long in flower, to 20 mm long in fruit |
|
15 | Leaf blades glabrous and shiny on both sides, rarely with a few appressed-ascending trichomes to 0.25 mm long; corolla stellate in outline, adaxially white with yellow-green or purple markings near the base; anthers yellow, the connective usually dark in color |
|
– | Leaf blades glabrous to moderately pubescent with appressed to spreading trichomes to 1.25 mm long; corolla entire to stellate in outline, adaxially white to pale purple, with or without markings near the base; anthers yellow to purple, the connective usually light in color |
|
16 | Leaf blades sparsely to densely pubescent, the trichomes collapsing when dry; berry orange to red at maturity; Mexico (state of Chiapas) and Guatemala |
|
– | Leaf blades glabrous to sparsely pubescent, the trichomes remaining conical and acute at the tip when dry; berry dark purple at maturity; Mexico (states of Veracruz and Oaxaca) |
|
17 | Trichomes usually simple (sometimes dendritic), often curling and crisped; calyx appendages to 0.5 mm long; corolla white adaxially, shallowly to deeply stellate in outline, up to 1 cm long; berry globose; Guatemala |
|
– | Trichomes simple, not curling or crisped; calyx appendages 3–6 mm long; corolla light purple with darker markings at base adaxially, entire to shallowly stellate in outline, up to 3 cm long; berry usually ovoid; Mexico (state of Chiapas) and Guatemala |
|
18 | Corolla pale to dark purple with green markings at base adaxially, stellate in outline, divided 1/3 to 2/3 of the way to the base; Mexico (state of Veracruz) |
|
– | Corolla white to light purple with dark purple ring and green markings at base adaxially, nearly entire in outline; Mexico (state of Oaxaca) |
|
19 | Calyx usually nearly glabrous, the calyx appendages less than 2 mm long in flower, less than 3 mm long in fruit; plants glabrous to sparsely pubescent, the trichomes less than 1 mm long; mature berry dark purple (if fruit not present, also try 19b); greater than 1000 m in elevation |
|
– | Calyx glabrous or pubescent, the calyx appendages often2 mm long or more in flower, 3 mm long or more in fruit; plants glabrous to densely pubescent, the trichomes often at least 1 mm long; mature berry orange to red; greater than or less than 1000 m in elevation |
|
20 | Plant usually glabrous, rarely with occasional tan to brown appressed trichomes to 0.25 mm long; calyx margin often torn, appearing lobed, the appendages in flower up to 1 mm long or absent; corolla stellate in outline, divided 1/2 to all of the way to the base; stamens equal to slightly unequal, the longest filament to 2 mm long, less than twice the length of the short filaments, the anthers 2–3 mm long, often with dark-colored connective; seeds unnotched; Mexico (widespread in the Sierra Madre del Sur from state of Jalisco to Chiapas) south to El Salvador |
|
– | Plant glabrous to sparsely pubescent, the trichomes to 0.75 mm long; calyx margin entire, not appearing lobed, the appendages in flower 1–1.5 mm long; corolla entire to shallowly stellate in outline, divided ca. 1/5 of the way to the base; stamens very unequal, the longest filament to 3 mm long, usually twice the length of the short filaments, the anthers 3–4 mm long, usually with light-colored connective; seeds with deep notch on one side; Mexico (state of Chiapas) and Guatemala |
|
21 | Shrub to large vine to 10 m tall; trichomes tan, yellow, or red-brown, simple to furcate, 1–4 mm long; calyx often densely pubescent, the appendages in flower 7–17 mm long; mature berry 15–30 mm in diameter |
|
– | Shrub, sometimes scandent, to 5 (7) m tall; trichomes white, off-white, tan, light yellow, brown or light purple, but never red-brown, simple, to 2.5 mm long; calyx glabrous to densely pubescent, the appendages in flower 0.5–9 mm long; mature berry less than 15 mm in diameter |
|
22 | Flowering calyx with well-developed rim 1–3 mm long, the appendages 0.4–4 mm long, connate at their bases, forming a continuous shelf of tissue (this feature especially visible in fruit); Mexico (states of Chiapas and Oaxaca) to Guatemala, above 1500 m |
|
– | Flowering calyx with rim 0.5 to 1 mm long, the appendages 0.25–11 mm long, free at their bases, not forming a continuous shelf of tissue; Mexico and Central America, 350–3000 m |
|
23 | Calyx glabrous or nearly so; abaxial surface of corolla glabrous or sparsely puberulent with trichomes to 0.1 mm long (difficult to see without magnification); stem trichomes usually appressed-antrorse; Mexico (states of Jalisco, Puebla, and Veracruz) |
|
– | Calyx and abaxial surface of the corolla lobes usually puberulent with dense trichomes to 2 mm (easily seen without magnification), best seen in bud (sometimes nearly glabrous in |
|
24 | Apex of berry sometimes apiculate due to persisting remnant of style base; calyx 1.5–2.5 mm long; Mexico (from northern state of Puebla and adjacent state of Veracruz) |
|
– | Apex of berry rounded; calyx 2–4 mm long; Mexico (state of Jalisco) |
|
25 | Corolla lobes and interpetalar membrane purple; stem trichomes usually densely matted; some leaf trichomes to 2 mm long; Mexico (state of Veracruz) |
|
– | Corolla lobes and interpetalar membrane white to lilac, often with darker purple or maroon stripes on the lobes; stem trichomes often dense but not matted, the individual trichomes spreading and separated from neighboring trichomes; leaf trichomes usually less than 1.5 mm long; southern Mexico (excluding the state of Veracruz), Guatemala, and south to Nicaragua |
|
26 | New growth angled or ribbed, somewhat compressed upon drying; largest leaves usually with blade greater than 16 cm long; longest appendages on calyx 4–7 mm long; 1700–2000 m; Mexico (southeastern state of Chiapas) |
|
– | New growth terete, not angled or ribbed, not much compressed upon drying; largest leaves usually with blade less than 16 cm long (if largest leaf blade greater than 16 cm, the calyx appendages usually < 4 mm long); 500–3000 m; southern Mexico to Nicaragua |
|
27 | Calyx appendages 5–9 mm long, with at least some appendages on single calyx 7–9 mm long; base of appendages somewhat to very flattened; 700–1000 m |
|
– | Calyx appendages usually < 5 mm long (rarely to 5 mm in |
|
28 | Pedicels of the oldest (third day) flowers mostly greater than 1.2 cm long; pedicels of fully developed fruits often greater than 2 cm long; calyx rim above the appendage insertion usually less than 0.5 mm long and covered by the slightly spreading appendages; mature seeds with notch; usually above 2000 m; southern Mexico, Guatemala, El Salvador |
|
– | Pedicels of the oldest flowers mostly less than 1.2 cm long; pedicels of fully developed fruits often less than 2 cm long; calyx rim above the appendage insertion usually greater than 0.5 mm long and exposed by the widely spreading appendages; mature seeds usually lacking notch; 500–3000 m; southern Mexico, Guatemala, to Nicaragua |
|
Multangulate- or geminate-stellate trichomes (with more than two rays at a node, the rays sometimes rebranched) forming the majority of indument on stems and leaves
1 | Stamens equal or nearly so |
|
– | Stamens obviously unequal, one filament much longer than the other four |
|
2 | Calyx lacking appendages, the rim often undulate, lobed or torn |
|
– | Calyx with appendages greater than or equal to 0.25 mm long, the rim rarely undulate, lobed or torn |
|
3 | Larger leaf blades 8–15.5 × 4.5–10 cm, the abaxial side densely pubescent with, but not always obscured by, short to long-stalked, red-brown, multangulate- and geminate-stellate trichomes 0.5–1 mm in diameter; calyx 6–7 mm long in flower, 9–17 mm in diameter in fruit; anthers ca. 6 mm long |
|
– | Larger leaf blades 3–13 × 2–5 cm, the abaxial side obscured by a dense tomentum of overlapping, short-stalked, white to tan, stellate or multangulate-stellate trichomes less than 0.25 mm in diameter; calyx 2.5–4.5 mm long in flower, 6–8 mm in diameter in fruit; anthers 3–4 mm long |
|
4 | Indument of multangulate-stellate trichomes with 3–5 rays at a node, the rays often rebranched, the individual trichomes with a branching tree-like appearance (easily seen on both sides of the leaf); calyx appendages linear, usually narrowed at the tip; corolla stellate in outline, usually divided to 1/2 (rarely 2/3) of the way to the base, with abundant interpetalar tissue connecting the lobes; stamens usually somewhat unequal with one stamen slightly longer than the other four |
|
– | Indument of geminate-stellate trichomes with 5–8 rays at a node mixed with multangulate-stellate trichomes, the rays sometimes rebranched, the individual trichomes with a bottlebrush appearance (most easily seen on the adaxial leaf surface); calyx appendages usually obovate, rounded at the tip; corolla stellate in outline, divided 1/2 to 2/3 of the way to the base, with scant interpetalar tissue present only at base of corolla lobes; stamens equal |
|
5 | Calyx appendages with large, oblong, glandular area at tip, this area turning black upon drying |
|
– | Calyx appendages lacking glandular area at tip, remaining green upon drying |
|
6 | Upper stem branching divaricate (strongly zigzagging) well below the branch tips; leaf blades 2–10 × 0.5–3.5 cm, narrowly ovate to elliptic, acuminate at the tip, coriaceous, the trichomes often obscuring the abaxial surface; endemic to Guatemala |
|
– | Upper stem branching only divaricate at the very tips of the branches, not strongly zigzagging below the tip; leave blades 2.5–15 × 1.5–8 cm, broadly ovate to elliptic, acute to acuminate at the tip, thick chartaceous, the trichomes rarely obscuring the abaxial surface; Mexico to Nicaragua |
|
7 | Trichomes on leaves and stems often a mixture of colors (off-white to red-brown) and forms (simple, long-stalked furcate, and stalked multangulate-stellate) on the same plant, 1–4 mm long; flowering calyx appendages 7–17 mm long; mature berry 15–30 mm in diameter |
|
– | Trichomes on leaves and stems of various colors (white, off-white, yellow, orange, brown, or red-brown), but markedly different colors not usually present on the same plant and multangulate-stellate trichomes always present, these sessile to stalked, sometimes mixed with furcate trichomes, 0.05–1.5 (2) mm long; flowering calyx appendages 0.25–6 (8) mm long; mature berry 4–20 mm in diameter |
|
8 | Indument of stalked multangulate-stellate trichomes, the rays straight and usually rebranched, sometimes repeatedly; states of Chiapas, Mexico and Baja Verapaz, Guatemala |
|
– | Indument of sessile to stalked multangulate-stellate and/or geminate-stellate trichomes, the rays rarely rebranched; widely distributed in Mexico and Guatemala |
|
9 | Corolla shallowly stellate in outline, usually divided to 1/2 (rarely 2/3) of the way to the base, white to pale lilac with darker purple color on the lobes adaxially; Mexico (state of Chiapas) |
|
– | Corolla entire in outline, pale white to lilac, without darker purple color on the lobes adaxially; Guatemala (state of Baja Verapaz) |
|
10 | Trichomes on leaves, stems, and calyx white to tan, not yellow, orange or brown |
|
– | At least some trichomes on leaves, stems, and calyx yellow, orange, or brown, sometimes mixed with tan trichomes |
|
11 | Multangulate-stellate trichomes of adaxial side of leaf blade sessile to very short-stalked, the rays laying on the leaf surface; corolla white, shallowly stellate in outline |
|
– | Multangulate-stellate trichomes of adaxial side of leaf blade sessile to stalked, the rays not laying on the leaf surface; corolla white to purple, entire to shallowly stellate in outline |
|
12 | Vine to scandent shrub; trichomes furcate to multangulate-stellate, those on calyx minute (less than 0.25 mm in diameter) and difficult to see without magnification; leaf blade apex rounded to acute, the leaf veins usually obscure and light green in color; corolla light purple; coastal areas, up to 1000 m in elevation, widespread in Mexico and Central America, especially on the Caribbean slope |
|
– | Shrub, rarely vine; trichomes multangulate- to geminate-stellate, those on the calyx not minute (usually more than 0.25 mm in diameter), the surface often obscured by indument; leaf blade apex acute to short-acuminate, the leaf veins often prominent and white in color; corolla white to pale lavender; coastal areas and adjacent mountains, up to 1300 m in elevation, Pacific slope of Mexico south to Central America |
|
13 | Multangulate-stellate trichomes of adaxial side of leaf blade sessile to very short-stalked, the rays laying on the leaf surface; corolla white, shallowly stellate in outline |
|
– | Multangulate-stellate trichomes of adaxial side of leaf blade sessile to stalked, the rays not laying on the leaf surface; corolla white, entire to shallowly stellate in outline |
|
14 | Upper dichasial branching widely divaricate (the branches often spreading at a 180 degree angle), not forming a continuous, sinuous axis; epidermis light brown, usually smooth upon drying; multangulate-stellate trichomes usually with 5–8 (10) rays per whorl, the area where they join often becoming enlarged and spherical; corolla white, shallowly stellate in outline; mainly the Yucatán Peninsula of Mexico and adjacent Guatemala and Belize, usually 0–500 m in elevation |
|
– | Upper dichasial branches usually forming a continuous, sinuous axis; epidermis dark brown, often longitudinally wrinkled upon drying; multangulate-stellate trichomes usually with 3–6 rays per whorl, the area where they join just slightly enlarged and not particularly spherical; corolla white, entire to shallowly stellate in outline; away from immediate coast of Mexico and Central America, (500) 900–2500 m in elevation |
|
15 | Calyx 4–5 mm long in flower, 6–8 mm long in fruit |
|
– | Calyx 2–3.5 mm long in flower, 2–4 mm long in fruit |
|
Based on
Image of herbarium specimen of
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
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.
Map of geographic distribution of
Mexico. Fruit edible; maravilla, huevo de cuervo, chimpin, tsibu (
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.
When
Based on
Image of herbarium specimen of
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.
Map of geographic distribution of
Guatemala: Alta Verapaz: kaki saki maï (
Flowering specimens have been collected March through November; fruiting specimens have been collected May through February.
Mexico. Veracruz: Río Blanco, Orizaba,
Image of herbarium specimen of
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.
