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Research Article
The genus Lycianthes (Solanaceae, Capsiceae) in Mexico and Guatemala
expand article infoEllen Dean, Jennifer Poore, Marco Antonio Anguiano-Constante§, Michael H. Nee|, Hannah Kang, Thomas Starbuck, Annamarie Rodrígues, Matthew Conner
‡ University of California, Davis, United States of America
§ Universidad de Guadalajara, Guadalajara, Mexico
| Unaffiliated, Richland Center, United States of America
Open Access

Abstract

Lycianthes, the third most species-rich genus in the Solanaceae, is distributed in both the New and Old Worlds and is especially diverse in Mexico. Here we provide an identification key, taxonomic descriptions, distribution maps, and illustrations of specimens, trichomes, flowers, and fruits for the 53 known Lycianthes taxa of Mexico and Guatemala. The new combination Lycianthes scandens (Mill.) M.Nee is made and replaces the name Lycianthes lenta (Cav.) Bitter, which is placed in synonymy. Within L. scandens, two varieties are recognized (Lycianthes scandens var. scandens and Lycianthes scandens var. flavicans (Bitter) J.Poore & E.Dean, comb. nov.). In addition, one new species (Lycianthes rafatorresii E.Dean, sp. nov.) is described from eastern Mexico, and 10 names (either recognized taxa or synonyms of recognized taxa) are lectotypified, including the names Solanum heteroclitum Sendtn., S. rantonnetii Carrière, and S. synantherum Sendtn. The species L. multiflora Bitter and L. synanthera (Sendtn.) Bitter are excluded from the treatment, as research indicates that they do not occur in Mexico and Guatemala, however full synonymy for both names is given.

Keywords

Guatemala, Lycianthes, Mexico, Neotropics, Solanaceae, Taxonomy

Introduction

Lycianthes (Dunal) Hassler (Capsiceae, Solanaceae) with approximately 150–200 species (D’Arcy 1991; Hunziker 2001) is the third largest genus in the Solanaceae, after Solanum L. and Cestrum L. (Hunziker 2001). Although Lycianthes is native to both the New and Old Worlds, the center of distribution of the genus, and the majority of Lycianthes taxa, are found in the New World (from northern Mexico to Argentina as well as the Caribbean). The genus is recognized by its combination of poricidal anthers and a distinctive calyx morphology in which the five calyx lobes are truncated into a sleeve-like rim; in many Lycianthes, appendages (sometimes called calyx teeth) protrude below the calyx rim (D’Arcy 1986a) (Fig. 1). Calyx appendages are also present in Lycianthes’s closest relative (and only other member of tribe Capsiceae) Capsicum L. (Bitter 1919; Särkinen et al. 2013). When in fruit (and anthers are lacking), Lycianthes can be confused with Capsicum, as well as other Solanaceous genera with truncated calyces and no appendages (Cuatresia Hunz. and Witheringia L’Hér.).

Figure 1. 

Lycianthes calyces A green, immature fruits and relatively long calyx appendages in L. mariovelizii E. Dean, field photo of Dean 9509 (DAV) B flower buds and relatively short calyx appendages in L. jalicensis E. Dean, field photo of Dean 248 (DAV) C green, immature fruit and calyx with long, sleeve-like calyx margin and connate appendage bases in L. connata J.L. Gentry, field photo of Dean 9530 (DAV) D flower bud and glandular pubescent calyx with very short, nearly absent appendages (appearing as oval bulges) and scarious calyx margin with undulating rim in L. pringlei (B.L.Rob. & Greenm.) Bitter, field photo of Dean 327 (DAV) E immature pale green to yellow fruits and calyces with obovate appendages with rounded tips in L. sideroxyloides (Schltdl.) Bitter, field photo of Dean 9526 (DAV) F mature fruit with calyx nearly lacking appendages (present as short bulges) in L. ceratocalycia (Donn.Sm.) Bitter, field photo of Dean 9532 (DAV). Scale bars: 5 mm.

The only taxonomic monograph of Lycianthes as a whole was written in the early 20th century by the German botanist Friedrich August Georg Bitter (1919); that work includes 189 terminal taxa (species, subspecies, and varieties) divided into subgenera, sections, and series, with no identification keys to assist in distinguishing the species. Although Bitter’s monograph was published in volume 24 of the journal Abhandlungen herausgegeban vom Naturwissenschaftlichen Verein zu Bremen with a volume date of 1920, both Band 1 and Band 2 of volume 24 were preprinted in November 1919 and the pagination throughout the volume is continuous (Tropicos 2020), leading to confusion as to the date to cite for the species described in his monograph (1919, 1920, or 1919(1920)). The correct date for publication of all names and combinations published therein is 1919.

Early taxonomic history of the genus Lycianthes

Georg Bitter’s 1919 monograph provides a detailed account of the taxonomic history of Lycianthes which was also recounted in an unpublished dissertation thesis by Dean (1995); we are publishing Dean’s 1995 summary here. The genus was included in Solanum until the early part of the 20th century. The first person to work with the species as a group was the French botanist Michel Félix Dunal, who monographed Solanum in the early 19th century (Dunal 1813, 1816). In his earliest work, he placed a number of present-day Lycianthes species together in his section Inermia, group Polymeris, based upon the presence of calyx appendages and the many separate axillary pedicels common in Lycianthes (Dunal 1813, 1816; Knapp 1983). His group Polymeris was given sectional status in Solanum by the German botanist Wilhelm Gerhard Walpers (1844), who added two more sections with future Lycianthes species: Lycioides and Holochlaina. In a much expanded treatment published in De Candolle’s Prodromus, Dunal combined Walpers’ three sections into his subsection Lycianthes (Dunal 1852). That subsection comprised two series (Meiomeris and Polymeris) and four subseries (Eulycianthes, Pseudolycianthes, Gonianthes, and Lobanthes) (Dunal 1852). The Austrian botanist Richard Von Wettstein, in his generic level treatment of Solanum for Die natürlichen Pflanzenfamilien (Von Wettstein 1895), raised Lycianthes to a section of Solanum, and Bitter accorded it subgeneric status in 1917 (Bitter 1917).

The Swiss botanist Emil Hassler raised Lycianthes to generic level in 1917, transferring three species of Dunal’s subseries Eulycianthes (Hassler 1917). The evidence used to justify the creation of the genus came from anatomical work performed by both Hassler and Bitter; all three of the transferred species have fruits with stone cells in their endocarps. In his later comprehensive monograph of Lycianthes, Bitter (1919) expanded the circumscription of the genus to include most members of Dunal’s subsection Lycianthes, asserting that the calyx of these species places them closer to Capsicum than to Solanum. During the latter part of the 20th century, some authors continued to treat Lycianthes as part of Solanum (Morton 1944, 1976; Hunziker 1979; Rzedowski 1985; Symon 1985a, b, 1987).

Two names for the concept of Lycianthes predate the name Lycianthes in the literature. The American botanist Constantine Samuel Rafinesque published the name Otilix (Rafinesque 1828) based on Solanum lycioides L., but this new genus was lost to obscurity and not utilized by subsequent authors. Subsequently, the French botanist Henri Ernest Baillon (1888) published the genus Parascopolia based upon a single collection made in Acapulco, Mexico by the phycologist Charles Thiébaut (P. acapulcensis Baill.). It is unclear why Baillon published this new genus since, as pointed out by Bitter (1919), Dunal had clearly accounted for similar species in his subsection Lycianthes. In his description of Parascopolia, Baillon noted the 8–10 calyx teeth, a feature that clearly indicates that this taxon belongs with other species of Lycianthes. His description of the anther dehiscence as “2-rimosis” was interpreted by most workers as meaning longitudinal anther dehiscence. Bitter assumed that P. acapulcensis belonged to Lycianthes but was not able to see the type at Paris (Bitter 1919). Unable to clear up the anther dehiscence confusion, he delayed synonymizing Lycianthes with Parascopolia (Bitter 1919). After seeing a photo of P. acapulcensis, William D’Arcy proposed the inclusion of this species in Lycianthes and the conservation of Lycianthes over Parascopolia and Otilix (D’Arcy 1972); this was approved in 1973 (McVaugh 1973). The anthers of the flowers of the type of Parascopolia acapulcensis were finally examined by Dean (1995), and she confirmed that the anthers had small, linear, terminal pores, not longitudinal slits.

Taxonomic work on New World Lycianthes since Bitter (1919)

Since Bitter’s monograph, many additional New World Lycianthes taxa have been described (e.g., Bitter 1924; Standley 1927a, b, 1940; Standley and Steyermark 1947; Gentry 1948; Gentry 1973; Dean 1994, 1998; Rodríguez and Vargas 2002; Dean 2004, 2014; Dean et al. 2017b, 2018b, c, 2019a) and transferred to the genus (e.g., Bitter 1922; D’Arcy 1986b). Keys and species-level treatments for the genus have been included in a number of floristic works, including those for parts of Mexico (Rzedowski 1985 [as Solanum]; Nee 1986), Guatemala (Gentry and Standley 1974), Nicaragua (D’Arcy 2001), Costa Rica (Standley and Morton 1938; Bohs 2015), Panama (Standley 1928; D’Arcy 1973a), Venezuela (Benítez de Rojas and D’Arcy 1997), Peru (MacBride 1962 [as Solanum]), and Argentina (Barboza and Hunziker 1992; Barboza 2013); in addition, an identification key for upcoming species treatments has been published recently for the species of Brazil (Costa-Silva and Agra 2018b), and a summary of the Brazilian species, including lectotypifications, has also been completed (Costa-Silva 2018; Costa-Silva and Agra 2018a).

Morphological revisions have been completed on five of Bitter’s Lycianthes sections or series. These are series Meizonodontae Bitter (Dean 1995, 2004), series Microlobae Bitter (Dean et al. 2007), section Synantheroides Bitter (Reyes Cornejo 2015), series Tricolores Bitter (Dean et al. 2017a, 2018c), and series Piliferae Bitter (Dean et al. 2019b). Phylogenetic analyses using morphological characters have been completed by Dean (1995) for series Meizonodontae and Reyes Cornejo (2015) for section Synantheroides. The published work based on morphology indicates that the species united by the sections and series proposed by Bitter may be closely related in a least two groups (series Meizonodontae and series Tricolores), while other groups (series Piliferae and section Synantheroides) contain a mixture of closely and distantly related species.

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 Lycianthes series Meizonodonatae using molecular markers and an analysis of the biogeography of series Meizonodonatae has been completed by the third author at the University of Guadalajara (Anguiano-Constante et al. 2018; Anguiano-Constante 2019). An investigation of the morphology and phylogeny (using molecular markers) of the mostly South American series Strigosulae Bitter is also ongoing at the Universidad Nacional Mayor de San Marcos in Peru (Cueva-Manchego no date). In addition, a number of species of Lycianthes have been included in phylogenetic studies of the Solanaceae (Särkinen et al. 2013), and this partial analysis of the species indicates that Lycianthes may be a paraphyletic group in relation to Capsicum (Spalink et al. 2018), and if monophyly is preferred, the genus may need to be broken into smaller groupings or combined with Capsicum.

Herbarium collections of many of the species of Lycianthes are not numerous, perhaps because many of the species have flowers that close during the day, making them inconspicuous (Nee 1981). In our experience, many Lycianthes species are difficult to locate, often occurring singly or in small populations, which may be another reason why they are poorly collected. This lack of material, along with the problems inherent in dealing with an older monograph such as Bitter’s, have perhaps contributed to the lack of attention given to Lycianthes. It is likely that there are many new species of Lycianthes awaiting discovery in herbaria and in the field.

Goals of this paper

Over the past four years, the first author has been investigating the Lycianthes of Mexico and Central America as part of National Science Foundation-funded research that will produce species descriptions that will be posted at the website Solanaceae Source (http://solanaceaesource.org/). Mexico, known for its high plant species diversity and endemism (Myers et al. 2000; Villaseñor 2016; Sosa et al. 2018), is especially rich in Lycianthes taxa (48 recognized here; Table 1), likely the highest number of Lycianthes taxa of any country in the Neotropics. However, a floristic treatment of the genus has never been completed for the entirety of Mexico, and a summary of what is known about the Mexican species is overdue. In this paper, we provide species descriptions and synonymy (as well as necessary lectotypifications) for all the Lycianthes taxa of Mexico, as well as an identification key, photographs of specimens, illustrations of trichomes, flowers, fruits, and seeds, and distribution maps. As the Mexican state of Chiapas shares many Lycianthes species with neighboring Guatemala, we are also treating the Guatemalan species (Table 1) and updating the treatment of Lycianthes completed for Guatemala over forty years ago (Gentry and Standley 1974). At the end of the treatment, we also summarize difficult to place specimens and excluded taxa.

Table 1.

Geographic distribution of the native Lycianthes of Mexico and Guatemala, excluding the cultivated species L. rantonnetii.

Species Distribution
L. acapulcensis (Baill.) D’Arcy Mexico to Costa Rica
L. amatitlanensis (J.M.Coult. & Donn.Sm.) Bitter Mexico to South America
L. anomala Bitter Mexico
L. armentalis J.L.Gentry Mexico, Guatemala, Belize
L. arrazolensis (J.M.Coult. & Donn.Sm.) Bitter Mexico to Nicaragua
L. barbatula Standl. & Steyerm. Mexico, Guatemala
L. breedlovei E.Dean Mexico
L. caeciliae Bitter Mexico
L. ceratocalycia (Donn.Sm.) Bitter Mexico, Guatemala
L. chiapensis (Brandegee) Standl. var. chiapensis Mexico, Guatemala
L. chiapensis (Brandegee) Standl. var. sparsistellata Standl. & Steyerm. Mexico to Nicaragua
L. ciliolata (M.Martens & Galeotti) Bitter Mexico, Guatemala
L. connata J.L.Gentry Mexico, Guatemala
L. cuchumatanensis J.L.Gentry Guatemala
L. dejecta (Fernald) Bitter Mexico
L. fredyclaudiae E.Dean Guatemala
L. geminiflora (M.Martens & Galeotti) Bitter Mexico
L. glabripetala E.Dean Mexico
L. gongylodes J.L.Gentry Guatemala
L. gorgonea Bitter Mexico, Guatemala, Belize
L. grandifolia E.Dean Mexico
L. heteroclita (Sendtn.) Bitter Mexico to South America
L. hintonii E.Dean Mexico
L. hypoleuca Standl. Mexico to Honduras
L. inconspicua Bitter Guatemala to Panama
L. jalicensis E.Dean Mexico
L. limitanea (Standl.) J.L.Gentry Mexico, Guatemala, Belize
L. manantlanensis Aarón Rodr. & O.Vargas Mexico to El Salvador
L. mariovelizii E.Dean Mexico to Nicaragua
L. michaelneei E.Dean Mexico
L. moziniana (Dunal) Bitter var. margaretiana E.Dean Mexico
L. moziniana (Dunal) Bitter var. moziniana Mexico
L. moziniana (Dunal) Bitter var. oaxacana E.Dean Mexico
L. nitida Bitter Mexico to Panama
L. ocellata (Donn.Sm.) C.V.Morton & Standl. Mexico and Guatemala
L. orogenes Standl. & Steyerm. Mexico and Guatemala
L. peduncularis (Schltdl.) Bitter Mexico
L. pilifera (Benth.) Bitter Mexico
L. pringlei (B.L.Rob. & Greenm.) Bitter Mexico
L. purpusii (Brandegee) Bitter Mexico to Honduras
L. quichensis (J.M.Coult. & Donn.Sm.) Bitter Mexico and Guatemala
L. rafatorresii E.Dean Mexico
L. rzedowskii E.Dean Mexico
L. scandens (Mill.) Nee var. flavicans (Bitter) J.Poore & E.Dean Mexico to Costa Rica
L. scandens (Mill.) Nee var. scandens Mexico to South America and the Caribbean
L. sideroxyloides (Schltdl.) Bitter Mexico to Nicargua
L. starbuckii E.Dean Mexico
L. stephanocalyx (Brandegee) Bitter Mexico to Honduras
L. surotatensis Gentry Mexico
L. textitlaniana E.Dean Mexico
L. tricolor (Dunal) Bitter Mexico to El Salvador
L. venturana E.Dean Mexico

Materials and methods

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, CR, DAV, DUKE, E, F, G, GBH, GH, GOET, HAL, HBG, HCIB, IBUG, IEB, INBIO, JE, K, LD, LE, LL, M, MA, MEXU, MICH, MO, MPU, MSB, MSC, NDG, NY, P, PH, SERO, TEX, U, UC, UCR, US, VT, W, WIS, WU, XAL, Z, and ZEA (herbarium codes follow Thiers 2019).

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 I. We also chose representative specimens from the Mexican/Guatemalan distribution of each taxon (one specimen from each state/department where it occurs) and listed them at the end of each taxon treatment. Specimens in these sections are organized alphabetically first by country, then by state/department, then alphabetically by collector’s surname, and then numerically by collection number. No barcode or accession numbers are cited. Because the first author saw and confirmed the identification of all specimens listed in Appendix I and the “Representative Specimens Examined” sections, we did not use exclamation points to distinguish the specimens that we examined.

In order to create maps of the Mexican species, specimens were georeferenced by using either Geolocate, an online software-mapping package (Rios and Bart 2010), or manually, using Google Earth Pro (Sullivan 2009); for Mexican specimens, the latter was often used in conjunction with location data found in the Mexican Archivo Histórico de Localidades [The Archive of Mexican Historical Locations] (INEGI 2010). In lists of specimens examined, when coordinates were added to a specimen record, we provide those coordinates in brackets; if the coordinates are not in brackets, they were provided on the specimen label.

The georeferenced location data were analyzed in order to provide conservation assessments for each species using GeoCAT (Bachman et al. 2011); we calculated the Extent of Occurrence (EOO) and the Area of Occupancy (AOO) in km2, assuming cells of 2 km on a side. Preliminary assessment categories were proposed following the criteria of the International Union for Conservation of Nature (IUCN 2019).

Our species concept is a morphological one (Cronquist 1978), based on measurement of herbarium specimens and field and greenhouse observations. In separating the species, we emphasize discontinuities in habit, pubescence, leaf, floral, fruit, and seed characters that are well preserved on herbarium specimens. The species recognized in this paper are well-defined entities. As discussed below, we are recognizing 53 taxa, including 49 species and seven varieties, from Mexico and Guatemala.

Morphology and distribution

Habit

Lycianthes life forms (all perennial) include herbs, vines, shrubs, and treelets, with a few species epiphytic (Fig. 2). Most species are shrubs or vines (Fig. 2A, B); many species are intermediate between the two life forms and described as scandent shrubs to vines. Some shrubs, such as L. jalicensis, are rhizomatous and form clonal thickets (illustrated in Dean et al. 2017a). A minority of the species are true herbs (Fig. 2C, D). The eight species of series Meizonodontae Bitter (L. acapulcensis, L. ciliolata (Fig. 2C), L. dejecta, L. hintonii, L. moziniana, L. peduncularis (Fig. 2D), L. rzedowskii, and L. starbuckii) are true herbs, arising from and dying back to tuberous roots annually, only appearing above-ground with the commencement of the rainy season (Dean 2004). Depending on climate and habitat, L. stephanocalyx can grow as a rhizomatous herb, dying back to rhizomes annually, or it can persist above ground from season to season becoming woody at the base (illustrated in Dean et al. 2019b). Lycianthes heteroclita (Fig. 2A), L. geminiflora, and L. ceratocalycia have green, herbaceous primary stems when young but become shrubs or treelets with age; thus those three species are sometimes described on specimen labels as herbs. Lycianthes amatitlanensis, L. glabripetala, and L. inconspicua are very short subshrubs that are often described as herbs, although they are generally woody and persist above ground. Two species in Mexico and Guatemala are nearly always epiphytic (L. anomala and L. nitida), and L. heteroclita sometimes sprouts on the branches of trees but is usually found rooted in the ground.

