Jaltomata aijana Mione & S. Leiva

revised March 2020 
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.
Pyrex Journal of Biodiversity
and Conservation 1(3): 35-43. 2016.
Link to the Jaltomata of Ancash, Peru
Link to local names including of this species
Replaced synonym: Hebecladus weberbaueri Dammer, Bot. Jahrb. Syst. 37(5): 638. 1906
[not J. weberbaueri (Dammer) Mione, Novon 2(4): 383. 1992; Basionym: Saracha weberbaueri Dammer, Bot. Jahrb. Syst. 37(5): 638. 1906].
This taxonomic revision corrects the erroneous placement of H. weberbaueri Dammer in synonymy with J. bicolor (Mione et al., 1993).  
Figure 1. Jaltomata aijana. A—Flower in pistillate phase, anthers are purple. B—Flower near center has two dehisced and three undehisced anthers showing nonsimultaneous anther dehiscence. The flower on the left opened more recently; we can infer this because the stamens are not yet exserted. 
—Corolla was opened longitudinally to show androecium and gynoecium; two of the five stamens were removed. DDistal end of flower, showing alternating lobes and lobules (10 total) and dehisced anthers. E—Woody stem, smallest units are mm. F—Calyx in overhead view, attached to fruit hidden under the calyx, units along bottom are mm.
All scale bars equal one cm. Photos are of Mione et al. 863 = Leiva G. et al. 5892 except A is Mione et al. 728.
Photos were taken by T. M. in the field except B was taken by S. L. G.
in the field.

Figure 2. Ripe fruits of Jaltomata aijana. Plant grown in CCSU greenhouse. Units are mm. One of the fruits measured 8.7 high by 13.5 wide, and the other 8.9 wide by x 14 mm wide. Mione et al. 872

Figure 3. Jaltomata aijana. A—Fruit in side-view, approximately half-hidden by calyx.
—Branch with inflorescence and leaf with an expanded view of a dendritic hair.
—Open flower. D—Anther in ventral view. E—Seed. F—Anther in dorsal view. G—Gynoecium.
H—Corolla with adnate stamens dissected open longitudinally. 
—Base of staminal filament in ventral view. J—Anther in lateral view. 
Illustration by Segundo Leiva González, S. Leiva G. et al. 5892

above, Figure 4. Flowers, leaves, Jaltomata aijana, numbered units are cm, photo by Thomas Mione in Peru, Mione et al. 863


Description of Jaltomata aijana based on numerous herbarium specimens from throughout the range of the species:

Character Description Figures on this page
Habit & Height
Shrub to 2 (3 m) high.  
Branches, young

younger stems 4- or 5-sided, purple on upper surfaces, green on lower surfaces, sparsely to densely pubescent with eglandular unpigmented mostly simple hairs 

older brown, terete, with lenticels, glabrous, to 15 mm diameter at the base  
Leaves, size
Basal leaves alternate, the distal geminate; the blade ovate, to 8 (14.5) cm long X 5 (12) cm wide, the younger membranous, the older somewhat coriaceous, the upper face sparsely pubescent to glabrous, the lower puberulent to glabrous, the apex usually acute,  
shape the base of smaller leaves at times cuneate, the base of the largest leaves truncate, the margin usually entire but the largest are sinuate to sinuate-dentate; Leaves single, paired or verticillate; usually ovate but variable in size, shape and margin.      
hairs glabrate to sparsely pubescent with finger-like and dendritic hairs hypocotyl and cotyledons of seedlings are pubescent with unpigmented conspicuously gland-tipped finger hairs
(Oct 2016, Mione et al. 872)
petiole the petiole to 6 mm (3 cm) long, with a conspicuous main vein abaxially  
flowers  25 (6) per inflorescence  
peduncle green, terete, sparsely pubescent, to 3.1 cm long  
pedicel purple, terete, sparsely pubescent, to 1.4 cm long  
Calyx during anthesis
abaxially green to purple, five-lobed (stellate in outline), 1.6 cm diameter, sparsely pubescent with simple to dendritic eglandular hairs, the lobes triangular, to 6.5 mm long X 5 mm wide, the main vein somewhat raised abaxially.  
shape / position when flowering planar to recurved (Figure 1A, B)  
at fruit maturity enclosing 1/3 to 1/2 of the fruit (Figure 2A),
to 3.1 cm diameter; 15 mm from pedicel to lobe tip
Corolla color
Corolla green  
green spots no  
purple ring no  
purple in base of corolla yes: at least some collections having purple at the base Figure 11
shape and size