Map of geographic distribution of
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 (
This species is similar to the Central American species
Mexico. Quintana Roo: Cobá, east of the ruins, in advanced deciduous forest, [
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Based on
Image of herbarium specimen of
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 (
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
Map of geographic distribution of
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.
This species is very similar to
There is a wide range of morphological variation within
Guatemala. Suchitepéquez: Volcán Santa Clara, between Finca El Naranjo and upper slopes, 1250–2650 m, 23 May 1942,
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Mexico. Chiapas: Mpio. La Independencia, third ridge along logging road from Las Margaritas to Campo Alegre, [
Image of isotype of
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
Map of geographic distribution of
Mexico. Chiapas: chichol mut (Tzeltal) (
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.
Mexico. Veracruz: District Cordoba, Cerro de Chocomán, 12 May 1907,
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
As detailed elsewhere (
Based on
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Based on
Guatemala. Suchitepéquez: Las Nubes, Nov 1875,
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Guatemala. Chiquimula: Tixixí (Tishishí), 3–5 miles north of Jocotán, 500–1500 m, 10 Nov 1939,
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Based on
Image of herbarium specimen of
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
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,
Map of geographic distribution of
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
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 (
Guatemala. Huehuetenango: Cruz de Limón, between San Mateo Ixtatán and Mucá, Sierra de los Cuchumatanes, 2600–3000 m, 31 Jul 1942,
Image of herbarium specimen of
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
Map of geographic distribution of
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.
Guatemala: Huehuetenango: between Xoxlac and Nucapuxlac, Sierra de los Cuchumatanes, 1600–2500 m, 17 Jul 1942,
Image of holotype of
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.
Map of geographic distribution of
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.
Although
Based on
Image of herbarium specimen of
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,
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 (
Map of geographic distribution of
Mexico. Trompeta, chichi de perra (
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
Guatemala. Baja Verapaz: Niño Perdido, Cerro Verde, east of km 150 of Cobán Road, in high forest, elevation not recorded, 3 Dec 1976,
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Based on
Image of herbarium specimen of
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.
Map of geographic distribution of
Mexico. Oaxaca: leaves used as an edible green (
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.
Mexico. Querétaro: Mpio. Landa, 10 km al noreste de Agua Zarca, sobre camino a Neblinas, 1100 m, 23 Jun 1988,
Image of holotype of
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
Map of geographic distribution of
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
Guatemala. Huehuetenango: Mpio. San Mateo Ixtatán, 4 miles east of San Mateo Ixtatán on road to Barillas, 8500 ft, 7 Feb 1965,
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
Guatemala. Sacluc, Aug 1877,
Image of herbarium specimen of
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.
Map of geographic distribution of
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 (
Mexico. Chiapas: [Mpio. Siltepec?] Letrero, near Siltepec [
Image of herbarium specimen of
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.
Map of geographic distribution of
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.
This species belongs to series
This species is poorly known and has been rarely collected. One collection from southeastern Chiapas that may belong to
Based on
Image of herbarium specimen of
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.
Map of geographic distribution of
Mexico. Chiapas: fruits and leaves eaten after cooking in water, yerbabuena (Spanish), ashinte (Tzeltal) (
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.
In Mexico,
Confusion about what name to use for this species exists in the literature. In his treatment of the
Where they co-occur,
The name
Mexico. Nuevo León: Mpio. Aramberri, Cerro El Viejo, 1200 m, 28 Jul1993,
Image of herbarium specimen of
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.
Map of geographic distribution of
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
Although this species has not been observed in the field, it is obviously related to the species of series
Belize. Orange Walk: 10 Oct 1926,
Image of herbarium specimen of
Scandent shrub to twining woody liana, 2.5–5 m tall (or taller, described as climbing high into the tree canopy). Indument white to tan, uniseriate, multicellular, sessile to short-stalked, stellate or multangulate-stellate, eglandular, spreading trichomes 0.1–0.2 mm long, 0.1–0.25 mm in diameter, the rays 3–6 per whorl, straight, not rebranched. Young stems greenish, sparsely to densely pubescent, compressed at the nodes when dried in a plant press, becoming dark reddish brown and woody with age; upper sympodial branching points monochasial and dichasial. Leaves simple, the leaves of the upper sympodia paired or not, the pairs unequal in size, the larger ones with blades 3–11 (12.8) × (1.8) 1.5–5 cm, the smaller ones (often not developing) with blades 2.5–6.2 × 1–3.6 cm, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, chartaceous, the two sides of the blade very different in color, the adaxial side green, glabrous, the abaxial side pale, densely pubescent with overlapping trichomes, the base cuneate to attenuate, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate, the petiole 0.2–2.5 cm long, sometimes absent, the larger leaf blades with 3–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–3, axillary, oriented horizontally; peduncles absent; pedicels 12–33 mm long and erect to arching in flower, 20–35 mm long (probably longer) and erect in fruit; calyx 2.5–4.5 mm long, 2.5–4 mm in diameter, campanulate, moderately to densely pubescent, the margin truncate, undulate or lobed, the appendages lacking; fruiting calyx enlarged, bowl-shaped to rotate, 2.5–4 mm long, 6–8 mm in diameter; corolla 0.8–1.2 cm long, rotate in orientation, entire to shallowly stellate in outline, divided ca. 1/4 of the way to the base), with abundant interpetalar tissue, white, adaxially sometimes with green markings at the base of the lobes, glabrous, abaxially moderately puberulent near the major veins; stamens equal to slightly unequal, straight, the four short filaments ca. 1 mm long, the one long filament 1–2 mm long, glabrous, the anthers 3–4 mm long, ovate to oblong, the tips narrowed, free from one another, yellow (drying brownish perhaps due to glandular exudate), bumpy in texture, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–9 mm long, linear, straight to curved, glabrous, the stigma capitate, slightly bilobed, decurrent down two sides. Fruit a berry, 6–12 mm long, 8–13 mm in diameter, depressed globose, orange to red when mature, glabrous, lacking sclerotic granules. Seeds 20–40 per fruit, 2.5–3 × 2–2.5 mm, flattened, oval in outline, with slightly thickened margin, yellow-orange to brown, the surface reticulum rough with indistinct serpentine pattern and deep luminae.
Unknown.
Mexico (Campeche, Chiapas, Quintana Roo), Guatemala (Petén), Belize, and Honduras, in primary or disturbed tropical moist forest and tropical dry forest, on slopes and ridges, in ravines, often on limestone, 0–800 m in elevation (Fig.
Map of geographic distribution of
Mexico. Chiapas: tumat zak (Tzeltal) (
Flowering specimens have been collected from May through October; specimens with mature fruits have been collected from June through October. In Belize, corollas open in the early morning (sometimes before sunrise) and close by sunset (
Guatemala. [Quetzaltenango]: [Volcán] Santa María, [
Image of herbarium specimen of
Subshrub, shrub or treelet, erect, 0.3–6 m tall. Indument pale yellow, uniseriate, multicellular, simple, eglandular, ascending-appressed to ascending trichomes 0.3–1.5 mm long, these usually remaining cylindrical and acute upon drying, the trichomes sometimes becoming less appressed on older stems. Stems green and slender when young, moderately to densely pubescent, sometimes glabrate with age, often compressed upon drying in a plant press, becoming woody with age; upper sympodial branching points usually monochasial with a few dichasial branching points, the branching angles not particularly apparent. Leaves simple, the leaves of the upper sympodia usually paired, usually conspicuously different in size and shape, the larger ones with blades 6–19.5 × 2–7 cm, narrowly elliptic, lanceolate, or oblanceolate, sometimes appearing slightly falcate, the smaller ones with blades 0.6–3.5 × 0.4–2.1 cm, ovate, the blades of both the large and small leaves chartaceous, moderately pubescent, the pubescence densest along the veins of the abaxial side, the trichomes along the midvein of the abaxial side appressed, the base of large blades cuneate, the base of small blades rounded, oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole to 0.6 (1.1) cm long, sometimes absent, the large leaf blades with (6) 8–12 primary veins on each side of the midvein. Flowers solitary or in groups of 2–3, axillary, nodding; peduncles absent; pedicels very slender, (11) 15–30 mm and straight to arching or deflexed in flower, to 36 mm long, arching or deflexed in fruit, moderately pubescent; 1.5–2.5 mm long, 2–3 mm in diameter, obconic to campanulate, moderately pubescent, the margin truncate, with 5–10 narrow, erect appendages 0.5–3 mm long, emerging ca. 0.25 mm below the calyx rim; fruiting calyx usually enlarged, widely campanulate to bowl-shaped, 1–2 mm long, 3.5–7 mm in diameter, the appendages 1–4 mm long, erect, sometimes withering; corolla 0.5–1 cm long, campanulate to rotate in orientation, stellate in outline, divided 1/2 to nearly all the way to the base, interpetalar tissue present near base, white, adaxial markings unknown, sparsely pubescent abaxially especially near the lobe tips; stamens equal, straight, the filaments 1–2 mm long, glabrous, the anthers 2–3 mm long, ovate, somewhat narrowed at the tip (the narrowed portion ca. 0.25 mm long), free of one another, yellow, glabrous, poricidal at the tips, the pores round, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 3.5–5 mm long, linear, straight, glabrous, the stigma capitate. Fruit a berry, 3.5–9 mm long, 3.5–11 mm in diameter, globose to ovoid, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 50–100 per fruit, 0.9–1.5 × 1 mm, compressed, but not flat, ridged, rhombic to triangular in outline, tan to orange, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Guatemala (Quetzaltenango, San Marcos, Suchitepéquez), Costa Rica, and Panama, in cloud forest, montane forest, and oak forest, sometimes on slopes in canyons and drainages, often near streams, 1250–1870 m in elevation (Fig.
Map of geographic distribution of
Guatemala. Chiltepe de montaña (
Flowering specimens have been collected in January, February, June, and July. Specimens with mature fruits have been collected in July, August, and February. Information about the diurnal movements of the corolla of this species has not been determined; the flowers on specimens range from fully open to somewhat closed (campanulate).
In Guatemala,
Mexico. Jalisco: S of Puerto Vallarta and N of El Tuito, along hwy. 200, 20.3 road km S of Playa Mismaloya, W side of the road, along footpath that follows small drainage, 500 m, 13 Aug. 1991,
Image of herbarium specimen of
Shrub, 0.3–2.2 m tall, from horizontal rhizomes. Indument white to tan, uniseriate, multicellular, simple, eglandular, appressed (usually appressed-ascending) trichomes 0.1–1.25 mm long. Stems green to purple (drying greenish tan) with purple (drying blackish) lenticular vertical striations and purplish nodes when young, glabrous to sparsely pubescent, not much compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points mostly monochasial, sometimes dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 5.5–15 (22) × 2.9–8.7 (11) cm, the smaller ones with blades 2–9 (12) × 1.3–4.9 (7) cm, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, chartaceous, glabrous to sparsely pubescent, the base cuneate to truncate (rounded on small leaves), sometimes oblique, the margin entire, usually irregularly undulate, the apex acuminate, the petiole 0.1–1.5 (2.5) cm long, the larger leaf blades with 5–7 primary veins on each side of the midvein. Flowers solitary or in groups of 2–7, axillary, oriented horizontally; peduncles absent; pedicels 5–30 mm long and erect in flower, 11–29 mm long and erect in fruit, glabrous (rarely sparsely pubescent); calyx 2.5–4 mm long, 2.5–5.5 mm in diameter, campanulate, glabrous to sparsely puberulent, the margin truncate, with 10 spreading, linear appendages 1–5.5 mm long emerging 0.5–1 mm below calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 1.5–3.5 mm long, (3.5) 5–8 mm in diameter, the appendages to 6 mm long; corolla (1) 1.4–2.3 cm long, campanulate in orientation (sometimes opening wider by tearing), mostly entire in outline (with shallow notches), with abundant interpetalar tissue, adaxially white and glabrous with no markings, the abaxial side of the lobes green, glabrous to sparsely puberulent; stamens unequal, straight, the four short filaments 0.5–2 mm long, the one long filament 4–7 mm long, glabrous, the anthers 4–6 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, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–11 mm long, linear, slightly curved downward, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 6–10 mm long, 7–12 mm in diameter, globose, red at maturity, glabrous, lacking sclerotic granules. Seeds 20–40 per fruit, 2–3 × 1.5–2 mm, flattened, depressed ovate in outline, tan to light brown, the surface reticulum with minutely pitted serpentine pattern and shallow luminae.
n = 12, from
Mexico (Jalisco) in tropical moist forest, tropical dry forest or in oak forest, often near drainages, 350–1350 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected from June through November (and March); specimens with mature fruits have been collected August through December (and March). Field observation of the corollas indicates that they are open in the early morning and closed by late morning (
Based on
Image of herbarium specimen of
Scandent shrub to woody vine, 3–4 (10) m tall, the lower stem to 2.5 cm in diameter. Indument of tan to orange, uniseriate, multicellular, short- to long-stalked, multangulate-stellate to geminate-stellate, eglandular, spreading trichomes 0.5–0.75 (1.2) mm long and in diameter, the rays 5–8 per whorl, straight, rarely rebranched. Stems greenish when young, densely pubescent, not compressed when dried in a plant press, becoming light brown and 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 8–15.5 × 4.5–10 cm, the smaller ones with blades 1.5–4.5 (9.5) × 1–3 (4) cm, the leaf pairs similar in shape, the blades ovate to elliptic (often widely so), coriaceous, adaxially sparsely pubescent, abaxially densely pubescent, the base truncate to rounded (rarely cuneate or slightly cordate), the margin entire, usually undulate, the apex acute to acuminate, the petiole (0.5) 1–3.7 cm long, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–12, axillary, nodding; peduncles absent; pedicels 7–12 mm long and reflexed in flower, to 28 mm long and erect in fruit, densely pubescent; calyx 6–7 mm long, 6–7 mm in diameter, campanulate, densely pubescent, the margin truncate, undulate, or lobed, the appendages not present; fruiting calyx enlarged, bowl-shaped, unevenly torn and lobed, 3–10 mm long, 9–17 mm in diameter; corolla 1–1.5 cm long, open corolla orientation not known, entire to shallowly stellate in outline, divided ca. 1/4 of the way to the base, with abundant interpetalar tissue, white, adaxially glabrous, abaxially densely puberulent on the lobes; stamens equal, the filaments ca. 1 mm long, glabrous, the anthers 5–6 mm long, lanceolate, free of one another, yellow, glabrous to sparsely pubescent, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 8–9 mm long, linear, straight to curved, glabrous, the stigma oblong, sometimes slightly bilobed, decurrent down two sides. Fruit a berry, 8–21 mm long, 7–22 mm in diameter, globose to depressed globose, green to white when immature, orange to red when mature, glabrous to sparsely pubescent, lacking sclerotic granules. Seeds 20–50 per fruit, 2.9–3.8 × 2.5–3.2 mm, flattened, with thickened margin, circular to depressed ovate in outline, yellow-orange to brown-orange, the surface reticulum in the center nearly smooth with indistinct serpentine pattern and shallow luminae, the texture on the margin wrinkled and rough.