Figure 2. 

Lycianthes habit A free-standing shrub habit in L. heteroclita (Sendtn.) Bitter, field photo taken by the first author at El Chocoyero, Nicaragua B vine habit in L. gorgonea Bitter, field photo of Dean 9528 (DAV) C upright herb habit in L. ciliolata (M.Martens & Galeotti) Bitter, field photo of Dean 225 (DAV) D prostrate herb habit in L. peduncularis (Schlechtd.) Bitter, field photo of Dean 230 (DAV). Scale bars: 10 cm.

The sympodial branching pattern of Lycianthes is similar to that of Solanum and has been described and illustrated by Child and Lester (1991), Bohs (1994), and Dean (2004). In both herbs and shrubs, initial stem growth begins with an upright sympodial unit (or trunk) with multiple spirally-arranged leaves; the trunk usually terminates in an inflorescence. Growth continues by the production of upper sympodial units that emerge either singly (monochasial branching) or in pairs (dichasial branching) beneath the inflorescence of the prior sympodial unit. If the plant senesces or is damaged, the entire growth cycle can begin again from axillary buds, with the first sympodial unit duplicating the form, leaf number, and leaf arrangement of the trunk. In the species descriptions, we provide information on the length and internode number of the first sympodial unit for the true herbs of series Meizonodontae, because it is of significant length and a highly visible portion of the mature plant in most of the species. In contrast, we only provide information on the upper sympodial growth of all the other species (shrubs and vines).

Indument

Trichome type is important in the identification of many Lycianthes species; trichome density, however, can be quite variable within a species. The trichome types of the Lycianthes of Mexico and Guatemala include simple (Fig. 3A–D), furcate (Fig. 3E), dendritic (Fig. 3F), true stellate (uncommon in the species covered in this paper and not illustrated here, however illustrations are available, for example in Payne (1978 [fig. 43]) or Harris and Harris (1994 [fig. 1492]), multangulate-stellate (Fig. 3G, H), and geminate-stellate (Fig. 3I)). All trichomes are uniseriate (one-cell wide). Trichome terms are taken from Roe (1968, 1971) and follow current usage in the Solanaceae (Sampaio et al. 2014), although some current papers use the term multiangulate trichome rather than multangulate trichome (used by Roe and used here).

Figure 3. 

Lycianthes trichomes A simple trichomes that do not flatten upon drying: left, in L. purpusii, from Beaman 5130 (NY); right, in L. pilifera, from Lorence 4035 (CAS) B simple trichomes that flatten upon drying: left, with flattened cells oriented alternately at right angles to one another in L. quichensis, from Breedlove 31746 (CAS); right, with flattened cells not oriented at right angles to one another in L. acapulcensis, from Dean 249 (DAV) C simple, curved trichome that does not flatten upon drying in L. tricolor, from Dean 297 (DAV) D glandular, simple trichomes: left, glandular trichome with ovoid glandular tip in L. textitlaniana, from Zarate Marcos AZM-274 (MEXU); right, glandular trichome with globose tip in L. surotatensis, from Rzedowski 43384 (DAV) E furcate trichome in L. purpusii, from Beaman 5130 (NY) F dendritic trichome in L. dejecta, from Dean 261 (DAV) G multangulate-stellate trichome in L. armentalis, from Breedlove 56055 (MO) H multangulate-stellate trichome with rebranched rays in L. breedlovei, from Breedlove 34793 (MO) I geminate-stellate trichome in L. sideroxyloides, from Dean 9526 (DAV). Scale bars: 0.25 mm.

Furcate (forked) trichomes (Fig. 3E) resemble simple trichomes but are forked at the tip. Dendritic trichomes (Fig. 3F) are repeatedly forked with no more than two branches per node. Stellate, multangulate-stellate (Fig. 3G, H), and geminate-stellate (Fig. 3I) trichomes have more than two branches per node, and these whorled branches are called rays. True stellate trichomes are uncommon in the species included here (mainly seen in Lycianthes hypoleuca); true stellate trichomes have all rays emerging at just one node, with the rays in one plane. Multangulate-stellate trichomes have all rays emerging at just one node, but the rays are in more than one plane. In several species, the rays of the multangulate-stellate trichomes may be rebranched (Fig. 3H) (Dean et al. 2019a). In the species that have multangulate-stellate trichomes with rebranched rays, trichomes with both unbranched and rebranched rays are present, and the intermediate forms can be seen, which is why we do not refer to these trichomes as dendritic. Geminate-stellate trichomes have rays emerging at numerous consecutive nodes; in the species covered in this paper, geminate-stellate trichomes often have numerous rays per node (more than five) and the rays are relatively short (Fig. 3I). We are using the term geminate-stellate trichomes, as we are following the terminology of Roe (1968, 1971); however, some authors refer to these trichomes as candelabra trichomes (Payne 1978).

In some species with simple trichomes, the cells collapse and flatten upon drying, appearing like a flattened ribbon (Fig. 3B); in some species, the adjoining flattened cells are oriented to one another at 90 degree angles (e.g. L. moziniana and L. quichensis) (Fig. 3B). In contrast, some species have trichomes that remain cylindrical with a pointed tip (e.g. L. amatitlanensis, L. inconspicua, L. glabripetala, and L. pilifera) (Fig. 3A). In most species with furcate, dendritic, and the various types of stellate trichomes, the cells do not collapse, but remain cylindrical upon drying. An exception to this is the dendritic to multangulate-stellate trichomes of L. dejecta, which do flatten upon drying.

Only four species included here have glandular trichomes (Fig. 3D) as a major component of stem, leaf, and/or inflorescence pubescence (L. surotatensis, L. textitlaniana, L. pringlei, L. gorgonea). In all cases, these species have simple trichomes with a globose (L. suratotensis, L. gorgonea) to ovoid (L. pringlei, L. textitlaniana) glandular tip. Minute glandular trichomes are sometimes located inside the calyces or at the tips of the corolla lobes in some species, but these can only be seen with high magnification.

Leaves

The upper sympodial units of the Lycianthes covered here are difoliate. In many species, the leaves are in geminate pairs, often with one leaf smaller than the other, sometimes with the leaves of very different sizes and shapes. In some species, the smaller leaf fails to develop, and the sympodia appear to be unifoliate. The leaves are simple, usually petiolate, and have mostly entire but undulate margins. In a few populations of a few species the margin can sometimes have irregular, sparse, coarse dentations (e.g. L. tricolor and L. surotatensis), but this characteristic is uncommon and unpredictable. The texture of the leaf blades ranges from membranaceous to coriaceous.

Inflorescence

The sympodial units of Lycianthes terminate in inflorescences. The majority of inflorescences are located on the upper sympodia and appear axillary. The peduncle in most Lycianthes is reduced and usually not visible, resulting in an umbellate inflorescence of one to many pedicelled flowers; in a minority of species (e.g. L. amatitlanensis, L. nitida), a short peduncle is present, often covered with the many pedicel scars of fallen flowers. In a minority of the species, the inflorescence is always a solitary flower (e.g. all the herb species of series Meizonodontae, L. stephanocalyx, L. textitlaniana, L. amatitlanensis, L. gorgonea); all the other species can have one to many flowers. However, usually only one to a few flowers at a time are present at a single axil.

Pedicels

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 Meizonodontae). In other species, the pedicel is completely erect, and the flower is oriented upwards as it develops (e.g. L. heteroclita). In L. amatitlanensis and its close relatives L. glabripetala and L. inconspicua, the pedicels are usually deflexed and held beneath the leaves with the flowers nodding beneath the plant.

Calyx

In all Lycianthes, there are no lobes on the synsepalous calyx; rather the calyx is a cup-shaped structure with a truncate rim. Below this rim, calyx appendages may emerge (Fig. 1A, B). In most species, the length of the calyx margin between the rim and the appendages is short, less than 0.5 mm long. In a few species, the margin is conspicuous, as a long as 2 mm long (e.g. Lycianthes connata [Fig. 1C]). A very few species have an undulate calyx rim, sometimes with a large papery margin, or a rim that tears with age (e.g. L. pringlei [Fig. 1D], L. manantlanensis, L. limitanea); in these cases, the margin can sometimes resemble unequal calyx lobes. Appendages are usually linear in shape, sometimes subulate (wider at the base and very pointed at the tip, e.g. L. pilifera [illustrated in Fig. 8B of Dean et al. 2019b]), more rarely widely obovate with a broadly rounded tip (e.g. L. sideroxyloides [Fig. 1E], L. ocellata) or linear with an enlarged and bulbous tip (e.g. L. rafatorresii). In a minority of the species, appendages are lacking completely (e.g. L. heteroclita, L. geminiflora, L. nitida) or can be reduced to mere bumps on the calyx (e.g. L. pringlei [Fig. 1D], L. ceratocalycia [Fig. 1F]). In two of the species included here (L. anomala, L. connata), the appendages are connected to one another at their base, forming a shelf that reflexes as a unit as the flower ages and the fruit develops (Fig. 1C).

Corolla

The corolla in Lycianthes is gamopetalous with a short tube that is inserted into the calyx. The limb is five-lobed, with the lobes usually connected to various degrees by thinner “interpetalar” tissue (Fig. 4A–D). In some species, the interpetalar tissue is lacking, and the limb is divided nearly to the tube (Fig. 4E, F). We use the term “stellate” for corollas that are divided shallowly to deeply and the term entire for corollas that are undivided. For consistency with past publications on Lycianthes (Dean 2014; Dean et al. 2007, 2017a, 2018c, 2019a, b), we are using the terms rotate, campanulate, and reflexed to indicate the orientation of the corolla limb in relation to the tube. We are using the term rotate as it is defined in Lawrence (1951, pg. 768) and Radford et al. (1974, pg. 104) to mean “… a gamopetalous corolla with a flat and circular limb at right angles to the short obsolete tube.” The illustrations in Lawrence (fig. 315, pg. 754) and Radford et al. (fig. 6.6 pg. 101) show a divided limb, indicating that the limb does not have to be entire to be rotate, and so we use the term rotate to mean a fully open wheel-shaped corolla, even if it is somewhat stellate. This usage of rotate differs from usage of the term in works on the genus Solanum to mean an unlobed, entire limb (see for example Spooner et al. 2004). We use the positional term campanulate for limbs that are oriented to the tube at greater than a right angle and the term reflexed for limbs that are oriented to the tube at less than a right angle.

Figure 4. 

Lycianthes corollas and stamens A flower of L. pringlei with corolla slightly campanulate to rotate in orientation, entire in outline, abundant interpetalar tissue, and green markings at the base of the lobes, the stamens unequal and free; field photo of Dean 327 (DAV) B flower of L. connata with corolla rotate in orientation, very shallowly stellate in outline, abundant interpetalar tissue (with the lobes protruding beyond the tissue), and purple markings at the base of the lobes, the stamens unequal and free; field photo of Dean 9530 (DAV) C flower of L. chiapensis (Brandeg.) Standl. var. sparsistellata Standl. & Steyerm. with corolla rotate in orientation, shallowly-stellate in outline, abundant interpetalar tissue, and green markings at the base of the lobes, the stamens unequal and free; field photo of Dean 9507 (DAV) D flower of L. sideroxyloides with corolla slightly campanulate to rotate in orientation, stellate in outline, and interpetalar tissue connecting just the lower portions of the lobes, the stamens equal and free; field photo of Dean 9512 (DAV) E flower of L. heteroclita with corolla slightly campanulate to rotate in orientation, deeply-stellate in outline, and no interpetalar tissue, the stamens equal with partially connate anthers; field photo of Dean 9500 (DAV) F flower of L. ceratocalycia (Donn.Sm.) Bitter with corolla rotate in orientation, deeply-stellate in outline, sparse interpetalar tissue at edges of lobes, and purple markings at the base of the lobes, the stamens equal and free; field photo of Dean 9532 (DAV). Scale bars: 1 cm.

The corollas of Lycianthes usually exhibit diurnal movements, opening and closing each day for several days in a row (Dean 2001). The corollas of many Mexican and Guatemalan species open at daybreak and close by late morning; herbarium specimens of species with this pattern often have closed corollas. Exceptions to this pattern are the morphologically similar and perhaps closely-related L. glabripetala, L. inconspicua, and L. amatitlanensis, as well as the horticultural species L. rantonnetii (Carr.) Bitter, which have flowers that stay open for most of the day and close at night; herbarium specimens of these species usually have open corollas. As explained elsewhere (Dean et al. 2017a), with only closed corollas available for study, it is often difficult to measure corolla diameter on specimens and most accurate to measure corolla length from the bottom of the calyx tube to the tip of the corolla lobe. It may also be difficult to know the typical corolla orientation (campanulate, rotate, reflexed), which can also affect a diameter measurement. Therefore, corolla size in this paper is usually just given as length. An exception to this is species that were studied extensively in the field and/or greenhouse (such as the herbs of series Meizonodontae); in those cases, floral diameter is also given.

The color of the corollas of Lycianthes included here range from white to purple on the adaxial side; the abaxial side may be the same color as the adaxial side, or the lobes may be green. As described elsewhere (Dean 2004; Dean et al. 2017a, 2019b), many Lycianthes have darker-colored markings on the adaxial side of the corolla. These markings range from purple stripes on the lobes, to a ring of purple color at the base, or green or yellow patches of color on the lobes (Fig. 4).

Stamens

Stamens in Lycianthes can be equal (Fig. 4D–F) or unequal (Fig. 4A–C). If unequal, differences in length are due to differences in filament length; anther length within a single flower is usually equal, or nearly so. In Lycianthes species with unequal stamens, the species native to Mexico and Guatemala have one long stamen and four shorter stamens (Fig. 4A–C); the one species included here that is non-native to the region (the cultivated L. rantonnetii) has three long stamens and two short stamens (illustrated in fig. 4 of Barboza and Hunziker 1992).

Anthers can be connate, connivent, or free. They can be glabrous or pubescent; most of the Lycianthes with multangulate- or geminate-stellate trichomes that are covered in this paper have sparsely pubescent anthers. Dehiscence is usually by pores at or near the anther tip, sometimes on the inner or outer face of the anther. An exception to this pattern is found in the Mexican endemic L. anomala. In L. anomala, the anthers are connate and dehisce at their edges, with the pores of one anther being coherent with those of the adjacent anther; these extremely large pores eventually become slits that extend half-way down the anther. This type of dehiscence is also found in the Central American species L. synanthera.

The pollen of most Lycianthes has not been investigated. As part of her monograph on the perennial herbs of series Meizonodonatae, the first author examined the pollen of all the species of the series (Dean 1995, 2004). She found that most species in the series have smooth, triangular, tricolporate pollen. However, two of the species (L. acapulcensis and L. rzedowskii) are dicolporate, one species (L. ciliolata) is dicolporate with a remnant third pore, and L. starbuckii has variable pollen ranging from tricolporate to dicolporate with a remnant third pore (Dean 2004). The first author completed an informal survey of the pollen of eight additional taxa included in this paper (L. anomala, L. geminiflora, L. jalicensis, L. pilifera, L. pringlei, L. scandens var. scandens, L. stephanocalyx, and L. tricolor). All seven taxa have pollen that is smooth and tricolporate, with the shape ranging from spherical to triangular.

Gynoecium

The gynoecium in the Lycianthes species included here is bicarpellate with a conic, ovoid, or globose ovary. The single linear style is straight to curved, and the stigma can be truncate, capitate, or oblong.

Fruits

The fruits of the Lycianthes treated here are berries ranging in shape from depressed-globose to ovoid (Fig. 5A, B). The exocarp can be yellow, orange, red, blue, or dark purple (nearly black) (Fig. 5A, B); immature fruits are usually green (Fig. 5A), but in a minority of species the color changes from green to white before changing to the mature color. In L. barbatula the mature fruit color is still unknown, but it has been listed as white on labels. In species with simple trichomes, the fruit exocarp is usually glabrous; in species with multangulate- or geminate-stellate trichomes, a few trichomes may be present on the exocarp. The mesocarp and placental areas of the fruits can be dry, powdery, or soft (fleshy or juicy). A minority are dry throughout (e.g. L. textitlaniana), while a few of the species in series Meizonodontae have a juicy mesocarp and a powdery placental area (L. acapulcensis, L. ciliolata, L. rzedowskii) (Fig. 5C). In two of the species treated here (L. peduncularis and L. rantonnetii), the mesocarp has numerous yellow sclerotic granules (also called sclereids or stone cells) (Fig. 5D) which sometimes attach to the seeds or are mistaken for seeds.

Figure 5. 

Lycianthes fruits A immature (green) and mature (orange to red) globose berries of L. heteroclita (note the calyx completely lacking appendages), field photo taken by the first author at El Chocoyero, Nicaragua B mature, ovoid, dark purple fruits of L. pilifera, photo of Dean 242 collected at the San Francisco Botanical Garden C longitudinal section of mature, ovoid, dark purple fruits of L. ciliolata showing the light purple, powdery placental area, greenhouse photo of Dean 295 (DAV) D dissected fruit of L. peduncularis showing the black seeds and yellow sclerotic granules, field photo of Dean 303 (DAV). Scale bars: 1 cm.

Seeds

Seed shape terminology used in this paper is taken from Radford et al. (1974); seed surface terminology is taken from Gunn and Gaffney (1974). The seeds of 15 of the species covered in this paper have been illustrated in previous publications (Barboza and Hunziker 1992; Dean 2004; Dean et al. 2007, 2017a, b, 2019b). Most Lycianthes species covered in this paper have compressed seeds, usually lenticular (completely flattened). A minority of species have seeds that are not compressed at all. This includes some of the species in series Meizonodonatae (L. acapulcensis, L. ciliolata, L. peduncularis, L. rzedowskii) and L. rantonnetii. Others are compressed but not lenticular (e.g. L. dejecta, L. moziniana, L. pringlei, L. textitlaniana).

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 (Dean 2004; Dean et al. 2019b), some seeds have fibrils that protrude from the periclinal walls of the reticulum (e.g. L. dejecta, L. pilifera) (Lester and Durrands 1984).

