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Jaltomata I: Circumscription, Description, and New Combinations for Five South American Species (Solaneae, Solanaceae)

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Jaltomata I: Circumscription, Description, and New Combinations for Five South American Species (Solaneae, Solanaceae) Jaltomata I: circumscription, description, and new combinations for five South American species (Solaneae, Solanaceae) THOMASMI ONE,GREGORY' J. ANDERSON, AN D MICHAELN EE Mione, Thomas, Gregory J. Anderson (Ecology and Evolutionary Biology, Uni- versity of Connecticut, Storrs, CT 06269, U.S.A.), and Michael Nee (The New York Botanical Garden, Bronx, New York 10458-5126, U.S.A.). Jaltomata I: circumscription, description, and new combinations for five South American spe- cies (Solaneae, Solanaceae). Brittonia 45: 138-145. 1993.-The genus Jaltomata (including Hebecladus) is described. Five Hebecladus species are transferred to Jaltomata. Jaltomata viridiflora is widespread, from western Venezuela through Ecuador; J. bicolor and J. propinqua occur in central Peru; J. umbellata of the Loma Formation of the Department Lima, Peru is rare; J. ventricosa is known only from the vicinity of La Libertad, Otuzco, Peru. All are montane except for J. umbellata. Included are short descriptions and illustrations. El gCnero Jaltomata (incluyendo Hebecladus) se describe en este trabajo. Cinco especies de Hebecladus son transferidas a Jaltomata. Jaltomata viridiflora esti difundida extensamente en 10s Andes desde el oeste de Venezuela hasta el Ecuador; J. bicolor y J. propinqua se encuentran en la region central del Perfi; J. umbellata de la Formaci6n Loma de Departamento Lima, Perfi, es rara; J. ventricosa es conocida solamente en la vecindad de La Libertad, Otuzco, Perk Las especies tratadas son de montaiia, exceptuando J. umbellata. Se incluyen descripciones cortas e ilustraciones. Key words: Andean flora, Hebecladus, Jaltomata, Solanaceae, Solaneae, Solanoi- deae. Prior to this study it was virtually im- were used for taxonomy, comparative mor- possible to identify species of South Amer- phology, and chloroplast DNA restriction ican Jaltomata because nearly all basio- site-based phylogeny construction. Here we nyms are in other genera and species treat five species, providing a brief descrip- descriptions were widely scattered in the lit- tion and an illustration of part of each. erature. Macbride (1962) treated the Peru- The new combinations presented here re- vian species of this group but commented sult from the current recognition among that, "The group needs to be studied by a systematists that Jaltomata Schldl. (1838) student who can study living plants; . ." should include most species previously as- Our revision of the taxonomy of this genus signed to Saracha (D'Arcy, 1973, 1987; derives from a study of about 400 living D'Arcy et al., 1992; Davis, 1980; Gentry, plants (greenhouse-grown and in the field) 1973, 1974) and all species of Hebecladus and herbarium specimens. The living plants Miers (1845) (Hunziker, 1979; Mione, 1992). Some of the latter were originally described as Atropa by Ruiz and Pav6n I Current address: Biological Sciences, Central Con- necticut State University, New Britain, CT 06050, (1799). This brings the total number of rec- U.S.A. ognizedJaltomata species to about 30. These Brittonia, 45(2), 1993, pp. 138-145. ISSUED: 24 May 1993 O 1993, by The New York Botanical Garden, Bronx, NY 10458-5126 19931 MIONE ET AL.: JALTOMATA 139 species are tropical and subtropical, peren- Atropa have undisputedly been considered nial herbs and small shrubs that are distrib- Hebecladus since 1845 and therefore do not uted from Arizona, U.S.A., to southern Bo- require separate justification for their inclu- livia (Mione, 1992; Morton, 1944), in the sion within Jaltomata. Greater Antilles (1 species, Adams, 1972; Earlier descriptions of the genus Jalto- Liogier & Martorell, 1982; Mione, 1992) mata are either based on species of only part and on the Galbpagos Islands (1 species, of the geographic distribution or represent D'Arcy, 1982). the earlier circumscription, i.e., without He- Traditionally, corolla form and habit were becladus species (e.g., Benitez de Rojas, 1974 used to distinguish between Jaltomata and [as Saracha]; Castillo Pinilla, 1974; D'Arcy, Hebecladus (Davis, 1980; Hunziker, 1979). 1973; Davis, 1980; Dodson & Gentry, 1978; Miers (1 845) described Hebecladus species Gentry & Standley, 1974; Kearney and Pee- as suffrutescent, bearing tubular or infun- bles, 195 l [as Saracha]; Macbride, 1962 [as dibular corollas, and residing in South Saracha]; Nee, 1986; Rzedowski & Rze- America, while Jaltomata species have been dowski, 1985; Shreve & Wiggins, 1964 [as described as herbs with rotate or broadly Saracha]). A generic synonymy and de- campanulate corollas, distributed through- scription of Jaltomata in the current sense out the combined range of the two genera follow. Hair terminology follows Seithe (Davis, 1980). Georg Bitter seems to have (1979). been the first to note (1924, p. 374) that these genera grade into each other. This con- JALTOMATA Schldl. tention was reiterated by Morton (1938) who wrote, "the genus [Jaltomata, as Saracha] Jaltomata Schldl., Index sem. hort. hall. 1838.8. 