Unknown.
Mexico (Chiapas), Guatemala (Huehuetenango, Izabal, Petén), and Belize, in tropical rain forest, pine forest, pine-oak forest, cloud forest, and tropical moist forest, both in primary forest and along road edges, on slopes and ridges, 80–1500 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected December through May; specimens with mature fruits have been collected from March through October. The diurnal movements of the corolla of this species are unknown. The five flowering collections that we examined had closed flowers, indicating that the flowers must open and close very early in the morning or at night.
Mexico. Jalisco: Mpio. Guautitlán de García Barragán, Majada de las Avellanas, comunidad indígena de Cuzalapa, 3–4 km al NNW de El Durazno, 800–1000 m, 6 Nov 1995,
Image of herbarium specimen of
Shrub to tree, 1.5–7 (10) m tall. Indument mostly lacking, rarely with a few tan to brown, uniseriate, multicellular, simple, eglandular, appressed-ascending trichomes 0.1–0.25 mm long. Stems green when young, usually glabrous, not compressed, but sometimes slightly 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 3–15 × 0.75–6 cm, the smaller ones with blades 1–6 × 0.3–3 cm, the leaf pairs usually similar in shape, the blades ovate (sometimes narrowly), elliptic, or obovate, coriaceous, glabrous and shiny on both sides, the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex obtuse to acute or acuminate, the petiole 0.2–1.5 cm long, the larger leaf blades with 5–8 primary veins on each side of the midvein. Flowers solitary or in groups of 2–5, axillary, oriented horizontally to nodding; peduncles absent; pedicels slender, 14–27 mm long and erect to arching in flower, to 35 mm long, arching to deflexed in fruit, usually glabrous; calyx 2–3.5 mm long, 3–4.5 mm in diameter, campanulate, glabrous or with a few small scattered trichomes, the margin truncate to shallowly lobed, often irregularly notched or torn, with 0–5 knob-like appendages 0.5–1 mm long, emerging 0.25 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, often torn, 1–3 mm long, 3.5–5.5 mm in diameter, the appendages usually not visible; corolla 0.6–1.3 cm long, rotate to campanulate in orientation, stellate in outline, divided 1/2 to nearly all of the way to the base, the lobes with interpetalar tissue, white both abaxially and adaxially, often with yellow-green or purple markings near the stamen insertion on the adaxial side, glabrous except for tiny trichomes along the margins of the lobes; stamens equal to unequal, straight, when unequal the four shorter filaments 1–1.5 mm long, the fifth filament 1.5–2 mm long, glabrous, the anthers 2.5–3 mm long, lanceolate, free of one another, yellow, sometimes with a brown connective, 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, the stigma capitate. Fruit a berry, 5–12 mm long, 5–13 mm in diameter, globose to depressed globose, green when immature, purple at maturity, glabrous, lacking sclerotic granules. Seeds 10–50 per fruit, 2.5–3 × 2–2.5 mm, flattened, depressed ovate to circular in outline, sometimes folded, brown, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Chiapas, Guerrero, Jalisco, Michoacán, Oaxaca), Guatemala (Quetzaltenango), and El Salvador, in primary or secondary forest, often in the transition between tropical dry forest and cloud forest, including tropical moist forest,
Map of geographic distribution of
Mexico. Naranjillo (
Flowering specimens have been collected June through December. Specimens with mature fruits have been collected December through June. The corollas on the specimens are often open; this indicates that the corollas must be open for a substantial amount of time each day.
Guatemala. Huehuetenango: Mpio. Santa Ana Huista, N of Parque Victoria aquatic center, Aldea El Tabacal, bank above the Río Santa Ana, upstream of the sumidero,
Image of isotype of
Shrub, 0.75–2 m tall, erect. Indument of light yellow (sometimes appearing tan or off-white), uniseriate, multicellular, simple, eglandular, appressed-ascending trichomes 0.1–1 (1.5) mm long. Stems green with small light green lenticular vertical striations when young, sparsely to moderately pubescent, not much compressed 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 5.5–14 × 2–7 cm, the smaller ones with blades 2–7 × 1–4.5 cm, the leaf pairs similar in shape, the blades ovate to elliptic, chartaceous, glabrous to sparsely (moderately) pubescent, the trichomes usually densely spreading outward (towards the margins) along the abaxial veins, especially at the base of the main vein, the base cuneate to attenuate, sometimes oblique, the margin entire, usually irregularly undulate, the apex acuminate, the petiole 0.2–1.5 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 2–3, axillary, oriented horizontally; peduncles absent; pedicels (8) 10–18 mm long and erect in flower, 15–25 mm long and erect in fruit, sparsely to moderately pubescent; calyx 2–3 mm long, 4–5 mm in diameter, obconic to campanulate, sparsely to moderately pubescent, the margin truncate, with 10 spreading linear appendages 5–11 mm long (appendages on the same calyx of varying lengths, but at least some appendages on the same calyx > 7 mm long), the base of the appendages flattened and 0.5–1 mm wide, emerging ca. 0.25–0.5 mm below the calyx rim; fruiting calyx enlarged, bowl-shaped to rotate, 2–3 mm long, 5–7 mm in diameter, the appendages 5–14 mm long, to 1.5 mm wide at the flattened base; corolla 0.9–1.8 cm long, rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white, the adaxial side with three green spots located between the short stamens, glabrous, the abaxial side of the lobes green, sparsely to moderately puberulent near the major veins; stamens unequal, straight, the four short filaments 1–2 mm long, the one long filament 3–5 mm long, glabrous, the anthers 4–6 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, not opening into longitudinal slits; pistil with glabrous ovary, the style 8–9 mm long, linear, glabrous, the stigma oblong, decurrent down two sides, slightly lobed. Fruit a berry, 10–11 mm long, 10–12 mm in diameter, globose to depressed globose, orange (red) at maturity, glabrous, lacking sclerotic granules. Seeds 10–40 per fruit, 2.5–3 × 2–2.5 mm, flattened, nearly circular to oval in outline, not obviously notched (if slightly indented, indentation is usually less than 0.3 mm), yellow-orange, the surface reticulum with indistinct serpentine pattern and shallow luminae.
Unknown.
Mexico (Chiapas, Oaxaca), Guatemala (Huehuetenango, El Progreso), El Salvador, and Nicaragua, in oak or oak-pine forest, tropical moist forest, tropical dry forest, often on calcareous soils, sometimes near drainages, 700–1000 m in elevation (1600 m in El Salvador) (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected from July through November; specimens with mature fruits have been collected September through December. The corollas of this species are open in the early morning and closed by late morning (
Mexico. Veracruz: Mpio. Calcahualco: 4.2 km W of Escola on road to Jacal, 17.5 km by road NW of Coscomatepec, 2,200 m, 12 Jan 1981,
Image of herbarium specimen of
Shrub, 2–4 m tall. Indument of light yellow, uniseriate, multicellular, simple, eglandular, spreading to appressed, weak, sometimes matted (on stem) trichomes, 0.25–2 mm long. Stems tan to purplish with vertical ridges when young (dark striations not evident on dried material), moderately to densely pubescent, not compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points mostly monochasial, sometimes dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 13.9–23.5 × 5.5–10.5 cm, the smaller ones with blades 5.7–8.5 × 2.6–4.7 cm, the leaf pairs similar in shape, ovate, elliptic, or obovate, chartaceous, densely pubescent, the base rounded to cuneate, sometimes oblique, the margin entire, usually undulate, the apex acuminate, the petiole 0.2–3.5 cm long, the larger leaf blades with 5–7 primary veins on each side of the midvein. Flowers solitary or in groups of 2–6 (10), axillary, oriented horizontally; peduncles absent; pedicels 15–28 mm long and erect in flower, 26–42 mm long and erect in fruit, densely pubescent; calyx 2–3 mm long, 2.5–3.5 mm in diameter, urceolate to campanulate, densely pubescent, the margin truncate, with 10 spreading, linear appendages 1–4 mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl- to plate-shaped, 1.5–3 mm long, 5.5–9 mm in diameter, the appendages to 6 mm long; corolla 1.1–1.6 cm long mm long, rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, purple adaxially, greenish-purple and densely pubescent near the major veins abaxially; stamens unequal, straight, the four short filaments 0.5–1 mm long, the one long filament 3.5–4 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 short stamens dehiscing away from the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–8.5 mm long, linear, straight to slightly curved, glabrous, the stigma capitate, unlobed. Fruit a berry, 6–9 mm long, 6–8 mm in diameter, globose, red at maturity, glabrous, lacking sclerotic granules. Seeds 20–30 per fruit, 2–3 × 2–2.5 mm, flattened, reniform in outline with notch on one side, brown to orange, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Veracruz), in oak-pine forest, tropical moist forest, or cloud forest, often in disturbed or secondary forest, in shady canyons and ravines, 1750–2600 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected from January to July; specimens with mature fruits have been collected June to January. The diurnal corolla movements of this species are not known. The corollas on specimens are usually closed, indicating that the flowers are probably only open in the early morning.
Based on
Perennial herb from fusiform storage roots, decumbent, ascending, to erect, usually recumbent with age, ca. 0.1–0.5 (0.9) m tall, dying back each season far beneath the soil surface. Indument of white, uniseriate, multicellular, simple, eglandular, spreading to appressed trichomes (0.1) 0.5–2 mm long, (in the state of Michoacán, these sometimes intermixed with forked or dendritically branched trichomes). Stems green to purple-green, moderately pubescent (rarely glabrate in age), compressed and ribbed when dried in a plant press, usually with little woody tissue; first stem (2) 5–35 (40) cm long to the first inflorescence, the internodes (3) 6–14 (21); first sympodial branching point usually dichasial, the subsequent sympodial branching points usually monochasial. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades 2–10.5 × 0.8–4.2 cm, the smaller ones with blades 1/4–7/8 the size of the larger, the leaf pairs similar in shape, the blades obovate, elliptic, ovate, or lanceolate, chartaceous, moderately pubescent, the primary veins 5–7 on either side of the midvein, the base rounded to cuneate, the margin entire, usually undulate to irregularly angled, the apex rounded to acute, the petioles 0.1–0.5 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels (30) 50–150 (180) mm and erect in flower, 54–170 mm long (probably longer) and deflexed in fruit, moderately pubescent with trichomes of two distinct lengths, the smaller 0.1–0.3 mm and appressed, the longer 0.5–1.5 mm and mostly spreading, rarely only the longer trichomes present; calyx 3–5.5 mm long, 3.5–7 mm in diameter, campanulate, densely pubescent, the margin truncate, with 10 linear, lax appendages laying closely against the corolla 2–10 mm long emerging ca. 0.5–1 mm below the calyx rim; fruiting calyx enlarged, 4–11 mm long, 9–22.5 mm in diameter, the appendages appressed to fruit, often broken, to 11.5 mm long; corolla 1.3–3.6 cm long (2.9–6.8 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, lilac to dark purple (very rarely white or very pale lilac), with darker purple stripes along the major veins adaxially, green and moderately pubescent near the major veins abaxially; stamens unequal, straight, the filaments of three lengths, the two shortest filaments 1–4.5 mm long, the two medium filaments 1.5–5.5 mm long, the one long filament (2.5) 3–8.25 mm long, the length of the longest filament 1.2–2 times that of the medium filaments, glabrous, the anthers 4.5–8.5 mm long, elliptic to ovate, free of one another, yellow, glabrous, poricidal at the tips, the pores obovate, dehiscing distally or toward the style, not opening into longitudinal slits; pollen grains tricolporate; pistil with glabrous ovary, the style 8–14.5 mm, linear, straight to slightly curved, glabrous, the stigma round to slightly bilobed. Fruit a berry, remaining attached to calyx at maturity, pendent or lying on the ground, 14–41.5 mm long, 12–28 mm diameter, round to ovoid, the exocarp green, glabrous, the mesocarp area green, soft and juicy, lacking sclerotic granules, placental area green, soft and juicy. Seeds (10) 40–139 per fruit, 2.3–2.8 × 1.7–2.5 mm, rounded, slightly compressed, reniform to depressed-obovate brownish-black, the surface reticulum with minute serpentine pattern and shallow luminae.
Image of herbarium specimen of
2n = 24 (
Mexico (Distrito Federal, Durango, Guanajato, Hidalgo, Jalisco, México, Michoacán, Nayarit, Puebla, Querétaro, San Luis Potosí, Tlaxcala, Veracruz, perhaps Zacatecas), mainly restricted to the volcanic soils of the transvolcanic belt in disturbed areas such as pastures, agricultural fields, along paths and roadsides, and in clearings in xerophilous schrub, oak and coniferous forest, 1600–3000 m in elevation (Fig.