Geographic distribution, habitat and elevation

Twenty-two taxa of Lycianthes are endemic to Mexico, three are endemic to Guatemala, and eight are found in Mexico and Guatemala and no other country (Table 1). The rest of the taxa included here have distributions that extend further into Central America, with a few extending to South America and one to the Caribbean (Table 1). Their elevational ranges include sea level (e.g. Lycianthes scandens var. scandens, L. armentalis) to nearly 4000 m in elevation (L. quichensis). Based on the vegetation classification of Rzedowski (1978), the habitats where these taxa are found include coniferous forest, deciduous forest (including oak forest), cloud forest, tropical dry forest, tropical moist forest, xerophilous scrub and agricultural areas; the most common habitat is oak forest. In common with many Solanaceae, many of the taxa included here are found in light gaps or edges, rather than dark forest.

Conservation status

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 2).

Table 2.

Preliminary conservation assessments for the native Lycianthes of Mexico and Guatemala calculated using GeoCAT.

Species Assessment
L. acapulcensis Least Concern
L. amatitlanensis Least Concern
L. anomala Endangered
L. armentalis Least Concern
L. arrazolensis Least Concern
L. barbatula Endangered
L. breedlovei Endangered
L. caeciliae Endangered
L. ceratocalycia Endangered
L. chiapensis var. chiapensis Endangered
L. chiapensis var. sparsistellata Least Concern
L. ciliolata Least Concern
L. connata Least Concern
L. cuchumatanensis Critically Endangered
L. dejecta Least Concern
L. fredyclaudiae Endangered
L. geminiflora Near Threatened
L. glabripetala Vulnerable
L. gongylodes Critically Endangered
L. gorgonea Least Concern
L. grandifolia Endangered
L. heteroclita Least Concern
L. hintonii Critically Endangered
L. hypoleuca Least Concern
L. inconspicua Endangered
L. jalicensis Vulnerable
L. limitanea Least Concern
L. manantlanensis Least Concern
L. mariovelizii Least Concern
L. michaelneei Endangered
L. moziniana var. margaretiana Near Threatened
L. moziniana var. moziniana Least Concern
L. moziniana var. oaxacana Vulnerable
L. nitida Least Concern
L. ocellata Endangered
L. orogenes Endangered
L. peduncularis Least Concern
L. pilifera Endangered
L. pringlei Endangered
L. purpusii Least Concern
L. quichensis Endangered
L. rafatorresii Least Concern
L. rzedowskii Endangered
L. scandens var. flavicans Least Concern
L. scandens var. scandens Least Concern
L. sideroxyloides Least Concern
L. starbuckii Endangered
L. stephanocalyx Least Concern
L. surotatensis Least Concern
L. textitlaniana Critically Endangered
L. tricolor Least Concern
L. venturana Endangered

Taxonomic treatment

Lycianthes (Dunal) Hassl., Annuaire Conserv. Jard. Bot. Genève 20: 180. 1917. Nom. conserv.

Otilix Raf., Medical Fl. 2: 87. 1830. Nom. rej. Type: Solanum lycioides L.

Solanum subsect. Lycianthes Dunal, Prodr. [A. P. de Candolle] 13(1): 29. 1852. Type: Solanum lycioides L. (designated by D’Arcy 1972, pg. 211)

Parascopolia Baill., Hist. Pl. 9: 338. 1888. Nom. rej. Type: P. acapulcensis Baill.

Solanum sect. Lycianthes (Dunal) Wettst., Nat. Pflanzenfam. 4(3b): 22. 1891. Type: Based on Solanum subsect. Lycianthes Dunal

Solanum subgenus Lycianthes (Dunal) Bitter, Bot. Jahrb. Syst. 54: 424. 1917. Type: Based on Solanum subsect. Lycianthes Dunal

Type

Based on Solanum subsect. Lycianthes Dunal

Description

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.

Discussion

Genus name based on the type species Solanum lycioides, named, presumably, for its thorny branches that resemble the genus Lycium L., which was first described from Lycia, in what is now Turkey.

Artificial key to Mexican and Guatemalan Lycianthes

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 Group 1
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
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 Group 2
Plant lacking obvious glandular trichomes, sometimes with small obscure glandular trichomes inside the calyx or elsewhere 3
3 Simple, furcate and/or dendritic trichomes forming a majority of the indument on stems and leaves, or plant completely glabrous Group 3
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) Group 4

Group 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.

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 L. dejecta
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
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 L. peduncularis
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 L. starbuckii); indument of simple or dendritic trichomes, these spreading or retrorsely appressed (rarely, antrorsely appressed in Chiapan or Guatemalan populations of L. ciliolata) 3
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 4
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 6
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 L. acapulcensis
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
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 L. rzedowskii
First sympodial unit c. 25 cm long, the internodes c. 13; lateral branching from the nodes of the first sympodial unit usually present at the time of flowering; subsequent sympodial growth about equal to the length of the first sympodial unit; pollen tricolporate; state of Nuevo León, on limestone soils L. hintonii
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 7
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 9
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 L. moziniana var. moziniana
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
8 Filaments glabrous; exocarp of berry green; placental area green and juicy; disturbed clearings and agricultural fields of mountains of state of Oaxaca L. moziniana var. oaxacana
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) L. moziniana var. margaretiana
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) L. starbuckii
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 L. ciliolata

Group 2

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 L. pringlei
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
2 Corolla purple; berry turbinate, the tip pointed, orange; seed surface with widely spaced serpentine cell pattern, noticeably pitted; state of Oaxaca L. textitlaniana
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
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 L. surotatensis
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 L. gorgonea

Group 3

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 L. rantonnetii
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
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 3
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 4
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 L. anomala
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 L. barbatula
4 Stamens equal in length or nearly so 5
Stamens unequal in length, one filament noticeably longer than the other four 19
5 Calyx lacking appendages; corolla shallowly to deeply stellate in outline, sometimes without interpetalar tissue 6
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 9
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 L. manantlanensis
Calyx margin entire, truncate, in flower and fruit; corolla stellate, deeply divided to the base, lacking interpetalar tissue; anthers without dark connective 7
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 L. nitida
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
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 L. geminiflora
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 L. heteroclita
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 L. ceratocalycia
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
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 11
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 13
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 L. amatitlanensis
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
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) L. glabripetala
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 L. inconspicua
13 Perennial herb to woody vine or shrub; mature berry red; anthers connate to connivent; usually occurring at or below 1000 m in elevation 14
Herb, shrub or treelet; mature berry orange, red or dark purple; anthers free from one another; usually occurring above 1000 m in elevation 15
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 L. stephanocalyx
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 L. gorgonea
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 L. manantlanensis
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
16 Leaf blades sparsely to densely pubescent, the trichomes collapsing when dry; berry orange to red at maturity; Mexico (state of Chiapas) and Guatemala 17
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) 18
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 L. gongylodes
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 L. quichensis
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) L. caeciliae
Corolla white to light purple with dark purple ring and green markings at base adaxially, nearly entire in outline; Mexico (state of Oaxaca) L. pilifera
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 20
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 21
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 L. manantlanensis
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 L. orogenes
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 L. purpusii
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
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 L. connata
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
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) 24
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 L. arrazolensis in the state of Oaxaca); stem trichomes spreading to appressed; many different regions of Mexico and Guatemala, south to Nicaragua 25
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) L. venturana
Apex of berry rounded; calyx 2–4 mm long; Mexico (state of Jalisco) L. jalicensis
25 Corolla lobes and interpetalar membrane purple; stem trichomes usually densely matted; some leaf trichomes to 2 mm long; Mexico (state of Veracruz) L. michaelneei
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
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) L. grandifolia
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
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 L. mariovelizii
Calyx appendages usually < 5 mm long (rarely to 5 mm in L. arrazolensis in state of Guerrero); base of appendages not flattened; 500–3000 m 28
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 L. tricolor
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 L. arrazolensis

Group 4

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 2
Stamens obviously unequal, one filament much longer than the other four 7
2 Calyx lacking appendages, the rim often undulate, lobed or torn 3
Calyx with appendages greater than or equal to 0.25 mm long, the rim rarely undulate, lobed or torn 4
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 L. limitanea
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 L. hypoleuca
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 L. breedlovei
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
5 Calyx appendages with large, oblong, glandular area at tip, this area turning black upon drying L. ocellata
Calyx appendages lacking glandular area at tip, remaining green upon drying 6
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 L. cuchumatanensis
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 L. sideroxyloides
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 L. purpusii
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
8 Indument of stalked multangulate-stellate trichomes, the rays straight and usually rebranched, sometimes repeatedly; states of Chiapas, Mexico and Baja Verapaz, Guatemala 9
Indument of sessile to stalked multangulate-stellate and/or geminate-stellate trichomes, the rays rarely rebranched; widely distributed in Mexico and Guatemala 10
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) L. breedlovei
Corolla entire in outline, pale white to lilac, without darker purple color on the lobes adaxially; Guatemala (state of Baja Verapaz) L. fredyclaudiae
10 Trichomes on leaves, stems, and calyx white to tan, not yellow, orange or brown 11
At least some trichomes on leaves, stems, and calyx yellow, orange, or brown, sometimes mixed with tan trichomes 13
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 L. rafatorresii
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
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 L. scandens var. scandens
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 L. scandens var. flavicans
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 L. rafatorresii
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
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 L. armentalis
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
15 Calyx 4–5 mm long in flower, 6–8 mm long in fruit L. chiapensis var. chiapensis
Calyx 2–3.5 mm long in flower, 2–4 mm long in fruit L. chiapensis var. sparsistellata

Species descriptions

Lycianthes acapulcensis (Baill.) D’Arcy, Solanaceae Newsl. 2(4): 23. 1986

Fig. 6

Parascopolia acapulcensis Baill., Hist. Pl. (Baillon) 9: 339. 1888. Type: Mexico. Guerrero: Acapulco, Punto Griffon, 1888, C. Thiébaut 1002 (lectotype, designated here: P [P00070403]).

Lycianthes grandifrons Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 418. 1919. Type: Costa Rica. San José: Llanos de Turrucares, 600 m, 18 Sep 1888, H. Pittier & T. Durand 478 (holotype: BR [000000552872]; isotypes: CR [mixed collection with Witheringia solanacea L’Hér.], US [00027878]).

Lycianthes guatemalensis Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 419. 1919. Type: Guatemala. Retalhuleu: Retalulëu [Retalhuleu], May 1877, K. Bernoulli & O. Cario 2384 (lectotype, designated by Dean 2004, pg. 393: GOET [GOET003442]).

Lycianthes somniculenta (Kunze ex Schltdl.) Bitter var. cladotricha Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 413. 1919. Type: Mexico. Morelos: Cuernavaca, in moist copses, 5000 ft, Jun–Jul 1896, C. Pringle 6399 (lectotype designated by Dean 2004 pg. 395: MEXU [00029023]; isolectotypes: B [not seen, cited by Bitter 1919, probably destroyed], BM [000514912], BR [000000552840], E [E00570140], G [G00343072, G00343073], GH [00021855], GOET [GOET003441, GOET003440], HBG [HBG-511362], JE [JE00004691], K [K000063119], M [M-0166091], MEXU [00029022, 00029023], MO [MO-153222], NDG [NDG45130], NY [00138707], PH [00016314], S [S-G-9982], UC [104211], W [acc. # 1897-4064], WRSL [cited by Bitter 1919, not seen], WU [acc. # 037953], Z [Z-000028495]).

Lycianthes somniculenta (Kunze ex Schltdl.) Bitter var. ramosipila Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 413. 1919. Type: Cultivated in Paris (?) “plaines de terre froide”, JDP (Jardin des Plantes) 82 (holotype: P [P00070402]).

Lycianthes villosula Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 420. 1919. Type: Costa Rica. Alajuela: El Brazil, gorges of Virilla River, 800 m, 14 Jul 1911, H. Pittier 3676 (holotype: US [00027904]).

Type

Based on Parascopolia acapulcensis Baill.

Figure 6. 

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

Description

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.

Chromosome number

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

Distribution and habitat

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

Figure 7. 

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

Common names and uses

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

Phenology

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.

Preliminary conservation status

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

Discussion

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

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

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

Representative specimens examined

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

Lycianthes amatitlanensis (J.M.Coult. & Donn.Sm.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 441. 1919

Fig. 8

Solanum amatitlanense J.M.Coult. & Donn.Sm., Bot. Gaz. 37: 420. 1904. Type: Guatemala. Alta Verapaz: Cubilqüitz [Cubilhuitz], [15.6675, -90.4293], 350 m, Feb 1903, H. von Tuerkheim 8488 (lectotype designated by Dean and Reyes 2018a, pg. 40: US [01269192]; isolectotypes: F [0073066F, acc. # 185826], M [M-0171813], NY [00138963, 00138964], US [01014253]).

Solanum sylvicola Brandegee, Univ. Calif. Publ. Bot. 6: 373. 1917. Type: Mexico. Chiapas: Finca Irlanda, Jun 1914, C. Purpus 7315 (holotype: UC [acc. # 173378]; isotypes: HBG [HBG-511491], M [M-0171815]).

Type

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

Figure 8. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 9. 

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

Common names and uses

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

Phenology

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

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes anomala Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 514. 1919

Fig. 10

Type

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

Figure 10. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 11. 

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

Common names and uses

None known.

Phenology

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

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes armentalis J.L.Gentry, Phytologia 26: 269. 1973

Fig. 12

Type

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

Figure 12. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 13. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

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

Fig. 14

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

Type

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

Figure 14. 

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

Description

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

Chromosome number

2n = 24 (Gentry and Pearce 1977).

Distribution and habitat

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

Figure 15. 

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

Common names and uses

None known.

Phenology

Flowering specimens have been collected from February through October; specimens with mature fruits have been collected January through December. The corollas of this species are open in the morning and closed by late morning.

Preliminary conservation status

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

Discussion

This species is very similar to Lycianthes tricolor. It can be easily distinguished from L. tricolor by seed shape. The seeds of L. tricolor have a definite sharp notch that is usually deeper than 0.5 mm, whereas the seeds of L. arrazolensis lack this notch. In some Mexican L. arrazolensis populations in the states of Morelos and México, the seeds are shallowly indented on one side, but this indentation is usually less than 0.25 mm and never sharply notched (Dean et al. 2017a). Non-fruiting specimens can be challenging to identify. If a specimen was collected below 1700 m in elevation, it is most likely to be Lycianthes arrazolensis. In addition, the following non-seed characters can be helpful: the calyx rim of L. arrazolensis tends to be more prominent, often protruding beyond the appendage insertion by over 0.5 mm, while the calyx rim of L. tricolor is usually less than 0.5 mm; the appendages of L. arrazolensis tend to bend away from the rim, exposing the rim, while the appendages of L. tricolor are oriented closer to the rim and corolla, hiding the rim; the pores of the short stamens in L. arrazolensis usually face away from the style, while those of L. tricolor usually face toward the style; the pedicels of the oldest, largest flowers and the pedicels of the most mature fruits of L. arrazolensis are usually shorter than those of L. tricolor, although there is overlap in this characteristic; and the leaves of typical L. arrazolensis tend to have obvious geminate leaf pairs with elliptic to obovate laminas and leaf bases often attenuate into the petiole, while in L. tricolor the small geminate leaf often abscises early, and the laminas are more ovate with a less-attenuate leaf base (Dean et al. 2017a).

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

Representative specimens examined

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

Lycianthes barbatula Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23(5): 228 1947

Fig. 16

Type

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

Figure 16. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 17. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes breedlovei E.Dean, Phytotaxa 409: 265. 2019

Fig. 18

Type

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

Figure 18. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 19. 

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

Common names and uses

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

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimen examined

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

Lycianthes caeciliae Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 429. 1919

Fig. 20

Type

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

Figure 20. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 21. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimen examined

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

Lycianthes ceratocalycia (Donn.Sm.) Bitter Abh. Naturwiss. Verein Bremen 24 [preprint]: 498. 1919

Fig. 22

Brachistus ceratocalycius Donn.Sm. Bot. Gaz. 48: 297. 1909. Type: Guatemala. Dept. Alta Verapaz: In silva montana prope Cobán, 1600 m, Jan 1908, H. von Tuerkheim II 2060 (lectotype designated by Dean and Reyes 2018a, pg.40: US [00624002]; isolectotypes: BM [000514916], BR, E [E00190707], F [0072757F, acc. # 680315], G [G00379123], M [M-0171534], NY [00007074], US [00624001], W [acc. # 1908-5994]).

Type

Based on Brachistus ceratocalycius Donn.Sm.

Figure 22. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 23. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes chiapensis (Brandegee) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 173. 1936

Solanum chiapense Brandegee, Univ. Calif. Publ. Bot. 6: 192. 1914. Type: Mexico. Chiapas: Finca Irlanda, Jun 1914, C. Purpus 7328 (holotype: UC [UC175061]; isotypes: BM [BM000775080]; GH [00077473]; MO [acc. # 765046]; NY [00138972]; US [00027504]).

Type

Based on Solanum chiapense Brandegee.

Lycianthes chiapensis var. chiapensis

Fig. 24

Lycianthes nyssifolia Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 366. 1919.

Type

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

Figure 24. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 25. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes chiapensis (Brandegee) Standl. var. sparsistellata Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 22(4): 274 1940

Fig. 26

Type

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

Figure 26. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 27. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes ciliolata (M.Martens & Galeotti) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 410. 1919

Fig. 28

Solanum ciliolatum M.Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12: 140. 1845. Type: Mexico. Oaxaca: Cordillère orientale de’Oaxaca, dans les bois de la Sierra de Capulalpan et de Llano Verde, Rocheu de Castrasana, de 6,000 à 7,000 pieds, Sep 1840, H. Galeotti 1230 (lectotype designated by Dean 2004 pg. 399: BR [000000552873]; isolectotypes: BR [000000552906, 000000552842]).

Solanum somniculentum Kunze ex Schltdl., Linnaea 19: 306. 1847. Type: Germany. Grown by G. Kunze in the Leipzig Botanical Garden from seed brought from Mexico by C.A. Ehrenberg, 1845, G. Kunze s.n. (lectotype designated by Dean 2004, pg. 399: W [acc. # 0074704]; isolectotype: HAL [acc. # 100603]).

Solanum andrieuxii Dunal, Prodr. [A. P. de Candolle] 13(1): 165. 1852. Type: Mexico. No exact locality (probably from southern Puebla or Oaxaca), 1832?, G. Andrieux 195 (holotype: G-DC [G00145630]; isotype: M [M0171802, mistakenly listed as the holotype in Dean (2004)]).

Lycianthes ciliolata (M.Martens & Galeotti) Bitter var. pratorum Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 411. 1919. Type: Guatemala. Baja Verapaz: Patal, 1600 m, H. von Tuerkheim II 2317 (lectotype designated by Dean 2004 pg. 399: W [acc. # 9874]; isolectotypes: BR [000000552875], E [E00570142], F [0072905F, acc. # 574699], G [G00343162], MO [acc. # 3834546], NY [00023650], US [00479453]).

Lycianthes mociniana Dunal var. andrieuxi (Dunal) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 409. 1919. Type: Based on Solanum andrieuxii.

Lycianthes somniculenta (Kunze ex Schltdl.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 411. 1919. Type: Based on Solanum somniculentum Kunze ex Schltdl.

Lycianthes somniculenta (Kunze ex Schltdl.) Bitter var. lanceolata Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 414. 1919. Type: Mexico. Oaxaca: no exact locality, 1842, Ghiesbrecht 81 (holotype: P [P00070404]).