1838. verges on . Hebecladus." Similarly, Mac- TYPE:Jaltomata edulis Schldl., which was preceded by a description of the same species, as Atropa pro- bride (1962) observed that the two genera cumbens Cav. The species that the type belongs to together formed a natural group, and thus is Jaltomata procumbens (Cav.) J . L. Gentry. included Hebecladus species in his key to Hebecladus Miers, London J . Bot. 4: 321. 1845. TYPE: the Peruvian species of Jaltomata (the latter Hebecladus viridiflorus (Humb., Bonpl. & Kunth) as Saracha). Furthermore, Hunziker (1 979) Miers, based on Atropa viridiflora Humb., Bonpl. & Kunth. treated both genera under Jaltomata and remarked, "I am unable to draw a line of Herbs to shrubs to 2 m high, sometimes separation between Jaltomata and Hebecla- scandent (without specialized structures), dus Miers. Both groups share a number of terrestrial, unarmed. Branches erect or common attributes . [and] there are in- spreading, 4- or 5-sided to terete, usually termediate [corolla] forms." Later, Nee hollow. Vesture of multicellular finger- or (1986), Hunziker again (1 987), Knapp et al. branchlet-hairs, gland-tipped or not. Leaves (1991) and Mione (1992) all supported the alternate or appearing opposite (rarely inclusion of Hebecladus within Jaltomata. appearing verticillate), simple, petiolate, es- In addition to the gradation in corolla tipulate, ovate, ovate-lanceolate or ovate- morphology and habit that prompted these acuminate, sometimes basally truncate, en- workers to suggest that Hebecladus and Jal- tire, subentire or toothed, often tapering tomata should be united, Mione et al. (1990) asymmetrically along distal portion of pet- presented chloroplast DNA restriction site iole. Inflorescence axillary or from a stem data showing that Jaltomata is strongly dichotomy (false dichotomy), pedunculate, paraphyletic without the inclusion of He- umbel-like (Fig. 1E ). Pedicels basally artic- becladus. Sixteen to 17 chloroplast DNA ulated after fruit ripens, sometimes angled. characters firmly place Hebecladus viridzjlo- Flowers five-merous; bisexual; regular. Ca- rus (H.B.K.) Miers and H. ventricosus Baker lyx enclosing the bud with valvate aesti- within Jaltomata (Mione, 1992). vation. Flowering calyx with sepals partially Based on the above evidence, we choose connate, lobes triangular, narrowly trian- to unify Jaltomata and Hebecladus. The gular or obtuse, rotate with a stellate outline species treated below with basionyms in (Fig. I), or the lobes reflexed. Fruiting calyx 140 BRITTONIA [VOL. 45 FIG. 1. Some South American Jaltomata. A. J. ventricosa (redrawn From portion of Baker, 1870, t. 208). B. J. viridiflora (portion of Humboldt, Bonpland and Kunth, 18 18, t. 196). C. J. bicolor (redrawn from Hooker, 1845, t. 4192). D. Pressed flower of J. propinqua (from Ochoa & Vilcapoma 279). E. Infructescence of J. prpinqua (redrawn from portion of Miers, 1849, t. 38). F. J. urnbellata (drawn from living material of Chavez s.n.). G. Typical gynoecium and calyx of Jaltomata; arrow points to the annular disk (redrawn from Hooker, 1.c.). conspicuously accrescent, green or purple, conspicuous. Corolla aestivation with five spreading behind berry, rotate (Fig. 1E) to valvate lobes, each lobe alternating with an reflexed. Most species having 10-lobed co- inwardly pleated and shorter lobule. Corolla rollas, the five larger lobes alternating with 1-5 cm diam., rotate, broadly inhdibular, 5 smaller lobules, the latter sometimes in- campanulate, tubular, in one species (J.ven- 19931 MIONE ET AL.: JALTOMATA 141 tricosa) the tube urceolate and the limb re- terete. Flowers nodding from 45" below hor- trorse (Fig. lA), if the corolla is non-rotate izontal to pendent. Flower buds triangular then the lobes flare. Filaments 5, filiform, when viewed perpendicular to bud's length. markedly enlarged/swollen at base where Flowering calyx shiny green abaxially, 18- adnate to base of corolla, the expanded base 22 mm diam., lobe radius 9.5-10 mm, sinus pubescent with finger hairs, slender part of radius 3-5 mm. Corolla, tube urceolate, limb filament pubescent with finger hairs (Fig. completely and outwardly retrorse and 10- 1D) or not, filament inserted obliquely into lobed, 8-10 mm long, 12-14.5 mm diam., ventral face of anthers in bud. Anthers de- whitish to pale yellow, remaining open at hiscing longitudinally. Ovary superior, bi- night. Base of the flower filling with orange- carpellate, ovules 42-4 12 (n = 128 ovaries) red nectar that is visible through the corolla. per ovary. Style slender (Fig. lG), exerted Stamens (not including expanded base) 14- or not. Stigma single, bilobed, slightly bi- 15 mm long, exserted beyond corolla 7-9 lobed, or grooved. Nectary disk annular mm. Filaments, slender part pubescent on around base of ovary (Fig. 1G). Nectar clear basal half, the hairs purple, branched with to yellowish drops at base of corolla and up to 4 termini, the hairs to 0.7 mm long. androecium, or, in several Peruvian and one Anthers, undehisced 2.6-3.0 mm long x 2.0- Bolivian species orange to red, filling base 2.2 mm wide, dehisced 2-2.3 mm long. Style of corolla and visible through corolla. Fruit 13.8-1 5.7 mm long. Stigma exserted be- a juicy globose or oblate berry, 4-25 mm yond anthers to 5 mm.
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