Map of geographic distribution of
Mexico. Berenjena, berenjito, chumpin, chimpina, huevo de gato, tintolón, tilindon, tlanoxtle, tlanochtle, tlalnonochtle, shimpe, tilapó, tochin, la chichi, chochocuero, coyotomate, purga de las animas, xipes, mazatlatlaixtli (
Flowering specimens have been collected June to October; specimens with mature fruit have been collected September to December. The first author observed in the field that the corollas open in the very early morning and close by noon. The pollen of this variety has a sweet scent. Solitary bees in the genus
This “weedy” variety has had an intimate relationship with the people of Mexico and may owe its current distribution to humans, who are probably its primary dispersal agents. The fruit of
In some areas of Jalisco and Michoacán, where this variety grows with
Mexico. Nuevo León: Mpio. Zaragoza, Cerro El Viejo, 2085 m, 17 Jun 1993,
Image of herbarium specimen of
Perennial herb, from fusiform storage roots, erect, 0.1–0.6 m tall, dying back each season. Indument of white, uniseriate, multicellular, simple, eglandular, spreading to appressed trichomes 0.1–1.5 mm long, these often of two distinct lengths, the shorter more numerous, 0.1–0.25 mm long and appressed, the longer less numerous, 0.5–1.5 mm long and spreading. Stems green to purple-green, sparsely to moderately pubescent, compressed and ribbed when dried in a plant press, usually with very little woody tissue except at the base; first stem 9–30 cm long to the first inflorescence, internodes (3) 6–14 ; first sympodial branching point usually dichasial, the subsequent sympodial branching points usually monochasial. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades 1.5–10 × 0.5–4.5 cm, the smaller ones with blades 1/3–9/10 the size of the larger, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, chartaceous, sparsely pubescent with trichomes similar to those of the stem, the primary veins 4–6 on either side of the midvein, the base cuneate, attenuate onto the petiole, the margin entire, usually irregularly undulate, the apex rounded to acute, the petioles to 1.5 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 30–130 (190) mm and erect in flower, 60–100 mm (probably longer) and deflexed in fruit, often looped or curved, moderately pubescent with trichomes of two distinct lengths, the smaller 0.1–0.3 mm and appressed-retrorse, the longer 0.5–1.5 mm and mostly spreading (slightly retrorse), rarely only the longer trichomes present; calyx 3–5 mm long, 4–7 mm in diameter, campanulate (conic), moderately pubescent (densest on the ribs), the margin truncate, with 10 linear, slightly spreading appendages 3–8 (12.5) mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, 8–9 mm long, 16–17.5 mm in diameter, the teeth spreading slightly, often broken, 3.5–9 mm long; corolla 1–2.8 cm long (2–4.6 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, lilac, with darker purple stripes on the major veins adaxially, green and glabrous to very sparsely pubescent near the major veins abaxially; stamens unequal, straight, the filaments unequal, the two shortest filaments 2–3 mm long, the two medium filaments 2–3.5 mm long, the one long filament 2.5–5.5 mm long, the length of the longest filament 1.5–2 times the length of the medium filaments, glabrous to pubescent; anthers 4–5 (6) mm long, elliptic to ovate, 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 9–12 mm, linear, straight to slightly curved, glabrous, the stigma round to slightly bilobed. Fruit a berry, remaining attached to calyx at maturity, pendent, 22–30 mm long, 11–19 mm in diameter, ovoid, the exocarp green with rose or tan blotches, glabrous, the mesocarp soft, juicy, lacking sclerotic granules, the placental area variable in texture, sometimes green and juicy, other times purplish and slightly powdery. Seeds ca. 50–100 per fruit per fruit, 2.3–2.8 × 1.7–2.5 mm, rounded, slightly compressed, reniform to depressed-obovate, brownish-black, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Sierra Madre Oriental of Mexico (Nuevo León, Querétaro, San Luis Potosí), in oak and pine forest that may be mixed with xerophilous scrub, on limestone soils, 900–2700 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected July and August (May in Tamaulipus); mature fruits of
The fruits of
Mexico. Oaxaca: Mpio. Santa María Jaltianguis, along hwy 175, ca. 5.0–7.2 rd mi N of Ixtlán de Juárez, N of turnoff to Sta. María Jaltianguis, W side of road, downslope along footpaths, 2439 m, 11 Oct 1991,
Image of herbarium specimen of
Perennial herb from fusiform roots, usually erect, ca. 0.1–0.4 m tall, dying back each season. Indument of white, uniseriate, multicellular, simple, eglandular, spreading to appressed trichomes 0.1–1 mm long, these often of two distinct lengths, the shorter more numerous, 0.1–0.25 mm long and appressed retrorse, the longer less numerous, 0.5–1 mm long and spreading, some populations lacking the longer trichomes. Stems green to purple-green, sparsely to moderately pubescent, only the youngest stems compressed and ribbed when dried in a plant press, woody with age; first stem 2.5–25 cm long to the first inflorescence, internodes 3–8, the first sympodial branching point dichasial or monochasial, the subsequent sympodial branching points monochasial. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades 2.5–7 × 1.5–3.5 cm, the smaller ones with blades 3/4 the size of the larger, the leaf pairs similar in shape, the blades ovate to obovate, chartaceous, the primary veins 4–6 on either side of the midvein, the base rounded to cuneate to shortly attenuate onto the petiole, the margin entire, usually irregularly undulate, the apex acute to acuminate, the petioles 0.5–1.5 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 50–140 mm and erect in flower, 50–90 mm long (probably longer) and deflexed in fruit, sparsely to moderately pubescent with trichomes of two distinct lengths, the smaller 0.1–0.25 mm long and appressed-retrorse, the longer 0.5–1 mm long and spreading; calyx 4–5 mm long, 4–5.5 mm in diameter, campanulate, the ribs pubescent with spreading trichomes, the margin truncate, with 10 linear, slightly spreading appendages 2–10 mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, 5.5–9 mm long, 11–15 mm diameter, the appendages spreading slightly, not lax, often broken, 7–10 mm long; corolla 1.2–2.3 cm long (2.3–4.5 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, lilac, with darker purple stripes along the major veins adaxially, green and moderately pubescent near the major veins abaxially; stamens unequal, straight, the filaments unequal, the two shortest filaments 1.5–3 mm long, the two medium filaments 2–3.5 mm long, the one long filament 3–5.5 mm long, the length of the longest filament 1.5–2 times the length of the medium filaments, glabrous; anthers 4–5.5 mm, ovate, 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 8–12 mm, linear, straight to slightly curved, glabrous, the stigma round. Fruit a berry, remaining attached to calyx at maturity, pendent, 22–28 mm long, 13–17 mm diameter, ovoid, the exocarp green, glabrous, the mesocarp and placental area soft and juicy, lacking sclerotic granules, the placental area soft and juicy. Seeds ca. 50–90 per fruit per fruit, 2.3–2.8 × 1.7–2.5 mm, rounded, slightly compressed, reniform to depressed-obovate, brownish-black, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Oaxaca), in oak, oak-pine, and pine forest, typically found in anthropogenically disturbed habitats such as roadsides, pastures, old fields and corn fields, 2100–2900 m in elevation (Fig.
Map of geographic distribution of
Mexico. Oaxaca: chichi de venado (
Flowering specimens have been collected June to July; specimens with mature fruits have been collected in October. The first author observed in the field that the corollas open in the very early morning and close by late morning. The pollen of this variety has a sweet scent.
Guatemala. [Alta Verapaz]: Cubilqüitz, [Cubilhuitz], [
Image of herbarium specimen of
Shrub, treelet, or woody vine, sometimes epiphytic, 2–6 m tall. Indument of tan to brownish, uniseriate, multicellular simple, eglandular, curved or spreading trichomes 0.1–0.5 mm long (mostly glabrous). Stems green when young, glabrous to very 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, the leaf pairs often conspicuously different in size and shape, the larger ones with blades 8–25 × 2–9 cm, ovate (usually narrowly so), lanceolate, elliptic, or oblanceolate, the smaller ones with blades 1.75–8 × 1.3–6.7 cm, suborbicular, ovate or obovate, the leaf pairs similar in texture, coriaceous, usually glabrous, the base rounded to cuneate (usually oblique on larger leaves), the margin entire, usually undulate, the apex acute to acuminate on larger leaves, acute to rounded on smaller leaves, the petiole to 3 cm long, sometimes absent, the larger leaf blades with 6–10 primary veins on each side of the midvein. Flowers solitary or in groups of 2–11 (30), axillary, erect; peduncles absent or present as a short stub 3–5 mm long, with many pedical scars; pedicels 4–18 mm and erect in flower, to 28 mm long and erect in fruit, glabrous; calyx 1.75–4 mm long, 2.5–5 mm in diameter, widely campanulate, glabrous, the margin truncate, the appendages lacking; fruiting calyx enlarged, widely bowl-shaped, 1–4 mm long, 5–9 mm in diameter; corolla 0.6–1.3 cm long, rotate to reflexed in orientation, stellate in outline, deeply divided to the base, lacking interpetalar tissue, adaxially blue to purple and glabrous, abaxially creamy white, pinkish, or pale green and glabrous, sometimes with a linear appendage to 1 mm long at the lobe tips; stamens equal, straight, the filaments 1–2 mm long, glabrous, the anthers 5–6.5 mm long, ovate, connivent at edges to adjacent anther, forming a cone, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 5–10 mm long, linear, straight, glabrous, the stigma capitate. Fruit a berry, 4–12 mm long, 5–10 mm in diameter, globose to depressed globose, green to white when immature, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 50–250 per fruit, 1–2 × 1–2 mm, flattened to slightly curved, triangular, rectangular, or depressed ovate in outline, yellow to yellow orange, sometimes the margin lighter in color than the center, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico, (Chiapas, Oaxaca, Veracruz), Guatemala (Alta Verapaz, Huehuetenango, Izabal, Petén), Belize, El Salvador, Honduras, Nicaragua, Costa Rica, and Panama in high forest, tropical moist forest, tropical rain forest, cloud forest, montane rain forest, tropical dry forest, and
Map of geographic distribution of
None known.
Flowering specimens and specimens with mature fruits have been collected March through December. Possibly flowering and fruiting throughout the year in some locations. Corollas opening at night (
Based on
Image of herbarium specimen of
Scandent shrub to vine, climbing to 10 m (or more) into the tree canopy. Indument of pale-yellow to reddish-brown, uniseriate, multicellular, sessile or stalked, multangulate-stellate to geminate-stellate, eglandular, spreading trichomes 0.1–0.5 (0.75) mm long, 0.5–0.75 mm in diameter, the rays 5–8 per whorl, straight, not rebranched. Stems pale green (drying tan) when young, sparsely 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 near the tips of the plant not divaricate. Leaves simple, the leaves of the upper sympodia usually unpaired, the blades 3–11 × 1.5–5 cm, ovate, elliptic, or obovate, chartaceous to thick chartaceous, glabrous (especially adaxially) to moderately pubescent, the base cuneate to rounded, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate, rarely obtuse, the petiole 0.5–2.5 cm long, the larger leaf blades with 3–5 primary veins on each side of the midvein. Flowers usually in groups of 4–20, axillary, erect; peduncles absent; pedicels 4–10 mm long and erect in flower, to 15 mm long and erect in fruit, moderately to densely pubescent (the surface often obscured); calyx 2–3.5 mm long, 3–4.5 mm in diameter, campanulate, densely pubescent, the margin truncate, with 10 small obovate appendages 0.5–1 mm long emerging 0.5–1 mm below the calyx rim, the appendages sticky glandular when fresh, drying black; fruiting calyx enlarged, widely bowl-shaped, 2–2.5 mm long, 5–6 mm in diameter, the appendages not enlarging; corolla 0.6–1.2 cm long, rotate to reflexed in orientation, stellate in outline, divided 2/3 of the way to the base, with scant interpetalar tissue present at the sides of the lobes, white (lilac) and glabrous to sparsely pubescent adaxially, densely and evenly pubescent on the lobes abaxially; stamens equal, straight, the filaments 0.5–1 mm long, glabrous or with scattered trichomes, the anthers 3–4 mm long, lanceolate, free of one another, yellow to reddish-yellow, glabrous or with scattered trichomes, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous to sparsely puberulent ovary, the style 6–8 mm long, linear, straight, glabrous, the stigma capitate, decurrent down the sides. Fruit a berry, 5–7 mm long, 5–7 mm in diameter, globose, green to whitish when immature, orange-red when mature, glabrous or with scattered trichomes, lacking sclerotic granules. Seeds 5–10 per fruit, ca. 2 × 3 mm, flattened, thickened on edges, circular, depressed ovate, or reniform in outline, yellow-orange to dark orange, surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Chiapas), Guatemala (Alta Verapaz, Baja Verapaz, Quiché), in montane rainforest, cloud forest, high forest, on slopes, 1300–1800 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected May to August; specimens with mature fruits have been collected June to August. Many specimens have closed corollas, indicating that the corollas are open for a short time during the day, probably during the morning. The first author observed that the corollas were closed in the afternoon in Guatemala.
Guatemala. Chimaltenango: southwestern slopes of Volcán de Fuego, above Finca Montevideo, along Barranco Espinazo, 1200–1600 m, 20 Sep 1942,
Image of herbarium specimen of
Shrub, erect, 1–3 m tall. Indument of white to light brown, uniseriate, multicellular, simple, eglandular, patent to curved and appressed-ascending trichomes 0.25–0.75 mm long. Stems green 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 8–18 × 2–6 cm, the smaller ones with blades 2–6.5 × 1–3 cm, the leaf pairs usually similar in shape, the blades ovate (sometimes narrowly) to elliptic, membranaceous to thin chartaceous, sometimes with purple color along the veins, glabrous to sparsely pubescent, the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole 0.2–3.3 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 2–7, axillary, erect to nodding; peduncles absent; pedicels 15–30 mm and erect to arching in flower, to 40 mm long, arching to deflexed in fruit, usually glabrous; calyx 1.5–2.5 mm long, 3–4 mm in diameter, campanulate, glabrous, the margin truncate, with 10 erect to slightly spreading, slightly flattened appendages 1–1.5 mm long emerging 0.25–0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 1.5–2.5 mm long, 6–7 mm in diameter, the appendages 1–2.5 mm long, spreading; corolla 0.7–1.6 cm long, campanulate to reflexed in orientation, entire to slightly stellate in outline, divided ca. 1/5 of the way to the base, with interpetalar tissue, white, sometimes with purple markings on the adaxial side near the stamen insertion, glabrous; stamens unequal, the four shorter filaments 1–1.5 mm long, the one long filament 2–3 mm long, glabrous, the anthers 3–4 mm long, lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores round, dehiscing distally or towards the pistil, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–7 mm long, linear, straight, glabrous, the stigma capitate to oblong, decurrent down two sides. Fruit a berry, 5–8 mm long, 5–10 mm in diameter, ovoid, globose, or depressed globose, green when immature, purple at maturity, glabrous, lacking sclerotic granules. Seeds 10–50 per fruit, 2–2.5 × 1.5–2 mm, flattened, reniform in outline with deep notch on one side, orange-brown in center with yellow-orange margin, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Chiapas) and Guatemala (Alta Verapaz, Baja Verapaz, Chimaltenango) in cloud forest, including oak forest, 1200–2300 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected in August and September. Specimens with mature fruits have been collected in February. Immature fruits have been collected in August. The phenological record is incomplete, due to the paucity of specimens, and there is very little information on the diurnal movements of the corollas.
