Inflorescences and leaves on a plant grown from seed of the type collection. Photo by Mione.
Above: Flower at left was pressed in Ecuador (type collection). Flower at right is fresh (not pressed) and was removed from a plant grown from seed of the type collection a moment before the photo was taken. The plant at left was probably competing for nutrients and sun while the plant at right was not competing for nutrients and sun. This should give us pause when we consider describing a new species from a single collection or from a single cultivated plant. Photo by Mione.
Above: Plant at right was pressed in Ecuador (type collection). Plant at left was pressed in Connecticut, after being grown from seed of the type collection. The plant at right was probably competing for nutrients and sun while the plant at left was not competing for nutrients and sun. This should give us pause when we consider describing a new species from a single collection or from a single cultivated plant. Smallest units on ruler shown in photo are mm. Photo by Mione.
FLORAL BIOLOGY
To look for structures that may release scent (osmophores), flowers were stained in 0.01% neutral red for several hours and then observed with bright field microscopy (Kearns and Inouye 1993). Observations on floral phenology were made in May of 1992 at UConn, and March to May of 1998 at CCSU. Chromosome counts were made with meiocytes of immature anthers stained with acetic carmine. Floral fragrance was evaluated by seven people who were blindfolded and asked to describe the fragrance of an empty jar and then a jar containing flowers. Jars were uncapped seconds before being placed in close proximity to the nose. To examine seed set flowers were manually pollinated during the pistillate and hermaphroditic phases. Undehisced anthers were removed at the time of pollination.
The stigma (including papillae) stained darkly with neutral red but the style did not. Anthers and pollen stained darkly while filaments did not. Multicellular glands, located on the abaxial face of the corolla and both faces of the calyx, stained darkly, except for the stalk cell. Similar glands have been described on the leaves of Solanum (Seithe 1979) and Physalis (Seithe and Sullivan 1990). Neither the corolla margin hairs nor the hairs of the filaments absorbed stain. On the multicellular finger hairs only the glandular tip, when present, absorbed stain. Hand-sections of epidermal tissue of the ovary disk, stained and then observed with a compound microscope, revealed that the nectary disk absorbs stain only in the immediate vicinity of the stomata. The guard cells stained deeply, and the cells surrounding the guard cells also absorbed stain but staining was less intense. Flowers produce a subtle fragrance that was described as licorice-like, vanilla-like, or faintly sweet by seven people polled (two of these people also detected a fragrance in the empty/control jar) and the authors.
Flowers remain open 5 to 7 days (mean 5.7 days, n = 9 flowers) and close each night. Within an inflorescence one to four flowers are open at a time. The corolla is pale-green prior to anthesis but after the corolla opens for the first time it becomes white and remains white. The anthers of a flower either dehisce a few at a time over the course of several hours, or two or three of the anthers dehisce one day and the others dehisce the next day. Anthers dehisced prior to 8:30 am. Some flowers exhibited one day of protogyny, with the stigma protruding through the partially open corolla. Two out of three flowers manually pollinated during the pistillate phase set seed, as did many (percentage not calculated) of the flowers that were manually pollinated during the hermaphroditic phase. In the greenhouse, plants did not set fruit unless hand-pollinated. This is likely due to herkogamy: the stigma is located 3 to 8 mm from the anthers. Nectar drops at the base of the corolla (alternating with the stamens) were large enough to be observed by eye. The broad, orange ovary disk (easily seen in the top photo) is concentric around the green ovary and increases the diameter of the ovary by 1 to 1.5 mm relative to the diameter the ovary would be without the disk. Nectar seems to be secreted by the ovarian disk but may also be secreted by the base of the corolla.
TAXONOMY
Jaltomata lojae (Solanaceae) of southern Ecuador and northern Peru is distinguished by the following features: finger hairs borne by the leaves and stems; a white, broadly infundibular to rotate corolla; exserted stigma; a broad nectary disk; and orange fruits. Corollas change from pale-green to white at anthesis. Flowers are self-compatible, herkogamous, and sometimes protogynous. Microscopic, densely staining, multicellular glands are located on the perianth. Berries are eaten by humans.
Jaltomata lojae is similar to J. sagastegui Mione and J. cajamarca Mione but J. lojae bears only finger hairs on the leaves and stems, the corolla is rotate when fully open, and the ovarian disk is broad, while the latter two species bear dendritic hairs on the leaves and stems, have short-tubular corollas, and narrower ovary disks.
