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Jaltomata I: Circumscription, Description, and New Combinations for Five South

American Species (Solaneae, Solanaceae)

Article  in  Brittonia · April 1993

DOI: 10.2307/2807496

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Jaltomata I: circumscription, description, and new
combinations for five South American species
(Solaneae, Solanaceae)
THOMAS J. ANDERSON,
MIONE,'GREGORY AND MICHAEL
NEE

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
I Current address: Biological Sciences, Central Con-
described as Atropa by Ruiz and Pav6n
necticut State University, New Britain, CT 06050, (1799). This brings the total number of rec-
U.S.A. ognized Jaltomata 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 (1845) 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 (1987), 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. Ovules 1 18-1 8 1 (n
diam., many-seeded, black, purple, green, = 3 ovaries) per ovary. Mature berries red
orange, red or yellow (also white w e Ruiz according to collector, orange in green-
& Pavbn, 1799). house. Description based on living plants
At this time we make the following trans- of Leiva 138 (provided by S. Leiva, A. Sa-
fers and synonymies. Calyx lobe radius is g6stegui A. and M. 0 . Dillon). Chromo-
the length from the center of the calyx to some number n = 12 (this study, meiosis
the tip of one of the calyx lobes. The calyx of anthers from plants from seeds of Leiva
sinus radius is the length from the center of 138). See Macbride (1962) and Baker (1.c.)
the calyx to one of the calyx sinuses. Label for other English descriptions.
data for specimens cited as providing local Distribution: Known only from the De-
names or use information are listed under partment of La Libertad, Province Otuzco,
specimens examined. Peru, alt. 2500-3200 m. Roadsides, rock
walls, and rocky areas.
Jaltomata ventricosa (Baker) Mione, comb. Local name: "sogorome" (Leiva 138, Sa-
nov. (Fig. 1A). ghstegui 11574); "sorogome" (Leiva & Lei-
va 118).
Hebecladus ventricosus Baker, Saund. Refug. Bot. 3: t. Uses: Berries are edible (Leiva I 18 & 138,
208. 1870. TYPE:PERU. Without locality, a culti-
vated specimen (LECTOTYPE, here designated: t. 208
Saghstegui 11574).
of J. G. Baker (I.c.), curators of K and WELT were Specimens examined: PERU. La Libertad: PROV.
unable to find any type specimens). O ~ u z c o :San Miguel (Camino Salpo-Samne), flores
amarillo-verdoso, 2500 m, S. Leiva & P. Leiva I18
Erect shrub to 1 m high, glabrate. Leaves (F);Salpo-Shitahuara (A1 norte de Salpo), flores blan-
ovate, acute, blade to 6.5 cm long by 4.5 quecinas, 3100 m, S. Leiva 138 (CONN, F); Cerro
cm wide, entire or less commonly sinuate- Ragache (Salpo), flores blancas y bayas globosas, 3200
dentate or toothed with no more than 3 teeth m, A. Sagcistegui A, et al. 11574 (NY).
per margin, the upper surface bright green.
Inflorescence usually 1-2(-4) flowered Jaltomata viridiflora (Humb., Bonpl. &
(sometimes branched on cultivated plants Kunth) M. Nee & Mione, comb. nov. (Fig.
from seed of Leiva 138 but not mentioned 1B).
in the original publication nor shown in the
lectotype). Peduncle 4-9 mm long, green, Atropa viridiflora Humb., Bonpl. & Kunth, nov. gen.
sp. 3: 11, t. 196. 1818.Hebecladus viridiflorus(H.B.K.)