Type

Based on Solanum ciliolatum M.Martens & Galeotti.

Figure 28. 

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

Description

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.

Chromosome number

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

Distribution and habitat

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

Figure 29. 

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

Phenology

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

Common names and uses

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

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes connata J.L.Gentry, Phytologia 26: 271, 1973

Fig. 30

Type

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

Figure 30. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 31. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes cuchumatanensis J.L.Gentry, Phytologia 26: 273. 1973

Fig. 32

Type

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

Figure 32. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 33. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes dejecta (Fernald) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 415. 1919

Fig. 34

Solanum dejectum Fernald, Proc. Amer. Acad. of Arts: 35: 569. 1900. Type: Mexico. Durango: near city of Durango, Iron Mountain and vicinity, rare in crevices of rocks, July 1896, E. Palmer 347 (lectotype designated by Dean 2004 pg. 404: GH [00077482]; isolectotypes: BM [000514923], C, CAS[acc. # 162966], E [E00526483], F [0073089F, acc. # 51446], G [G00343134], K [K000063110], MEXU [MEXU00029058], MO [acc. # 2495231, acc. # 2495232, acc. # 2495233], NY [00214383], S [acc. # S-G-9980], UC [acc. # 104212, acc. # 124634], US [00027543]).

Lycianthes dejecta (Fernald) Bitter var. palmeri Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 416. 1919, nom. illeg. Type: as above, E. Palmer 347 (holotype: B [not found, cited by Bitter (1919), probably destroyed]; isotypes as listed above).

Type

Based on Solanum dejectum Fernald.

Figure 34. 

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

Description

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.

Chromosome number

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

Distribution and habitat

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

Figure 35. 

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

Common names and uses

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

Phenology

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

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes fredyclaudiae E.Dean, Phytotaxa 409: 268. 1919

Fig. 36

Type

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

Figure 36. 

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

Description

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

Chromosome number

Unknown.

Distribution and habitat

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

Figure 37. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimen examined

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

Lycianthes geminiflora (M.Martens & Galeotti) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 497. 1919

Fig. 38

Solanum geminiflorum M.Martens & Galeotti, Bull. Acad. Brux. 12 (1): 142. 1845. Type: Mexico. Oaxaca: Chinantla (Toavela), 3000 ft, H. Galeotti 1242 (lectotype designated by Dean and Reyes 2018a, pg. 42: BR [000000552845]; isolectotypes: BR [000000552904], LE [LE00016926]).

Type

Based on Solanum geminiflorum M.Martens & Galeotti.

Figure 38. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 39. 

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

Common names and uses

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

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes glabripetala E.Dean, Phytologia 100: 28, 2018

Fig. 40

Type

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

Figure 40. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 41. 

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

Common names and uses

None known.

Phenology

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

Preliminary conservation status

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

Discussion

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

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

Representative specimens examined

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

Lycianthes gongylodes J.L.Gentry, Phytologia 26: 274. 1973

Fig. 42

Type

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

Figure 42. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 43. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes gorgonea Bitter, Repert. Spec. Nov. Regni Veg. 20: 364. 1924

Fig. 44

Solanum cuspidatum C.V.Morton, Contr. Univ. Michigan Herb. 4: 25. 1940. Type: Belize, El Cayo District, on Arenal-Valentín road along roadside through high forest, 20–21 Jun 1936, C. Lundell 6172 (holotype: US [acc. # 1688328]; isotypes: GH [00077476], LL [00372872], MI [1109941], MO [acc. # 1278667], NY [00138976], S [acc. # 04-2899]), TEX [00372873]).

Lycianthes cuspidata (C.V.Morton) Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 18. 1943. Type: Based on Solanum cuspidatum C.V.Morton.

Type

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

Figure 44. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 45. 

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

Common names and uses

None known.

Phenology

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

Preliminary conservation status

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

Discussion

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

Representative specimens examined

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

Lycianthes grandifolia E.Dean, Brittonia 70: 479. 2018

Fig. 46

Type

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

Figure 46. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 47. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

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

Discussion

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

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

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

Representative specimen examined

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

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

Fig. 48

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

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

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

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

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

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

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

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

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

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

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

Type

Based on Solanum heteroclitum Sendtn.

Figure 48. 

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

Description

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

Chromosome number

Unknown.

Distribution and habitat

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

Figure 49. 

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

Common names and uses

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

Phenology

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

Preliminary conservation status

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

Discussion

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

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

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

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

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

Representative specimens examined

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

Lycianthes hintonii E.Dean, Bot. J. Linn. Soc. 145: 407. 2004

Fig. 50

Type

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

Figure 50. 

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

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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

Figure 51. 

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

Common names and uses

None known.

Phenology

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

Preliminary conservation status

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

Discussion

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

Representative specimen examined

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

Lycianthes hypoleuca Standl., Trop. Woods 9: 12. 1927

Fig. 52

Solanum hypoleucum (Standl.) C.V.Morton, Contr. Univ. Michigan Herb. 4: 27. 1940. Type: Based on Lycianthes hypoleuca Standl.

Type

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

Figure 52. 

Image of herbarium specimen of L. hypoleuca, Mendéz 9067 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 53).

Figure 53. 

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

Common names and uses

Mexico. Chiapas: tumat zak (Tzeltal) (A. Méndez G. 9067).

Phenology

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 (Smith and Knapp 2002).

Preliminary conservation status

Lycianthes hypoleuca is a species ranging from southern Mexico to Honduras, represented by 33 collections and occurring in four protected areas. The EOO is 78,970.5 km2, and the AOO is 124 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes hypoleuca is a distinctive species of lowland Caribbean forest. It is easily identified based on its leaves. The whitish, tomentose, stellate pubescence of the underside of the leaf surface makes the underside much paler than the upper side. The pollination of this species was studied in Belize by Smith and Knapp (2002) and they found that a number of different bee species visit the flowers.

Representative specimens examined

Guatemala. Petén: 2 mi S of entrance of Tikal National Park, [17.2107, -89.6247], 500 ft, 19 Jun 1973, T.B. Croat 24707 (MO). Mexico. Campeche: Mpio. Calakmul, N de Rancho Ek Sacrificio, camino a nuevo centro de población Ejidal Ley de Fomento Agropecuario, 17.9897, -89.3944, 61 m, 5 Aug 1997, E.M. Martínez S. 28113 (NY). Chiapas: al SW de Santo Domingo, [17.0458, -91.4317], 30 Jul 1982, J.M. Quintanilla 14 (MO). Quintana Roo: a 2 km al norte de Estero Franco, sobre la carretera La Unión-Ucum, [17.9512, -88.8769], 20 Aug 1983, E. Cabrera-C. 5444 (MO, NY).

Lycianthes inconspicua Bitter, Repert. Spec. Nov. Regni Veg. 20: 368, 1924

Fig. 54

Lycianthes storkii C.V.Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1061. 1938. Type: Costa Rica, 0.5 mile south of Santa María, 5700 ft, 8 Aug 1932, H. Stork 3138 (holotype: F [0072922F, acc. # 672865]).

Type

Guatemala. [Quetzaltenango]: [Volcán] Santa María, [14.7559, -91.5537], Dec 1877, K. Bernoulli & O. Cario 2373 (holotype: GOET [GOET003447]).

Figure 54. 

Image of herbarium specimen of L. inconspicua in Guatemala, Croat 40950 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 55).

Figure 55. 

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

Common names and uses

Guatemala. Chiltepe de montaña (Gentry and Standley 1974).

Phenology

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).

Preliminary conservation status

In Guatemala, Lycianthes inconspicua is represented by only three collections, although it is more common in Costa Rica and Panama, and present in one protected area in Panama (Cerros de La Carpintera, Panama). The EOO is 27,148.801 km2, and the AOO is 32 km2. Based on the IUCN (2019) criteria, the preliminary assessment category for Guatemala is Endangered (EN).

Discussion

Lycianthes inconspicua is an uncommon species of high elevation areas in Central America. It is uncommon in Guatemala, but more common in high elevation areas of Costa Rica and Panama. It is morphologically similar to three other Mexican and Central American species: L. glabripetala, L. amatitlanensis and L. inaequilatera. Lycianthes amatitlanensis differs from L. inconspicua in having shorter pedicels (4–12 mm in flower and 6–16 mm in fruit), long, coarse trichomes that spread away from the midvein on the abaxial side of the leaf (usually with some trichomes at an angle close to ninety degrees), and lanceolate anthers with a longer attenuate portion at the tip (ca. 0.5 mm long). Lycianthes inaequilatera and L. glabripetala are more similar to L. inconspicua in terms of having appressed trichomes along the leaf blade midvein, but they tend to have shorter pedicels (15 mm or less in flower); in addition, L. glabripetala has larger corollas (1–1.2 cm long).

Representative specimens examined

Guatemala. [Quetzaltenango]: [Volcán] Santa María, [14.7559, -91.5537], Dec 1877, K. Bernoulli & O. Cario 2373 (GOET). San Marcos: Finca Armenia, Rafael de Cuesta, San Marcos, 5000 ft, 6–7 Jul 1977, J.D. Dwyer 14408 (LL , MO). Suchitepéquez: Mpio. San Francisco Zapotitlán, Reserva Natural Privada Las Nubes, 14.7061, -90.9744, 1600 m, 3 Jul 2014, B. Escobar 171 (BIGU).

Lycianthes jalicensis E.Dean, Novon 8: 133. 1998

Fig. 56

Type

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, E. Dean 248 (holotype: DAV [DAV158081]; isotypes: IEB [000183677], MEXU [MEXU01195794], NY [00687933], UC [1797878], XAL [XAL0106678]).

Figure 56. 

Image of herbarium specimen of L. jalicensis, Iltis 29181 (NY). Specimen used with permission from the William and Lynda Steere Herbarium, New York Botanical Garden.

Description

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.

Chromosome number

n = 12, from Iltis 29181, count reported on herbarium label as done by Leslaw Przywara, July 1984, apparently unpublished.

Distribution and habitat

Mexico (Jalisco) in tropical moist forest, tropical dry forest or in oak forest, often near drainages, 350–1350 m in elevation (Fig. 57).

Figure 57. 

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

Common names and uses

None known.

Phenology

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 (Dean et al. 2017a).

Preliminary conservation status

Lycianthes jalicensis is currently only known from the state of Jalisco, Mexico, represented by 27 collections, only two from protected areas. The EOO is 6,855.367 km2, and the AOO is 92 km2. Unfortunately, the lands where this species grows are vulnerable due to recent land use changes. Based on the IUCN (2019) criteria, the preliminary assessment category is Vulnerable (VU).

Discussion

Lycianthes jalicensis occurs at the low elevations and in habitats that can also be inhabited by the widely distributed L. arrazolensis; some populations of L. arrazolensis also share the white, campanulate corollas and seeds found in L. jalicensis. The two species differ in that L. jalicensis has longer pedicels (the length more similar to that of L. tricolor), has glabrous calyces and corollas, and usually has larger floral dimensions than L. arrazolensis. Lycianthes jalicensis can be distinguished from L. tricolor by having unnotched seeds, more glabrous calyces and flowers, and lower elevational range (Dean et al. 2017a).

Representative specimen examined

Mexico. Jalisco: streamside bottoms and steep lower erosive wooded slopes of La Calera, a deep narrow valley cut into the SW-facing slope of Sierra de Manantlán Occidental, just NW of km 188 marker on Autlán-Manzanillo hwy (Mex 80), 9 km (by air) NNE of La Resolano (Casimiro Castillo) and ca 16 km SSE of Autlán, [19.6778, -104.4084], 800–1100 m, 10 Mar 1992, Iltis 31037 (DAV, WIS).

Lycianthes limitanea (Standl.) J.L. Gentry, Phytologia 26: 275. 1973

Fig. 58

Solanum limitaneum Standl., Publ. Carnegie Inst. Wash. 461 (4): 85. 1935. Type: Belize. Camp 33 (on Guatemala-Belize boundary), 2850 feet [869 m], 24 Apr 1934, W.A. Schipp S-681 (holotype: F [0073115F, acc. # 733642]).

Type

Based on Solanum limitaneum Standl.

Figure 58. 

Image of herbarium specimen of L. limitanea, Cabrera 6091 (WIS). Specimen used with permission from Wisconsin State Herbarium, University of Wisconsin, Madison.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 59).

Figure 59. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes limitanea is an uncommon species of southern Mexico, Guatemala, and Belize, represented by 17 collections and occurring in two protected areas (Columbia River, Belize and Río Dulce, Guatemala). The EOO is 69,835.282 km2, and the AOO is 68 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes limitanea is a distinctive, but poorly known and rarely collected, species of the Caribbean slope ranging from southern Mexico to Belize and Guatemala. It is easily identified based on its dense, tan to orange, multangulate-stellate trichomes, relatively large leaves, large calyces which lack appendages, equal stamens, and large, round fruits. Until this paper, the flowers had not been described in the literature, and of the 17 collections we examined, only five collections were in bud and three collections were in flower. All the other collections were in fruit. Therefore, the flower measurements in this description are based on dissection of a single flower.

Representative specimens examined

Guatemala. Huehuetenango: Nentón, along the road from Nuevo San José Frontera to Las Palmas, 16.0333, -91.55, 900–1200 m, 17 Mar 2009, M.J.M. Christenhusz 5619 (NY). Izabal: El Estor, La Llorona, 15.5142, -89.4236, 500 m, 30 Aug 1998, M. Véliz 6652 (BIGU). Petén: on Melchor de Mencos Road, 8 May 1967, E. Contreras 6873 (MO, NY, LL). Mexico. Chiapas: Mpio. Palenque, 6–12 km south of Palenque on road to Ocosingo, [17.4468, -91.9623], 300 m, 10 May 1973, D. Breedlove 35007 (MO).

Lycianthes manantlanensis Aarón Rodr. & O.Vargas, Novon 12: 245. 2002

Fig. 60

Type

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, R. Cuevas 5009 (holotype: IBUG (not seen); isotypes: ENCB (not seen), IBUG [IBUG0157395], MEXU (not seen), MO (not seen), WIS (not seen), ZEA).

Figure 60. 

Image of herbarium specimen of L. manantlanensis, Alvarado 575 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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, Liquidambar, oak, or pine-oak forest, rarely in fir forest, in canyons or on slopes, 1200–2500 m in elevation (Fig. 61).

Figure 61. 

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

Common names and uses

Mexico. Naranjillo (Rodríguez and Vargas 2002). El Salvador. Chiltepe morado (Reyna 1492).

Phenology

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.

Preliminary conservation status

Lycianthes manantlanensis is a widespread species ranging from western Mexico to El Salvador, represented by 29 collections and occurring in three Mexican protected areas (Sierra Manantlán, La Sepultura, and El Triunfo). The EOO is 198,390.562 km2, and the AOO is 104 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes manantlanensis is sometimes confused with L. orogenes and L. barbatula, and it is probable that the three are closely related, because they share similarities in floral structure, such as white, stellate corollas and yellow anthers that sometimes have a brownish connective. Lycianthes barbatula differs from the other two species in having tufts of trichomes in the vein axils on the abaxial leaf side. Lycianthes manantlanensis differs from L. orogenes in having a calyx that nearly lacks calyx appendages and often tears in fruit, as well as having equal to slightly unequal stamens. It should be noted that the original description of L. manantlanensis incorrectly stated that the mature fruits are green (Rodríguez and Vargas 2002). The mature fruits are purple; the immature fruits are green. In addition, the original description states that the stamens are equal. However, the stamens are not always equal; they are often slightly unequal, with one stamen slightly longer than the other four. In the preparation of this treatment, we realized that this species is not endemic to Mexico but ranges to El Salvador. The specimens collected in El Salvador and Guatemala had been identified originally as L. orogenes, and many of the specimens were collections made by Standley or Steyermark, the original authors of that species. However, examination of the type material of L. manantlanensis and L. orogenes shows differences in the calyx structure that allows us to assign some of the Guatemalan and all of the El Salvadorian material to L. manantlanensis.

Representative specimens examined

Guatemala. Quezaltenango: along road between Finca Pirineos and Patzulín, 1200–1400 m, 9 Feb 1941, P.C. Standley 87013 (US). Mexico. Chiapas: 5 km del camino Ejido Las Golondrinas a Rosario Sacatonal, 15.4436, -92.6856, 1400 m, 9 Mar 2006, R. Martínez-Camilo 936 (MO). Guerrero: 33.3 km SW of Filo de Caballo, on Chilpancingo-Atoyac road, [17.6405, -99.7304], 2000 m, 7 Nov 1999, T. Yahara 1925, (MEXU). Jalisco: Mpio. Cuautitlán de García Barragán, Majada de las Avellanas, comunidad indígena de Cuzalapa, 3–4 km al NNW de El Durazno, [19.4988, -104.3182], 800–1000 m, 6 Nov 1995, R. Cuevas 5009 (IBUG). Michoacán: Dto. Coalcoman, Naranjillo, [18.7624, -103.1397], 1400 m, 2 Aug 1941, Hinton 15942 (F, NY). Oaxaca: cañada al N de Cerro de la Leona (cerro al NE de Cerro Quetzal y ca. 7–9 km al N de Cerro Guayabitos), ca. 46 km en línea recta al N de San Pedro Tapantepec, 16.7833, -94.1833, 1300 m, 28 Feb 1987 S. Maya J. 4226 (MEXU).

Lycianthes mariovelizii E.Dean, Brittonia 70: 482. 2018

Fig. 62

Type

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, 15.6949, -91.8721, 753 m, 15 Aug 2017, E. Dean 9509 (holotype: BIGU [acc. # 76960]; isotypes: BIGU [acc. # 76959], DAV [acc. # 221941, acc. # 221937], MEXU, NY).

Figure 62. 

Image of isotype of L. mariovelizii, Dean 9509 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 63).

Figure 63. 

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

Common names and uses

None known.

Phenology

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 (Dean et al. 2018c).

Preliminary conservation status

Lycianthes mariovelizii is a widespread but poorly collected species ranging from southern Mexico to Nicaragua, represented by only nine collections and occurring in eight protected areas. The EOO is 87,389.609 km2, and the AOO is 32 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes mariovelizii belongs to series Tricolores and is distinguished from most of the other species in the series by long calyx appendages (to 11 mm in flower and 14 mm in fruit) that are somewhat flattened and relatively wide at the base (appearing very flattened when dried). Within a calyx, the appendages often vary in length, but at least some of the appendages on a calyx are greater than 7 mm long and 1 mm wide at the base. The only species in series Tricolores with appendages similarly long and wide is L. surotatensis, a species with glandular trichomes (Dean et al. 2018c). Within the series, L. mariovelizii is most similar to L. arrazolensis, L. jalicensis, and L. surotatensis, three species with which L. mariovelizii shares the following characters: unnotched seeds, white corollas, and occurrence mostly below 1000 m. Lycianthes mariovelizii differs from L. surotatensis by eglandular trichomes. It differs from L. arrazolensis and L. jalicensis by longer calyx appendages inserted < 0.5 mm below the calyx rim, versus > 0.5 mm. The appendages of L. arrazolensis are typically < 2.5 mm long, rarely reaching 5 mm in Guerrero; those of L. jalicensis are typically < 5 mm long. Also, the calyx and corolla of L. jalicensis are usually glabrous or nearly so, whereas those of L. mariovelizii are pubescent (Dean et al. 2018c). A collection of L. mariovelizii (Steyermark 51177) was listed by Gentry and Standley (1974) as a difficult to place collection and unnamed taxon at the end of their treatment of Lycianthes in Flora of Guatemala; Dean et al. (2017a) listed a different specimen of L. mariovelizii in their Specimen Group E in their section on difficult to place collections at the end of their paper on series Tricolores (E. M. Martínez S. 23237 – listed in the paper as Droege 23237).

