Description, habitat and geographic distribution of the genus Jaltomata (Solanaceae)

revised 27 Feb 2016

Link to Jaltomata
homepage
The information on this page may be cited as a communication with professor Thomas Mione, Central Connecticut State University, Biology Department, Copernicus Hall, 1615 Stanley Street, New Britain, Connecticut 06050-4010, United States of America.
Literature Cited
Jaltomata is a diverse solanaceous genus of approximately 60 species. Flowers range from rotate to tubular, from under one cm to six cm across, and flowers per inflorescence range from 1–41. Habitats range from full sun in the desert (Mione, Leiva & Yacher 2014) to shaded trail edges in rain forest (Mione, field work). Species of this genus naturally grow at altitudes from near sea level to over 4,100 m, and are distributed from Arizona, USA to Boliva and on the Galápagos and the Greater Antilles. Berries of at least 40 species are eaten by people (Mione, unpublished data), are fleshy, and orange, red, purple/black or green at maturity. Hummingbirds and bees have been seen visiting the flowers of a few species, and among various species the nectar color ranges from transparent to blood-red (Mione & Anderson 1996, Hansen et al. 2007). In this genus both widespread species and narrow endemics are known: for example J. repandidentata (Dunal) Hunz. grows from Mexico to Bolivia (Mione & Yacher 2005) but J. atiquipa Mione & S. Leiva is known only from the coast of Peru near the top of one small mountain receiving all of its moisture from fog (Mione et al. 2011). This genus is of interest for studies of ethnobotany (Davis & Bye 1982, Williams 1990), floral biology and colored nectar (Mione et al., 2001, Hansen et al. 2007), seed germination (Saldívar-Iglesias et al. 2010), biogeography (Dillon 2005), phylogeny (Mione et al. 1994, Miller et al. 2011), taxonomy (Mione, Anderson & Nee 1993, Mione, Leiva & Yacher 2007, 2014, Leiva et al. 2010, Leiva, Mione & Yacher 2013), ecological genomics (Haak et al., 2014), and the genetic basis of morphological evolution (Mione & Anderson 2017).   

 

 

Description of Jaltomata (Solanaceae): A diverse neotropical genus
author Schlechtendal  
bibliographic citation Index Sem. (Halle) 8. 1838  
vernacular names follow this link  
derivation of genus name Jaltomata may be a variant of Jaltomate, the name of a small pueblo in Zacatecas, Mexico (ca. 22 10' N, 101 47'W; Davis & Bye 1982, page 231). Gledhill (2008, page 220) lists Jaltomata as a "a Mexican vernacular name for False Holly."
major synonyms all Hebecladus (Andes), many Saracha  
growth form perennial; most species are shrubs, some are herbaceous, usually to about 1 or 1.5 m high (-- 4 m when scandent without specialized structures); unarmed.   
roots tap root
(expanded and therefore resembling a tuber in some collections of J. procumbens)
photo of a tap root on the J. procumbens web page
pubescence / vestiture, trichomes Overall, plants are glabrous to densely pubescent.
The different kinds of hairs that one may encounter on surfaces of Jaltomata species are shown on another page, one can reach by following the link at right.
follow this link to see trichomes of Jaltomata
branches erect or spreading, 4- or 5-sided to terete, often hollow, to 4 cm diam but usually smaller. One species is consistently procumbent.  
Leaves alternate or appearing opposite (rarely appearing verticillate), simple, petiolate, estipulate, ovate, ovate-lanceolate or ovate-acuminate, sometimes basally truncate, entire, subentire or toothed; sometimes tapering asymmetrically along distal portion of petiole. The leaves of at least one species are deciduous!  
Inflorescence

umbel-like (Figure 1) or branched in a few species (Figure 2), axillary or from a stem dichotomy (false dichotomy), flowers per inflorescence 1 - 41 (the high value of this range was recorded on a J. darycana plant cultivated for research and included both flowers and very small flower buds)

pedunculate (exceptions follow: J. andersonii has an unarticulated axis joining the flower and the plant, by definition a peduncle though the peduncle may have been lost by reduction, and J. diversa lacks a peduncle -- the umbel-like inflorescence is sessile). 

pedicel basally articulated after fruit ripens

pedicels and peduncles often angled in cross section

Figures 1 & 2
Flowers 5-merous, actinomorphic/regular, bisexual/perfect, usually protogynous  
Calyx

enclosing the bud with valvate aestivation

after flower fully opens: planar or lobes reflexed or bowl-shaped; sepals partially fused, stellate in outline, the lobes triangular, narrowly triangular or obtuse

with fruit: conspicuously accrescent; green or purple; bowl-shaped around berry or planar or reflexed

Figures 1 & 2
Corolla color

Pale-green, whitish, white, blueish, blue-purple, purple, violet, some species red in base due to nectar and/or pigmentation in the base of corolla.