terete. Pedicel 7-1 6 mm long, always longer Miers, London J. Bot. 4: 321. 1845. TYPE:COLOM-
than peduncle to which it is attached, green, BIA. Crescit in convalli Guaitarensi, inter urbem
 ONIA IVOL. 45
Pasto et Chilanquer, Humboldt & Bonpland 2175 Salento, path to Romerales, Amarguras, Finca Sta. Inks,
(HOLOTYPE: P!;ISOTYPE: B destroyed, photo of B spec- 3200 m, J. G. Hawkes 422 (K); Nariiio: N end of La-
imen, F neg. 253 1 at GH! and NY!). guna de la Cocha, weed on cultivated slopes, 2850 m,
Hebecladus lanceolatus Miers, London J. Bot. 4: 323. F. R. Fosberg 20426 (NY); Santander: quebrada de
1845. TYPE: ECUADOR. Hacienda de Pinantura, Pais, N of La Baja, ca. 3200 m, E. P. Killip & A. C.
prope Quito, Hartweg 1301 (HOLOTYPE: K!; ISOTYPE: Smith 18764 (GH, NY); without locality, 2300 m, M.
LD!). K0ie 5258 (C). ECUADOR. Azuay: Nudo de Portete,
Hebecladus mollis Miers, London J . Bot. 7: 352, t. 33. pass between headwaters of the 150s Tarqui and Giron,
1848. TYPE:COLOMBIA. Plages de Combayma, 2744 m, W . H. Camp E-2181 (NY); Carchi: common
Goudot s.n. (HOLOTYPE: K!). The seemingly distinc- along Tulcan-El Carmelo road, 3290 m, T. Mione &
tive toothed leaf margin of this holotype was seen C. McQueen 458 and 459 (CONN), and direct (dirt)
on plants also having entire leaves, which are char- road between Tulcan and El Angel, 3300 m, 460
acteristic of the isotype of A. viridzyora. (CONN); Tulcan-Pun road bank, 3260 m, Y. Mexia
7586 (K); Cotopaxi: 12.7 km from gasoline station at
Herb to shrub, to 2 m high, sometimes N end of Pilalo, on rd. to La Mana, 3 180 m, D. Spooner
scandent. Leaves and young axes velutinous et al. 5091 (herbarium of T. Mione); Pichincha: declive
oriental del VolcLn Pichincha, near summit of Cruz
to less commonly puberulent with multi- Loma, 3400 m, J. A. Steyermark 52383 (NY); Napo:
cellular, uniseriate, unbranched finger hairs, 10 km E of Papallacta on road to Baeza, 2850 m, L.
sometimes to mostly gland-tipped. Leaves Holm-Nielsen et al. 6814 (NY);Chiborazo-Caiiar bor-
entire to toothed, ovate to ovate-lanceolate, der: Between Sta. Rosa and Joyagshi, 2637 m, W. H.
Camp E-4014 (NY).
the apex acuminate. Inflorescence (1-)2(-4)
flowered. Corolla tubular, the tube (1-)2 cm Jaltomata bicolor (Ruiz L6pez & Pav6n)
long, pale green, less commonly white or
Mione, comb. nov. (Fig. 1C).
pale yellow, the five larger lobes alternate
with five smaller lobules, all flaring. Anthers Atropa bicolor Ruiz Lopez & Pav6n, FI. peruv. 2: 45.
1.7-2.2 mm long. Stigma and anthers some- 1799. Kukolis bicolor (Ruiz Lopez & Pav6n) Raf.,
times differing in height by up to 5 mm, Sylva tellur. 55. 1838. Hebecladus bicolor (Ruiz L6pez
& Pavon) Miers, London J. Bot. 4: 322. 1845. TYPE:
either may be exserted. Mature berries or- PERU. Haurocheri versus S. Matthaei, Surco et S.
ange ("reddish" on Fosberg 20426), broader Joannis de Matucana vicos, Ruiz LBpez & PavBn s. n.
than high (oblate spheroid) and transversely (HOLOTYPE: perhaps MA, not seen). No type speci-
oblate, 13-19 mm diam., 10-12 mm high mens (or photographs) were present in a loan of spec-
imens from MA. We postpone lectotypification until
(Mione 458-460). Chromosome number n any specimens at MA that may not have been loaned
= 12 (Heiser, 1963). Description based on are seen. We are confident about this new combi-
numerous herbarium and field-collected nation without having seen a type specimen because
specimens from throughout the range of the there are no Atropa species in the New World, the
tubular, bicolored corolla described in the original
species. publication of A. bicolor Ruiz L6pez & Pav6n is
Distribution: Andes of western Venezue- unique, and the Jaltomata specimens having this
la through Ecuador, alt. 2500-3400 m. Sub- feature were collected in the region specified in the
paramo and disturbed habitats. original publication of A. bicolor.
