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Brittonia
ISSN 0007-196X
Volume 67
Number 2
Brittonia (2015) 67:105-112
DOI 10.1007/s12228-014-9360-2
Two new Peruvian species of Jaltomata
(Solanaceae, Solaneae) with red floral
nectar
Thomas Mione, Segundo Leiva González
& Leon Yacher
1 23
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Two new Peruvian species of Jaltomata (Solanaceae, Solaneae) with red
floral nectar
THOMAS MIONE
1
,SEGUNDO LEIVA GONZÁLEZ
2
,AND LEON YACHER
3
1
Biology Department, Central Connecticut State University, New Britain, CT 06050-4010, U.S.A.;
e-mail: MioneT@ccsu.edu
2
Museo de Historia Natural, Universidad Antenor Orrego, Avenida America Sur 3145, Trujillo,
Peru; e-mail: segundo_leiva@hotmail.com
3
Department of Geography, Southern Connecticut State University, New Haven, CT 06515-1355,
U.S.A.; e-mail: yacherl1@southernct.edu
Abstract. Two new Jaltomata species (Solanaceae) of Peru that produce red floral nectar are
described. Jaltomata neei of Department Cajamarca has 1–4 flowers per inflorescence, the
campanulate corolla is green and then changes to blue, corolla lobes (5) and lobules (5) are
equally prominent, a corona is lacking, five radial staminal-corolla thickenings create nectar
troughs between, and the stigma is capitate. Jaltomata quipuscoae of Department Arequipa
has solitary flowers, a purple, broadly crateriform, 5-lobed corolla, a corona on which nectar
pools, a punctiform stigma, lacks corolla thickenings, and the mature fruit is whitish.
Photographs, illustrations and tables are included that allow comparison with closely related
species.
Key Words: Arequipa, Cajamarca, corolla color change, corona, Jaltomata neei,Jaltomata
quipuscoae, protogyny, Peru, red nectar..
The genus Jaltomata (Solanaceae) includes
about 70 species of herbs and shrubs, distributed
from Arizona in the United States to Bolivia, and
there are two insular species (J. antillana [Krug &
Urban] D’Arcy, Greater Antilles; J. werffii
D’Arcy, Galápagos Islands). Jaltomata can be
found throughout Andean Peru from near sea-
level to about 4000 m of elevation. There are three
major clades in the genus. Species of the orange-
fruited clade (Miller et al., 2011) grow in South
America, are mostly woody, and express the full
range of corolla forms and colors (described be-
low). The species of the red-fruited clade grow in
South America and the Greater Antilles, are her-
baceous to suffrutescent, and have rotate, white to
pale-yellow corollas. In contrast, the species of
the black-fruited clade grow mostly in Mexico
and Central America, are mostly herbaceous,
and have only rotate, greenish corollas. Green
fruits, characteristic of few species, evolved inde-
pendently in the orange-fruited and the black-
fruited clades (Miller et al., 2011). This paper is
a contribution to ongoing taxonomic studies of
the genus (Mione, 1992; Leiva González &
Mione, 1999;Mione&Serazo,1999;Mione
et al., 2000;Mioneetal.,2004,2011,2013;
Leiva González et al., 2010).
The species of Jaltomata have simple, alternate
(often geminate) leaves and pentamerous flowers
with five corolla lobes, or ten total lobes plus
lobules. The corolla form varies greatly, depend-
ing on the species (tubular, urceolate, crateriform,
campanulate-rotate, or rotate). The inflorescence
is umbellate. A subset of the species of the
orange-fruited clade have radially oriented
staminal-corolla thickenings, adnate to the base
of each stamen and extending toward the corolla
lobule (or to the area where the lobule, when
lacking, would be). Between the radially oriented
staminal-corolla thickenings are troughs, visible
to the unaided eye, that hold nectar at the base of
the corolla (Bitter, 1924;Fig.2B). The ovary of
all species has a basal disk, and anthers dehisce
longitudinally. Filaments of all but three species
insert into the lower ventral face of the anther. The
calyx is accrescent but never encloses the berry.