Based on
Image of herbarium specimen of
Perennial herb from large fusiform storage roots, prostate to decumbent, to 0.25 m tall and 0.5 m in diameter, dying back each season. Indument of white, uniseriate, multicellular, simple (rarely a few dendritically branched), curved, eglandular, usually appressed-ascending trichomes, 0.1–1.25 mm long. Stems green with darker green and purple striations, moderately pubescent, much compressed upon drying in a plant press, usually nonwoody; first stem 0.5–7 cm long to the first inflorescence, the internodes 4–6 (10); first sympodial branching point dichasial, followed by a mixture of monochasial and dichasial branching, this branching extensive, usually resting on the soil surface. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades (1) 1.5–9 (14) × (0.5) 0.7–4 (7) cm, the smaller ones with blades 1/8–3/4 the size of the larger, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, thick chartaceous, sparsely to moderately pubescent, the primary veins 3–5 on either side of the midvein, the base truncate to cuneate, short to long attenuate onto the petiole, sometimes oblique, the margin entire, usually slightly undulate, the apex short acuminate to rounded, the petioles winged and poorly defined, to 2 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels (20) 30–115 mm and erect in flower, 33–180 mm long, deflexed and undulate in fruit, sparsely to moderately pubescent; calyx (1.5) 2–3.5 (4.5) mm long, (2.5) 3.25–4.5 mm in diameter, broadly cupulate, moderately pubescent, the margin truncate, with 10 linear, spreading to reflexed appendages (0.5) 1–4.5 (6) mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, 5.5–12.5 mm long, (9.5) 12–25 mm in diameter, the teeth recurved, often making a complete loop, often broken, 1–12.5 mm long; corolla 1.1–2.5 cm long (2.2–4.9 cm in diameter) , rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white to lilac, with violet stripes along the major veins adaxially, green and moderately pubescent near the major veins abaxially; stamens unequal, curved, the filaments of three different lengths, the two shortest filaments (1.25) 1.75–3 (4) mm long, the two medium filaments (1.5) 2.25–3.75 (5) mm long, the one long filament 2.5–5.5 (8.5) mm long, the length of the long filament 1.2–1.8 (2.5) times that of the medium-short filaments, glabrous; anthers (2.5) 3–5.5 (6.5) mm long, elliptic to ovate, rarely lanceolate or oblong, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate to slit-like, dehiscing distally or toward the style, not opening into longitudinal slits; pollen tricolporate; pistil with glabrous ovary, the style 6–10.5 (12) mm, linear, curved downward, the stigma round to shallowly lobed. Fruit a berry, remaining attached to the calyx at maturity (the fruit matures while lying on the ground), 9–24 mm long, 9–26 mm in diameter, round to ovoid, the exocarp green with purple or black lines (becoming yellowish in age), the mesocarp thin, green and juicy, with profuse sclerotic granules, 32–83 per fruit, round to angular, yellow, the placental area narrow, greenish-white, juicy. Seeds (12) 20–100 (143) per fruit, (2.5) 3–4 × 2.2–3.1 (3.7) mm, not compressed, oblong to depressed-ovate, smooth, ridged, dark brown to black, surface reticulum with loose serpentine pattern with deep luminae and microscopic fibrils protruding from the cell walls.
2n = 24,
Mexico (Guanajuato, Hidalgo, México, Oaxaca, Puebla, Querétaro), in xerophilous scrub, rarely pine/juniper woodland, on rocky, limestone soils, often near a drainage or in a wash or canyon, often on an eroded floodplain, sometimes within or at the sides of agricultural fields or pastures, 770–2500 m in elevation (Fig.
Map of geographic distribution of
Mexico. Trompeta, berenjena, chichi de perra, ojo de venado, tonchichi, tomatillo del monte (
Flowering specimens have been collected June through August; specimens with mature fruits have been collected August to October. The first author has observed that the corollas open in the very early morning and close by noon. The pollen has a sweet scent. Solitary bees in the genera
This species was widely cultivated in German botanical gardens in the 19th and early 20th centuries, and an interesting article was written by the German botanist Purpus on how to cultivate the species (
At the time Dean published the neotypification for
Because Schlechtendal placed primary importance on the horticultural syntype from the Halle Botanical Garden, we assume that he took his description from that material, material that was shared with other botanical gardens, such as the Leipzig Botanical Garden from which the neotype was collected. The characters in the protologue description of
Based on
Image of herbarium specimen of
Shrub, 1–4 m tall. Indument of brown, uniseriate, multicellular, simple, acute, eglandular, appressed to spreading trichomes to 1.25 mm long, these usually remaining cylindrical and acute upon drying. Stems green to purple-green, glabrous to densely pubescent, not much compressed upon drying in a plant press, brown and woody with age; upper sympodial branching points mostly monochasial, some dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades (3) 6–15 × (1) 2–6.5 cm, elliptic to obovate (sometimes narrowly so), the smaller ones with blades 1–6 × 0.6–3 cm, suborbicular, ovate, elliptic or obovate, the blades of both the large and small leaves chartaceous to subcoriaceous, glabrous to moderately pubescent (denser on the veins), the base cuneate (sometimes rounded in smaller leaves), sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole to 1 (3) cm long, sometimes absent, the larger leaf blades with 4–7 primary veins on each side of the midvein. Flowers solitary or in groups of 2–6, axillary, oriented horizontally to nodding; peduncles absent; pedicels 15–60 mm long and arching in flower, to 30–55 mm long (probably longer) and arching in fruit, glabrous to densely pubescent; calyx 2–3 mm long, 3–4.5 mm in diameter, campanulate, often purplish in color, glabrous to densely pubescent, the margin truncate, with 10 spreading, linear-subulate appendages 2–9 mm long emerging 0.5–1 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 1.5–4 mm long, 7–9 mm in diameter, the appendages up to 15 mm long; corolla 0.8–2.1 cm long, campanulate in orientation, entire to shallowly stellate in outline, with abundant interpetalar tissue, adaxially white to light purple with darker purple ring near the base (sometimes with a green ring or spots at base below the purple ring), glabrous, abaxially white to purple, sometimes green near the major veins, nearly glabrous; stamens equal, straight, the filaments 1–2 mm long, glabrous, the anthers 5–6 mm long, ovate to lanceolate, free of one another, yellow-purple to purple, glabrous, poricidal at the tips, the pores ovate, dehiscing toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 10–11 mm long, linear, glabrous, the stigma capitate. Fruit a berry, (6) 12–18 mm long, 9–15 mm in diameter, ovoid, dark purple at maturity, glabrous, lacking sclerotic granules. Seeds 10–30 per fruit, 2.5–4 × 2–2.5 mm, compressed but not flat, ridged on one side or near the center, irregular in outline (shallowly crescent-shaped, semi-circular, depressed ovate, rhombic, or reniform with small notch), medium-brown to nearly black, the surface reticulum with a serpentine to honeycomb pattern with deep luminae, appearing pitted, with fibrils protruding from the cell walls.
Unknown.
Mexico (Oaxaca), in cloud forest, tropical moist forest, including pine-oak, oak, and mixed forest with
Map of geographic distribution of
Mexico. Oaxaca: monte agua zapote (
Flowering specimens and specimens with mature fruits have been collected most months of the year. The corollas are at least partially open on many specimens, indicating that they are open for much of the day.
Based on
Image of herbarium specimen of
Shrub, to 2 m tall. Indument of grey, uniseriate, multicellular, simple, glandular and eglandular, spreading trichomes (0.25) 0.5–1.5 (2.5), often glabrate with age. Stems pale green-brown and herbaceous when young, moderately pubescent, not much compressed when dried in a plant press, becoming brown and woody with age, often glabrate; upper sympodial branching points dichasial and monochasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades (6) 8–14 (18.5) × 3–9 cm, the smaller ones with blades 3–10 × 1.9–5.6 cm, the leaf pairs similar in shape, the blades widely ovate, elliptic, or obovate, chartaceous, moderately pubescent, the base truncate to short-attenuate, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate, the petiole (0.2) 1–4 (5) cm long, the larger leaf blades with 4–6 (7) primary veins on each side of the midvein. Flowers solitary or in groups of 2–3, axillary, oriented horizontally; peduncles absent; pedicels (5) 20–35 (45) mm long and erect to arching in flower, 12–35 mm long and erect to arching in fruit, moderately pubescent; calyx 3–7.5 mm long, 3–7.5 (11) mm in diameter, urceolate to campanulate, mostly glabrous except for spreading trichomes at the base near the juncture with the pedicel, the margin undulate, sometimes torn and the calyx appearing lobed, with 10 small triangular appendages 0.25–1.5 mm emerging 1–2 mm below calyx rim, these often reduced to oval protuberances 0.25–1 mm in diameter; fruiting calyx enlarged, widely bowl-shaped, 3–4 mm long, 7–12 mm in diameter, the appendages not enlarging, darkening in color; corolla 0.7–2 cm long (1.4–2.5 cm in diameter), slightly campanulate to rotate in orientation, entire in outline, with abundant interpetalar tissue, adaxially pale lilac to blue-purple with darker purple markings on the lobes, glabrous, abaxially pale lilac to blue-purple, glabrous; stamens unequal, straight, the four short filaments 1–4 mm long, the one long filament 3–7 mm long, glabrous, the anthers 3–3.5 mm long, elliptic, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–10 mm long, linear, slightly curved, glabrous, the stigma capitate, slightly bilobed. Fruit a berry, 7–20 mm long, 5–12 mm in diameter, ovoid, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 90–200 per fruit, 0.8–1.7 × 0.6–1.5 mm, flattened but not flat, rounded on the edges, depressed ovate to round in outline, tan-orange to reddish brown, the surface reticulum pitted with loose serpentine pattern and deep luminae.
Unknown.
Mexico (Guerrero, Jalisco, México, Michoacán), in oak, oak-pine, tropical dry forest, and xerophilous scrub, often in moist or seasonally wet habitat, on volcanic soils, 1750–2100 m in elevation (Fig.
Map of geographic distribution of
None known.
Specimens with flowers have been collected from June through January; specimens with mature fruits have been collected from August through March. The first author observed in the field that the corollas are open in the morning and closed by afternoon.
The calyx of
Based on
Image of herbarium specimen of
Shrub to woody vine, 2–10 m tall. Indument of long, pale yellow, orange, or red-brown, uniseriate, multicellular, simple, dendritically branched or long-stalked multangulate-stellate, eglandular, spreading trichomes 1–4 mm long, 0.75–1.4 mm in diameter, the rays of the multangulate trichomes 3–5 per whorl, straight, rarely rebranched, sometimes with an enlarged sphere where the rays join, the trichomes on an individual sometimes a mixture of colors and textures. 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 usually monochasial, sometimes dichasial, the branching sometimes zigzagging but not strongly divaricate. Leaves simple, the leaves of the upper sympodia sometimes paired and unequal in size, the larger ones with blades 3.5–15 × 2–8 cm, the smaller ones with blades 1.5–4.5 × 0.5–3 cm, ovate, elliptic or obovate (sometimes the small geminate leaf nearly orbicular), thick chartaceous, moderately to densely pubescent, the base cuneate to rounded, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate (rarely rounded on smaller leaves), the petiole 0.3–1.5 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 (4) 8–20 (30) mm and erect to arching in flower, to 30 mm long and erect to arching in fruit, moderately to densely pubescent; calyx 3–4 mm long, 3.5–4.5 mm in diameter, campanulate, moderately to densely pubescent (sometimes nearly obscured), the margin truncate, with 10 very long spreading linear appendages 7–17 mm long emerging 0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 5–6 mm long, 11–14 mm in diameter, the appendages to 20 mm long, often curling over the fruit as it develops and then spreading when the fruit is mature; corolla 1.1–1.6 (2) cm long, rotate in orientation, nearly entire in outline, with very shallow lobes (divided about 3 mm toward the base), with abundant interpetalar tissue, white to pale blue-violet, glabrous adaxially, moderately pubescent with short trichomes abaxially near the veins; stamens unequal, the four short filaments 1–1.5 mm long, the one long filament 4–5 mm long, glabrous, the anthers 5–6 mm long, narrowly oblong to lanceolate, the anthers of some of the short anthers sometimes connivent at their edges to adjacent anther, yellow, glabrous, poricidal at the tips, the pores obovate, those of the shorter stamens dehiscing away from the style, those of the long stamen dehiscing toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 10–12 mm long, linear, straight to slightly curved, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 15–30 mm long, 15–30 mm in diameter, globose to depressed globose, green to white when immature, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 20–50 per fruit, 3.5–4 × 3–3.5 mm, flattened, with thickened rim, depressed ovate in outline, tan to dark brown, the surface reticulum with minute serpentine pattern with shallow luminae.
Unknown.