tubular, the tube and limb together to 3.5 cm long, the tube narrowing slightly from proximal to distal, 610 mm diameter at midlength

lobes / lobules 10 total: the lobes alternate with the inconspicuous lobules.  
hairs the exterior of the corolla densely pilose, the hairs simple or sometimes dendritic, the interior of the tube glabrous, the limb pubescent on both faces  
Stamen length including anther
5, adnate to base of corolla, connivent, hidden inside corolla tube when corolla opens, elongating at unequal rates but finally equal, 3.13.5 cm long including anther, exserted beyond mouth of corolla  
length stamens exserted beyond distal end of corolla (applicable if corolla has a well-defined tube)
0 - 8 mm in Department Ancash
radial thickenings
726, 728, 872 were dissected carefully
base expanded laterally? yes Figure 2I and dissection microscope observations by Mione September 2017
filaments pale-green, the proximal 10% villous with unpigmented, eglandular, simple hairs
anther color anthers dark purple, the connective greenish Figures 1A, 1C, 13
anther size sagittate after dehiscence, glabrous, conspicuously longer than wide, 3-3.5 mm long X 1.8-2 mm wide (fresh), 2.5-3 mm long X 1-1.5 mm wide (pressed specimens), finally in close proximity with stigma or to 2 mm beyond stigma  
anther mucronate?

yes: mucronulate
(not showing in Figure 2)

dissecting microcope observations by Mione on fresh anthers from cultivated plant of Mione et al. 872
insertion of filament into anther directly under anther Figure 13
anthers of a flower with temporally staggered dehiscence? yes Figure 1B
pollen grain size    
corona? no  
capitate (Figure 1C, 2G), dark green, exserted, shallowly bilobed, 0.5–1 mm diameter (pressed specimens)  
Style straight, filiform, pale-green, lighter than the stigma, 2.3–2.9 cm long   
Ovary 2.4 mm high X 3.4 mm wide including the annular disk, containing 236 ovules, the annular disk pentagonal, pale-orange and approximately 60 % of the height of the ovary  
Probably translucent most of the time in the field because during field collections in Peru I never noticed colored nectar. Pale-orange during both pistillate and hermaphroditic phases (Jamie Kostyun, personal communication May 2013, cultivated plants)  
Protogyny yes Mione 872, cultivated
Fruit color (at maturity)
Figure 2
Fruit Size and Seeds per fruit

9 mm high X 13.5 mm wide containing 82 seeds (Mione et al. 872);
11 mm (pole to pole) X 13 mm diam not ripe but may represent full size (Mione et al.863);
22, 30, 51 seeds per fruit (Mione et al. 863)
206 seeds per fruit

Data from the field is not comprable with data from the greenhouse: for example,
. aijana collection Mione et al. 872 had 82 seeds in one fruit in the field (year 2016), and 38 to 40 seeds per full-size, ripe fruit (weeks after manual self-pollinations) in the greenhouse (year 2018).

Berry globose, compressed at the poles, to 11 mm (pole to pole) X 13 mm diameter, the style persistent
Seeds brown, sub-orbicular to sub-triangular, the surface reticulate (Fig. 2E),
1.55–1.7 X 1.25–1.4 X 0.5–0.6 mm thick
Mione's collection numbers of this species 625, 627, 726, 728, 863, 870, 871, 872  
For other descriptions see
protologue and description of Hebecladus weberbaueri Dammer in Macbride (1962)  


Character, Jaltomata aijana    
Growability in Connecticut, USA
Will grow and flower in a greenhouse, and in a window of an air conditioned building, in Connecticut Mione et al. 872
How long does it take from flower to ripe fruit?
5 weeks and 2 days (two fruits, greenhouse) Mione et al. 872
Flowers Closing For The Night?
No Mione 872 cultivated at CCSU
Mione et al. 872 at CCSU
Seed Germination
26 days from planting to first above-ground signs of germination, seeds in cup of potting mix on warm mat, no GA
Mione et al. 872
Pollen quantity per flower 65,450 and 79,063 Mione et al. 728, n = 2 flowers
93,250 Mione et al. 726, n = 1 flower
flowers collected in Peru (not grown for study),
counts by Elisabeth dos Santos and Emmett P. Varricchio
Ovules per ovary
236 Mione et al. 728, n = 1 flower count by Thomas Mione, the value shown at left is double the count from one locule
Ratio of pollen to ovules
277 (65,450 / 236 ovules) Mione et al. 728, n = 1 flower
Chromosome number
no data
above, Figure 5. A flower bud was pulled open longitudinally to allow view of stamens. Anthers are dark purple. Given that stamens of open flowers of this collection were 31 to 35.5 mm long, and filaments in this photo are approx 1 mm long, we can conclude that filaments elongate tremendously between the floral bud stage (showing) and the flower being functional. Photo by Thomas Mione in the field in Peru, Mione et al. 863. above, Figure 6. Flower with petal removed to show stamens and style, Jaltomata aijana, photo by Thomas Mione in Peru, Mione et al. 863
abaove, Figure 7. Distal end of flower, Jaltomata aijana, photo by Thomas Mione in Peru, Mione et al. 863 above, Figure 8. Flower opened recently, probably within the last few hours (we can infer this because the stamens have not yet elongated enough for us to see them in a view perpendicular to the corolla tube). Photo by T. Mione in Peru, Mione et al. 863.