Representative specimens examined

Guatemala. Huehuetenango: Paso del Boquerón, Río Trapichillo, below La Libertad [15.5174, 91.8385], 1200–1300 m, 21 Aug 1942, J.A. Steyermark 51177 (F); El Progreso: a 2 km al N de Los Leones, camino El Rancho-Cobán, 14.95, -90.2, 4 Aug 1988, E.M. Martínez S. 23237 (MEXU, MO, NY). Mexico. Chiapas: Mpio. Frontera Comalapa, 6–8 km E of Frontera Comalapa along road to Ciudad Cuahtémoc, [15.6916, -92.0871], 1000 m, 15 Aug 1972, D. Breedlove 27009 (CAS, MEXU, MO). Oaxaca: Dto. Pochutla, Mpio. San Miguel del Puerto, en el Cafetal Arroyo Arena, 15.9778, -96.1006, 700 m, 16 Nov 2003, A. Nava-Zafra 205 (DAV).

Lycianthes michaelneei E.Dean, Phytoneuron 2014–42: 4 (2 Apr 2014)

Fig. 64

Type

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, M. H. Nee & G. Schatz 19791 (holotype: WIS; isotypes: CAS [483750, acc. # 648091, MEXU [acc. # 303241]).

Figure 64. 

Image of herbarium specimen of L. michaelneei, Martinez 1227 (NY). Specimen used with permission from the William and Lynda Steere Herbarium, New York Botanical Garden.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 65).

Figure 65. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes michaelneei is a rarely collected species of eastern Mexico, represented by only six collections, only one from a protected area (Pico de Orizaba). The EOO is 181.413 km2, and the AOO is 24 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Discussion

Lycianthes michaelneei is a member of series Tricolores (Dean et al. 2017a) and was provisionally identified as L. pilosissima (M.Martens & Galeotti) Bitter (a synonym of L. tricolor) in the Flora of Veracruz (Nee 1986). Lycianthes michaelneei is similar to L. tricolor in having notched seeds. It differs from L. tricolor in having an entirely purple corolla (lobes and membrane) rather than a white membrane with some purple on the lobes, and it may lack the green, glandular spots on the corolla (although more fieldwork is needed to determine this). In general, this species is more robust than similar species such as L. tricolor and L. arrazolensis. Another distinctive difference is the matted stem trichomes, which are not found in closely related species (Dean et al. 2017a).

Representative specimen examined

Mexico. Veracruz: Mpio. Calcahualco, 1 km al S de Escola, 19.1167, -97.1333, 1950 m, 24 Jul 1986, J. L. Martínez 1227 (NY, XAL).

Lycianthes moziniana (Dunal) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 408, 1919

Solanum mozinianum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 757. 1814. Type: Painting made during the Royal Botanical Expedition to New Spain (1787–1803) under the direction of Martín de Sessé y Lacasta (lectotype designated by Dean 1997, pg. 193: Hunt Institute for Botanical Documentation, catalogue number 6331.0121).

Type

Based on Solanum mozinianum Dunal.

Lycianthes moziniana var. moziniana

Fig. 66

Solanum uniflorum Sessé ex Lag., Gen. Sp. Pl. [Lagasca]: 10. 1816. Type: Painting made during the Royal Botanical Expedition to New Spain (1787–1803) under the direction of Martín de Sessé y Lacasta (lectotype designated by Dean 1997, pg. 196: Hunt Institute for Botanical Documentation, catalogue number 6331.0025).

Solanum monanthum Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes]. 4: 608. 1819. Type: Based on Solanum uniflorum Sessé ex Lag.

Solanum mocinianum Dunal forma luteiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 164. 1852. Type: Based on Solanum uniflorum Sessé ex Lag.

Solanum uniflorum Moçiño & Sessé, Pl. Nov. Hisp.: 35. 1888. Type: Based on Solanum uniflorum Sessé ex Lag.

Description

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.

Figure 66. 

Image of herbarium specimen of L. moziniana var. moziniana, Dean 216 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Chromosome number

2n = 24 (Williams 1993); 2n = 24, Dean 300, 306 (Dean 2004).

Distribution and habitat

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. 67).

Figure 67. 

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

Common names and uses

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 (Dean 2004). Used medicinally as a purgative in the state of México (Altimirano 72).

Phenology

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 Thygator visit this species (Dean 2001).

Preliminary conservation status

Lycianthes moziniana var. moziniana is a common variety of the Trans-Mexican Volcanic Belt of Mexico, represented by 181 collections and occurring in 12 Mexican protected areas. The EOO is 264,912.814 km2, and the AOO is 688 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes moziniana var. moziniana is separated from closely related taxa by its combination of green fruits, small smooth seeds (without fibrils on cell walls), a tendency toward having only one dichasial or no dichasial branching points, very elongate pedicels, and abaxially pubescent corolla lobes. Variety moziniana is separated from the other varieties of the species in having fruiting calyx teeth that are lax in flower and stay appressed to the fruit, cuneate (rather than attenuate) leaf bases, relatively dense and long stem and leaf pubescence, consistently pubescent abaxial corolla lobes, and an affinity for the soils of the transvolcanic belt (Dean 2004).

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 L. moziniana var. moziniana is edible and in past decades was gathered for sale at markets. Some researchers believe this variety was once a domesticated plant that has since reverted to a weed of agricultural areas (Williams 1993). It persists under traditional agricultural practices by sprouting from its underground root system as well as from seed dispersed by humans (Williams 1993). With the advent of herbicide usage in agricultural fields, changes in field preparation techniques, and lack of usage by humans, this once plentiful variety is becoming rare in Mexico (Dean 2004).

In some areas of Jalisco and Michoacán, where this variety grows with L. rzedowskii or L. acapulcensis, plants with intermediate trichome, leaf and floral color characteristics have been collected or observed. Limited crossing experiments indicate that L. moziniana var. moziniana is capable of crossing with L. moziniana var. oaxacana, although with reduced fertility; crosses between this variety and mature accessions of L. moziniana var. margaretiana have not yet been performed (Dean 2004).

Representative specimens examined

Mexico. Distrito Federal: Limbo, delegación de Álvaro Obregón, [19.3247, -99.2633], 2700 m, 4 Aug 1979, Á. Ventura A. 3477 (ENCB, F, MEX). Durango: 1 km al noroeste de Santa María Ocotán, Mezquital, [24.1, -104.6], 13 Jul 1984, M. González Espinosa 1378 (MEXU). Hidalgo: Mpio. Acatlán, small town of Los Reyes, ca. 9 rd. mi from Acatlán, along road to Huasca, NW of Tulancingo, [20.1888, -98.4489], 7200 ft, 21 Sep 1991, E. Dean 259 (DAV, IEB). Jalisco: 1 km al suroeste de Nueva Colonia, Santa Catarina, [22.1554, -104.1165], 2200 m, 20 Jul 1992, J.J. Reynoso-Dueñas 929 (IBUG, IEB). México: Valle de México, Cuajimalpa a Río Hondo, [19.4315, -99.2919], 2400 m, 9 Sep 1951, Matuda 21819 (MEXU). Michoacán: lado sureste del Cerro El Aguila, subiendo por el poblado de Huatzanguio, 19.6064, -101.3792, 2530 m, 14 Aug 2008, G. Cornejo-Tenorio 2840 (IEB). Nayarit: Sierra Madre, territory of Tepic, between Sta. Gertrudis and Sta. Teresa, [21.7167, -104.7333], 8 Aug 1897, Rose 2068 (GH). Puebla: Mpio. Xochiapulco, Rosa Chica, en un sitio llamado El Plan, cerca de la escuela primaria, 19.7950, -97.6568, 2041 m, 25 Jun 2015, M. Jiménez -Chimil 30760 (DAV). Querétaro: El Picacho, desviación San Pedro Tenango, 3 km al sureste de Amealco, [20.1374, -100.1152], 2650 m, 20 Jul 2003, V. Serrano-Serrano 121 (IEB). San Luis Potosí: chiefly in the region of San Luis Potosí, 6000–8000 ft, 1878, Parry 662 (GH, MO, NY [the K duplicate of this collection number is L. moziniana var. margaretiana]). Tlaxcala: Mpio. Espanita, 0.95 rd mi from intersection with Hwy 136, along road to Espanita, [19.4931, -98.4599], 8920 ft, 28 Oct 1991, E. Dean 302 (DAV). Veracruz: camino a Zacatonal, 18.7869, -97.2767, 9 Jul 2013, A.F. Vargas-Rueda 637 (MEXU). Zacatecas: 2 km al oeste de Monte Escobedo, 22.3253, -103.5829, 2376 m, 28 Aug 2005, A. Rodríguez 4462 (IBUG).

Lycianthes moziniana (Dunal) Bitter var. margaretiana E.Dean, Bot. J. Linn. Soc. 145: 413. 2004

Fig. 68

Type

Mexico. Nuevo León: Mpio. Zaragoza, Cerro El Viejo, 2085 m, 17 Jun 1993, Hinton et al. 22937 (holotype: DAV [DAV155246]; isotypes: GBH [GBH022937], TEX [00208090], XAL [XAL0106696]).

Figure 68. 

Image of herbarium specimen of L. moziniana var. margaretiana, Carranza 6365 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 69).

Figure 69. 

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

Common names and uses

None known.

Phenology

Flowering specimens have been collected July and August (May in Tamaulipus); mature fruits of Lycianthes moziniana var. margaretiana have not yet been collected from the field. As discussed below, intermediates between L. moziniana var. margaretiana and Lycianthes ciliolata are known from several locations, and specimens with mature fruits have been collected from those populations in November and December. The diurnal corolla movements of this variety were observed in the greenhouse by the first author; the corolla opens in the very early morning and close by late morning. The pollen has a sweet scent.

Preliminary conservation status

Lycianthes moziniana var. margaretiana is an uncommon variety of the Sierra Madre Oriental of Mexico, represented by 19 collections, two of which are from the protected area of the Sierra de Álvarez. The EOO is 36,938.334 km2, and the AOO is 76 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Near Threatened (NT).

Discussion

Lycianthes moziniana var. margaretiana is closely related to L. moziniana var. oaxacana E. Dean based on DNA sequence data, and it shares the attenuate leaf bases and spreading calyx teeth (in fruit) of that variety. However, its distribution is disjunct from var. oaxacana and presumably the two varieties have been separated for quite some time. Variety margaretiana has several characteristics that differ from the other two varieties of L. moziniana: 1) upon maturity, the fruit exocarp can have tan or purple patches of color; 2) the placental area of the fruits may be purple, sometimes even of a powdery texture similar to the fruits of L. ciliolata; 3) the pollen is somewhat larger than that of the other two varieties; 4) the stamen filaments can sometimes be pubescent; 5) the abaxial sides of the corolla lobes can sometimes be glabrous; and 6) this variety does not appear to be a weed of agricultural situations (Dean 2004).

The fruits of L. moziniana var. margaretiana in Nuevo León are green with tan or rose blotches with seeds typical in shape and size for L. moziniana var. moziniana. The placental area of the Nuevo León fruits can be intermediate between L. moziniana var. moziniana and L. ciliolata. Lycianthes moziniana var. margaretiana may be an evolutionary transition between L. ciliolata and L. moziniana var. moziniana. Intermediates between var. margaretiana and L. ciliolata are found in San Luis Potosí, Guanajuato, and Querétaro. The intermediate plants resemble var. margaretiana in vegetative and floral characters, however the branching pattern and pollen morphology are intermediate between the two taxa. The fruits of the intermediates are large, up to 50 mm long, the exocarp is rose-colored with the grainy light purple placental area typical of L. ciliolata, but the fruit shape and the shape of the fruiting calyx is that of L. moziniana var. margaretiana. Finally, the seeds of the intermediates have the texture and shape of L. moziniana var. margaretiana but are much larger (4–5 mm long) than usually found in that variety. More study is needed to understand the relationship between L. moziniana var. margaretiana and L. ciliolata in northern Mexico (Dean 2004).

Representative specimens examined

Mexico. Nuevo León: Mpio. Galeana, E of the town of Pablillo, San Francisco Canyon, [24.5666, -99.9666], 4 Sep 1993, Dean 360 (DAV, XAL, ANSM). Querétaro: aproximadamente al oeste de La Veracruz, carretera a San Joaquín, 20.9008, -99.5311, 2350 m, 6 Jul 2002, E. Carranza 6365 (DAV, IEB, IBUG). San Luis Potosí: Hwy 86, 25 mi from Juárez Circle, beyond Xoconostle, 22.2, -100.9667, 9000 ft, 5 Jul 1971, M. Andreasen 544 (MO).

Lycianthes moziniana (Dunal) Bitter var. oaxacana E.Dean, Bot. J. Linn. Soc. 145: 415. 2004

Fig. 70

Type

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, E. Dean 285 (holotype: DAV [DAV172076]; isotypes: NY [00687932], XAL [XAL0106697]).

Figure 70. 

Image of herbarium specimen of L. moziniana var. oaxacana, Cervantes 220 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 71).

Figure 71. 

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

Common names and uses

Mexico. Oaxaca: chichi de venado (Dean 2004).

Phenology

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.

Preliminary conservation status

Lycianthes moziniana var. oaxacana is an uncommon variety of Oaxaca, Mexico, represented by 19 collections, none of which are from protected areas. The EOO is 12,876.905 km2, and the AOO is 72 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Vulnerable (VU).

Discussion

Lycianthes moziniana var. oaxacana differs from var. moziniana in having spreading calyx appendages in fruit and in having attenuate leaf bases. It is closely related to L. moziniana var. margaretiana (a northern variety found in the Sierra Madre Oriental) based on unpublished DNA sequence data, however it differs from var. margaretiana in lacking tan or purple blotches on the fruit exocarp, not having a purple, powdery placental area, and in always having glabrous stamen filaments (Dean 2004). This variety was described from the Sierra de Juárez in Oaxaca. Since that time, many more populations of L. moziniana var. oaxacana have been discovered in Oaxaca, and it is possible that some of them are the more widespread var. moziniana. Lycianthes moziniana var. oaxacana is similar to L. ciliolata, with which it overlaps in distribution. It differs from L. ciliolata in having pubescence on the abaxial side of the corolla lobes (versus no pubescence), having the length of the longest stamen filament 1.5–2 times the length of the medium filaments (vs 1.5–3 times), having trichomes of two distinct lengths on the pedicels (vs trichomes of one length), and having tricolporate pollen grains (vs grains with two pores and a remnant third pore) (Dean 2004).

Representative specimen examined

Mexico. Oaxaca: Llano de las flores, on the Oaxaca-Valle Nacional Highway, 20 km E of Ixtlán, 2870 m, 22 Jul 1960, J. Beaman 3703 (GH, LL).

Lycianthes nitida Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 501. 1919

Fig. 72

Solanum calochromum S.F.Blake, Contr. U.S. Natl. Herb. 24: 21. 1922. Type: Honduras. Between Hacienda El Limon and El Paraiso, 12 May 1919, S. Blake 7370 (holotype: US [00027489]).

Type

Guatemala. [Alta Verapaz]: Cubilqüitz, [Cubilhuitz], [15.6675, -90.4293], 350 m, Aug 1907, H. von Tuerkheim II 59 (lectotype designated by Dean and Reyes 2018a, pg. 43: BR [000000552878]; isolectotypes E [E00190704], GH [00936250], GH [00936251], M, NY [00007318, 00007334], U [U-0113931], US [00027489, 00624006], WIS).

Figure 72. 

Image of herbarium specimen of L. nitida, S. Sinaca C. 53 (WIS). Specimen used with permission from Wisconsin State Herbarium, University of Wisconsin, Madison.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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 Liquidambar forest, sometimes in forest clearings or disturbed areas, including agricultural areas, or along drainages or on slopes or ridges, sometimes on limestone, 200–1000 m in elevation (Fig. 73).

Figure 73. 

Map of geographic distribution of L. nitida from Mexico to Costa Rica based on herbarium specimen data.

Common names and uses

None known.

Phenology

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 (Nee 1986) or in the morning, closed in the afternoon (from Nee 18808).

Preliminary conservation status

Lycianthes nitida is a widespread species ranging from southern Mexico to Costa Rica, represented by 94 collections and occurring in 10 protected areas. The EOO is 564,238.851 km2, and the AOO is 352 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes nitida is a relatively common and widely distributed (southern Mexico through Central America) epiphytic herb or shrub with calyces lacking appendages, purple, stellate corollas, and equal stamens. Its distinctive shiny, glabrous, coriaceous leaves, in which the geminate leaf pairs are of very different shapes and sizes (the smaller leaf much shorter and rounder than the larger) makes this species difficult to confuse with similar species that lack calyx appendages, such as L. heteroclita and L. synanthera. Lycianthes nitida may be confused with L. anomala, another epiphyte with stellate corollas and equal stamens, but L. anomala has short appendages on its calyx and tufts of trichomes in the vein axils of the underside of the leaves.

Representative specimens examined

Guatemala. Alta Verapaz: 7 miles up road to Oxec along road which turns off Highway 7E between Tucúru and El Estor CA 6 km NE of Panzós, 700 m, 20 Jul 1977, T.B. Croat 41622 (MO). Huehuetenango: between Ixcan and Río Ixcan, Sierra de los Cuchumatanes, bordering Río Lacandón, 150–200 m, 23 Jul 1942, J.A. Steyermark 49352 (NY). Izabal: Mpio. Puerto Barrios, en la torre de Guatel, Sierra del Mico, 940 m, 8 Sep 1988, E.M. Martínez S. 23554 (MO). Petén: El Petén, La Cumbre on las Cañas, on 142/143 km of El Petén/ Izabal road, 6 Mar 1975, C.L. Lundell 19056 (DUKE, MO). Mexico. Chiapas: Mpio. Ocosingo, al N de la Estación Chajul, 16.0833, -90.4167, 180 m, 23 Jun 2000, S. Sinaca-C. 2548 (XAL). Oaxaca: Mpio. Santa María Chimalapa, San Antonio Nuevo Paraíso, a 3 km al W, Plan de la Ceiba, 17.1625, -94.3711, 250 m, 21 Sep 1997, E. Torres 1353 (IEB, BIGU, XAL). Veracruz: Rancho “El Milagro,” 5 km en línea recta al sureste de la colonia Nueva Tabasquenia, 17.53, -94.0289, 115 m, 5 Aug 2002, E. López 195 (XAL).