Mexico (Chiapas, Oaxaca, Puebla, Veracruz), Guatemala (Baja Verapaz, Izabal, Petén), Belize, and Honduras, in primary or secondary high forest or tropical dry forest, very rarely in cloud forest at the upper part of its elevational range, often on limestone, 80–1000 (1500) m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected February to October. Specimens with mature fruits have been collected throughout the year. The corollas have been reported as opening at night (
Based on
Image of herbarium specimen of
Shrub, 1–5 m tall. Indument of white to pale yellow, uniseriate, multicellular, simple, acute, eglandular, appressed to spreading trichomes 0.1–1 mm long, the larger trichomes becoming flattened upon drying. Stems green when young, sparsely to densely pubescent, not much compressed upon drying in a plant press, brown and woody with age; upper sympodial branching points mostly monochasial, some dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 4–15 (25) × 2.5–9 (14) cm, ovate to elliptic (sometimes very widely so), rarely lanceolate, the smaller ones with blades 1.5–9 × 1–5 cm, ovate, the blades of both the large and small leaves chartaceous, sparsely to moderately pubescent, denser along the veins and in the abaxial leaf axils, the base cuneate to truncate or rounded, sometimes oblique, the margin entire, usually undulate, the apex acute to acuminate, the petiole to 4 (6) cm long, sometimes absent, the larger leaf blades with 4–7 primary veins on each side of the midvein. Flowers solitary or in groups of 2, axillary, oriented horizontally; peduncles absent; pedicels 20–60 mm long, erect to arching in flower, to 70 mm long (probably longer), deflexed and arching in fruit, sparsely to densely pubescent; calyx 3–4 mm long, 3.5–6 mm in diameter, campanulate, often purplish in color, sparsely to densely pubescent, the margin truncate, with 10 spreading, linear-subulate appendages 3–6 mm long emerging 0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 2–5 mm long, 7–13 mm in diameter, the appendages not greatly enlarging; corolla 1.2–3 cm long, campanulate to rotate in orientation, entire to shallowly stellate in outline, with abundant interpetalar tissue, adaxially light purple with darker purple ring near the base and green markings at very base, glabrous, abaxially light purple, sometimes paler or white on the lobes, sparsely to densely pubescent with small trichomes near the veins; stamens equal, straight, the filaments 1.5–2 mm long, glabrous, the anthers 5–6 mm long, 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 10–12 mm long, linear, glabrous, the stigma capitate. Fruit a berry, 9–17 mm long, 7–15 mm in diameter, usually ovoid (round), red at maturity (drying dark purple on herbarium sheets), glabrous, lacking sclerotic granules. Seeds 20–40 per fruit, 2.5–3 × 2–2.5 mm, compressed but not flat, ridged on one side or near the center, irregular in outline (shallowly crescent-shaped, triangular, or semi-circular), orange-brown, the surface reticulum with a serpentine to honeycomb pattern with deep luminae, appearing pitted, with fibrils protruding from the cell walls.
Unknown.
Mexico (Chiapas) and Guatemala (Chimaltenango, Huehuetenango, Quetzaltenango, Quiché, Sacatepéquez, San Marcos, Sololá, Totonicapán), tropical moist forests or thickets in cloud forest and oak-pine forest with
Map of geographic distribution of
Guatemala. Chilete, choshel, coxel, flor de rosa, quilete, tomatillo blanco (
Flowering specimens and specimens with mature fruits have been collected during most months of the year. Many herbarium specimens of this species have flowers with open corollas, indicating that the flowers must stay open until at least noon, if not after.
México. Oaxaca: Distrito Juchitán, Mpio. Santa María Chimalapa, San Antonio Nuevo Paraiso, a 2 km en línea recta al W, en Cerro Camedor,
Image of herbarium specimen of
Very similar to
Shrub (sometimes scandent), treelet, to vine, 1–5 (20) m tall. Indument of off-white, grey, pale yellow, or yellow-orange, uniseriate, multicellular, sessile to short-stalked, multangulate-stellate, eglandular, spreading trichomes 0.1–0.25 mm long, 0.25–0.75 mm in diameter, the rays 5–8 per whorl, usually straight (not crisped), not rebranched, the center of the trichome where the rays meet sometimes enlarged and spherical, the trichomes of the adaxial leaf surface often sessile and appressed to the leaf surface. Stems light green when young, (drying tan to brown), moderately to densely pubescent (appearing like dense felt), slightly compressed and ribbed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points a mixture of monochasial and dichasial branching (often monochasial), the branching not divaricate. Leaves simple, the leaves of the upper sympodia usually paired, the pairs unequal in size, the larger ones with blades 4.5–13 × 2.5–6 cm, the smaller ones with blades 1.5–7.6 × 1–3.8 cm, the leaf pairs similar in shape, the blades ovate to elliptic, chartaceous to subcoriaceous, sparsely to moderately pubescent, the base cuneate (sometimes widely so), rarely rounded, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate, the petiole 0.3–2.5 cm long, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–6 (–10), axillary, oriented horizontally; peduncles usually absent, sometimes a small pad forming with numerous pedicel scars, to 2 mm long; pedicels 4–10 mm long and erect in flower, to 17 mm long and erect in fruit, moderately to densely pubescent; calyx 2–4 mm long, 3–4 mm in diameter, campanulate, moderately to densely pubescent, the margin truncate, with 10 erect linear appendages with widened, bulbous apex, 1–4 mm long, emerging 0.5–1 mm below the calyx rim, the rim membranaceous and papery; fruiting calyx enlarged, widely bowl-shaped to rotate, 2–4 mm long, 6–10 mm in diameter, the appendages to 5 mm long; corolla 0.7–1.5 cm long, orientation of open corolla unknown (most likely campanulate or rotate), shallowly stellate in outline, divided 1/4 to 1/2 of the way to the base (the division increasing by tearing as the flower ages), with abundant interpetalar tissue, the adaxial side white, glabrous, the abaxial side of the lobes of unknown color, densely pubescent with multangulate-stellate, appressed trichomes; stamens unequal, straight, the four short filaments 0.5–1 mm long, the one long filament 2–3.5 mm long, glabrous, the anthers 4–5 mm long, lanceolate, narrowed at the tip, 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, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 8–15 mm long, 8–15 mm in diameter, globose, red-orange when mature, glabrous, lacking sclerotic granules. Seeds 40–100 per fruit, 2–3 × 2–3 mm, flattened, circular to depressed ovate in outline, sometimes shallowly indented on one side (< 0.5 mm), thickened on the margin, yellow-orange to orange-brown, the margin darker in color than the center, the surface reticulum nearly smooth in the center with indistinct serpentine pattern and shallow luminae, the luminae much deeper on the margin.
Unknown.
Mexico, Caribbean slope (Oaxaca, Puebla, Veracruz), in tropical moist forest, tropical dry forest, and cloud forest, in both primary forest and disturbed areas, such as coffee plantations and along roadsides, often growing with
Map of geographic distribution of
Mexico. Oaxaca: paniui poj (Zoque) (
Flowering specimens have been collected March through September; specimens with mature fruits have been collected May through November. The corollas on specimens of this species are usually closed, indicating that the corollas are open for only a short time period during the day, most likely in morning.
This species is named for Mexican botanist Rafael Torres Colín (Rafa), an expert on the flora of Oaxaca, who led us into the field in 2017, when we attempted to locate this species.
(full list of paratypes in Appendix
Based on
Image of herbarium specimen of
Shrub, 1–3 (4) m tall, multiple stems emerging at the soil level. Indument of whitish, multicellular, simple, furcate, or dendritically branched, eglandular, spreading trichomes to 0.5 mm long, as well as sparse simple, glandular hairs. Stems green with yellow striations when young, sparsely to moderately pubescent, not compressed, but sometimes angled, upon drying in a plant press, brown and woody with angled ridges with age, becoming glabrate; upper sympodial branching usually dichasial, sometimes monochasial. Leaves simple, the leaves of the upper sympodia usually unpaired, when paired often equal in size, the blades 1–15.5 × 0.5–7.5 cm, widely ovate, elliptic, rhombic-lanceolate, or narrowly lanceolate, chartaceous, moderately pubescent, the base cuneate, attenuate into the petiole, sometimes oblique, the margin entire, irregularly undulate, the apex acute or acuminate (sometimes obtuse at the very tip), the petiole (0.1–) 0.8–2.5 (–4) cm long, winged toward the apex, the larger leaf blades with 3–7 primary veins on each side of the midvein. Flowers solitary or in groups of 2–7, axillary, erect to ascending; peduncles absent; pedicels slender, 5–25 mm long and erect to arching in flower, to 25 mm long (probably longer), arching to deflexed fruit, moderately pubescent; calyx 1.5–4 mm long, 2.5–4.5 mm in diameter, obconic to campanulate, puberulent, the margin truncate, with 5–10 spreading, linear-subulate appendages of two different lengths, the five longer appendages 2–5.2 mm long, emerging at the calyx rim, the five shorter appendages 0.25–2 mm long (these sometimes lacking), emerging ca. 0.25 mm below the calyx rim; fruiting calyx slightly enlarged, widely bowl- or plate-shaped, 3–5 mm long, 9–11 mm in diameter, the appendages not elongating, often withering; corolla 1–1.2 (2) cm long, rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, adaxially deep violet-purple with yellow center, glabrous, abaxially deep violet-purple, glabrous; stamens unequal, curved, the two short filaments 0.8–1.5 mm long, the three long filaments 2–3 mm long, sparsely to densely pubescent on inner face at juncture with corolla, the anthers 2.5–4 mm long, elliptic to oblong, free of one another, yellow to orange, glabrous, poricidal at the tips, the pores opening horizontally, dehiscing toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 3.5–5.5 mm long, linear, slightly curved, widening near the stigma, glabrous, the stigma truncate, slightly bilobed. Fruit a berry, 10–20 (35) mm long, 8–15 (35) mm in diameter, subglobose to ellipsoid, green (sometimes with dark lines when immature), yellow to orange when mature, glabrous, with abundant (often more than 20) tan, irregularly shaped sclerotic granules, 1–2 (3) mm long, these sometimes attached to the seeds. Seeds ca. 3–12 per fruit in horticultural plants, possibly more in native habitat, (1.75) 2.25–3.5 × (1.5) 3–3.3 mm, compressed but not flat, round elliptic to reniform in outline, dark brown, the surface reticulum with loose serpentine pattern with deep luminae.
2n = 24, Gerasimenko and Reznikova 1968, cited in
Native to South America (Argentina, Bolivia, Paraguay, and southern Brazil) in thickets and woodlands, weedy in disturbed areas along roadsides, 100–2000 m in elevation. Horticulturally, widely distributed worldwide, including Mexico and Guatemala (no distribution map completed for this species).
Argentina. Meloncillo del aire (
This species flowers most of the year in cultivation and may be similar in its native habitat. Fruiting period not known. Corollas open in the morning, closing by late afternoon or evening.
As this species is a horticultural plant in our floristic region, we are not providing a conservation assessment.
In the protologue,
The protologue for
In his protologue for
In his protologue for
México. Michoacán: Mpio. Charo, along hwy 15, 20 rd km E of Morelia, just E of Pontezuelas, 2165 m, 13 Nov 1991,
Image of herbarium specimen of
Perennial herb from fusiform storage roots, usually erect, often recumbent with age, 0.17–1.1 m tall, dying back each season. Indument of white, uniseriate, multicellular, simple or dendritically branched, eglandular, spreading to appressed, sometimes crisped, trichomes, 0.1–1.5 (2) mm long. Stems green to reddish-purple, glabrous to sparsely pubescent, usually compressed and ribbed when dried in a plant press, somewhat woody with age, especially at base of plant; first stem 7–90 cm long to first inflorescence, the internodes (6) 10–21; first two sympodial branching points usually dichasial, usually followed by monochasial branching, this branching usually limited. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades (2.5) 5–10 (15) × 1–6 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, sparsely to moderately pubescent, the primary veins 5–7 on either side of the midvein, the base truncate or cuneate, attenuate onto the petiole, sometimes oblique, the margin entire, usually irregularly undulate, the apex acuminate, the petioles of larger leaves winged and poorly defined, 0.1–2.8 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally, and somewhat nodding; peduncles absent; pedicels 19–86 mm and erect in flower, 30–110 mm long and deflexed in fruit, glabrous to moderately pubescent; calyx 3–6 mm long, 3–6.5 mm in diameter, campanulate, glabrous to moderately pubescent, the margin truncate, with 10 knob-like to linear, slightly spreading or erect appendages 1–7.25 mm long emerging ca. 1 mm below the calyx rim; fruiting calyx enlarged, 2–10 mm long, 5–14.5 mm in diameter, the appendages stout, stiff, remaining appressed to fruit or somewhat spreading, often broken, 1–7.5 mm long; corolla 1–2.5 cm long (2–4.7 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, white (rarely tinged lilac), with violet stripes near the major veins adaxially, green near the major veins abaxially, usually glabrous; stamens unequal, the filaments of three lengths, the two shortest filaments 1–4 mm long, the two medium filaments 2–4 mm long, the one long filament 2.5–6.25 mm long, the length of the long filament always less than 2 times that of the medium filaments, glabrous, the anthers 3–5.5 mm long, lanceolate to elliptic, free of one another, yellow, glabrous, poricidal at the tips, the pores round to oval, dehiscing distally, not opening into longitudinal slits; pollen grains dicolporate; pistil with glabrous ovary, the style 7–11 mm, linear, straight to slightly curved, the stigma round, rarely somewhat lobed. Fruit a berry, usually remaining attached to calyx at maturity, pendent, 19–75 mm long, 8–19 mm in diameter, turbinate, elongate, the exocarp dull light purple to black at maturity (green with light or dark longitudinal lines when immature), glabrous, the mesocarp dark purple, soft and juicy, lacking sclerotic granules, the placental area light purple, powdery in texture. Seeds 3–38 per fruit, 3.5–5 × 3.2–4.7 mm, not compressed, depressed obovate, ridged and blistered along one side, black in color, the surface reticulum rough in texture with loose serpentine pattern with deep luminae.
2n = 24,
Mexico (México, Michoacán, Morelos) in oak, oak-pine, pine-oak, and fir forests, often on forested slopes near drainages, on volcanic soils, 1794–2645 m in elevation (Fig.
Map of geographic distribution of
Mexico. Chilillo (
Flowering specimens have been collected June to October; specimens with mature fruits have been collected in late November and December. The first author has observed in the field that the corollas open in very early morning and closed in the late morning. The pollen of this species has a sweet, powdery fragrance.