Dendritic hairs are uniseriate, many-celled and
have three or more branches emerging at different
Brittonia 67(2): 105Y112 (2015), DOI 10.1007/s12228-014-9360-2
ISSN: 0007-196X (print) ISSN: 1938-436X (electronic)
© 2014, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
Published Online: 16 October 2014
Author's personal copy
levels. Forked hairs are also uniseriate and many-
celled but have only two termini. Finger hairs are
uniseriate and unbranched. In the following de-
scriptions, hairs are not gland-tipped and are
unpigmented unless indicated otherwise. Pollen
size is based on 30 grains in polar view stained
with analine blue in lactophenol.
During fieldwork in 2007, 2010 and 2013 we
collected the following two new species.
Jaltomata neei Mione & S. Leiva, sp. nov. Type:
Peru. Dept. Cajamarca: Prov. San Miguel, 7
o
00.744 S, 78
o
50.188 W, road from San Miguel
de Pallaques to Llapa, 2645 m, 27 Mar 2013,
S. Leiva G., T. Mione & L. Yacher 5426
(holotype: HUT; isotype: NY). (Figs. 1,2)
Diagnosis: Inflorescence 1–4 flowered, corolla campanulate
and green changing with age to blue, corolla lobes and lobules
equally prominent, radial staminal-corolla thickenings (five) cre-
ating nectar troughs (five) between, corona lacking, stigma
capitate.
Shrub 1–2 m high, the young branches green,
pubescent with erect, dendritic and forked hairs,
somewhat angular to nearly terete, the older stems
brown, glabrous, with lenticels, hollow, terete,
and to 12 mm diameter. Leaves alternate or gem-
inate (Fig. 1D), the blade to 8 cm long ×4.8 cm
wide, darker green above, ovate to ovate-
lanceolate, the apex acute, the base of some leaves
somewhat truncate, both faces densely pubescent,
the hairs erect and dendritic; petiole 1.1–3.9 cm
long. Inflorescence 1–4 flowered. Peduncle 4–
8 mm long, green at flowering, nearly terete,
pubescent with erect, dendritic and forked hairs;
pedicel 11–18 mm long, green at flowering, an-
gular in cross section at the distal end, pubescent
with erect, dendritic and forked hairs. Calyx
green, 16–18 mm diameter, subplanar, glabrous
internally, pubescent externally, especially on the
veins, the hairs dendritic, at fruit maturity hiding
1/3 of the fruit in side view, at least 25 mm diam-
eter probably somewhat larger (fruits were not
ripe at the time of collection). Corolla green, later
turning blue, the red nectar showing through the
wall of the corolla (Fig. 2A), and purple pigmen-
tation of the corolla showing at the bottom of the
nectar troughs after nectar is removed and in side-
view of flower, campanulate with a planar limb,
8.5–13 mm long × 25–39 mm diameter, the lobes
alternating with lobules together totaling 10
(Fig. 2B), externally with dendritic hairs on veins
and glabrous between veins, internally the hairs
simple, erect, gland-tipped, and evenly distribut-
ed. Stamens 11–12 mm long, exserted beyond
distal end of corolla 0–3 mm, the radially oriented
staminal-corolla thickenings extending toward
corolla lobule with copious red nectar pooling in
troughs between thickenings, the filament base
expanded laterally (ventral view), the filaments
connivent, pale-green, pilose only at the base
(Fig. 1J), the anther yellow with a green connec-
tive prior to dehiscence, 2.5 mm (herbarium spec-
imen) to 3 mm (fresh) long, mucronulate (evident
only with a dissecting microscope prior to dehis-
cence), the filament inserting into ventral face of
the anther, the anthers of a flower opening asyn-
chronously; pollen grains 32.75–38.75 μm(mean
35.34) diameter. Stigma capitate, bilobed, darker
green than the style (Fig. 2A); style 11–13 mm
long, straight, pale-green, exserted 5 mm beyond
mouth of corolla; ovary green,3 mm high ×3 mm
wide, the disk orange and half as high as the ovary
(Fig. 1E); ovules per ovary 170–196 (n = 2
flowers). Fruit 9–11 × 13–16 mm; color (at matu-
rity) not seen, almost certainly orange.
Additional specimens examined. PERU. Cajamarca: Prov.