Local names: "uchuva blanca" (Hawkes Atropa biflora Ruiz L6pez & Pavbn, F1. peruv. 2: 44.
t. 18 1b. 1799. Ulticona bzflora (Ruiz L6pez & Pav6n)
422); "chantilly" ( K ~ i e5258); "uvilla co- Raf., Sylva tellur. 55. 1838. Hebecladus biflorus (Ruiz
mun" (Mione 458); "yerba-mora" (Luteyn L6pez & Pav6n) Miers, London J. Bot. 4: 322. 1845.
6476 & 6497); "uvilla de conejo," "la vieja" TYPE:PERU. Tarmae, Cantae et Huarocheri versus
(Mexia 7586). Huassahuassi, Culluay et Div. Joannis de Matucana
vicos, Ruiz Lbpez & PavBn s.n. (HOLOTYPE: likely
Uses: Fruits not eaten (Hawkes 422, MA; ISOTYPE: B destroyed, photo of B specimen, F
Mione 458). neg. 2530 at GH!) [Note: t. 181b is misleading be-
cause it does not show the corolla vesture that is
Representative specimens: VENEZUELA. Merida: shown on the lower part of the corolla on the photo
Rangel, Parte baja de la caiiada de La Mucuchache, la of the isotype].
quebrada hombnima, afluente del n o Chama, 3400- Hebecladus intermedius Miers, London J. Bot. 4: 323.
3300 m, L. Ruiz-Tercin & M . Lopez-Figueiras 287 1845. TYPE: PERU. Puruchuca, Mathews 524
(MERF); trail leading from La Negrita to the Boquer6n (HOLOTYPE: K!).
of the Quebrada de las Caiias, 2990-3300 m, J. L. Hebecladus weberbaueri Dammer, Bot. Jahrb. Syst. 37:
Luteyn et al. 6122 ( N Y ) . COLOMBIA. Antioquia: 638. 1906. TYPE: PERU. Department of Ancash,
Municipio Jardin, Alto de Ventanas, 2400-2800 m, R. Province Cajatambo, 3200-3400 m, Weberbauer
Callejas et al. 391 7 (NY);Boyaca: hacienda Retacuba, 2652 (HOLOTYPE: B destroyed, photo of B specimen,
3600 m, P. J. Grubb et al. 244 (K, US);Caldas: above F neg. 2533 at GH!).
19931 MIONE ET AL.: JALTOMATA 143

Woody shrub to 1 m. Young axes with Suffrutescent to small shrub, to 1.2 m.

