Jaltomata biflora (R. & P.) Benítez
Peru
revised 26 July 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.
Rev. Fac. Agron (Maracay), IX (1): 91 - 92.
Link to list of Jaltomata species having edible fruits, including this species
Link to Jaltomata of northern Peru
(does not include this species)
Link to local names including of this species
Link to list of Jaltomata species having red / orange nectar

Figure 1.

Three anthers have dehsiced,
two (of shorter stamens)
have not yet
dehisced.

Photo by Thomas Mione

Figure 2.

The above photo and the photo on the right
are of Daniel Mugaburu's collection,
grown as Mione 608
and photographed in Connecticut
by Thomas Mione.

 

Mione is indebted to Daniel Mugaburu
for seeds and pressed specimens
of this species
collected at the type locality.

Figure 3.

Photo of the type specimen
housed in Paris.


Smallest units on ruler are mm.

 


Illuminated with four incandescent bulbs
photo by Thomas Mione



 

Saracha biflora Ruiz & Pav. Flora Peruviana 2: 42-43, t. 179. 1797-99 [not Atropa biflora, a synonym of J. bicolor].

 

Character Description of Jaltomata biflora Figures on this page
Habit & Height

sprawling woody shrub
Macbride (1962 page 31, as Saracha biflora) incorrectly described this species as herbaceous.
Branches, young
very densely hairy, gland-tipped finger hairs  
older
   
Leaves, size
   
shape    
hairs    
petiole    
Inflorescence
1 - 2 flowered (my observations)
"2- or rarely 3-flowered" (Macbride 1962, page 31, as Saracha biflora)
 
peduncle
green, to 15 mm  
pedicel
green, to 15 mm  
Calyx at flowering
green, dense indument of gland-tipped finger hairs  
shape / position when flowering rotate 6
at fruit maturity    
Corolla color
green  
shape and size
urceolate, 1.3 cm long, the base to 1.4 cm in diameter, the limb becoming revolute  
lobes/lobules
10-total: narrowly triangular lobes alternating with inconspicuous lobules  
hairs
dense indument of gland-tipped finger hairs  
Stamen length including anther
22 - 23 mm 7
length stamens exserted beyond distal end of corolla (applicable if corolla has a well-defined tube) 7.5 - 14 mm based on both plants I grew and photos of the type specimen at P
yes 4
base expanded laterally? yes, as seen in ventral view 5
filaments pubescent on proximal 36 - 39 % of length 5, 7
anther color purple before dehiscence  
anther size    
anther mucronate / mucronulate no  
insertion of filament into anther    
anthers of a flower open simultaneously? During April, two to four of the anthers of a flower dehisced one day and the others dehisced the morning of the next day (n = six flowers). In May, however, when daytime temperatures in the greenhouse were noticeably warmer, all of the anthers of a flower dehisced in one day by 10:00 am (n = four flowers). Mione et al. 2001
pollen grain size    
corona no  
Stigma
shallow medial groove, shallowly bilobed  
Style
12 -- 18 mm, with the extremes of this range present on the same plant at the same time 4
Ovary
ovarian disk 4
Nectar
Nectar was clear at first, and although inconspicuous, could be seen through the corolla wall. Nectar became more conspicuous toward the end of the hermaphroditic phase: it turned amber to orange in color, as viewed both through the corolla wall and when removed with a clear pipette tip. Mione et al. 2001
Herkogamy yes prior to anther dehiscence  
Protogyny yes  
Fruit color (at maturity) and size
orange, up to 14 mm diam in greenhouse;
 11 X 16 mm containing 119 seeds in wild (Mione et al., 876)
 
Seeds per fruit
161 and 166
"over 300" (Mugaburu, personal communication)
n = 2 fruits collected in Peru, Mione et al. 2001
Seed Size
   
Character Description of Jaltomata biflora Figures on this page

 

Character Description of J. biflora  
Growability in Connecticut, USA
Easy to grow Mione et al. 2001
How long does it take from flower to ripe fruit?
no data
 
Flowers Closing For The Night?
no Mione et al. 2001
Self-Compatible?
yes Mione et al. 2001
Seed Germination
After no germination for weeks while being kept moist, I let the potting mix (in a cup) containing the seeds dry out completely and then put the cup in the mist room, and then several seeds germinated!  
Pollen quantity per flower
104,167 n = 1 flower, Elisabeth dos Santos 2011

n=1 flower, Kenneth C. Plourd 2016
167,500 grains
Ovules per ovary
174 the ovules of one locule were counted by T. M. and then doubled
Ratio of pollen to ovules
963 167,500 grains/174 ovules from the same flower
Chromosome number
no data
 
Character Jaltomata biflora  

 

 

 

 

Figure 4. Longitudinally dissected flower of Jaltomata biflora; dissection and photo by Melissa Luna, D. Mugaburu 5 grown as Mione 608, flower from a plant grown in Connecticut and then preserved in 70% ethanol.