San Miguel, same locality and date as the type specimen, Mione
et al. 833 (F); Visita Alegre, Niepos, 2400 m, “1-11-1985,”
S. Llatas Quiroz 1575 (F); 6° 59.001 S, 78° 48.237 W, 2919 m,
20 Mar 2007, S. Leiva G. et al. 3643 (HAO), Mione et al. 740 (F);
6° 58’25”S, 78° 47’58”W, 2840 m, 18 June 1999, S. Leiva G.
et al. 2370 (HAO), Mione et al. 669 (CCSU).
Jaltomata neei grows in Peru, Department
Cajamarca, Province San Miguel, between 2400
and 2920 m in the native vegetation along road-
sides. The flowers are protogynous (Fig. 2B).
Younger flowers have a green corolla and are
pistillate: they are fully open but anthers have
not yet dehisced. Older flowers have a blueish
corolla and dehisced anthers, and thus are
functionally hermaphroditic. The dehiscence of
the anthers and the corolla color change from
green to blue are not always synchronous; some
flowers in the field had both a bluish corolla and
anthers that had not yet dehisced. Red nectar is
produced by flowers in both the pistillate and
hermaphroditic phases. In the phylogeny of
Miller et al. (2011) this species was referred to
as J.“rednectar.”
Table Icompares Jaltomata neei with similar
congeneric species: shrubs having a campanulate
corolla, copious red/orange nectar, radial
staminal-corolla thickenings creating five nectar
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FIG. 1. Jaltomata neei. A. Flower, side view, with dendritic hairs expanded. B. Berry with calyx, side view. C. Seed. D. Branch with
leaves flowers and fruit. E. Gynoecium including basal disk. F. Ovary, cross section. G. Anther, lateral view. H. Corolla dissected to show
the insertion of the stamens. I. Anther, dorsal view. J. Stamen, ventral view. (Drawn from S. Leiva G. et al. 5426).
107MIONE ET AL.: JALTOMATA (SOLANACEAE)2015]
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troughs (one trough between each pair of thick-
enings) and no corona. Jaltomata leivae Mione
and J. ventricosa (Baker) Mione were omitted
from Table Ibecause the corolla is urceolate,
and J. alviteziana S. Leiva was omitted because
its corolla is tubular, but these species otherwise
share these features. Jaltomata umbellata (Ruiz &
Pav.) Mione & M. Nee also has copious red
nectar, but was omitted from Table Ibecause it
has a tubular corolla and lacks radial staminal-
corolla thickenings (Mione et al., 1993).
The name of this species was chosen to honor
Dr. Michael Nee, eminent Solanologist.
Jaltomata quipuscoae Mione & S. Leiva, sp. nov.
Type: Peru. Arequipa: Prov. Arequipa, Distrito
Yarabamba, 16° 33’58.6”S, 71° 25’40.6”W,
alrededores de Petroglifos de Sogay, ladera
arbustiva, rocosa, 2625 m, 11 Feb 2006,
V. Quipuscoa S., G. Castillo P., K. Arce C. 3352
(holotype: HUSA; isotype: NY). (Figs. 2,3)
Diagnosis: Flowers solitary, corolla purple, broadly
crateriform and 5-lobed, nectar pools on the corona, lacking
radial staminal-corolla thickenings, stigma punctiform, mature
fruits whitish.
Much branching, perennial herb to 70 cm, the
younger branches green with a dense pubescence
of gland-tipped finger hairs, the older branches
brown-purple, lacking lenticles. Leaves alternate,
sometimes geminate, the blades ovate to elliptical,
to 4.8 cm wide ×7.5 cm long, the base symmetri-
cal or nearlyso, adaxially and abaxially pubescent
with gland-tipped finger hairs, the margin nearly
FIG. 2. Jaltomata neei (A,B)andJ. quipuscoae (C–E).A. Flower, side view. B. Flowers, front view, pistillate phase (anthers
undehisced) on left, andhermaphroditic phase (anthers dehisced) on right. A,B: Units along bottom of photos are mm; photos by
T. Mione. C. Flower, all anthers dehisced. D. Fruit. E. Flower, side view. C–E: All scale bars represent 0.5 cm;
photos by Victor Quipuscoa S.