Some species have green spots at the base of the corolla, sometimes visible through the corolla, see for example J. calliantha and figure 3 of the J. mionei web page.

The corolla changes color after it opens in at least one species, from green (pistillate phase) to pale-blue (hermaphroditic phase) in the Jaltomata shown on the right collected in northern Peru (J. neei).

purple
corolla
ring
Some species of section Hebecladus, but no species of sections
Jaltomata
and Modillonia, have this. J. mionei to the right shows the purple corolla ring.
Corolla

Actinomorphic; rotate, broadly infundibular, campanulate, tubular, urceolate and the limb revolute; 5-lobed, or lobes alternating with lobules totalling 10 lobes plus lobules; 1 - 6 cm in diameter

aestivation with five valvate lobes, each lobe alternating with an inwardly pleated and shorter lobule

 
Corona

The corona is a bowl of tissue on which nectar pools. Developmentally it could be:
a) formed of the bases of the stamens (staminal tissue),
b) petaloid and adnate to the bases of the stamens,
c) formed of both staminal and petaloid tissue
d) neither staminal nor petaloid.

Only the three species of section Modillonia have a corona:
J. aspera, J. calliantha,
J. quipuscoae
Stamens epipetalous at base of corolla, usually equal in length but there are exceptions  
Radial thickenings
from base of stamen toward corolla lobule

Some South American species have radially oriented thickenings that extend from the base of each stamen toward the corolla lobule (or where the lobule would be if present) as shown in the photo of Jaltomata herrerae to the right). Where present, these create troughs (between radial thickenings) that hold nectar at the base of the corolla.

On some species radial thickenings are short, merely an expansion of the base of the stamen in an outward direction (away from the stamen and toward the corolla lobule, see for example figure 12 of the J. spooneri webpage). In other species these expansions continue as part of the corolla, as show to the right, and see for example J. weberbaueri. The color of the radial thickenings contrasts with the color of the corolla in J. weberbaueri and J. alviteziana (see figure 6 of the J. alviteziana web page) where the thickenings are whitish. Radial thickenings show in the photo to the right as green radially oriented lines in the base of the corolla.

Filaments Filiform; markedly enlarged / swollen at base where adnate to base of corolla.
The expanded base of the filament is usually pubescent with finger hairs; the slender part of the filament is often but is not always pubescent with finger hairs.
 
Anthers Ovate to linear; the filament is inserted either on the lower ventral face of the anther or basifixed; to 5 mm long; dehiscing longitudinally; connivent in distantly related species (J. procumbens race 321, J. paneroi).  
Ovary Superior, with an annular disk(1) around the base of the ovary, bicarpellate (rarely tricarpellate in J. auriculata, and J. repandidentata I grew)  
Ovules per ovary 42 to about 700  
Style Straight (exceptions: the style of both J. darcyana and J. repandidentata are curved);
exserted or not;
slender (the styles of the three species that have a corona are thickest at the base and gradually decrease in diameter proximal to distal) 
 
Stigma Single, shallowly bilobed or bilobed;
capitate (punctiform on the three species of section Modillonia);
self-compatible in the species studied to date
 
Fruits juicy globose or oblate berry; 4 - 25 mm in diameter; many-seeded;
the berries of at least one species split open at maturity (J. chotanae)
 
Fruit Color black/purple, green, orange, red or nearly white  
Seeds: to 395, the quantity per fruit varies among species, ovate to reniform  
Chromosome base number
12
Davis 1975, Mione 1992
Number of species worldwide
70
 
Nectar

Color & Relative Volume - clear (footnote 2) and produced in small volume, or in contrast red / orange and produced in much greater volume - filling the base of corolla and sometimes visible through the corolla. 

Follow this link to see a list of the species having red / orange nectar, a subset of the Peruvian and Bolivian species.

Where does nectar collect? Nectar pools on the base of the corolla and around the base of the androecium in all species except three: In the species of section Modillonia nectar pools on the corona (see above, on this page, for more information about the corona).