Lycianthes ocellata (Donn.Sm.) C.V.Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 274. 1940

Fig. 74

Solanum sideroxyloides Schltdl. var. ocellatum Donn.Sm., Bot. Gaz. 14(2): 28. 1889. Type: Guatemala. Department Alta Verapaz: Pansamalá, 3800 ft, May 1887, H. von Tuerkheim 1155 (holotype: US [00027797]; isotypes: GH [00934885], NY [00139029], LE [LE0017035], US [00027798].

Lycianthes sideroxyloides (Schltdl.) Bitter ssp. ocellata Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 405. 1919. Type: Based on Solanum sideroxyloides Schltdl. var. ocellatum Donn.Sm.

Type

Based on Solanum sideroxyloides Schltdl. var. ocellatum Donn.Sm.

Figure 74. 

Image of herbarium specimen of L. ocellata, Dean 9504 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

Mexico (Chiapas), Guatemala (Alta Verapaz, Baja Verapaz, Quiché), in montane rainforest, cloud forest, high forest, on slopes, 1300–1800 m in elevation (Fig. 75).

Figure 75. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes ocellata is an uncommon species of Guatemala and immediately adjacent areas in Mexico represented by only six collections, two of which are from protected areas (Lagos de Moreno, Mexico and Mario Dary Rivera, Guatemala). The EOO is 2,567.916 km2, and the AOO is 24 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Commentary

Lycianthes ocellata is closely related to L. sideroxyloides with which it shares obovate calyx appendages, stellate corollas, and yellow to orange, geminate-stellate trichomes. It differs from that species in having glandular appendages that are shiny and sticky when fresh and turn black upon drying. The corolla and anthers are also somewhat glandular and bruise easily when fresh, turning a reddish color. Lycianthes ocellata is mainly restricted to the cloud forests of Guatemala; however, there is at least one collection from Chiapas at the border with Guatemala that has calyx appendages with a black area.

Representative specimens examined

Guatemala. Alta Verapaz: Cobán, 1350 m, Jun 1907, H. von Tuerkheim 1810 (GH, NY). Baja Verapaz: along Cobán-Guatemala Highway 14 near Biotopo del Quetzal Reserve, 15.2107, -90.2169, 1800 m, 9 Aug 2017, E. Dean 9505 (DAV). Quiché: Cerro Putul, “Zona Reyna,” 5300 ft, 3 Dec 1934, A.F. Skutch 1835 (A). Mexico. Chiapas: E of Laguna Tzikaw, Monte Bello National Park, [16.0873, -91.6625], 1300 m, 13 May 1973, D. Breedlove 35135 (MEXU, MO).

Lycianthes orogenes Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23(5): 229. 1947

Fig. 76

Type

Guatemala. Chimaltenango: southwestern slopes of Volcán de Fuego, above Finca Montevideo, along Barranco Espinazo, 1200–1600 m, 20 Sep 1942, J.A. Steyermark 52104 (holotype: F [0072921F, acc. # 1148518]; isotype: US [00027892]).

Figure 76. 

Image of herbarium specimen of L. orogenes, Cabrera 5773 (MEXU). Specimen used with permission from the Herbario Nacional de México, Universidad Autónoma de México.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

Mexico (Chiapas) and Guatemala (Alta Verapaz, Baja Verapaz, Chimaltenango) in cloud forest, including oak forest, 1200–2300 m in elevation (Fig. 77).

Figure 77. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes orogenes is a rarely collected species of Mexico and Guatemala, represented by only five collections, none of which is from a protected area. The EOO is 19,783.690 km2, and the AOO is 20 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Discussion

Lycianthes orogenes is an undercollected species of cloud forests in Chiapas and Guatemala that is morphologically similar to L. manantlanensis and L. barbatula, with which it shares slender elongate pedicels and white flowers. Lycianthes orogenes has calyx appendages that are intermediate in length between the two other species, and it lacks the trichomes in the leaf axils that are present in L. barbatula. Although the mature fruits have been reported in the literature as green (Gentry and Standley 1974; Rodríguez and Vargas 2002), they are reported as purple on specimen labels, and the immature fruits are green. Specimens of L. manantlanensis from some areas of Guatemala and El Salvador have sometimes been misidentified as L. orogenes.

Representative specimens examined

Guatemala. Alta Verapaz: San Cristóbal, Finca Pamac II, 15.3980, -90.5883, 2179 m, 16 Aug 2015, E. Car 35 (BIGU). Baja Verapaz: Unión Barrios, top of hill, W of km 153/154, 16 Aug 1975, C.L. Lundell 19655 (LL, F, MO). Chimaltenango: lower and middle southwestern slopes of Volcán Fuego, above Finca Montevideo, along Barranco Espinazo and tributary of Río Pantaleón, 1200–1600 m, 20 Sep 1942, J.A. Steyermark 52104 (US). Mexico. Chiapas: 3 km al O de la carretera San Cristóbal de las Casas-Tenejapa, sobre el camino a Matzala, [16.6651, -92.5487], [2500 m], 29 Sep 1983, E. Cabrera C. 5773 (MEXU).

Lycianthes peduncularis (Schltdl.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 416. 1919

Fig. 78

Solanum pedunculare Schltdl., Linnaea 19: 305. 1847. Type: Germany. Leipzig Botanical Garden, 1842, G. Kunze s.n. (neotype designated by Dean 2004, pg. 416: W [acc. # 292213] see discussion in commentary below).

Type

Based on Solanum pedunculare Schltdl.

Figure 78. 

Image of herbarium specimen of L. peduncularis, Dean 226 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

2n = 24, Dean 303 (Dean 2004)

Distribution and habitat

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. 79).

Figure 79. 

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

Common names and uses

Mexico. Trompeta, berenjena, chichi de perra, ojo de venado, tonchichi, tomatillo del monte (Dean 2004).

Phenology

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 Exomalopsis and Ptiloglossa visit this species (Dean 2001).

Preliminary conservation status

Lycianthes peduncularis is a common species ranging from northern to southern Mexico, represented by 64 collections and occurring in two protected areas (Yagul and Tehuacán-Cuicatlán Valley). The conservation status of this species was evaluated by Anguiano-Constante et al. (2018) and found to be Least Concern (LC).

Discussion

Lycianthes peduncularis is recognized by its combination of prostrate to decumbent habit, simple, ascending-appressed trichomes, small calyces, and round, green fruit with maroon to black striations. The fruits have yellow sclerotic granules in the mesocarp. This species may once have had a broader and more continuous distribution on limestone soils. It is currently restricted to limestone soils on either side of the transvolcanic belt and to some eroded volcanic areas within the transvolcanic belt (rarely on rhyolite) (Rzedowski 1986). In addition, this species may tolerate other, more unusual, substrates. In Oaxaca, L. peduncularis has been collected near onyx and marble quarries. Several other localities are in or near mining areas. Some of the populations in Oaxaca are atypical in the size and shape of their leaves, the long length of the longest stamen filament, and the straight style (Dean 2004).

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 (Purpus 1923). Currently, the type of Solanum pedunculare is a neotype at W from the Leipzig Botanical Garden designated by Dean (2004). In his monograph of Lycianthes, Bitter (1919) mentioned that the type material that he studied of S. pedunculare were mixed collections representing both L. moziniana and L. peduncularis (Bitter 1919). As previously detailed in Dean (2004), Schlechtendal cites three syntypes. One is based on cultivated material from the Halle Botanical Garden (which he indicates is of primary importance, because he cites it after the species epithet as H. Hal) and two herbarium specimens from B, cited near the end of the protologue: Ehrenberg 81 from Hidalgo, Mexico; Schiede s.n. from Michoacan, Mexico.

At the time Dean published the neotypification for S. pedunculare, no authentic material seen by Schlechtendal was available. Recently, the Ehrenberg specimen seen by Schlechtendal has been located at HAL and seen by the first author, and the specimen matches L. moziniana var. moziniana. This specimen is annotated by Schlechtendal as “S. pedunculare, S. mocinianum?” This indicates that he was uncertain that the specimen was S. pedunculare. The specimen is also annotated by Bitter as L. mociniana, indicating that this is the syntype he saw. The entire Solanaceae collections at GOET, HAL, M, and W were searched in 2019, and no other material of Solanum pedunculare seen by Schlechtendal was located. However, the original material from the Halle Botanical Garden existed at the time Bitter (1919) completed his monograph, and he was clear that that the Halle Botanical Garden material belonged to a different taxon than S. moziniana and matched other material that he annotated as L. peduncularis; the specimens and material he examined may have been lost in the destruction of the Berlin herbarium during World War II (Hiepko 1978, 1987; Vorontsova and Knapp 2010).

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 S. pedunculare do not match the Ehrenberg specimen in several important ways. First, the protologue says that S. pedunculare is a branching, prostrate plant, while the Ehrenberg specimen is clearly erect and not highly branched. Second, the protologue describes the root as thick, whereas only narrow underground stems (which lead to a root far beneath the soil surface) are visible on the Ehrenberg specimen. Third, the protologue mentions rhombic-shaped leaves with cuneate bases which are not present on the Ehrenberg specimens. In addition, although the protologue does not mention the density of pubescence, Schlechtendal adds a comment at the end of the protologue that says that the plants in their natural habitat have denser pubescence. By this comment, Schlechtendal acknowledges that the pubescence of the syntypes from Mexico do not match that of the horticultural material from the Halle Botanical Garden. Lycianthes peduncularis as recognized by Bitter (1919), by Dean (2004), and in this treatment is a prostrate, highly branched plant with a very thick root that is very close to the soil surface, rhombic to widely oblanceolate leaves with cuneate bases, and short, sometimes sparse trichomes. In the future, if more authentic material seen by Schlechtendal becomes available, the name S. pedunculare may need to be retypified, however we are not doing so at this time.

Representative specimens examined

Mexico. Guanajuato: Mpio. San José Iturbide, near Rancho El Guajolote, SW of San José Iturbide, one hwy exit S of exit to San José, dirt rd that goes W to large drainage, farm of Margarita Vargaz Fuentes de Acosta, [20.9019, -100.4215], 6000 ft, 31 Oct 1991, E. Dean 308 (DAV). Hidalgo: cañada de Arrollo Hondo, 25.9 km al noreste de Ixmiquilpan, carretera a Tolatongo, 20.6320, -99.0268, 1870 m, 17 Jun 2000, R. Cruz-Durán 4674 (MEXU). México: N of Huehuetoca along the road to Apaxco, ca. 4.2 road mi from building Los Arcos in dowtown Huehuetoca, E side of rd, near where RR tracks come close to rd, [19.8896, -99.2083], 2134 m, 3 Aug 1991, E. Dean 244 (DAV, MEXU). Oaxaca: Mpio. Mitla, La Colorada, 16.9261, -96.3950, 1767 m, 2 Jun 2009, H. Hernández O. 137 (DAV). Puebla: Mpio. Zapotitlán de las Salinas, San Antonio Texcala, along hwy 125 S of Tehuacán, canyon with onyx mine just N of town, [18.4004, -97.4465], 1677 m, 22 Oct 1991, E. Dean 298 (DAV). Querétaro: alrededores de Bernal, [20.7423, -99.9567], 2200 m, 5 Jun 1992, R. Hernández-Magaña 9905 (MEXU).

Lycianthes pilifera (Benth.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 427. 1919

Fig. 80

Solanum piliferum Benth., Pl. Hartw. 68. 1840. Type: México. Oaxaca: Llano Verde, 1839, C. T. Hartweg 499 (lectotype designated by Dean and Reyes 2018a, pg. 44: K [K000585745]); isolectotype: LD [1212266]).

Solanum pilosiusculum M.Martens & Galeotti, Bull. Acad. Brux. 12(1): 136. 1845. Type: México. Oaxaca: Cerro del Malacate (Pelado Capulalpan and Llano Verde), near Villa Alta, 7500 ft, Nov–Apr 1840, H. Galeotti 1171 (lectotype designated by Dean and Reyes 2018a, pg. 44: BR [000000552882]; isolectotypes: BR [000000552849, 000000552911], G [G00343182], LE [LE00017009], NY [00139019], US [00027745], W [acc. # 1889-156335, acc. # 0004160]).

Lycianthes pilifera (Benth.) Bitter var. pilosiuscula (M.Martens & Galeotti) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 428. 1919. Type: Based on Solanum pilosiusculum M.Martens & Galeotti.

Type

Based on Solanum piliferum Benth.

Figure 80. 

Image of herbarium specimen of L. pilifera, Breedlove 66854 (NY). Specimen used with permission from the William and Lynda Steere Herbarium, New York Botanical Garden.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

Mexico (Oaxaca), in cloud forest, tropical moist forest, including pine-oak, oak, and mixed forest with Ilex, Podocarpus, Weinmannia, Persea, Ocotea, Oreomunnea, Taxus and/or Cupressus, in shady canyons, slopes, and drainages, 1800–3050 m in elevation (Fig. 81).

Figure 81. 

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

Common names and uses

Mexico. Oaxaca: monte agua zapote (J. Rivera-Reyes 2609); rojo monte papel (J. Rivera-Reyes 3141).

Phenology

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.

Preliminary conservation status

Lycianthes pilifera is a common species of the cloud forests of Oaxaca, represented by 61 collections, none of which is from a protected area. The EOO is 4,808.353 km2, and the AOO is 192 km2. Based on the EOO and AOO areas, and following the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Discussion

Lycianthes pilifera is extremely variable in terms of width of leaves, size of flowers and calyx appendages, as well as amount of pubescence. The type material has nearly glabrous, relatively narrow leaves and relatively short calyx appendages. Morphological forms with longer calyx appendages are found in Oaxaca below 2000 m, and forms with shorter appendages are found above that elevation. Very small-leaved, small-flowered, and small-fruited forms have been collected from near Conception Papalo, Oaxaca, at 2700 m (Dean et al. 2019b). This species has been sometimes confused with L. stephanocalyx and L. quichensis, both of which can have one to two-flowered inflorescences and flowers with equal stamens. Unlike L. pilifera, both of those species have red fruits. Lycianthes quichensis is only found in Chiapas and Guatemala and does not overlap in distribution with L. pilifera. Lycianthes stephanocalyx does overlap in distribution with L. pilifera and differs in having red fruit, connate anthers (which are usually yellow), and small, whitish, curved trichomes. Lycianthes pilifera also resembles L. caeciliae, an endemic of Veracruz, in having purple flowers with equal stamens and dark purple fruit with large, dark seeds, however L. caeciliae differs in having dark purple, stellate corollas and dark purple anthers (Dean et al. 2019b).

Representative specimen examined

Mexico. Oaxaca: Sierra de Juárez, Mpio. San Pedro Yólox, along Hwy 175 to the NE of the turnoff to Comaltepec and NE of the cabins and restaurant of Mirador, along old undeveloped road to Yólox (just E of new turnoff to Yólox), 17.6028, -96.4175, 2022 m, 10 Sep 2017, E. Dean 9522 (DAV225278).

Lycianthes pringlei (B.L.Rob. & Greenm.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 421. 1919

Fig. 82

Solanum pringlei B.L.Rob. & Greenm., Amer. J. Sci. ser. 3, 50: 160. 1895. Type: México. Jalisco: mountain cañons near Guadalajara [isotypes say Sierras near L. Chapala], 18 Nov 1892, C. Pringle 5343 (holotype: GH [00077531]; isotypes: MEXU [MEXU00028862, MEXU00029069], US [00027755], VT [UVMVT026445]).

Type

Based on Solanum pringlei B.L.Rob. & Greenm.

Figure 82. 

Image of herbarium specimen of L. pringlei, Dean 327 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 83).

Figure 83. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes pringlei is a rarely collected species of western Mexico, represented by 36 collections, none of which is from a protected area. The EOO is 49,603.413 km2, and the AOO is 128 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Discussion

The calyx of Lycianthes pringlei, with its undulate margin that may look bilabiate and poorly developed appendages which often appear as elongated dark bumps, is uncommon in the genus. This feature, in combination with the very glandular pubescence and reddish-orange ovoid fruits, differentiate this shrub from all other species of Lycianthes in the flora area (Dean et al. 2007).

Representative specimens examined

Mexico. Guerrero: 4 km al sur de Tetipac, sobre el camino Tetipac-Taxco, [17.6355, -99.6487], 1820 m, 5 Dec 1982, E.M. Martínez-Salas 2863 (MEXU, NY). Jalisco: Área Natural Protegida Piedras Bola, 20.6489, -104.0435, 1947 m, 29 Oct 2011, M.A. García Martínez 128 (IBUG, MEXU). México: Nanchititla, [18.8346, -100.4071], 27 Nov 1935, Hinton 8750 (CAS, GH, NY). Michoacán: La Alberca de Teremendo de los Reyes, 19.8064, -101.4556, 2072 m, 15 Oct 2013, J. Contreras L. 93 (MEXU).

Lycianthes purpusii (Brandegee) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 382. 1919

Fig. 84

Solanum purpusii Brandegee, Univ. Calif. Publ. Bot. 6: 62. 1914. Type: México. Chiapas: Finca Mexiquito, C. Purpus 7011 (holotype: UC [acc. # 173078]; isotypes: F [0073141F, acc. # 415779]; NY [00139025]; US [00027766]).

Lycianthes purpusii (Brandegee) Bitter var. extensidentata Bitter, Repert. Spec. Nov. Regni Veg. 20: 365. 1924. Type: Guatemala. San Francisco Miramare, Apr 1878, K. Bernoulli & O. Cario 2334 (holotype: GOET [GOET003449]).

Lycianthes vulpina Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 321. 1929. Type: Honduras, Atlántida: Lancetilla Valley, 11 Jan 1928, P. Standley 54356 (holotype: F [0072925F, acc. # 583595]; isotypes: A [00934883], US [00027906]).

Type

Based on Solanum purpusii Brandegee.

Figure 84. 

Image of herbarium specimen of L. purpusii, Nee 18846 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 85).

Figure 85. 

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

Common names and uses

None known.

Phenology

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 (Nee 1986), and all flowering specimens have closed corollas.

Preliminary conservation status

Lycianthes purpusii is a widespread species ranging from southern Mexico to Honduras, represented by 92 collections and occurring in eight protected areas. The EOO is 91,196.026 km2, and the AOO is 328 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes purpusii is a wide-ranging species of tall tropical forest and relatively low elevations. It is distinguished by very long calyx appendages (7–17 mm long in flower) and dendritically branched or long-stalked multangulate-stellate trichomes. The species is variable in leaf arrangement; the leaves are usually paired (geminate) in Honduras but unpaired in many places in Mexico. It is also variable in the length of the pedicels (sometimes becoming unusually long in Guatemala) and the color and density of the pubescence. This species is somewhat similar to L. furcatistellata Bitter of Costa Rica but differs from that species in habitat preference (L. furcatistellata occurs in upper elevations, often in cloud forest), length of the pedicels (L. purpusii often has flowering pedicels less than 20 mm long, while L. furcatistellata usually has pedicels greater than 20 mm long), and calyx appendage length (L. furcatistellata has appendages on the flowering calyxes of 4 mm or less). The two species do not have overlapping distributions. Several specimens with dense, soft, very branched calyx trichomes, very short pedicels, and flowers in very tight groupings are included in this species circumscription, and they might prove to be a separate species.