Figure 9. Note the orange nectar inside the corolla tube, evident only in cultivated plants, not seen in the wild. Plants grown by and photo by Jamie Kostyun, Mione et al. 726. Figure 10. Note the orange nectar inside the corolla tube, evident only in cultivated plants, not seen in the wild. Plants grown by and photo by Jamie Kostyun, Mione et al. 726.


We collected specimens of the genus Jaltomata in Peru in 1998, 1999, 2005, 2007, 2008, 2010, 2011, 2013, 2015 and 2016. Specimens have been deposited at CONN and HAO. Herbarium specimens were borrowed from or studied at BH, CONN, F, G, GH, K, M, MA, MO, MOL, NY, P, US, USM, VT and WIS with herbarium acronyms as in Thiers (continuously updated). Seeds of Mione et al. 872 were measured and Leiva González et al. 5892 was used for the seed count. Other characters were measured on all of the specimens we collected in Department Ancash (Table 1).



Jaltomata aijana flowers November through June. Flowers are protogynous: both open flowers having undehisced anthers (pistillate phase) and open flowers having dehisced anthers (hermaphroditic phase) were seen on the same plant at the same time (Figure 1B) both in the field on plants cultivated in Connecticut, USA. Anther dehiscence involves anther color change from purple (undehisced) to greenish (dehisced), and is temporally staggered within a flower (Figure 1B). This species is self-compatible and flowers do not close for the night (observations made on plants cultivated in Connecticut, USA). In the greenhouse, during February of 2020 I tagged a flower to observe it daily: The stigma protrudes from the closed corolla for one day; the following day the corolla is open but the anthers remained undehisced (pistillate phase); the following day all anthers dehisced by 9 am (temperature 17.5 C) and the flower remained functionally hermaphroditic for the remainder of this day and the next; the following day the corolla-androecium abscised by 10 am. From pollination (by hand to record the date) to ripe fruit takes 5 weeks and 2 days (Mione et al. 872, 2 fruits 20 Jan to 26 Feb 2018, and one fruit 27 Jan to 4 March 2020).


The habitat of Jaltomata aijana is open and shrub-covered hillsides, grazed rocky slopes, moist ravines, crevices in stone walls, along edge of agricultural field where it meets a river, roadsides.  It lives in association with plants of Eucalyptus amygdalina Labill. “eucalipto” (Myrtaceae), Bidens pilosa L. “cadillo,” Viguiera weberbaueri S. F. Blake “suncho,” Ambrosia peruviana Willd. “marco,” Tagetes minuta L. “chiche,” Mutisia acuminata R. & P. (Asteraceae), Jaltomata weberbaueri (Dammer) Mione “frutilla,” Brugmansia sanguinea (R. & P.) D. Don “floripondio rojo” (Solanaceae), Pisum sativum L. “arveja,” Otholobium pubescens (Poir) J. W. Grimes “culén” (Fabaceae), Rumex crispus L. “acelga” (Polygonaceae), Alnus acuminata Kunth “aliso” (Betulaceae), Passiflora tripartita (Juss.) Poir. “puro puro” (Passifloraceae), Sambucus peruviana Kunth “saúco” (Adoxaceae), Alonsoa meridionalis (L. f.) Kuntze (Scrophulariaceae), and the genera Urtica L. (Urticaceae), Cestrum L., Solanum L., Salpichroa Miers (Solanaceae), Calceolaria L. (Calceolariaceae), Barnadesia Mutis ex L. f. (Asteraceae) among others.