dendritic hairs (sparse to dense); branches Young branches, leaves and the abaxial face
glabrate. Leaves single, paired or verticil- of the corolla and calyx covered with mul-
late; usually ovate but variable in size, shape ticellular, erect finger hairs (sometimes
and margin. Inforescence 2-4(-5) flowered. gland-tipped, uniseriate but sometimes
Corolla straight-tubular; bicolored, purple branched on calyx). Leaves single or paired,
(rarely pink or yellow-green) proximal 2/3and lanceolate with the apex acuminate, to ovate
pale-green or cream distal l/3; 2-3 cm long; with the apex obtuse, entire to sinuate-cre-
with 5 narrowly triangular flaring lobes 3- nate, to 9.5 cm long by 5.5 cm wide but
5 mm long, vesture of tube externally vari- usually somewhat smaller. Inflorescence 4-
able, dense to sparse, trichomes to 1.1 mm 7(-10) flowered, rarely branched. Corolla
long uniseriate or dendritic (all conditions with a short tube (5 mm, evident only on
seen on Saunders 1383). Anthers blue or some herbarium specimens) and reflexed
black. Style and stamens usually exerted, limb (Fig. ID) 2 cm diam., center dark vi-
both up to 1 cm beyond corolla tube. Ac- olet to blue, paler peripherally. Filaments
cording to Macbride (1962) berry white or densely villous proximally with uniseriate
rarely blue. Description based on numerous finger hairs, glabrous distally (Fig. ID). An-
herbarium specimens from throughout the thers 1.3-2 mm long, dark. Style twice as
range of the species. For other English de- long as stamens. Description based on the
scriptions see Hooker (1845, H. biflorus with herbarium specimens cited below. See Bit-
plate) and Macbride (1962, H. bicolor, H. ter (1921) and Macbride (1962) for other
interrnedius and H. weberbaueri). descriptions, in Latin and English, respec-
Distribution: Central Peru, alt. 3000-5000 tively.
m. Open and shrub-covered hillsides, grazed Distribution: Peru, Department of Lima,
rocky slopes, moist ravines and crevices in mostly prov. Huarochiri, alt. usually 2000-
stone walls. 3000 m (extremes 1800 and 3800 m). Rocky
Local name: "shuplac" (L6pez et al. 7601). and gravelly places, dry open hillsides, and
Uses: ". . . fruit . . . eaten by natives" river and stream banks.
(Metcalf30270). Local name and uses: none mentioned on
herbarium specimens.
Representative specimens: PERU. Ancash: Bolog-
nesi, 9.7 km SW of highest point on road between
Palivilca and Huaraz, 4000-5000 m, W. D. Stevens Representative specimens: PERU. Lima: Province
21952 (NY); Recuay, Km 107 between Recuay and Canta, Arahuay, ladera rocosa, 2600 m, C. Ochoa &
Pativilca, 3300 m, M. Dillon et a/. 3178 (BH, F);Aija, G. Vilcapoma 279 (MO); Arahuay, a orillas del no,
en la bajada de Tranca hacia Huayan, 3200-3000 m, 3000 m, G. Vilcapoma S. 55 (US); Province Huaro-
C. Ochoa & A . Salas 15168 ( N Y ) ;Recuay, Culgumarca chin, Km 82 carretera Lima-Huancayo pasando de
(Pativilca-Conococha), 3600 m, '4. L6pez M. et a/. 7601 Matucana hacia Ticlio, 2600 m, C. Ochoa & A. Salas
(NY); Huancavelicia: Castrovirreina, near Cbrdova, 15207 (F, NY); Encima de Langa, 2000 m, C. Ochoa
along rock wall at side of stream in shade, 3 175 m, R. 14610 (NY); Cerca de Pachacosa, 3800 m, C. Ochoa
D. Metcalf 30270 (GH); Lima: Huaros, Canta, hills 7425 (F, US); Matucana, Km 83 Canetera Central,
about 1 km above Huaros, 3700 m, S. G. Saunders 8500-9500 ft, S . G. E. Saunders 303 (GH); Santa Eu-
1383 ( K ) ; Huarochiri, San Mateo, 200 yds up valley lalia valley, Rio Chira, at Pariagancha, 2600 m, P. C.
of Rio Atacra, from junction with Rio Rimac, Km 1 10 Hutchison & C. Saravia 7107 (K, MO, NY); distrito
Central Hwy, 3476 m, S. G. E. Saunders 821 ( K ) ;Rio Santiago de Tuna, frente a1 Monte Zarate, a1 norte de
Blanco, 3000-3500 m, E. P. Killip & A. C. Smith 21558 Lucmani, 3200 m, C. Ochoa & A. Salas 15103 (US);
(NY). Huquicha a m b a de Surco, pradera con arbustos, 3000
m, E. Cerrate 5852 (MO); Huaquicha, a m b a de Surco,
Jaltomata propinqua (Miers) Mione & M. 1800 m, C. Ochoa 1159 (IVY);quebrada San Juan,
Nee, comb. nov. (Fig. ID, E). entre Surco y Matucana, pedregoso, falda cerro, 2300
m, R. Ferreyra 5422 (US); Surco, 2034 m, J. Soukup
Sarachapropinqua Miers, Ann. Mag. Nat. Hist. ser. 2, 3731 (F, US); Surco, shore of rivulet, c. 2000 m, E.