Jaltomata lojae Mione. TYPE: Ecuador. Prov. Loja: by guard station on the E side of Celica, 1950 m, growing along roadside, 2 May 1991, D. M . Spooner, R. Castillo, and L. López 5037 (Holotype: CONN).
The following description is based on the holotype and plants grown, from seed of the type collection, in the University of Connecticut (UConn) and Central Connecticut State University (CCSU) greenhouses.
Perennial shrub; branches and leaves densely villous, bearing finger hairs, these sometimes gland-tipped, 0.15 - 6 mm long, becoming less villous with age. Leaves alternate, often geminate, ovate, to 15 cm long X 6 cm wide, the margin entire to sinuate-dentate, undulate, the petiole to 4.5 cm long. Inflorescence umbelliform, to 9-flowered. Peduncle 5 - 9 mm long (at flowering), pedicel 8 - 11 mm long (at flowering), both having a dense covering of erect finger hairs to 1.5 mm long, some hairs gland-tipped. Calyx (at flowering) light green, 9.5 mm in diameter, strongly reflexed, abaxially villous with gland-tipped finger hairs, forked hairs, and dendritic hairs all 1 - 2 mm long, and abaxially and adaxially with stalked multicellular glands 62 - 80 µm long (Figure 2), at fruit maturity calyx diameter to 12 mm. Corolla infundibular when partially open, rotate when fully open, white with two green, proximally positioned maculae straddling the radial vein to each corolla lobe (Figure 1), 5 prominent lobes alternating with 5 small lobules, 25 - 27 mm in diameter on plants we grew, 18 mm in diameter on the holotype, adaxially glabrous, abaxially with sparsely but evenly distributed stalked multicellular glands 62 - 80 µm long. Two classes of hairs extend out from the corolla margin: marginal hairs to 110 µm long, and submarginal (attached abaxially) hairs to 0.5 mm long. Stamens 5, 5 mm long; filaments on living plants angling away from style, and slightly curved outward, villous on basal 1/4 of the length with unpigmented finger hairs to 1.2 mm long; anthers 2.1 - 2.5 mm long prior to dehiscing, 1.6 - 1.8 mm long after dehiscing, when corolla is fully open exserted out of corolla 2 mm, otherwise included. Pollen grains 32.5 µm average diameter (n = 22 grains). Ovarian nectar disk orange (visible by eye on living flowers). Style 9 - 11 mm long, slender and straight (Figure 1), exserted 3 - 8 mm beyond the anthers; stigma shallowly bilobed, 0.4 to 0.75 mm wide on pressed specimens, broader than the style (Figure 1), the papillae 30 - 60 µm long. Mature fruits orange. Seeds numerous, ovate to reniform, 1.3 - 1.6 mm long. Chromosome number n = 12 (nine counts).
Other Specimens Examined:
specimens made from plants grown from seeds of the type collection Mione 560 (CONN, MO, NY, QCA).
Ecuador. Prov. Chimborazo: Cañon on the Río Chanchan, directly above the village of Huigra, 29-31 May 1945, Camp E-3498 (NY);
Prov. Loja: Lugma Huycu 12 km north of Saraguro, 19 Jan 1989, Ellemann 66799 (AAU not seen, NY); Cerro Sozoranga, Colaisaca-Utuana, km 0.5, 24 Apr 1994, Jorgensen et al. 567 (MO not seen, NY);
Sevillán, 26 Mar 1995, Van den Eynden and Cueva 342 (LOJA not seen, NY).
Peru. Dept. Piura, prov. Huancabamba: Abra de Porculla, entre Olmos y Jaén, ladera con monte bajo, 22 Apr 1964, Ferreyra 15667 (K, US);
Carretera entre Canchaque y Huancabamba, km del 16 al 25 desde Canchaque, 17 Apr 1987, Díaz and Baldeón 2395 (MO, NY);
Perculla, 2 May 1981, Llatas and Laes 631 (HUT not seen, MO, NY).
Acknowledgments. I thank David M. Spooner for seeds and review, Clinton E. Morse for care of living plants, Kancheepuram N. Gandhi for Latin translation, and Janet R. Sullivan and Gregory J. Anderson for review.