 Jaltomata aijana
was included in the molecular phylogeny of Miller et al. (2011) as “J. bicolor 726” where it was found to be most closely related to J. umbellata (R. & P.) Mione & M. Nee in both the strict consensus and the majority-rule consensus trees. We now understand that the circumscription of J. bicolor was artificial when Miller et al. (2011) published, and that J. bicolor and J. aijana are distinct and defendable species (Table 2). 


Local names



Table 1. All specimens of Jaltomata aijana examined, all collected in Department Ancash, Peru.

Province. Locality




collector (herbarium)

See footnote 1.

3200 3400

interwoven grasses and bushes ("graminosis fruticibus intertextis")2

27 March 1903

Type of Hebecladus weberbaueri Dammer.
Weberbauer 2652: Isotype G!, photos of B specimen (destroyed) GH!, NY!

Huari. road from San Marcos to Chavín de Huántar. Río Mosna valley near Olayan


Low but closed tropical dry forest vegetation with Arnaldoa, Llangunoa

13 Mar 2001

M. Weigend et al. 5151 (M)

Huaraz. km 186 Recuay - Huaraz


on dry ground along bank of tributary to Río Santa

Jan 1998

T. Mione, S. Leiva G. & L. Yacher 627

Huaraz. 15 km S of Huaraz, 77 29' W, 9 41' S


brush borders in agricultural land

27 Jan 1985

D. N. Smith et al. 9388 (NY)

Bolognesi. km 107, poblado Sta. Rosa (ruta Pativilca-Recuay)


borde de río entre Ambrosia sp., Barnadesia

18 Jan 1998

S. Leiva, T. Mione & L. Yacher 2133 (F)
T. Mione, S. Leiva G. & L. Yacher 625

Bolognesi. Huacacorral, 22.4 km E of Conococha, 77.2 W, 10.1 S


growing among Opuntia in clay soil

3 Apr 1999

D. M. Spooner et al. 7356a (WIS)

Bolognesi. 9 57' 23.5" S, 77 05' 57.0" W


steep roadside

20 May 2016

T. Mione, S. Leiva G. & L. Yacher 870
S. Leiva, T. Mione & L. Yacher 6048

Bolognesi. Arriba de Chiquián


falda pedregosa

31 Mar 1957

R. Ferreyra & E. Cerrate 12133 (USM)

Bolognesi. Chiquián


fields, pastures and quebrada above village

5 Feb2 Apr 1997

M. Weigend & N. Dostert 97/185 (F)

Bolognesi. Below Chiquián
10 08' 16.3" S, 77 09' 43.2" W



20 May 2016

T. Mione, S. Leiva G. & L. Yacher 871
S. Leiva, T. Mione & L. Yacher 6049

Bolognesi. Chiquián


hedgerows, trail edges

20 May 2016

T. Mione, S. Leiva G. & L. Yacher 872
not pressed, seeds only 

Bolognesi. highway Pativilca - Recuay, 
10 9' 47" S, 77 20' 16" W


steep roadside

15 Jun 2005

T. Mione, S. Leiva G. & L. Yacher 728
digital photos only, not pressed

Recuay. Bosque de Pararín

2800 3000


24 May 1988

N. Valencia 2168 (NY)

Recuay. km 110 road from Lima to Huaraz, 19.7 km N of Cajacay, 77.3 W, 10.2 S


growing among Opuntia and shrubs

3 Apr 1999

D. M. Spooner et al. 7359a (BH, P, WIS)

Recuay. km 107 between Recuay and Pativilca


moist ravines

30 Jan 1983

M. Dillon et al. 3178 (BH, F)

Recuay. ca. Laguna Conococha, km 107 (Pativilca-Recuay)


quebrada de arbustos

17 Nov 1995

S. Leiva G. 1738 (HAO)

Recuay. Culquimarca (Pativilca-Conococha)


en ladera de arbustos

27 May 1970

A. López M. et al. 7601 (US)

Aija. En la bajada de Tranca hacia Huayán


no data

23 Apr 1983

C. Ochoa & A. Salas 15,168 (F)

Aija. 7 km E of town of Aija, canyon San Juan (containing Río San Juan), at Cuesta de Melliso, 77.6 W, 9.8 S


on a rocky slope with grasses, Calceolaria, Solanum hirsutum

1 April 1999

D. M. Spooner et al. 7352c (WIS)

Aija. 6 km down from Aija
9 48' 18" S, 77 36' 18" W


along river and edge of agricultural field

14 Jun 2005

T. Mione, S. Leiva G. & L. Yacher 726

Aija. Road from Aija to Huarmey



20 May 2015

T. Mione, S. Leiva G. & L. Yacher 863
S. Leiva G., T. Mione & L. Yacher 5892 (HAO)


1. In the protologue the type locality is given as “Prov. Cajatambo, dep. Ancachs: Ocros 3200-3400 m.” On modern maps Ocros is the capital town of the province Ocros, at the south end of the department of Ancash, while province Cajatambo is at the north end of the department of Lima, just south of the department of Ancash. 