Representative specimens examined

Guatemala. Baja Verapaz: Purulhá, Biótopo del Quetzal, 15.21306, -90.22, 400 m, 18 Oct 1995, A. Cahuec s.n. (BIGU). Izabal: Montañas del Mico, 11 km W of Santo Tomás de Castillas, microwave tower, [15.6719, -88.6929], 940 m, 8 Sep 1988, W. Stevens 25496 (NY). Petén: 19 km N of Modesto Méndez, 200 m, 21 Jun 1971, W.E. Harmon 5855 (MO). Mexico. Chiapas: Mpio. Barriozábal, along road from Berriozábal to Las Maravillas, ca. 1.4 km S of the town of Efraín A. Gutiérrez, remnant of tall forest called La Mata Café, 16.8711, -93.2956, 1005 m, 12 Sep 2017, E. Dean 9527 (DAV). Oaxaca: Nueva Santa Flora, 17.9275, -96.4608, 700 m, 22 Dec 1992, G. Ibarra-Manríquez 3781 (XAL). Puebla: Mpio. Hueytamalco, 1 km hacia el Oeste de las instalaciones del Campo Experimental “Las Margaritas,” Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP), 20.0044, -97.3167, 550 m, 19 Nov 2007, B. Gómez-Chagala 349 (IEB, MEXU). Veracruz: Rancho “El Milagro,” 5 km en línea recta al sureste de la colonia Nueva Tabasquenia, 17.53, -94.0289, 115 m, 5 Aug 2002, E. López 192 (IEB, XAL).

Lycianthes quichensis (J.M.Coult. & Donn.Sm.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 428. 1919

Fig. 86

Solanum quichense J.M.Coult. & Donn.Sm., Bot. Gaz. 37: 422. 1904. Type: Guatemala. Quiché: Chiul, Apr 1892, E. T. Heyde & E. Lux 3450 (holotype: F [0073142F, acc. # 264727]; isotypes: G [G00379126], GH [GH00077532], M [M-0171931], NY [00139026], US [00624010], US [00027769].

Lycianthes obliquifolia Standl., Field Mus. Nat. Hist., Bot. Ser. 22: 101. 1940. Type: México. Chiapas: Volcán de Tacaná, 30 Mar 1939, E. Matuda 2938 (holotype: F [0072919F, acc. # 980247]; isotypes: A [00077523], MEXU [MEXU00082111], MICH [1109852], NY [00138706]).

Type

Based on Solanum quichense Coult. & Donn.Sm.

Figure 86. 

Image of herbarium specimen of L. quichensis, Breedlove 40387 (NY). Specimen used with permission from the William and Lynda Steere Herbarium, New York Botanical Garden.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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 Chiranthodendron, Symplocos, Drimys, and Clethra (in Guatemala often in Cupressus or Abies forests), 2200–3900 m in elevation (Fig. 87).

Figure 87. 

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

Common names and uses

Guatemala. Chilete, choshel, coxel, flor de rosa, quilete, tomatillo blanco (Gentry and Standley 1974)

Phenology

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.

Preliminary conservation status

Lycianthes quichensis is a rare species of Mexico and Guatemala, represented by 38 collections and occurring in seven protected areas. The EOO is 22,627.703 km2, and the AOO is 148 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Endangered (EN).

Discussion

Lycianthes quichensis is an attractive shrub that horticulturalists are trying to bring into cultivation and is sometimes available in seed catalogues. It has relatively large purple flowers with distinctive dark purple and green markings and widely ovate leaves. It has sometimes been confused with L. pilifera, a species of Oaxaca, Mexico. The two species differ in fruit color (L. pilifera has purple-black fruits) and pubescence (the trichomes of L. pilifera are darker and more tubular-conical upon drying than those of L. quichensis which are pale yellow to white and flattened upon drying) (Dean et al. 2019b).

Representative specimens examined

Guatemala. Chimaltenango: Volcán de Acatenango, [14.5255, -90.8753], 2800 m, 22 Apr 1999, M. Véliz 7020 (BIGU, MEXU). Huehuetenango: Mpio. La Libertad, Peña Blanca, 15.5075, -91.9158, 3193 m, 14 Dec 2000, M. Véliz 10846 (BIGU, CAS). Quetzaltenango: Cumbre de Alaska, [14.7560, -91.4705], 3100 m, 10 Sep 1999, M. Véliz 7285 (BIGU, MEXU). Quiché: Chiul, Apr 1892, E.T. Heyde & E. Lux 3450 (F, GH, M, NY, US). Sacatepéquez: Volcán de Agua, 14.4769, -90.7322, 2852 m, 6 Feb 2006, M. Véliz 16671 (BIGU, TEX). San Marcos: Mpio. San José Ojetenam, [15.2342, -91.9736], 3100 m, 26 Nov 2009, F. Pérez 18 (BIGU). Sololá: San Pedro La Laguna, Volcán San Pedro, ladera noreste del volcán, 14.6572, -91.2664, 3006 m, 28 Jan 2005, P. Pardo 27 (CAS). Totonicapán: María Tecún, [14.8341, -91.2174], 3000–3600 m, 12-–23 Jan 1966, A. Molina R. 16399 (CAS-DS, NY). Mexico. Chiapas: Mpio. Chamula, Tzontehuitz, 16.8108, -92.5775, 2740 m, 20 Apr 1999, L.Y. Domínguez-Torres 70 (DAV, MEXU).

Lycianthes rafatorresii E. Dean, sp. nov.

Fig. 88

Type

México. Oaxaca: Distrito Juchitán, Mpio. Santa María Chimalapa, San Antonio Nuevo Paraiso, a 2 km en línea recta al W, en Cerro Camedor, 17.1444, -94.3577, C. Perret 352 (holotype: MEXU [acc. # 1143589]; isotypes: IEB [acc. # 210462, acc. # 209686], SERO).

Figure 88. 

Image of herbarium specimen of L. rafatorresii, Perret 352 (MEXU). Specimen used with permission from the Herbario Nacional de México, Universidad Autónoma de México.

Diagnosis

Very similar to Lycianthes multiflora Bitter of Central America, L. sideroxyloides of Mexico and Central America, and L. armentalis of the Yucatán Peninsula. Differing from those species in having the following combination of characters: multangulate-stellate (not geminate-stellate) trichomes with straight (not crisped) trichome rays, branching not widely divaricate, inflorescences that are not restricted to the last few terminal, leafless, sympodial units of the plant, calyx appendages with a widened bulbous apex (rather than acute), stellate corollas divided one quarter to approximately half way to the base with abundant interpetalar tissue, unequal stamens, and few, large fruits, 8–15 mm in diameter with 40–100 seeds per fruit, the seeds usually slightly notched on one side.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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 Quercus or Liquidambar, sometimes on steep slopes or along drainages, sometimes on limestone, 200–1600 m in elevation (Fig. 89).

Figure 89. 

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

Common names and uses

Mexico. Oaxaca: paniui poj (Zoque) (Hernández G. 1234).

Phenology

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.

Preliminary conservation status

Lycianthes rafatorresii is a rare shrub with a discontinuous distribution represented by 22 collections, and it is only present in one protected area (Los Tuxtlas). The size of the EOO area (51,517.851 km2) suggests a preliminary conservation assessment of Least Concern (LC). In contrast, the size of the AOO area (76 km2) suggests an Endangered (EN) status.

Etymology

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.

Discussion

Lycianthes rafatorresii is a Mexican endemic of the Caribbean slope of Oaxaca, Puebla, and Veracruz. Morphologically, it is close to L. multiflora Bitter of Central America (which does not occur in Guatemala or Mexico), and it has been misidentified as L. sideroxyloides (with which it overlaps in distribution) and L. armentalis (with which it does not overlap) (Nee 1986). It differs from L. multiflora in having straight to curved rays on the trichomes (rather than crisped rays in L. multiflora), having the inflorescences and fruits on leafy stems that are not restricted to the last few terminal sympodial units of the plant (rather than having flowers and fruits on mostly leafless terminal sympodia (so that they appear to be in a large panicle), having calyx appendages with a widened bulbous apex (rather than acute appendages), and having stellate corollas (rather than entire corollas). Lycianthes rafatorresii differs from L. sideroxyloides in lacking geminate-stellate trichomes, by having a corolla divided just half way to the base (or less) with abundant interpetalar tissue (versus a deeply stellate corolla with scant interpetalar tissue), by having unequal stamens (versus equal stamens), and by having fewer, larger mature fruits (8–15 mm in diameter versus 2–7 mm in diameter in L. sideroxyloides). It differs from L. armentalis in lacking widely divaricate branching, having appendages with a widened bulbous apex (versus acute appendages), having more seeds per fruit (40–100 in L. rafatorresii versus 20–30 in L. armentalis), occurring at higher elevations (to 1600 m in L. rafatorresii versus to 500 m in L. armentalis), and not occurring in the lowlands of the Yucatán Peninsula. More field study of L. rafatorresii is needed to better separate it from these similar species and understand its distribution, which currently does not appear to be continuous. Attempts by the first and third authors to locate this species in the field in Oaxaca in 2017 were unsuccessful.

Representative specimens examined

(full list of paratypes in Appendix I). Mexico. Oaxaca: Dto. de Ixtlán, 3 km al S de Metates, carr. Tuxtepec-Oaxaca, [17.5872, -96.5065], no elevation, 10 Sep 1985, R. Torres C. 7270 (MO). Puebla: Limonateno, [19.9110, -97.3076], 1000 m, 12 May 1970, F. Ventura A. 1077 (IEB). Veracruz: Bastonal, 3–5 km adelante, camino a la Sierra de Santa Marta, 18.40, -94.95, 25 Nov 1985, G. Castillo-Campos 4417 (XAL).

Lycianthes rantonnetii (Carrière) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 332. 1919

Fig. 90

Solanum rantonnetii Carrière, Rev. Hort. [Paris] 32: 135. 1859, as rantonnei. Type: Illustration (Rev. Hort. [Paris] 32: fig. 32, page 135 (lectotype designated here). Orthographic variant correcting epithet to “rantonnetii” published by Lescuyer, Hort. Français, sér. II. i (9): 197. Late 1859 or 1860.

Solanum corniculatum Hiern, Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn (1877–78): 45. 1877. Type: Brazil, Río de Janeiro, 1867. A. Glaziou 1078 (lectotype designated here: C [10019192]; isolectotypes: BR [00000552267, 00000552234]).

Solanum urbanum Morong, Ann. New York Acad. Sci. 7: 177. 1893. Type: Paraguay. Asunción, streets of Asunción, Nov 1888, T. Morong 147 (lectotype designated by Barboza 2013, pg. 29: NY [00172225]; isolectotypes: MO [acc. # 2495263], NDG [NDG45160], PH [00030498], US [00027839], WIS [v0004256WIS]).

Solanum muticum N.E.Br., Bull. Misc. Inform. Kew 85: 6. 1894. Type: Uruguay. Montevideo “Originaire du Paraguay, cultivé à Montevideo comme plante d’ornament,” Mar 1858, Gibert 56 (lectotype designated by Barboza 2013, pg. 29: K [K000585755]).

Solanum urbanum Morong var. foliosum Chodat, Bull. Soc. Bot. Genève, ser. 2, 8: 152, fig. 47. 1916. Type: Paraguay. Cerros de Paraguarí, R. Chodat & W. Vischer 60 (holotype: G [G00392293]).

Solanum urbanum Morong var. nervosum Chodat, Bull. Soc. Bot. Genève, ser. 2, 8: 152, fig. 47. 1916. Type: Paraguay. No exact location, Jan 1900, É. Hassler 7024 (lectotype designated here: G [G00390048]; isolectotypes: G [G00392288, G00392285, G00392290], P [P03852955], W [acc. # 1904-804].

Solanum urbanum Morong var. subtomentosum Chodat, Bull. Soc. Bot. Genève, ser. 2, 8: 152, fig. 47. 1916. Type: Paraguay. Misiones: San Ignacio, Oct 1914, R. Chodat & W. Vischer 61 (holotype: G [G00392295]).

Solanum urbanum Morong var. typicum Chodat, Bull. Soc. Bot. Genève, ser. 2, 8: 151. 1916. Nom. illeg. Type: Paraguay. Dans les jardines de l’Assumption, 9 Apr 1875, B. Balansa 2104 (lectotype designated here: G [G00392296]; isolectotype: G [G00392297]).

Type

Based on Solanum rantonnetii Carrière

Figure 90. 

Image of herbarium specimen of L. rantonnetii, McCaskill 612 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

2n = 24, Gerasimenko and Reznikova 1968, cited in D’Arcy (1974). Acosta et al. (2005).

Distribution and habitat

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).

Common names and uses

Argentina. Meloncillo del aire (Barboza and Hunziker 1992).

Phenology

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.

Preliminary conservation status

As this species is a horticultural plant in our floristic region, we are not providing a conservation assessment.

Discussion

Lycianthes rantonnetii is a popular horticultural plant, often flowering prolifically throughout the year (depending on climate) and dying back to near the ground in cold temperatures. The species is widely planted in Mexico and California. Viable seeds are produced in Mexican plants, but in California, the fruits are often sterile, without viable seeds; it has been documented as persisting in the wild in southern California, but it is unknown whether viable seeds are being produced (Dean in press). The species was named for M. Victor Rantonnet, 19th century French horticulturist of Hyères, a town on the Mediterranean coast of southern France (Carrière 1859). The spelling of the epithet was first published as “rantonnei” in March of 1859 (Carrière 1859) and changed to rantonnetii by Lescuyer in the later months of 1859 (D’Arcy 1974). Retaining the spelling as L. rantonnetii was upheld by recent editors of the International Plants Name Index citing Art. 60.7 of the International Code of Botanical Nomenclature (Shaw and Shaw 2004).

In the protologue, Carrière (1859) described Solanum rantonnetii from horticultural material grown in Hyères. He says that the seeds were brought to France in ca. 1849 by a naval officer from the region of La Plata [assumed to be Argentina], and after being grown at Toulon, plants were then shared with Rantonnet. No herbarium specimens were cited by Carrière, but the protologue does have a detailed illustration (Carrière 1859, fig. 32, pg. 135) which can serve as a type, and we are lectotypifying that illustration here.

The protologue for Solanum corniculatum is part of an article edited by Warming but published by Hiern (a British botanist) on the Solanaceae of Brazil (Hiern 1877). The protologue cites one collection (Glazou 1078) but does not mention where the collection was seen or deposited. We located one duplicate of Glazou 1078 at C [10019192] and two at BR [00000552267, 00000552234]. The specimen at C is from the Warming Herbarium. Those at BR are from the Martius herbarium. Therefore, the duplicate from the Warming Herbarium at C was chosen as the lectotype: C [10019192].

In his protologue for Solanum urbanum Morong var. nervosum, Chodat and Vischer (1916) cite three collections from Paraguay which we located at G: É. Hassler 7024 [G00390048, G00392288, G00392285, G00392290], B. Balansa 2121 [G00390049], and É. Hassler 6728 [G00392289]. From these specimens we chose a duplicate of É. Hassler 7024 [G00390048] as the lectotype.

In his protologue for Solanum urbanum Morong var. typicum, Chodat and Vischer (1916) cite one collection from Paraguay at G: B. Balansa 2104. We located two duplicate syntypes of the collection at G [G00392296, G00392297]. From these we chose G00392296 as the lectotype.

Lycianthes rzedowskii E.Dean, Novon 4: 327, 1994

Fig. 91

Type

México. Michoacán: Mpio. Charo, along hwy 15, 20 rd km E of Morelia, just E of Pontezuelas, 2165 m, 13 Nov 1991, E. Dean 322a (holotype: UC [UC1797879]; isotypes: DAV [DAV158015], NY [00687931], XAL [XAL0106679]).

Figure 91. 

Image of herbarium specimen of L. rzedowskii, Dean 320 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

2n = 24, Dean 212, 317, 336 (Dean 2004).

Distribution and habitat

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. 92).

Figure 92. 

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

Common names and uses

Mexico. Chilillo (Dean 2004).

Phenology

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.

Preliminary conservation status

Lycianthes rzedowskii is a locally common species of central/western Mexico, represented by 43 collections and occurring in six protected areas. Unfortunately, the habitat of this species is vulnerable due to urban development. The conservation status of this species was assessed by Anguiano-Constante et al. (2018) and their preliminary assessment was Least Concern. The EOO is 24,120.227 km2, and the AOO is 160 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is now Endangered (EN).

Discussion

Lycianthes rzedowskii may be confused with L. acapulcensis and L. ciliolata. It can be distinguished from those species by the many internodes on its first stem, the usually poorly developed sympodial branching in flowering plants, smooth, white (rarely pale lilac) corollas, and broad sweetly scented anthers with terminal, round anther pores. In addition, one of the best ways to distinguish L. rzedowskii from the other two species is to look at the relative lengths of the stamen filaments. In L. rzedowskii the length of the longest filament is never more than twice that of the medium-short filaments, while in the other two species, the length of the longest filament is almost always more than twice that of the medium-short. Lycianthes rzedowskii may hybridize with L. acapulcensis and L. starbuckii E.Dean where they occur together (Dean 2004).

Representative specimens examined

Mexico. México: Avandaro, Cerro Gordo, 19.1142, -100.1341, 2301 m, 22 Jun 2011, L. Corral 1801 (MEXU). Michoacán: San Miguel Chichimequillas La Mesa, 19.4065, -100.3631, 2018 m, 15 Jul 2007, L. Corral 376 (MEXU). Morelos: Sierra de Morelos, Cuernavaca, [18.9794, -99.2841], 2050 m, 26 Jul 1969, Hinton 17221 (NY).

Lycianthes scandens (Mill.) M. Nee, comb. nov.

Solanum scandens Mill., Gard. Dict. ed. 8, no. 19. 1768. Type: Mexico. Vera Cruce [Veracruz], 1730, W. Houstoun s.n. (holotype: BM [BM000514911].

Type

Based on Solanum scandens Mill.

Lycianthes scandens var. scandens

Fig. 93

Solanum lentum Cav., Icon. [Cavanilles] 4: 4, tab. 308. 1797. Type: Spain. ex h. r. Mat., Oct 1795, A. Cavanilles s.n. (lectotype designated by Knapp 2007, pg. 198: MA [MA 476355]).

Solanum axilliflorum Dunal, Hist. Solan. 238. 1813. Nom. Illeg. New name for Solanum scandens Mill. Type: Based on same type as Solanum scandens Mill.

Solanum affine Dunal, in Prodr. [A. P. de Candolle] 13(1): 168. 1852. Type: Cuba, 1829, R. de la Sagra 231 (lectotype designated here: G-DC [G00145629]).

Solanum fugax Bert. ex Dunal, in Prodr. [A. P. de Candolle] 13(1): 170. 1852. Type: Naranjilla en S. Martha, 1822, C. Bertero s.n. (holotype: G-DC [G00145637]).

Solanum neglectum Bert. ex Dunal, in Prodr. [A. P. de Candolle] 13(1): 170. 1852. Type: [Colombia]. Santa Martha, 1822, C. Bertero s.n. (holotype: G-DC [G00145631]).

Solanum decemfidum Pav. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 173. 1852. Type: Nueva España [Mexico]. No date, Herb. Pavón s.n. (holotype: G [G00379128]).