Based on
Scandent shrub to vine, 0.5–3 (6) m tall. Indument of white to tan, uniseriate, multicellular, sessile to stalked, forked to multangulate-stellate, eglandular, spreading trichomes 0.05–0.25 (0.7) mm long and in diameter, the rays 3–4 per node, straight, not rebranched (sometimes mixed with simple trichomes). Stems green to light brown when young, sparsely to densely pubescent, rarely compressed when dried in a plant press, becoming woody with age; upper sympodial branching points mostly monochasial, the branching not widely divaricate, the branch segments shallowly zigzagging or sinuate. Leaves simple, the leaves of the upper sympodia sometimes paired, the pairs equal or unequal in size, the larger ones with blades 2.5–10 (11.2) × 1.6–5 (8.4) cm, the smaller ones with blades 1.6–7 (10.5) × 0.9–3.4 (7.5) cm, the leaf pairs similar in shape, the blades ovate to elliptic, chartaceous, sparsely to moderately pubescent (denser abaxially, sometimes nearly glabrous adaxially), the base cuneate, rounded, or truncate, sometimes oblique, the margin entire, usually irregularly undulate, the apex usually rounded to acute, the petiole 0.3–2 (2.6) cm long, the larger leaf blades with 3–5 primary veins on each side of the midvein, these not usually whitish or prominent. Flowers solitary or in groups of 2–6, axillary, erect; peduncles absent; pedicels 8–20 (26) mm long and erect in flower, 10–26 (30) mm long and erect in fruit, sparsely to moderately pubescent; calyx 2.5–4 mm long, 3–6 mm in diameter, campanulate, sparsely pubescent (the individual trichomes often difficult to see with the naked eye, up to 0.1 mm long), the margin truncate, with 10 spreading linear appendages (0.75) 1–3.5 (5) mm long emerging 0.5–1 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 3–4 mm long, 6–9 mm in diameter, the appendages to 9 mm; corolla 0.7–1.9 cm long, campanulate to rotate in orientation, entire in outline, with abundant interpetalar tissue, adaxially light purple, sometimes with green at the base of the lobes, nearly glabrous, abaxially green and sparsely to moderately puberulent on the lobes, especially at the distal end, this more evident in bud; stamens unequal, the four short filaments 0.5–1 (2) mm long, the one long filament (1) 2–4 (6.5) mm long, glabrous, the anthers 4–6 mm long, narrowly ovate to lanceolate, free of one another, yellow, sparsely pubescent on the inner face, poricidal at the tips, the pores ovate, dehiscing distally or the short stamens dehiscing away from the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 8.5–9 (11) mm long, linear, straight to curved, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 6–14 mm long, 7–17 mm in diameter, globose to depressed globose, red-orange at maturity, glabrous to sparsely pubescent, lacking sclerotic granules. Seeds 50–70 per fruit, 2–3 × 1.8–2 mm, flattened, circular, triangular, depressed ovate, or slightly notched and reniform in outline, yellow-orange to tan, the surface reticulum smooth, with indistinct, tight serpentine pattern with shallow luminae, the seed margin slightly thickened and lighter in color.
Image of herbarium specimen of
Unknown.
Mexico (Campeche, Chiapas, Jalisco, Nayarit, Oaxaca, Quintana Roo, Tabasco, Tamaulipas, Veracruz, Yucatán), Guatemala (Chiquimula, Izabal, Petén, Sacatepéquez), Belize, Honduras, the Caribbean, and northern South America, in open or disturbed areas, roadsides, fencerows, thickets, dunes or forest edges, sometimes seasonally flooded (in mangroves), sometimes on limestone, within or near tropical dry forest, 0–400 (915) m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected most months of the year (from January to November); specimens with mature fruits have been collected all months of the year. The first author observed the corolla open in the field between 5 am and 5:30 am. The corollas closed at 8 am.
In this treatment, we are replacing the widely used name
In his protologue for
In his protologue for
In his protologue for
Based on
Image of herbarium specimen of
Clambering or erect shrub, sometimes a vine, 1–4 m tall. Indument of white to tan, uniseriate, multicellular, sessile to stalked, furcate, multangulate-stellate, and geminate-stellate, eglandular, spreading trichomes 0.05–0.75 mm long and 0.2–0.75 in diameter, the rays of the stellate trichomes 3–5 per whorl, straight, rarely rebranched. Stems green to light brown when young, sparsely to densely pubescent, not compressed when dried in a plant press, becoming woody with age; upper sympodial branching points dichasial or monochasial, the branching not widely divaricate, the segments shallowly zigzagging. Leaves simple, the leaves of the upper sympodia usually paired, the pairs equal or unequal in size, the larger ones with blades (2.3) 4–13.2 × (1.5) 3–9 cm, the smaller ones with blades (1.3) 2.7–7.5 × (0.7) 1.2–5 cm, the leaf pairs similar in shape, the blades ovate to elliptic, chartaceous, moderately to densely pubescent (denser abaxially, sometimes nearly glabrous adaxially), the base cuneate, rounded, or truncate, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to short acuminate, the petiole 0.3–5.4 (6) cm long, the larger leaf blades with 3–5 primary veins on each side of the midvein, these usually pale in color and prominent. Flowers solitary or in groups of 2–7, axillary, erect; peduncles absent; pedicels 7–31 mm long and erect in flower, 8–38 mm long and erect in fruit, moderately to densely pubescent; calyx 2.5–4.5 mm long, 3–7 mm in diameter, campanulate, moderately to densely pubescent (the surface sometimes obscured), the margin truncate, with 10 spreading linear appendages 1.5–6 (8) mm long emerging 0.5–1.0 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 2.5–4.5 mm long, 7–12 mm in diameter, the appendages to 9 mm; corolla 1–1.8 (2.1) cm long, campanulate to rotate in orientation, entire in outline, with abundant interpetalar tissue, adaxially white to pale lavender and nearly glabrous, abaxially green and sparsely to moderately puberulent on the lobes, especially at the distal end, this more evident in bud; stamens unequal, the four short filaments 1–3 mm long, the one long filament 3–7 mm long, glabrous, the anthers 4–5.5 mm long, narrowly ovate to lanceolate, free of one another, yellow, very sparsely pubescent on the inner face, poricidal at the tips, the pores ovate, dehiscing distally or the short stamens dehiscing away from the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–12 mm long, linear, straight to curved, glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 7–17 mm long, 8–17 mm diameter, usually globose to depressed globose, red-orange at maturity, glabrous to very sparsely pubescent, lacking sclerotic granules. Seeds 40–80 per fruit, 2–3 × 1.5–2.5 mm, flattened, circular, triangular, depressed ovate, or very slightly notched in outline, yellow to yellow-orange, the surface reticulum with tight serpentine pattern with shallow lumina, the seed margin slightly thickened, lighter or darker in color.
Unknown.
Mexico (Chiapas, Colima, Jalisco, Michoacán, Oaxaca, Sinaloa), Guatemala (El Progreso, Huehuetenango, Zacapa), El Salvador, Honduras, Nicaragua, and Costa Rica, in oak forest, pine forest, and tropical dry forest, in open disturbed areas, riparian areas, road edges, canyons and ravines, thickets and hedges, sometimes on limestone, 0–1300 m in elevation (Fig.
Image of herbarium specimen of
Map of geographic distribution of
None known.
Flowering specimens and specimens with mature fruits have been collected all months of the year. The corollas of this variety have not been observed in the field by the authors, but it is probable that the diurnal movements are similar to those of var.
The type collection from Jalisco, Mexico upon which this variety is based (
Based on
Image of herbarium specimen of
Scandent shrub to vine, 1–10 m tall. Indument of pale yellow to reddish-brown, uniseriate, multicellular, stalked or sessile, multangulate-stellate or geminate-stellate, eglandular, spreading trichomes 0.1–0.5 (0.75) mm long, 0.25–0.75 mm in diameter, the rays 5–8 rays 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 a mixture of monochasial and dichasial, the branching near the tips of the plant divaricate (diverging at wide angles). Leaves simple, the leaves of the upper sympodia usually unpaired, the blades 2.5–15 × 1.5–8 cm, ovate to elliptic, chartaceous, thick chartaceous, or subcoriaceous, sparsely pubescent adaxially (often shiny, with trichomes just concentrated along the veins), moderately to densely pubescent abaxially with surface sometimes obscured, the base cuneate to rounded, sometimes oblique, the margin entire, usually irregularly undulate, the apex acute to acuminate, rarely obtuse, the petiole 0.5–3 cm long, the larger leaf blades with 4–7 primary veins on each side of the midvein. Flowers usually in groups of 4–30, (the densest groupings spherical in shape), axillary, erect; peduncles absent; pedicels 3–10 mm long and erect in flower, to 15 mm long and erect in fruit, densely pubescent (the surface often obscured); calyx 1–2 mm long, 2.5–3.5 mm in diameter, campanulate, densely pubescent, the margin truncate, with 10 obovate appendages, sometimes just small protuberances, 0.5–1.5 (2) mm long emerging 0.3–0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 1.5–2.5 mm long, 4–6 mm in diameter, the appendages not enlarging; corolla 0.5–1.1 cm long, campanulate to reflexed in orientation, stellate in outline, divided 1/2 to 2/3 of the way to the base, with scant interpetalar tissue present connecting the base of the lobes, white (lilac) and glabrous to sparsely pubescent adaxially, pale green to whitish and densely and evenly pubescent on the lobes abaxially; stamens equal, straight, the filaments ca. 0.5 mm long, glabrous, the anthers 2.5–3 mm long, elliptic to lanceolate, free of one another, yellow to purple-yellow, glabrous or with scattered trichomes, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 5–6 mm long, linear, straight to curved, glabrous, the stigma truncate, decurrent down the sides. Fruit a berry, 2–7 mm long, 2–7 mm in diameter, globose, green to whitish when immature, orange-red when mature, glabrous or with scattered trichomes, lacking sclerotic granules. Seeds 15–40 per fruit, 1.5–2 × 1.5 mm, flattened, thickened on edges, circular, depressed ovate, or reniform in outline, yellow-orange to dark orange, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Chiapas, Guerrero, Oaxaca, Veracruz), Guatemala (Alta Verapaz, Huehuetenango), El Salvador, Honduras, and Nicaragua, in montane rainforest, tropical moist forest, tropical dry forest, oak forest, and pine-oak forest, sometimes on slopes or in disturbed forest, along roadsides or in coffee plantations, often on limestone, 500–1850 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected from March to November; specimens with mature fruits have been collected May to December. Many specimens have closed flowers, indicating that the flowers are open for a short time during the day, probably during the morning. In the field in Guatemala, in cloud forest on an overcast day, the first author observed that the newest flowers were open midday, while older flowers were already closed.
In the protologue for
The status of
México. México: Sierra de Nanchititla, oak forest across the reservoir from the town of Nanchititla, 1945 m, 8 September 1991,
Image of isotype of
Perennial herb, from fusiform storage roots, usually prostrate to ascending, to 0.15 m tall, dying back each season. Indument of white, uniseriate, multicellular, simple, eglandular, spreading to appressed trichomes, 0.1–0.5 mm long. Stems greenish-purple, moderately to densely pubescent, usually compressed when dried in a plant press, with very little woody tissue; first stem 0.5–3 cm long to the first inflorescence, the internodes 2–5; first sympodial branching point usually dichasial, followed by a mixture of monochasial and dichasial branching, this branching extensive, usually resting on the soil surface. Leaves simple, those of the upper sympodia usually paired and unequal in size, the larger ones with blades 3–6 × 1.25–3 cm, the smaller ones with blades 1/4 to 1/2 the size of the larger, the leaf pairs similar in shape, the blades obovate, oblanceolate, ovate, broadly elliptic, or rhombic, thick chartaceous, sparsely to moderately pubescent, the primary veins 3–5 on either side of the midvein, the base cuneate, sometimes attenuate onto the petiole, sometimes oblique, the margin entire, usually irregularly undulate, the apex broadly acute to rounded, the petioles winged and poorly defined, to 1.8 cm long, sometimes absent. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels 42–86 mm and erect in flower, 52–122 mm long, deflexed, and undulate in fruit, moderately pubescent with spreading trichomes of two distinct lengths, the shorter to 0.25 mm long and the longer to 0.5 mm long; calyx 3–4 mm long, 4–5.5 mm in diameter, narrowly to broadly conic, moderately pubescent, the margin truncate, with 10 linear appendages lying laxly near the corolla surface 2–6.5 mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, 2–5.5 mm long, 6–12.5 mm in diameter, the appendages spreading to reflexed, often broken, 2–6.5 mm long; corolla 1–2 cm long (2.1–3.8 cm in diameter), rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, lilac, with violet stripes near the major veins adaxially, green and moderately pubescent near the major veins abaxially; stamens unequal, the filaments of three lengths, the two shortest filaments 1.25–3.25 mm long, the two medium filaments 1.5–3.75 mm long, the one long filament 2–5.25 mm long, the length of the long filament usually less than 2 times that of the medium filaments, glabrous, the anthers 3.25–5.25 mm, lanceolate to elliptic, free of one another, yellow, glabrous, poricidal at the tips, the pores round, dehiscing distally, not opening into longitudinal slits; pollen grains irregular in shape and number of pores; pistil with glabrous ovary, the style 7–10 mm, linear, straight to slightly curved, glabrous, the stigma round, shallowly lobed. Fruit a berry, separating from calyx at maturity and matures lying on the ground, 17–22 mm long, 7–17 mm in diameter, ovoid, the exocarp dull dark purple at maturity, glabrous, the mesocarp dark purple, soft and juicy, lacking sclerotic granules, the placental area light purple, powdery in texture. Seeds 2–29 per fruit, 3.5–4.5 × 3–4.3 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,
Mexico, endemic to southwestern state of México, Sierra de Nanchititla, on level oak forest floor, 1945 m in elevation (Fig.
Map of geographic distribution of
Mexico. México: chilillo (
Flowering specimens have been collected July and August; specimens with mature fruits have been collected in October and November. The first author has observed in the field that the corollas open in the very early morning and close by late morning. The pollen of this species has a sweet fragrance.