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TABLE I
COMPARISONOF JALTOMATA NEEI WITH OTHER SHRUBBY JALTOMATA HAVI N G A CAMPANULATE COROLLA,COPIOUS RED/ORANGE NECTAR,AND RADIAL STAMINAL-COROLLATHICKENINGS CREATING
FIVE NECTAR TROUGHS,ONE TROUGH BETWEEN EACH PAIR OF THICKENINGS.
Species J. neei Mione
&S.Leiva
J. paneroi Mione
& S. Leiva
J. herrerae (C. V.
Morton) Mione
J. dendroidea S.
Leiva & Mione
J. grandibaccata
S. Leiva & Mione
J. weberbaueri
(Dammer) Mione
Distribution Peru: Cajamarca Peru: Cajamarca Peru: Cuzco, Apurimac,
Ayacucho. Bolivia:
La Paz.
Peru: La Libertad Peru: La Libertad Peru: Ancash
Altitude (m) 2400–2920 3240–3500 3000–3750 3100–3360 3450–3680 2800–3795
Calyx when flowering,
diameter (mm)
16–18 13 21–26 17–21 23–35 29–41
Calyx lobes as long as wide as long as wide longer than wide as long as wide as long as wide longer than wide
Calyx when flowering, having
an abaxial keel where sepals
meet
no no no no yes yes
Corolla color green turning to blue green green green blue-green to blue purple to violet
Corolla length, proximal to
distal (mm)
8.5–13 5–10 10–15 10–13 20 20–35
Corolla diameter, lobe tip to
lobe tip (mm)
25–39 23–35 30–40 23–30 35–47 40–60
Alternating corolla lobes and
lobules
lobes (5) and lobules
(5) equally
prominent
lobes (5) conspicuously
longer than lobules
(5)
lobes (5) conspicuously
longer than lobules
(5)
lobes (5) conspicuously
longer than lobules
(5)
lobes (5) and lobules
(5) nearly equally
prominent
lobes (5), lobules
(0)
Style length (mm) 11–13 11–14 14 12–17 19–23 15–22
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FIG. 3. Jaltomata quipuscoae. A. Embryo. B. Berry with calyx, side view. C. Flower, side view, with hairs expanded. D.
Seed. E. Anther, lateral view. F. Branch with leaves flowers and fruit. G. Anther, dorsal view. H. Gynoecium including basal disk.
I. Corolla dissected to show the insertion of the stamens. J. Stamen, ventral view. (From S. Leiva G. et al. 4654).
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entire, ciliate; petiole to 3.5 cm long. Flowers
solitary (Fig. 3F); peduncle to 9 mm long, terete;
pedicel to 10 mm long, 5-sided or subterete, both
pedicel and peduncle green to purple and densely
pubescent, the hairs gland-tipped. Calyx during
anthesis: green, planar, sepals 14–15 mm long,
the lobes triangular, abaxially densely pubescent
with gland-tipped finger hairs; calyx with fruit:
green, infundibular, hiding berry in side view, and
to 25 mm across. Corolla broadly crateriform
(Fig. 2C, E), 5-lobed, lacking lobules, to 3.3 cm
diameter, purple with green ring at the base, the
inner surface puberulent with hairs 0.06 mm long,
the hairs of the outer surface concentrated on
veins, the margin ciliate. Nectar copious, red to
purple, pooling on corona. Stamens 5, 13 mm
long, not exserted beyond the distal end of the
corolla, the filaments dark purple proximally fad-
ing to light purple distally, nearly glabrous but
having a few short (0.25 mm long) finger hairs
at the base, the base of the filament appressed to
the ovarian disk; anthers yellow prior to dehis-
cence, 3.4 mm long (dehsiced, pressed),
basifixed, emucronate, those of a flower dehisc-
ing asynchronously; pollen grains 25–27.5 μm
(mean 25.5) diameter. Stigma punctiform
(Fig. 3H), not bilobed, darker green than the style;
style straight, pale-green, 1 cm long, widest at
base and gradually narrowing toward apex; ovary
pale-green, 3 mm wide × 2.8 mm high, the disk
one-third as high as the ovary. Berries whitish to
pale-green at maturity, 15–18 mm across, hidden
by the calyx in side-view (Fig. 2D).
