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. D—Distal 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

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:

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).

Phenology

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).

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

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.

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.

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