Based on
Image of herbarium specimen of
Perennial herb to climbing shrub, erect, often recumbent with age, to 2 (3) m tall, dying back to rhizomes. Indument of small, white, uniseriate, multicellular, simple, curved, eglandular, appressed-ascending trichomes 0.1–0.6 mm long. Stems green when young, glabrous to sparsely pubescent, compressed and ribbed upon drying in a plant press, 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 3.5–15 × 1.5–6.2 cm, the smaller ones with blades 0.7–6.5 (10.5) × 0.5–3.1 (5) cm, the leaf pairs similar in shape, the blades ovate (sometimes narrowly), elliptic, or obovate, the blades of both the large and small leaves chartaceous to thick chartaceous, glabrous to sparsely pubescent, the base truncate, cuneate, or attenuate, sometimes oblique, the margin entire, the apex acute to acuminate, the petiole 0.1–0.9 (2) cm long, sometimes absent, the large leaf blades with 3–6 primary veins on each side of the midvein. Flowers solitary, axillary, pendent; peduncles absent; pedicels 12–45 mm and arching to deflexed in flower, to 53 mm long and deflexed in fruit, glabrous to sparsely pubescent; calyx 1.5–3 (4) mm long, 3–4 mm in diameter, obconic to campanulate, glabrous to sparsely pubescent, the margin truncate, with 10 linear, spreading to reflexed appendages 1.5–5 mm long emerging 0.5 mm below the calyx rim; fruiting calyx usually enlarged, widely campanulate to bowl-shaped, 1.5–4 mm long, 3–8 mm in diameter, the appendages 2–8 mm long, spreading; corolla 0.5–1.4 cm long, campanulate to reflexed in orientation, stellate in outline, divided 1/3–2/3 of the way to the base, (lobes shallow on first day that the flower opens, becoming deeper each subsequent day that the flower opens), with interpetalar tissue, adaxially and abaxially white to light purple, glabrous; stamens equal, straight, the filaments 1–1.5 mm long, glabrous, the anthers 4.5–7 mm long, lanceolate, connivent to connate at edges to adjacent anther, yellow, glabrous, poricidal at the tip, the pores round, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–9 mm long, linear, straight, glabrous, the stigma truncate. Fruit a berry, 3–10 (17) mm long, 3–9 (12) mm in diameter, globose to ovoid, orange to red at maturity, glabrous, lacking sclerotic granules. Seeds 7–60 per fruit, 1.5–3 × 1.5–2 mm, flattened, depressed ovate in outline, tan to orange, the surface reticulum with a tight serpentine pattern with shallow luminae.
Unknown.
Mexico (Chiapas, Hidalgo, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tabasco, Veracruz), Guatemala (Huehuetenango, Izabal, Petén), Belize, and Honduras (and possibly further south in Central America), in tropical rainforest, tropical dry forest, tropical moist forest, and cloud forest, sometimes in coffee plantations or disturbed forest, near rivers or waterfalls, in gorges, or on the sides of canyons, 30–1050 m in elevation (Fig.
Map of geographic distribution of
Mexico. San Luis Potosi: tomatillo, arrete de la virgin, flor de mariposa (
Flowering specimens have been collected May to September. Fruiting specimens have been collected September to December. It is possible this species flowers and fruits throughout the year in some locations. The first author observed in the field in Mexico that the corollas are open in the very early morning and closed by late morning.
Mexico. Sinaloa: Sierra Surotato, Las Mesas, 15 Sep 1941,
Image of herbarium specimen of
Shrub, (0.6) 1–3 m tall. Indument of off-white to tan (purple), uniseriate, multicellular, simple, glandular and eglandular, spreading to appressed trichomes 0.2–2.1 mm long. Stems pale green (drying tan) with dark lenticular vertical striations when young, moderately pubescent, not much compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points dichasial and monochasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 8.1–16.5 × 3.4–9.5 cm, the smaller ones with blades 1.6–7.8 × 0.6–4.2 cm, the leaf pairs similar in shape, the blades ovate to elliptic, thin-chartaceous, moderately pubescent, the base cuneate to attenuate, sometimes oblique, the margin entire, usually irregularly undulate (rarely remotely coarsely dentate), the apex acuminate, the petiole 0.7–2.2 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 2–3 (5), axillary, oriented horizontally; peduncles absent; pedicels 9–33 mm long and erect in flower, 11–29 mm long and erect in fruit, moderately pubescent; calyx 2–4 mm long, 2.5–6 mm in diameter, widely obconic to widely campanulate, densely puberulent, the margin truncate, with 10 spreading linear appendages 2–10 mm long, emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped to rotate, 1–2 (4) mm long, (3) 5–9 mm in diameter, the appendages 6–11 mm long (probably longer), 0.5–1 mm wide at the base, spreading; corolla 0.9–1.8 cm long, rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, adaxially white, sometimes with three green spots near the insertion of the stamen filaments, abaxially green and puberulent with purple trichomes near the major veins; stamens unequal, straight, the four short filaments 1–2 mm long, the one long filament 3.5–4.5 mm long, glabrous, the anthers 3.5–5 mm long, elliptic, lanceolate, or oblong, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–10 mm long, linear, slightly curved, glabrous, the stigma capitate, decurrent down two sides, sometimes slightly bilobed. Fruit a berry, 4–11 mm long, 4–12 mm in diameter, globose to depressed globose, red at maturity, glabrous, lacking sclerotic granules. Seeds 40–137 per fruit, 1.75–2.25 × 1.5 mm, flattened, reniform to triangular in outline, tan, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Colima, Guerrero, Jalisco, Michoacán, Nayarit, Oaxaca, Quéretaro, Sinaloa, and Sonora) in broadleaved forest, oak forest, pine-oak forest, tropical dry forest, and riparian forest (including
Map of geographic distribution of
None known.
Flowering specimens have been collected from March to December; specimens with mature fruits have been collected February to December. The first author observed in the field that the corollas are open in the morning and closed by afternoon.
Mexico: State of Oaxaca, Dto. Sola de Vega, Mpio. Santiago Textitlán, Colonia Nueva, 18 Aug 2006,
Image of holotype of
Small shrub, ca. 0.5 m tall. Indument of clear to white, uniseriate, multicellular, simple, crisped, glandular (glandular tip golden-yellow to grey), spreading trichomes 0.25–1 (1.5) mm long. Stems pale green (drying tan) when young, moderately to densely pubescent, not much compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points dichasial and monochasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 3–7 × 1.5–3 cm, the smaller ones with blades 1.5–5 × 0.5–2 cm, the leaf pairs similar in shape, the blades ovate, to elliptic, chartaceous, moderately to densely pubescent, the base truncate, cuneate, or attenuate, sometimes oblique, the margin entire, usually irregularly undulate, the apex acuminate, the petiole 0.1–1 cm long, sometimes absent, the larger leaf blades with 4–5 primary veins on each side of the midvein. Flowers solitary, axillary, oriented horizontally; peduncles absent; pedicels ca. 20–25 mm long and erect in flower, 20–30 mm long and erect to arching in fruit, moderately to densely pubescent; calyx 3–3.5 mm long, 3.5–4 mm in diameter, campanulate, moderately to densely puberulent, the margin truncate, with 10 spreading linear, basally flattened appendages 7–10 mm long emerging ca. 0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, 4–6 mm long, 9–12 mm in diameter, appendages to 15 mm long; corolla 1.2–1.5 cm long, rotate in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, purple and glabrous adaxially, color of abaxial side unknown, glabrous; stamens unequal, straight, the four short filaments 1–2 mm long, the one long filament 2–3 mm long, glabrous, the anthers 3.5–4 mm long, elliptic to lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing distally, not opening into longitudinal slits; pistil with glabrous ovary, the style ca. 7 mm long, linear, glabrous, the stigma capitate, decurrent down two sides. Fruit a dry berry lacking juicy mesocarp, 13–15 mm long, 9–12 mm in diameter, turbinate, the tip apiculate, pale greenish orange when mature, glabrous, lacking sclerotic granules. Seeds 50–80 per fruit, 1.75–2 × 1.25–1.5 mm, somewhat compressed, round-edged, reniform in outline, tan-orange, surface pitted.
Unknown.
Mexico (Oaxaca), in pine-oak forest, ca. 1500 m in elevation (Fig.
Map of geographic distribution of
None known.
A flowering and fruiting specimen has been collected in July. The timing of the diurnal corolla movements is unknown; however, the corollas are open on the two specimens of the type collection, indicating that the flowers are open for part of the day.
This species is only known from the one type specimen cited above.
Based on
Image of herbarium specimen of
Shrub, 1.2–4.5 m tall, sometimes vining or arching through neighboring vegetation. Indument of light yellow, uniseriate, multicellular, simple, often curly, eglandular, spreading to appressed trichomes 0.2–1.75 mm long, sometimes with very small glistening bumps between longer trichomes. Stems green to purple (drying tan) with dark lenticular vertical striations when young, sparsely to densely pubescent, not much compressed when dried in a plant press, becoming light brown and woody with age; upper sympodial branching points monochasial and dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades (4.6) 6.3–11.8 (15) × (1.9) 2.5–6.5 (8.8) cm, the smaller ones with blades 1.7–5.2 (8.5) × 0.9–3.9 (6.3) cm, the leaf pairs similar in shape, the blades ovate to elliptic, chartaceous, moderately pubescent, the trichomes usually densely spreading outward (towards the margins) along the abaxial veins, especially at the base of the main vein, the base usually cuneate (rarely attenuate), sometimes oblique, the margin entire, usually irregularly undulate (rarely coarsely dentate), the apex acuminate, the petiole 0.5–3.3 cm long, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–7 (10), axillary, oriented horizontally; peduncles absent; pedicels (8) 13–33 mm long and erect in flower, (15) 18–33 (40) mm long and erect in fruit, moderately pubescent; calyx 2.3–3.5 (4.1) mm long, 2.5–3.5 (4) mm in diameter, obconic to campanulate, moderately to densely pubescent, the margin truncate, with 10 erect, linear appendages (0.5) 1–4.3 mm long emerging 0.25–0.5 mm below the calyx rim; fruiting calyx enlarged, bowl-shaped to rotate, 1.5–3.2 mm long, 5–8.5 mm in diameter, the appendages to 6 mm long; corolla (0.7) 0.9–1.8 cm long, rotate in orientation, entire in outline, with abundant interpetalar tissue, adaxially white to pale violet with purple stripes along the major veins, with three green spots located between the short stamens, glabrous, abaxially green on the lobes, moderately puberulent near the veins; stamens unequal, straight, the four short filaments 1–2 (3) mm long, the one long filament (2.75) 3.5–4.2 (5) mm long, glabrous, the anthers 2.5–4.5 mm long, lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, dehiscing toward the style or distally, not opening into longitudinal slits; pistil with glabrous ovary, the style 6–9 mm long, linear (sometimes curved upward at the tip), glabrous, the stigma oblong, decurrent down two sides. Fruit a berry, 6–11 mm long, 6–10 (15) mm in diameter, globose, red at maturity, glabrous, lacking sclerotic granules. Seeds 10–30 per fruit, (2) 2.5–4 × 2–3 mm, flattened, reniform in outline, with deep notch on side (usually more than 0.5 mm deep), yellow-orange, the surface reticulum with minute serpentine pattern and shallow luminae.
Unknown.
Mexico (Chiapas, Guerrero, Jalisco, Michoacán, Oaxaca), Guatemala (Chimaltenango, Escuintla, Guatemala, Huehuetenango, Quetzaltenango, Quiché, Sacatepéquez, San Marocis, Sololá), and El Salvador on steep slopes, in ravines, and in semi-disturbed areas such as roadside thickets, path edges, and disturbed agricultural areas in mixed broadleaved forest, oak forest, oak/pine forest, fir forest, and cloud forest, 2100–3000 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected from February through October; specimens with mature fruits have been collected January through December. Field observation of the corollas indicates that the corollas are open in the early morning and closed by late morning (
This species is very similar to
México. Puebla. Mpio. Teziutlán: Agua de Obispo, bosque de encino en cañada, [
Image of herbarium specimen of
Shrub, 0.5–1.8 m tall. Indument of tan, uniseriate, multicellular, simple, eglandular, appressed-ascending to ascending trichomes 0.25–1.25 mm long. Stems green with pale vertical lenticular streaks when young, glabrous to sparsely pubescent, not much compressed when dried in a plant press, becoming brown and woody with age; upper sympodial branching points mostly monochasial, some dichasial. Leaves simple, the leaves of the upper sympodia usually paired and unequal in size, the larger ones with blades 6.5–13.3 × 3.7–6.5 cm, the smaller ones with blades 4–8.5 × 1.9–4.9 cm, the leaf pairs similar in shape, the blades ovate, elliptic, or obovate, thin chartaceous, glabrous to sparsely pubescent (densest along the veins), the base cuneate to attenuate, sometimes oblique, the margin entire, usually undulate, the apex acuminate, the petiole 0.2–1.5 (2) cm long, the larger leaf blades with 4–6 primary veins on each side of the midvein. Flowers solitary or in groups of 2–6 (10), axillary, oriented horizontally; peduncles absent; pedicels 13–28 mm long and erect in flower, 18–25 mm long (probably longer) and erect in fruit, glabrous to sparsely pubescent; calyx 1.5–2.5 mm long, 1.75–2.5 mm in diameter, urceolate to campanulate, glabrous to sparsely pubescent, the margin truncate, with 10 spreading linear appendages 1–3.5 mm long emerging 0.3–0.5 mm below the calyx rim; fruiting calyx enlarged, widely bowl-shaped, ca. 2 mm long, 4 mm in diameter, the appendages to 4 mm long (possibly longer); corolla (0.5) 0.9–1.4 cm long, campanulate to reflexed in orientation, mostly entire in outline (with shallow notches), with abundant interpetalar tissue, sometimes the interpetalar tissue tearing as the corolla opens, adaxially white, glabrous, abaxially color unknown, sparsely pubescent near the major veins; stamens unequal, straight, the four short filaments 1.25–1.5 mm long, the one long filament 4–5 mm long, glabrous, the anthers 3.5–4 mm long, lanceolate, free of one another, yellow, glabrous, poricidal at the tips, the pores ovate, all dehiscing toward the style, not opening into longitudinal slits; pistil with glabrous ovary, the style 7–9 mm long, linear, curved upward at tip, glabrous, the stigma oblong-capitate, slightly bilobed. Fruit a berry, ca. 6 mm long, 4 mm in diameter (probably larger), ovoid, usually apiculate due to persistent style base, color unknown, glabrous, lacking sclerotic granules. Seeds not yet seen.
Unknown.
Mexico (Puebla, Veracruz) in disturbed broadleaved cloud forest and oak forest, in shady canyons and on mountainsides, 1250–1450 m in elevation (Fig.
Map of geographic distribution of
None known.
Flowering specimens have been collected from April to November; specimens with mature fruit have been collected in November. The diurnal corolla movements are not known, however the corollas on specimens are usually closed, indicating that the flowers are probably only open in the early morning.
Costa Rica. Cartago: Las Vueltas, Tucurrique, 635 m, Jan 1899,
We have not been able to confirm the existence of the shrub/vine
Based on
We have not been able to confirm the existence of the usually epiphytic shrub/vine
In his protologue for
In addition to the above specimen groups, there are difficult to place specimens among the herbs of series
Finally, at the end of their paper on series
We thank the curators of A, ANSM, ARIZ, ASU, BIGU, BR, BREM, BRIT, BM, C, CAS, CIIDIR,