Jaltomata aijana Mione & S. Leiva

Peru
revised 22 Sep 2017 
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. 
C
—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. Jaltomata aijana. A—Fruit in side-view, approximately half-hidden by calyx.
B
—Branch with inflorescence and leaf with an expanded view of a dendritic hair.
C
—Open flower. D—Anther in ventral view. E—Seed. F—Anther in dorsal view. G—Gynoecium.
H—Corolla with adnate stamens dissected open longitudinally. 
I
—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 3. Flowers, leaves, Jaltomata aijana, numbered units are cm, photo by Thomas Mione in Peru, Mione et al. 863
above, Figure 4. 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.

 

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  
Inflorescence
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
no
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  
Stigma
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  
Nectar
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)  
Herkogamy    
Protogyny yes Mione 872, cultivated
Fruit color (at maturity)
The fruit color was not mentioned in the protologue, and was not mentioned by Macbride (1962, p. 40) for Hebecladus weberbaueri. Jamie Kostyun grew this species (Mione et al. 726) and after an inquiry about mature fruit color responded (personal communication, 2016) that “I remember that ripe fruits were orange.”  Orange fruits would be predicted given the phylogenetic position of Jaltomata aijana in “Clade 3 orange fruits” (Miller et al. 2011).  Surprisingly, we 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.
Berry likely orange at maturity
Fruit Size and Seeds per fruit

9 mm high X 13.5 mm wide containing 82 seeds;
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

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?
no data
 
Flowers Closing For The Night?
No Mione 872 cultivated at CCSU
Self-Compatible?
Yes
Jamie Kostyun, personal communication 2013, Mione 726 cultivated
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
 
zxcv
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
photo.of.Mione863
abaove, Figure 7. Distal end of flower, Jaltomata aijana, photo by Thomas Mione in Peru, Mione et al. 863 above, Figure 8. Flower openned 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).

 

Phenology

Jaltomata aijana flowers November through June. Flowers of J. aijana 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 on plants cultivated in Connecticut, USA. Anther dehiscence within a flower is temporally staggered (Figure 1B). This species is self-compatible (Kostyun, personal communication, 2014) and flowers do not close for the night (observations made on plants cultivated in Connecticut, USA).

 

Habitat
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.

 

Phylogeny
 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

elevation
m

habitat

date

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

2800

Low but closed tropical dry forest vegetation with Arnaldoa, Llangunoa

13 Mar 2001

M. Weigend et al. 5151 (M)

Huaraz. km 186 Recuay - Huaraz

3150

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

3450

brush borders in agricultural land

27 Jan 1985

D. N. Smith et al. 9388 (NY)

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

3300

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

3820

growing among Opuntia in clay soil

3 Apr 1999

D. M. Spooner et al. 7356a

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

3971

steep roadside

20 May 2016

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

Bolognesi. Arriba de Chiquián

35003600

falda pedregosa

31 Mar 1957

R. Ferreyra & E. Cerrate 12133 (USM)

Bolognesi. Chiquián

34003900

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

3338

roadside

20 May 2016

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

Bolognesi. Chiquián

3338

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

3645

steep roadside

15 Jun 2005

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

Recuay. Bosque de Pararín

2800 3000

bosque

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

3600

growing among Opuntia and shrubs

3 Apr 1999

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

Recuay. km 107 between Recuay and Pativilca

3300

moist ravines

30 Jan 1983

M. Dillon et al. 3178 (BH, F)

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

3500

quebrada de arbustos

17 Nov 1995

S. Leiva G. 1738 (HAO)

Recuay. Culquimarca (Pativilca-Conococha)

3600

en ladera de arbustos

27 May 1970

A. López M. et al. 7601 (US)

Aija. En la bajada de Tranca hacia Huayán

30003200

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

2945

on a rocky slope with grasses, Calceolaria, Solanum hirsutum

1 April 1999

D. M. Spooner et al. 7352c (MOL)

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

3309

along river and edge of agricultural field

14 Jun 2005

T. Mione, S. Leiva G. & L. Yacher 726

Aija. Road from Aija to Huarmey

3474

roadside

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.

 

character

J. aijana Mione & S. Leiva

J. bicolor (R. & P.) Mione

J. biflora (R. & P.) Benítez

Distribution

Peru: Department Ancash

Peru: Departments Lima, Huancavalica, and Moquegua

Peru: Department Junín

Altitude (m)

28003970

21003800

27003200

Corolla color

green

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

green

Corolla shape

straight-tubular

urceolate-tubular

urceolate

Indument of external face of corolla tube

unpigmented

purple

unpigmented

Radial thickenings in corolla1

no

no

yes

Nectar (floral) color when plants are cultivated

clear, turning orange with age2

clear

clear, turning orange with age3

Fruit, at maturity, hidden by calyx?

upper ¼ to 1/2 of fruit hidden by calyx

no

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)

206

196, 216

119, 161, 166

Fruits eaten by humans

no data

Yes4

Yes4

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. 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.

 

 

Observations in Connecticut, Outdoor-grown plants, September 2017.
Bees (unidentified) climb into the corolla tube 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, at the time of this observation had no nectar. Presumably, the bees that climb into the corolla tube had been consuming the nectar. Mione et al. 872.

 

 

 

ACKNOWLEDGEMENTS

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