 

 

 Footnotes for the above: 1) The annular disk around the ovary (ovarian disk, the expanded base of the ovary) has been described as a nectary disk ("disco nectarífero" Benítez de Rojas 2010) but there is no evidence that the ovarian disk makes nectar.

2) the nectar produced in J. darcyana, J. procumbens, J. repandidentata, and J. sinuosa isn't actually clear, but rather a very dilute pale yellow (J. Kostyun, personal communication, 2014).

Similar Genera

     Jaltomata differs from Physalis by the former having a fruiting calyx that does not enclose the berry, while the fruiting calyx of Physalis encloses the berry.  Jaltomata differs from Solanum by the former having longitudinal anther dehiscence and an ovarian nectar disk, while Solanum has anthers with terminal pores and no ovarian disk nor nectar.  Jaltomata differs from Cuatresia by the former having basal pedicel articulation while the latter lacks pedicel articulation (Hunziker).

Habitat:

I have collected Jaltomata species most commonly along roadsides where the native vegetation is still present to some extent. If the native vegetation has been entirely eliminated for agriculture I usually can't find Jaltomata. In other words, most species are tolerant of a considerable amount of open sun, but not completely open sun. However, J. nigricolor and J. salpoensis are found in completely open sun, with no shade whatsoever. In the Andes, rock walls (J. sanchez-vegae) and steep roadside embankments are common habitats of Jaltomata species. The easiest way to find J. repandidentata is in coffee plantations that have not been weeded (Mexico to Bolivia), the coffee plants providing considerable shade. Even in the forest, when I have found Jaltomata it was along a trail or gravel road (Costa Rica) where light was entering through the canopy at least part of the day. In Mexico, the easiest way to find J. procumbens is in corn fields that have not been weeded, the corn plants usually providing shade depending on the height of the corn. Some Jaltomata species do not stand alone, and require other plants to grown among (J. dendroidea and J. pallascana, for example); here the physical support of other plants and the shade of other plants are inseparable. One species grows, with few exceptions, in between or very near the spiny leaves of Puya.

Most Jaltomata species are somewhere in the middle of the spectrum of dry to moist habitats. On the wet end of the spectrum there are ecotypes of J. procumbens that reside in the rainforest, for example in Costa Rica at high elevation, including an epiphyte at Monteverde. As well, J. werffii grows on wet slopes (Galápagos Islands). On the dry end of the spectrum are the species that reside in the lomas. And J. calliantha (red/orange nectar and green fruits) dies back to the ground for several months of the year; its habitat is seasonally quite dry. One of the lomas species has deciduous leaves, and it may receive no rainfall for months, and then when the fog returns, approximately in August, the leaves emerge again.

Geographic Distribution:

The approximately 60 species of Jaltomata are distributed from southeastern Arizona, USA (Kearney and Peebles, 1951) to southern Bolivia (Morton, 1944), on three of the Galápagos Islands (D'Arcy, 1982), and on the Greater Antilles (Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico; Adams, 1972; Davis and Bye, 1982; Liogier and Martorell, 1982). The majority of the species are Andean but one of the most common species, J. repandidentata, extends throughout much of the continental range of the genus, from Mexico to Bolivia (Mione and Yacher, 2005). The USDA Plants Profile (web site) for Jaltomata procumbens shows this species in Maryland, U.S.A. I have not seen collection(s) made from plants growing in Maryland. Highest Collection: J. yungayensis at 4,150 m of elevation (J. Mostacero L, S. Leiva G. et al. 1897).

In the department of Cajamarca, Peru, there are over 20 distinct and defendable species!

In the department of La Libertad, Peru there are so many species!

Link to Jaltomata of Arizona, Mexico and Central America       

In Connecticut, USA, where I have grown outdoor plants of J. procumbens, seeds in fruits dropped to the ground in the fall have germinated during the following spring, becoming weeds in the garden where they were grown the year before. This suggests that the northern end of the range of this genus is limited by dispersal and / or successful establishement among competitors, not by sensitivity to cold.

Literature 

Figure 1. Inflorescences are normally umbel-like in the genus Jaltomata
(J. procumbens, photo by T. Mione)
Figure 2. A few species of the genus Jaltomata have branched-umbel-like inflorescences (J. bernardelloana photo by T. Mione in Peru).