 

 

Country, Department Province

Locality elevation m habitat date collector Data Entry
Peru, Junín,  Tarma "huasa-huasi" on the label of the types and "Huassahuassi" in the protologue. Modern: Huasahuasi       Dombey
April 2009
Peru, Junín,  Tarma Carpapata, above Huacapistana 2700 - 3200 edge of forest
June 1929
E. P. Killip & A. C. Smith 24396 (NY, US)
April 2009
Peru, Junín, Tarma walked up mountain from town of Huasahuasi 2965 - 3030 sunny, among shrubs
June 1996
Daniel Mugaburu 5 & 6, grown as Mione 608
March 2009
Peru, Junín, Tarma up mountain from town of Huasahuasi
11 14 58.9" S, 75 40' 57.4" W
3109 among shrubs 23 May 2016 T. Mione, S. Leiva G. & L. Yacher 876 = S. Leiva G., T. Mione & L. Yacher 6071
July 2016

 

In the The Journals of Hipólito Ruiz Ruiz wrote (page 113) "During the 11 months that we remained in the province of Tarma [one of the provinces of the department of Junin, Peru], exploring the area and carrying out botanical excursions in all directions, we discovered in those gorges and at the edge of the forest a considerable number of trees, shrubs, and herbs that had familiar uses, medicinal properties, or economic potential. The following are those that I collected, dried, and described; illustrations were done for nearly all of them." And then he lists numerous species, including (on page 114) Jaltomata biflora (as Saracha biflora). He makes no mention of human uses of this species in this work.

 

The following is modified only slightly from Mione et al. 2001:

The small fruits of Jaltomata biflora are at least occasionally eaten. Until this study Jaltomata biflora (R. & P.) Benítez was known almost exclusively from the type specimens collected by Dombey in the late 1700s and the inaccurate illustration in the protologue (tab. 179a). The lectotype (MA; F neg. 29720: photos at GH, WIS) lacks flowers and fruit, but the isolectotype at P bears intact flowers. Specimen 8422/28 at G may be a type given that it is conspecific and a label on it indicates “Herb. Pavon,” but neither collection number nor locality are present.

Other than the types, only a few collections of J. biflora have been seen among the hundreds of specimens studied at B, BH, BM, C, COL, COLO, CONN, G, F, GH, K, LD, MA, MO, NY, P, US, USM, VT, WIS.

To track down the type locality we noted that it was given as "huasa-huasi" on the label of the types and "Huassahuassi" in the protologue. Also, Saracha biflora (the basionym) was listed by Ruiz among the species discovered near Tarma. With this information we consulted maps and learned that Huasahuasi is a small town in Peru, department Junín, province Tarma.

Daniel Mugaburu traveled to Huasahuasi to: 1) determine if Jaltomata biflora exists there, and if so, 2) assess whether it is common or rare, 3) learn about the habitat of the species, and 4) describe human uses, if any. With the help of a local farmer who served as a guide, D. M. found J. biflora twice at approximately 3000 m of elevation within a few hours hike of Huasahuasi. Its habitat was the narrow corridor of disturbed vegetation between a foot road and an agricultural field. Although the steep slopes of the mountains surrounding Huasahuasi are grazed and have been cleared for cultivation, this sprawling shrub appears to not be rare because it tolerates disturbance.

At least some locals currently call it "capulí silvestre" and "capulí de campo," and the local name was given as "tomatillo cimarron" in the protologue and as "tomate cimaron" on the type. Ruiz (1931) mentioned that fruits were eaten by children, and Daniel Mugaburu's guide said that fruits are occasionally eaten when found along the hillsides, and that the plants are grazed by livestock. Pressed specimens (D. Mugaburu 5 & 6) were deposited at MO and MOL.