Additional specimens examined. PERU. Arequipa:
Prov. Arequipa, Dist. Yarabamba, alrededores de los
Petroglifos de Sogay, 16° 33’58.6”S, 71° 25’40.6”W, 2607
m, 7 Jan 2010, S. Leiva G. et al. 4654 (HAO), Mione et al. 797
(798 seeds only) (HUSA). Prov. Caraveli, km 586
Panamericana Sur, 6 km north of Atiquipa turnoff on
Panamerican Highway, 300–860 m, 3 Oct 1997, M. Weigend
& H. Förther 97/701 (M).
Jaltomata quipuscoae and the other two spe-
cies of J. section Modillonia (Leiva González
et al., 2010) share the following characters: they
are endemic to Peru, they are herbaceous and
have solitary flowers with a 5-lobed corolla, a
corona (a structure on which nectar pools),
basifixed anthers (the filament is inserted on the
lower ventral face of the anther in other sections
of this genus), and a stigma no wider than the top
of the style (punctiform, not capitate). They also
produce copious red-orange nectar, and the fruits
are green to whitish (except for one report of
orange fruits in J. aspera, Table II). Jaltomata
quipuscoae was not included in the recent molec-
ular phylogeny of Jaltomata (Miller et al., 2011).
However, the other two species of section
Modillonia are together monophyletic in both
the strict consensus tree and the majority-rule
consensus tree.
The three species of Jaltomata section
Modillonia (Table II) individually have narrow
allopatric distributions but together are widely
distributed in Peru: J. aspera (Ruiz & Pav.)
Mione grows almost exclusively in the
Department Lima (Mione & Coe, 1992),
J. calliantha S. Leiva & Mione grows in the
Department La Libertad (Leiva Gonzales et al.,
2010), and J. quipuscoae (here described) grows
in the Department Arequipa.
The Weigend & Förther specimen (97/701, M)
was identified by T. M. as Jaltomata quipuscoae
based on morphological characteristics, but was
collected in a different habitat (lomas) some 2000
m lower than the other collections, in west-facing
sandy and rocky slopes. Similarly, J. aspera
TABLE II
COMPARISON OF ALL JALTOMATA SPECIES HAV IN G A FLORAL CORONA.
J. aspera J. calliantha J. quipuscoae
Corolla color greenish-yellow or yellow-green green purple
Ring of green at base of corolla no yes yes
Filament color whitish to pale yellowish purple purple
Base of filament pubescent glabrous nearly glabrous
Mature fruit color white (Macbride, 1962); orange
(A. Granda 926,MOL)
very pale-green whitish to pale-green
Elevation m Lomas: 150–600. Andes: 1420–1870. Lomas 300–860.
Andes: 1600–2550. Andes: 2607–2625.
Distribution (Peruvian Departments) Primarily Lima La Libertad Arequipa
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grows both in the lomas habitat and the Andes
(Mione & Coe, 1992). Andean populations and
lomas populations of both J. aspera and
J. quipuscoae flower during different months of
the year: in the lomas habitat flowering occurs
August through October; however in the Andes,
flowering occurs January through early May.
Lomas and Andean populations of these spe-
cies are thus apparently reproductively isolat-
ed, and additional study may reveal that
Andean and lomas populations within each
species as here circumscribed are actually
sibling species.
Both Jaltomata neei and J. quipuscoae are
protogynous; in the field we saw at the same time
open flowers having undehisced anthers and other
flowers having dehisced anthers. Protogyny is
widespread in the genus Jaltomata.
The name of this species was chosen to honor
Victor Quipuscoa Silvestre, discoverer of this
species.
Acknowledgments
Segundo Leiva G. drew the illustrations. We
thank Michael Nee, David M. Spooner, Gregory
J. Anderson, and Michael O. Dillon for review,
Nathaniel T. Mione for preparation of color fig-
ures, the administration of Universidad Antenor
Orrego, Trujillo, Peru, and the curators of F, K, M,
MO, NY, US for their loan of specimens. Support
from research grants from the Connecticut Uni-
versity System was provided to T. M.
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