2. from the protologue.

above, Figure 11. Two corollas of Jaltomata aijana, removed from flowers and inverted to show purple pigmentation in base of corolla.
Photo by Thomas Mione in Peru, Mione et al. 863.


Table 2. Comparison of Jaltomata species most similar to J. aijana. All three species are shrubs having 10-lobed corollas (the lobes and lobules alternating),
purple anthers, unpigmented floral nectar (in the field) and orange fruits.



J. aijana Mione & S. Leiva

J. bicolor (R. & P.) Mione

J. biflora (R. & P.) Benítez


Peru: Department Ancash

Peru: Departments Lima, Huancavalica, and Moquegua

Peru: Department Junín

Altitude (m)




Corolla color


proximal 2/3 intensely purple, distal 1/3 green


Corolla shape




Indument of external face of corolla tube




Radial thickenings in corolla1




Nectar (floral) color when plants are cultivated

clear, turning orange with age2


clear, turning orange with age3

Fruit, at maturity, hidden by calyx?

upper ¼ to 1/2 of fruit hidden by calyx


upper ¼ to 1/3 of fruit hidden by calyx

Fruit size (mm)(measured in the field, not cultivated)

9 x 13.5
11 x 13

8.5 x 18

11 x 16

Seeds per fruit (fruits collected in the field)


196, 216

119, 161, 166

Fruits eaten by humans

no data



Synonyms and page number(s) in Macbride (1962)

Hebecladus weberbaueri Dammer, p. 40

Hebecladus bicolor (R. & P.) Miers, p. 30; Atropa biflora R. & P., p. 30; H. intermedius Miers, p. 35.

Saracha biflora R. & P., p. 31

Author's collection numbers

625, 627, 726, 728, 863, 870, 871, 872

612, 617, 795, 880

608, 876

1. A subset of species of section Hebecladus have radially oriented thickenings that extend from the base of each stamen toward the corolla lobule (or where
the lobule would be if present); where present, these create troughs (between radial thickenings) that hold nectar at the base of the corolla. 
2. Nectar turning orange with age has been observed in the CCSU greenhouse on plants grown by T.M., and by Jamie Kostyun, personal communication (2013), from observations on cultivated plants.
3. Mione et al., (2001).
4. Leiva González et al., (2016a, pp. 49
50) and fieldwork by the authors.





Figure 12. Jaltomata aijana in Department Ancash, Peru.  Leiva G., Mione & Yacher 5892 = Mione, Leiva G. & Yacher 863, Photo by Segundo Leiva G.
Figure 13. Ventral (inner) face of anther on left, and dorsal (outer) face on right, of J. aijana. Photo by T. Mione, cultivated plant of Mione et al. 872. Units vertically on right are mm. Two photos, with a slight focus adjustment between, were stacked in photoshop.
above, Figure 14. Infructescence (the fruit showing is unripe), pedicels and peduncle of Jaltomata aijana. Units along bottom are mm. Photo by Thomas Mione in Peru, Mione et al. 863.



Surprisingly, Segundo Leiva G. and T. M. made a single observation (20 May 2015, Mione et al. 863) of fruits of J. aijana with purple pigmentation. This was the first observation of purple pigmentation in the fruits of any species of section Hebecladus (Mione, 1992) and we suspect that the purple fruits we saw on J. aijana were a developmental error and/or atavism.



Observations in Connecticut, Outdoor-grown plants, September 2017.
Bees (unidentified) climb into the tubular corolla of J. aijana and remain in the tube for several seconds before backing out. Flowers harvested and then torn open longitudinally and observed with a dissecting microscope, immediately after this observation, had no nectar. Presumably, the bees that climb into the corolla tube had been consuming the nectar. Mione et al. 872.





I thank David Spooner for sending his specimens to me, Lindsay T. Kmietek for preparation of figure 1, Kanchi Natarajan Gandhi for assistance with nomenclature, and Gregory J. Anderson and Gabriel Bernardello for providing an environment conducive to the birth of this project.

Literature Cited