3: 446. Miers, Illustr. S. Amer. P1. t. 38 1849. He- Asplund 11061 (US); quebrada Ayas b San Juan, entre
becladus propinquus (Miers) Bitter, Feddes Repert. Surco y Matucana, 2300 m, R. Ferreyra 5417 (US);
Spec. Nov. Regni Veg. 17: 246. 192 1. TYPE:PERU. quebrada San Juan, entre Surco y Matucana, 2200-
Lima. Cuesto d e Puruchuco, Mathews 774 2300 m, R. Ferreyra 5429 (MO, US); valley Rio Rimac,
(HOLOTYPE: K!; ISOTYPE: W, photo of W sheet, F neg. near Lima-Oroya highway at Km 81 east of Lima,
33 136 at GH! and WIS!). 2250 m, T. H. Goodspeed & R. D. Metcalf30242 (US).
 'ONIA IVOL. 45

Jaltomata umbellata (Ruiz L6pez & Pav6n) Local name and uses: none mentioned on
Mione & M. Nee, comb. nov. (Fig. IF). herbarium specimens.
Atropa umbellata Ruiz L6pez & Pav6n, Fl. Peruv. 2: Specimens examined: PERU. Lima: Chancay, Lo-
44, t. 18 1, fig. a. 1799. Hebecladus urnbellatus (Ruiz mas of Lachay, ca. 105 km N of Lima on Pan Am
L6pez & Pav6n) Miers, London J. Bot. 4: 322. 1845. Hwy, 300-500 m, M. 0. Dillon et a/. 3626 (BH, N Y ) ;
TYPE:PERU. Chancay, Limae et Lurin, Dombey s.n. Atacongo Lomas, in the sheltered loam pockets, 32 km
(LECTOTYPE, here designated: P!). S of Lima, 8 km east of Pachacamac Ruins, not com-
Hebecladus turneri Miers, London J. Bot. 4: 323. 1845. mon here, 400 m, H. E. Stork et a/. 9294 (GH, K);
TYPE:Without locality, evidently from a cultivated hills of San Agustin (Loma-Formation), 350 m, A. We-
specimenwe Miers, D. Turner s.n. (HOLOTYPE: K!). berbauer 5223 (GH); and same locality A. Weberbauer
5228 (GH); sandy hills of Lachay-Lima, dry and hot
Shrub to 1 m. Axes and abaxial face of desert, R. Chavez s.n. (plants grown from seeds, field
calyx bearing both gland-tipped finger hairs voucher not seen).
and shorter, non-glandular dendritic hairs
interspersed among each other. Leaf blades Acknowledgments
typically entire, ovate, the apex acute, less
than 11 cm long and 8 cm wide. Infores- We thank Tilton Davis IV, the herbarium
cence 4-9 flowered. Peduncle 4.5-1 1 mm curators at B, BH, BM, C, COLO, F, GH,
long. Pedicel 6-9 mm long, with dense gland- HNMN, K, LD, LPB, MERF, MA, MO,
tipped finger hairs. Lobes of calyx and co- NY, P, QCA, US, USD, WELT and WIS
rolla triangular to narrowly triangular. for loan of specimens and/or accommodat-
Flowering clayx 8-9 mm diam., lobe radius ing visits, Rene Chavez, Michael 0. Dillon,
4-5 mm long, sinus radius 2 mm long. Co- S. Leiva, Abundio Saghstegui A. and David
rolla tubular with a broad limb. Corolla tube Spooner for seeds, Mary Jane Spring for the
6.5-8 mm long, partially to nearly com- illustration, Lisa M. Campbell, William G.