S. lentum Cav. var. echinatum Dunal, Prodr. [A. P. de Candolle] 13(1): 173. 1852. Type: Nueva España [Mexico]. No date, Herb. Pavón s.n. (holotype: G [G00379129]).

Solanum nocturnum Fernald, Proc. Amer. Acad. Arts 35: 570. 1900. Type: Mexico. Guerrero: Acapulco, Jan 1895, E. Palmer 533 (lectotype designated here: GH [00077521]; isolectotype: US [00027708]).

Solanum virgatum Lam. var. lentum (Cav.) O. Schulz, in Urb., Symb. Antill. (Urban) 6: 189. 1909. Type: Based on Solanum lentum Cav.

Lycianthes lenta (Cav.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 364. 1919. Type: Based on Solanum lentum Cav.

Lycianthes lenta (Cav.) Bitter var. utrinquemollis Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 366. 1919. Type: Guatemala [Nicaragua]. Grenada [Granada], 1841, E. Von Friedrichsthal 940 (lectotype designated here: W [acc. # 0003076]; isolectotype: F [barcode F0072916F, acc. # 875961]).

Lycianthes nocturna (Fernald) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 368. 1919. Type: Based on Solanum nocturnum Fernald.

Lycianthes recticarpa Rusby, Bull. Torrey Bot. Club 53: 210. 1926. Type: Colombia. Santa Marta: Quebrada del Cabo, 100 ft, 26 Aug 1876, H. H. Smith 1876 (holotype: NY [00007274]).

Lycianthes variifolia Standl., Publ. Field Mus. Nat. Hist. Chicago, Bot. Ser. 4: 259. 1929. Type: Belize. [Orange Walk or Corozal]: Tower Hill, 1928, J. Karling 13 (holotype F [acc. # 579929]; isotype G [G00379131]).

Description

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.

Figure 93. 

Image of herbarium specimen of L. scandens var. scandens, Dean 238 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Chromosome number

Unknown.

Distribution and habitat

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. 94).

Figure 94. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes scandens var. scandens is a widespread variety ranging from Mexico to the Caribbean and northern South America, represented by many collections and occurring in ten protected areas in our region. The EOO is at least 1,162,412.966 km2. Based on the size of the EOO, and following the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

In this treatment, we are replacing the widely used name L. lenta with the new combination L. scandens var. scandens; this name change is due to the discovery of an older name for this taxon made during relatively recent work on the genus Solanum by the fourth author. Lycianthes scandens var. scandens is distinguished from the less common var. flavicans by having smaller sparser trichomes (usually less than 0.25 mm in diameter versus more than 0.25 mm in diameter in var. flavicans); the trichomes on the calyx are often so small that they cannot be seen with the naked eye. In addition, the leaf veins are obscure in var. scandens (versus prominent and whitish in var. flavicans), the corolla is purple (versus pale violet to white in var. flavicans), and the habit is usually a trailing vine (often an upright shrub in var. flavicans). This variety is also found at lower elevations (usually below 500 m) than var. flavicans (to 1300 m), and it is very common along the Caribbean slope of Mexico and Central America, while var. flavicans is more common along the Pacific slope. The two varieties can be difficult to distinguish in certain regions.

In his protologue for Solanum affine, Dunal (1852) cites two syntypes from the De Candolle herbarium, both of which were located and examined at G: R. de la Sagre 231 (G-DC [G00145629]) and R. de la Sagra 394 (G-DC [G00145619]). Both specimens match the protologue description, but only the R. de la Sagre 231 specimen has location information (Cuba), and it has better vegetative and fertile material. Therefore, we have chosen R. de la Sagre 231 (G-DC [G00145629]) as the lectotype.

In his protologue for Solanum nocturnum, Fernald (1900), cites two syntypes both of which have been located and examined at GH: E. Palmer 533 from Guerrero, Mexico (GH [00077521]) and E. Seler 1625 from Oaxaca, Mexico (GH [00077522]). As the Palmer 533 collection has both flowers and fruits and matches the protologue description more fully, we have chosen the GH specimen of Palmer 533 (GH [00077521]) as the lectotype.

In his protologue for Lycianthes lenta var. utrinquemollis, Bitter (1919) cites two syntypes both from Grenada [Granada], Nicaragua, and these have been located and examined at W and G: E. von Friedrichsthal 940 (W [acc. # 0003076]) and P. Levy 239 (G [G00379125]). We have chosen the W specimen of E. von Friedrichsthal 940 (W [acc. # 0003076]) as the lectotype.

Representative specimens examined

Guatemala. Chiquimula: vicinity of Chiquimula town, 16.1227, -90.9299, 400 m, 4 Dec 1969, A. Molina R. 25111 (NY). Izabal: Finca Mucielago [Murciélago], 22 Jun 1967, S.C. Snedaker 178 (NY). Petén: NW-Ufer des Lago Petén Itzá, 16.9917, -89.8983, 120–140 m, 30 Nov 1994, B. Wallnöfer 9577 (NY). Sacatepéquez: Agua Caliente, 28 Mar 1922, J.M. Greenman 5944 (MO). Mexico. Campeche: en la Ciudad del Carmen, sobre Avenida Camarón, [18.6632, -91.8145], 23 Nov 1987, E. Cabrera C. 14856 (MEXU, MO). Chiapas: Mpio. Ocosingo, Estación Chajul, sobre el Río Lacantun, [16.1227, -90.9300], 150 m, 9 Sep 1992, E. Martínez S. 25292 (MEXU). Jalisco: Arroyo Seco, [19.211, -104.6577], 30 m, 28 May 1990, A. Rodríguez 2077 (WIS). Nayarit: vicinity of Chacala, ca. 5 miles west of Las Varas, 25–50 m, 14 Sep 1960, R. McVaugh 19019 (NY). Oaxaca: Dto. Tehuantepec, Hacienda del Rosario, Río Seco, 15.9367, -95.7719, 33 m, 22 Aug 2010, R. García S. (ROG) 226 (DAV). Quintana Roo: Mpio. José María Morelos, a 3.4 km al SE de Sabana San Francisco, 19.52, -89.075, 86 m, 3 Sep 2004, D. Álvarez 10473 (DAV). Tabasco: Mpio. Comalcalco, Reyes Hernández, [18.2445, -93.2829], 0 m, 10 Sep 1984, F. Ventura A. 21240 (NY). Tamaulipas: vicinity of Tampico, 15 m, 27–30 Apr 1910, E. Palmer 334 (MO, NY). Veracruz: vicinity of Pozarica, [20.5669, -97.4332], 6 June 1987, T.B. Croat 66115 (NY). Yucatán: Pixoy, camino rumbo a San Lorenzo, Valladolid, 20.7147, -88.2625, 22 m, 13 Jul 1988, G. Remmers 30 (MO).

Lycianthes scandens (Mill.) M.Nee var. flavicans (Bitter) J.Poore & E.Dean, comb. nov.

Figs 95, 96

Solanum lambii Fernald, Botanical Gazette 20: 536. 1895. Type: Mexico. Sinaloa: Villa Union, Jan 1895, F. Lamb 446 (holotype: GH [00077501]; isotypes: CAS [0004496], G [G00442761], K [K000063068], MSC [MSC0092874], NDG [NDG45061], NY [00138999], US [00027643]).

Lycianthes lenta (Cav.) Bitter var. flavicans Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 366. 1919. Type: Mexico. Jalisco: Tequila, Mar 184X (year is missing the last digit), H. Galeotti 1225F (holotype: BR [000000563130]; isotypes: NY [00138704], US [01107998]).

Lycianthes lenta (Cav.) Bitter ssp. lambii (Fernald) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 368. 1919. Type: Based on Solanum lambii Fernald.

Type

Based on Lycianthes lenta (Cav.) Bitter var. flavicans Bitter.

Figure 95. 

Image of herbarium specimen of L. scandens var. flavicans, Reyes-Garcia 1267 (MEXU). Specimen used with permission from the Herbario Nacional de México, Universidad Autónoma de México.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 97).

Figure 96. 

Image of herbarium specimen of L. scandens var. flavicans, Lott 2799 (MO). Specimen used with permission from the Missouri Botanical Garden (http://www.tropicos.org).

Figure 97. 

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

Common names and uses

None known.

Phenology

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. scandens, opening and closing in the very early morning.

Preliminary conservation status

Lycianthes scandens var. flavicans is a widespread variety of western Mexico and Central America, represented by 111 collections and occurring in four protected areas. The EOO is 1,500,273.062 km2, and the AOO is 408 km2. Based on the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes scandens var. flavicans is distinguished from the more common var. scandens by having larger and denser trichomes (usually greater than 0.25 mm in diameter versus less than 0.25 mm in diameter in var. scandens) which often make the calyx look woolly; in addition, the leaf veins are often prominent and white in color (versus obscure in var. scandens), the corolla is often white (versus purple in var. scandens), and the habit is often more of a shrub than a trailing vine (the common habit in var. scandens). This variety is also found at higher elevations (to 1300 m) than var. scandens (usually less than 500 m), and it is usually along the Pacific slope of Mexico and Central America, while var. scandens is more common along the Caribbean slope. The two varieties can be difficult to distinguish in certain regions, especially Nicaragua.

The type collection from Jalisco, Mexico upon which this variety is based (H. Galeotti 1225F) has very small leaves, a form which is found in certain areas of the Pacific slope of Mexico. Other populations have much larger leaves. To illustrate this morphological variation, we have provided two figures (Figs 93, 94) for this taxon.

Representative specimens examined

Guatemala. El Progreso: Tulumaje, [14.9286, -90.0313], 346 m, 23 Oct 2003, R. Avila 71 (MO, MEXU). Huehuetenango: Between Santa Ana Huista and woods of Rancho Lucas, Sierra de los Cuchumatanes, 800–900 m, 26 Aug 1942, J.A. Steyermark 51365 (MO1294998). Zacapan: San Agustín Acasaguastlan, [14.9480, -89.9647], 300 m, 8 Feb 2003, F. Ramírez 254 (MEXU). Mexico. Chiapas: roadside on road MEX190 from Tuxtla Gutiérrez to San Fernando, about 8 km NW from the center of Tuxtla Gutiérrez (in a straight line), 16.7944, -93.1836, 733 m, 29 Nov 2012, L. Bohs 3920 (DAV, MEXU). Colima: 17–18 km al NW de Colima; 1 km al sur de Campo Cuatro, [19.3138, -103.7604], 1300 m, 15 Aug 1991, F.J. Santana Michael 5281 (MEXU). Jalisco: Rancho Cuixmala, along the Río Cuitzmala near the ranch headquarters, from the road crossing to Zapata downstream about 1 km, [19.38, -104.98], 10 m, 16 Mar 1991, A. Sanders 10509 (MO, NY). Michoacán: Cuatrocaminos, 4 km al N de Playa Azul, rumbo a la desviación a Lázaro Cárdenas-Tecomán, 18.0196, -102.3414, 50 m, 21 Jun 1998, P. Tenorio L. 19750 (MEXU). Oaxaca: Dto. Yautepec, Toma de Agua, 16.4433, -90.0044, 1036 m, 24 May 2012, D. López P. 3039 (DAV). Sinaloa: Culiacán, 27 Aug–15 Sep 1981, E. Palmer 1502 (NY).

Lycianthes sideroxyloides (Schltdl.) Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 403. 1919

Fig. 98

Solanum sideroxyloides Schltdl., Linnaea 8: 253. 1833. Type: México. Veracruz: Hacienda de la Laguna, July 1829, C. J. W. Schiede 135 (lectotype designated here: HAL [acc. # 100610]; isolectotypes: E [E00190770], G [G00379130], GOET [GOET003590], LE [LE00017034], MO [acc. # 2090811], NY [00139028], P [00371472], W [acc. # 1889-292190]).

Type

Based on Solanum sideroxyloides Schltdl.

Figure 98. 

Image of herbarium specimen of L. sideroxyloides, Pascual 1524 (MEXU). Specimen used with permission from the Herbario Nacional de México, Universidad Autónoma de México.

Description

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.

Chromosome number

Unknown.

Distribution and habitat

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. 99).

Figure 99. 

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

Common names and uses

None known.

Phenology

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.

Preliminary conservation status

Lycianthes sideroxyloides is a widespread species ranging from southern Mexico to Nicaragua, represented by 49 collections and occurring in seven protected areas. The EOO is 534,220.001 km2. Based on the EOO and the number of locations, and following the IUCN (2019) criteria, the preliminary assessment category is Least Concern (LC).

Discussion

Lycianthes sideroxyloides is a wide-ranging species with obovate calyx appendages that are noticeably thickened at the tips. It has yellow to orange, geminate-stellate (multistoried) trichomes with 5–8 rays in a whorl. The obovate calyx appendages are similar to those of the wide-ranging L. pauciflora (Vahl) Bitter of Central and South America (Benítez de Rojas and D’Arcy 1997) and its relatives. Lycianthes sideroxyloides exhibits variation in flower size and flower number per axil, length of calyx appendages, as well as fruit size. Most populations have very short calyx appendages, however there are populations with appendages as long as 2 mm long. In populations with small flowers, the number of flowers per axil can be as many as 30 with the inflorescence appearing spherical. In populations with few flowers per axil, the flower size is larger. Similarly, in populations with few, large flowers, the fruits are larger. Seed size, however, is consistent throughout the range and is the same in fruits of varying sizes; the number of seeds per fruit is greater in larger fruits, with larger fruits having twice as many seeds as small fruits. Lycianthes sideroxyloides is morphologically very similar to Lycianthes ocellata, which differs in having glandular appendages that dry black, and to L. cuchumatanensis, which differs in having thicker leaves with denser pubescence and larger seeds (2.5–3 mm, according to the protologue).

In the protologue for Solanum sideroxyloides, Schlechtendal (1833) cited one collection, Schiede 135, but did not cite a specific specimen or herbarium. We are designating the duplicate at HAL [acc. # 100610] as the lectotype, chosen from the many duplicates of Schiede 135 at multiple institutions listed above.

The status of L. sideroxyloides var. transitoria Bitter (Abh. Naturwiss. Verein Bremen 24 [preprint]: 405. 1919. Type: Guatemala, Depto. Alta Verapaz, Pansamalá, 1200 m, Jun, H. von Tuerkheim 923 [holotype: B]) is unknown, as the one specimen at B upon which it was based is lost and no photo negative is available at F. In the protologue, Bitter (1919) commented that the pubescence of the underside of the leaf in var. transitoria is between the densely felted pubescence of typical L. sideroxyloides in Mexico and the sparsely hairy leaf underside of L. sideroxyloides ssp. ocellata, however, var. transitoria lacks the dark spots at the upper end of the short calyx, which defines L. sideroxyloides ssp. ocellata. Therefore, Bitter’s description of the variety matches a sparsely pubescent L. sideroxyloides. Gentry and Standley (1974) synonymized var. transitoria with L. ocellata, because they did not recognize that L. sideroxyloides occurs in Guatemala.

Representative specimens examined

Guatemala. Alta Verapaz: on Cobán Road, between Chiracte and Chapultepec Farm, in clearing betweeen km 284/285, 19 May 1964, E. Contreras 4726 (MO); Huehuetenango: northern region, along road from San Ramón to Barillas, 15.8626, -91.2147, 790 m, 15 Aug 2017, Dean 9515 (DAV). Mexico. Chiapas: Mpio. Tuxtla Gutiérrez, Mirador El Roblar, Parque Nacional Cañón del Sumidero, 16.7972, -93.0897, 940 m, 23 Aug 2007, J.A. Espinosa J. 273 (MEXU). Guerrero: Mpio. Atoyac, cuenca del Río Balsas y Sierra Madre del Sur, a 15 km al NE de El Paraíso, [17.3574, -100.2194], 1100 m, 19 Aug 1985, J.C. Soto N. 10103 (MEXU). Oaxaca: Mpio. San Miguel del Puerto, El Enjambre, camino a la Constancia, 15.9771, -96.1264, 1412 m, 27 May 2005, J. Pascual 1524 (IEB, MEXU). Veracruz: Cerro Buenavista, 18.8944, -97.0375, 1255 m, 31 Aug 1995, B. Juárez L. 719 (MEXU, XAL).

Lycianthes starbuckii E.Dean, Novon 4: 324, 1994

Fig. 100

Type

México. México: Sierra de Nanchititla, oak forest across the reservoir from the town of Nanchititla, 1945 m, 8 September 1991, E. Dean 315 (holotype: UC [UC1862224]; isotypes: BM [001000924], DAV [DAV158254, DAV158084, DAV158083, DAV158082], ENCB, MEXU [MEXU01195795], MO [2246353], NY [00687930], XAL [XAL0106673, XAL0106672].

Figure 100. 

Image of isotype of L. starbuckii, Dean 315 (DAV). Image used with permission of the UC Davis Center for Plant Diversity.

Description

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.

Chromosome number

2n = 24, Dean 315 (Dean 2004)

Distribution and habitat

Mexico, endemic to southwestern state of México, Sierra de Nanchititla, on level oak forest floor, 1945 m in elevation (Fig. 101).

Figure 101. 

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

Common names and uses

Mexico. México: chilillo (Dean 2004).

Phenology

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.

Preliminary conservation status

Lycianthes starbuckii is a rare species of central Mexico, represented by only four collections from the type location, which is not a protected area. The conservation status of L. starbuckii was investigated by Anguiano-Constante et al. (2018), and their preliminary conservation assessment for this species was Endangered.

Discussion

Lycianthes starbuckii can be distinguished from other species of series Meizonodontae by its combination of prostrate to ascending habit, densely pubescent stems, thick-chartaceous leaves, lax calyx teeth at anthesis, moderately pubescent corolla lobes, and dark purple fruits with large brown to black seeds. Lycianthes starbuckii is unusual in the L. ciliolata complex in its habit and pubescent corolla lobes (Dean 2004). In these characteristics, it is much closer to such species as L. moziniana and L. peduncularis. Lycianthes starbuckii may hybridize with L. rzedowskii, which grows in drainage areas near Nanchititla (Dean 2004). There are specimens collected in southern Guerrero that strongly resemble L. starbuckii, but they have a more erect habit, rather than prostrate; these collections are discussed below under Difficult to Place Specimens. Field work is needed to investigate these populations.

Representative specimen examined

Mexico. México: km 15 carretera El Estado-cañadas de Nanchititla, camino Torrecillas, 18.874, -100.3326, 1934 m, 27 Jul 2010, A. Rodríguez 6083 (IBUG, IEB, MEXU).

Lycianthes stephanocalyx (Brandegee) Bitter, Repert. Spec. Nov. Regni Veg. 18: 315. 1922

Fig. 102

Solanum stephanocalyx Brandegee, Univ. Calif. Publ. Bot. 6: 374. 1917. Type: Mexico. Veracruz: Zacuapan, Jul 1915, C. Purpus 7519 (holotype: UC [UC178649]; isotypes: GH [00077535], NY [00139030]).

Lycianthes symphyandra Bitter, Abh. Naturwiss. Verein Bremen 24 [preprint]: 430. 1919. Type: Mexico. Veracruz: Mirador, 1842, F. Liebmann 1456 (lectotype designated by Dean and Reyes 2018a