For this study of floral biology, seeds were collected at or near the type locality. Plants were grown at the University of Connecticut in a greenhouse with minimum evening temperatures of 7-10 C, and daytime temperatures of 15-29 C depending on cloud cover/sunshine; plants did not flower under warmer conditions present in other greenhouse rooms. Two fruits, collected in Peru, were orange at maturity and had 161 and 166 seeds each. Other observations were made in the greenhouse during April and May of 2000. The corolla (Fig. 1, 2), 1.3 cm long, is green and urceolate with a narrow mouth and broader base 1.4 cm in diameter. The limb (ultimately revolute) is made up of five narrowly triangular lobes alternating with inconspicuous lobules. The stamens and style are exserted beyond the mouth of the corolla on mature flowers. The anthers are purple before dehiscence. The calyx and corolla bear a dense indument of gland-tipped finger hairs. Stalked glands were present but rare on the abaxial face of the calyx. Neutral red stained both the multicellular head of these glands and the tip of the glandular finger hairs.

Flowers remained open six to nine days (mean 7.25 days, n = 10 flowers), and remained open at night. The filaments and the style elongated during the first few days a flower was open, and only after a stamen elongated did its anther dehisce. During April, two to four of the anthers of a flower dehisced one day and the others dehisced the morning of the next day (n = six flowers). In May, however, when daytime temperatures in the greenhouse were noticeably warmer, all of the anthers of a flower dehisced in one day by 10:00 am (n = four flowers). The pistillate phase lasted two to three days (mean 2.3 days, n = 10 flowers) with the stigma protruding through the partially open corolla on the first day. This species is self-compatible. All three of the flowers manually emasculated and self-pollinated during the pistillate phase set fruit, as did all four of the flowers that were manually self-pollinated during the hermaphroditic phase. Many fruits were spontaneously set in the greenhouse in the absence of pollinators. These were orange at maturity, subspherical, and up to 14 mm in diameter.

Nectar was sweet to the taste. A refractometer was used to estimate percent sugars from destructively sampled flowers. Average sugar concentration during the pistillate phase was 14.2 % (range 14 to 14.4, n = 2) and 38.9% for the hermaphroditic phase (range 26 to 57.6 percent, n = 5). Given that the pistillate phase precedes the hermaphroditic phase, and no nectar was removed by pollinators, the higher sugar concentration during the latter phase may have been due to continual nectar secretion and/or evaporation of water from the nectar. Flowers produced 0 to 47 µl of nectar (n = eight flowers). Nectar was clear at first, and although inconspicuous, could be seen through the corolla wall. Nectar became more conspicuous toward the end of the hermaphroditic phase: it turned amber to orange in color, as viewed both through the corolla wall and when removed with a clear pipette tip.

 

Greenhouse observations: Styles measured during the hermaphroditic phase ranged in length from 12 to 18 mm, with the extremes of this range present on the same plant at the same time.   After two to three days of being pistillate, flowers with longer styles automatically self-pollinated because the stigma came in contact with the dehisced anthers.   Flowers having shorter styles were similarly protogynous but were also herkogamous: dehisced anthers and the stigma remained separated by 3 mm and the stigma remained free of pollen in the absence of pollinators.   Style length variation data are preliminary given low sample size (n = 7 measurements) and because for this study T.M. used three plants and did not know if these were from separate seeds, or cuttings of one or two individuals (seeds were sown in 1996 but these observations were not made until the year 2000).  
Figure 5. Stamens (two) attached to base of corolla, showing laterally expanded base of stamen in ventral view, Jaltomata biflora; photo by Melissa Luna, Mione 608, flower from a plant grown in Connecticut and then preserved in 70% ethanol.
Jaltomata biflora held by Peruvian man
Figure 6. Branches, leaves and inflorescenes of Jaltomata biflora. The corolla-androecium has dropped off of the flower on the right, revealing the calyx, yellowish ovarian disk and ovary. Photo by Daniel Mugaburu in Peru at or near type locality.

Figure 7.

Stamens (two,
three were
removed)
attached to
base of corolla

Jaltomata biflora










Photo by Melissa Luna, T. Mione 608, flower from a plant grown in Connecticut and then preserved in 70% ethanol.

 

Literature Cited