pletely filled with red nectar on living ma- D'Arcy, Kent E. Holsinger and Sandra
terial. Corolla limb rotate to campanulate, Knapp for helpful comments, William Cul-
cream or pale-green, 14-23 mm diam., re- lina and Sandra Ek for care of living plants,
maining open at night. Stamens (including and especially Armando T. Hunziker and
expanded base) 9.2-1 0.9 mm long, exserted Luis Bernardello who suggested this group
from corolla up to 7 mm. Filaments, slender for study. Support was from fellowships
part with a few to several finger trichomes from the Graduate School and the Research
to 0.5 mm long on basal l/4-'/3 of the length. Foundation of the University of Connecti-
Anthers undehisced 1.9-2.1 mm long x 1.4- cut to T.M., an N.S.F. doctoral dissertation
1.5 mm wide, dehisced 1.2-1.3 mm long. grant to T.M. and G.J.A., and an N.S.F.
Style 8-13.3 mm long. Stigma exserted be- grant to G.J.A.
yond anthers up to 8 mm but sometimes
not exserted, overhead dimensions 0.46- Literature Cited
0.56 mm by 0.4-0.46 mm, often protruding Adams, C. D. 1972. Flowering plants of Jamaica.
through the otherwise closed corolla prior University of West Indies. Indies Mona, Jamaica
to anthesis. Ovules 70-78 per ovary. Fruit- [Jaltomata antillana as Saracha a.].
ing calyx-lobe radius 6-8 mm. Mature berry Benitez de Rojas, C. E. 1974. Los gttneros de las
Solanaceae de Venezuela. Revista Fac. Agron. (Ma-
orange in greenhouse. Seed (field collected) racay) VII(3): 25-108 [Jaltomata as Saracha].
1.36-1.48 mm long. Chromosome number Bitter, G. 192 1. Eine verkannte Hebecladus-Art und
2n = 24 (Diers, 196I), n = 12 (this study, ihre Bedeutung fur die Stellung der Galtung in der
meiosis of anthers from plants from seeds Tribus der Solaneae. Repert. Spec. Nov. Regni Veg.
of Chavez s. n.). Description based on living 17: 246-25 1.
. 1924. Weitere Untersuchungen iiber Hebe-
plants of Chavez s.n., provided by R. Cha- cladus I. Repert. Spec. Nov. Regni Veg. 20: 372-
vez and T. Davis IV. See Bitter (1924) and 376.
Macbride (1962) for other descriptions, in Castillo Pinilla, R. 1974. Sinopsis de la familia So-
Latin and English, respectively. lanaceae en Colombia. Doctoral dissertation, Uni-
versidad Nacional de Colombia [available at Har-
Distribution: Peru, Department of Lima, vard University].
coastal desert (Loma formation), alt. 300- D'Arcy, W. G. 1973. Solanaceae. In: Flora of Pan-
500 m, apparently rare. ama. Ann. Missouri Bot. Gard. 60: 573-780.
 19931 MIONE ET AL.: JALTOMATA 145

-. 1982. Jaltomata werfli (Solanaceae), a new to Rico and adjacent islands: a systematic synopsis.
species from the Archipelago de Colon (Galapagos Editorial de la Universidad de Puerto Rico, Rio
Islands). Phytologia 52: 9-10. Piedras.
-. 1987. Saracha spinosa-a new combination Macbride, J. F. 1962. Solanaceae. Field Mus. Nat.
in Peruvian Solanaceae. Ann. Missouri Bot. Gard. Hist., Bot. Ser. 13, part V-B, No. 1 [Jaltomata as
74: 674-675. Saracha and Hebecladus].
-, T. Mione & T. Davis. 1992. Jaltomata gran- Miers, J. 1845. Contributions to the botany of South
dljlora (Solanaceae): a rare mexican species. Novon America. London J. Bot. 4: 319-325.
2: 190-192. Mione, T. 1992. the systematics and evolution of
Davis, T., IV. 1980. The generic relationship of Sara- Jaltomata (Solanaceae). Doctoral dissertation. The
cha and Jaltornata (Solanaceae; Solaneae). Rho- University of Connecticut, Storrs, Connecticut,
dora 82: 345-352. U.S.A.
Diers, L. 196 1. Der anteil an polyploiden in den ve- -, R. G. Olmstead, J. D. Palmer, R. K. Jansen &
getationsgiirteln der westkordillere Perus. Z. Bot. G. J. Anderson. 1990. Phylogenetic relations
49: 437-488. among Jaltomata, Hebecladus and related genera
Dodson, C. D. & A. H. Gentry. 1978. Flora of the (Solanaceae). Amer. J. Bot. 77(supplement 6): 145-
Rio Palenque Science Center, Los Rios Province, 146 [abstract].
Ecuador. Selbyana 4: 1-628 [Jaltomata pp. 545- Morton, C. V. 1938. Notes on the genus Saracha.
5461. Proc. Biol. Soc. Wash. 51: 75-78 [Jaltomata as Sa-
Gentry, J. L., Jr. 1973. Restoration of the genus Jal- racha].
tomata (Solanaceae). Phytologia 27: 286-288.
. 1974. The generic name Saracha Ruiz &
-. 1944. Una nueva especie peruana de Sara-
cha. Revista Univ. (Cuzco).
Pav6n (Solanaceae). Fieldiana, Bot. 36: 69-72.
-& P. C. Standley. 1974. Solanaceae. In: Flora Nee, M. 1986. Solanaceae I. Flora de Veracruz, Fas-
ciculo 49. Instituto Nacional de Investigaciones
of Guatemala. Fieldiana, Bot. 24, part X, Nos. 1 &
sobre Recursos Bibticos, Xalapa, Veracruz, MCxi-
2. 1-151.
co.
Heiser, C. B., Jr. 1963. Numeraci6n cromos6mica
de plantas Ecuatorianas. Ci. & Nat. 6: 2-6. Ruiz, H. & J. Pavon. 1799. Flora Peruviana et Chi-
Hooker, W. J. 1845. Hebecladus biflorus. Twin-flow- lensis 2. Gabrielis de Sancha, Madrid.
ered Hebecladus. Bot. Mag. 71: t. 4192. Rzedowski, J. & G. C. de Rzedowski. 1985. Flora
Hunziker, A. T. 1979. South American Solanaceae: fanerogamica del Valle de MCxico, Vol. 11. Instituto
a synoptic survey. Pages 49-85. In: J. G. Hawkes, de Ecologia MCxico, D. F.
R. N. Lester & A. D. Skelding, editors. The biology Schlechtendal, D. F. L. 1838. Index seminum horto
and taxonomy of the Solanaceae. Academic Press, academic0 halensi 1838 collectorurn p. 8.
Dorchester, Great Britain. Seithe, A. 1979. Hair types as taxonomic characters
-. 1987. Studies on Solanaceae. XXI. A prelim- in Solanurn. Pages 307-319. In: J. G. Hawkes, R.
inary synopsis of Cuatresia. Opera Bot. 92: 73-82. N. Lester & A. D. Skelding, editors. The biology
Kearney, T. H. & R. H. Peebles. 195 1. Arizona flora. and taxonomy of the Solanaceae. Academic Press,
University of California Press, Berkeley. Dorchester, Great Britain.
Knapp, S., T. Mione & A. Sagastegui A. 199 1. A new Shreve, R. & I. L. Wiggins. 1964. Vegetation and
species of Jaltornata (Solanaceae) from northwest- flora of the Sonoran Desert, Vol. 11. Stanford Uni-
ern Peru. Brittonia 43: 181-184. versity Press, Stanford, California [Jaltornata as
Liogier, H. A. & L. F. Martorell. 1982. Flora of Puer- Saracha].

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