THREE NEW SPECIES OF SOLANUM SECTION HERPYSTICHUM (SOLANACEAE) FROM ECUADOR

2009·14.5 MB·English

by  Eric J Tepe

NEW
SOLANUM
THREE
OF SECTION HERPYSTICHUM
SPECIES
FROM ECUADOR
(SOLANACEAE)
EricJ.Tepeand Lynn Bohs
Department 257
of 1400
Biology,
S. E.
USA.
Lake Utah 84
Salt
City,
12,
1
Author
correspondence: erictepe@ utah.edu
for
INTRODUCTION
The genus Solanum has been the focus of intense research because contains several important crop spe-
it
cies, including the potato tuberosum), tomato lycopersicum), and eggplant melongend). Despite the
(S. (5. (S.
many
among
economic importance of the genus, however, groups 1500 species remain poorly known.
its ca.
TBI
The
Solanum: a world-wide treatment" project (www.solanaceaesource.org), an NSF-funded
initiative
and
provide online descriptions keys Solanum has allowed increased exploration and
to to all taxa, for dis-
new home
covery of Solanum species. Ecuador 183 species of Solanum, including 31-34 endemics, and
to
is
&
one of the highest centers of diversity for the genus (J0rgensen Leon-Yanes 1999; Montufar 2000). This
is
&
Stem
with funded by (Knapp Bohs
paper, along three others this project 2007, 2008; 2009), adds seven
new
which
species, five of are endemic, to the of Ecuadorian Solanum.
list
Solanum sect. Herpystichum Bitter is a group of ca. ten species found from Central America to northern
&
member
and informally-named (Weese Bohs
Peru, a of the "Potato clade" 2007). Molecular phylogenetic
is
analyses indicate that sect. Herpystichum constitutes a monophyletic group sister to sect. Pteroidea Dunal
& &
(Weese Bohs 2007; Tepe Bohs, unpublished manuscript). Species Herpystichum
of are
sect. trailing,
woody
They
herbaceous
terrestrial or climbing, to vines rooting at the nodes. are primarily plants of the
but found open
rainforest understory, several species are also in pastures or other Like
areas. Pteroidea,
sect.
they have perfect flowers, but can be distinguished by the usually extra-axillary inflorescences (those in
sect.
Some
Pteroidea are axillary in position). species have flattened fruits that are unique in Solanum. This group
has never been revised and has received attention since Bitter described most of species in 1912
little its
and
1913 (Bitter 1912, 1913a, Detailed study of sect. Herpystichum has revealed three additional species,
b).
Two
described herein, that have been collected since focus on the genus. of the species are endemic
Bitter's
512 Journal of the Botanical Research Institute of Texas 3(2)
Ecuador and occur in small reserves in areas under intense pressure from large-scale agricultural or oil
to
exploration. They should both be added to the Red List for Ecuadorian Endemics (Knapp et 2007).
al.
Maximum
parsimony sequence from nuclear ITS and GBSSI waxy) sequences place
analysis of data (or
&
and
unpublished Solanum
Herpystichum (Tepe Bohs, manuscript). crassinervium
three species in
sect.
all
from
with Greenm., widespread occurs Central
species that
pacificum are in a clade evolvulifolium a
S. S.
America and Andean South America and loxophyllum Bitter from the Pacific lowlands of Ecuador. Solanum
5.
American
limoncochaense in a clade with the morphologically similar Central species phaseoloides Polak.
is S.
We
known
and Dunal, high-elevation species only from Bolivar Province, Ecuador. were unable
trijolium a
S.
and
obtain material molecular analysis of dalibardijorme Bitter dolichorhachis Bitter, the species
to for S. S.
and
most similar morphologically to limoncochaense pacificum, respectively.
S. S.
Solanum
pacificum ECUADOR.
Tepe, sp. nov. (Fig. 1). Los Rios: Centro Cientifico Rio Palenque, in secondary forest,
Type.
Vine, climbing other vegetation via adventitious roots the nodes. Stems slender, weakly herbaceous,
at
glabrous, densely gland-dotted; branch tips pendent. Sympodial units plurifoliate, not geminate.
fertile
membranaceous
Leaves simple, 14-19 x 4.5-8 cm, 2-3 times as long as wide, lanceolate to ovate, to thinly
more
and rounded
densely gland-dotted, the base obtuse, or
chartaceous, glabrous adaxially abaxially, to
symmetrical, the margin the apex acuminate; petioles 1-1.5 cm, glabrous, densely gland-dotted.
less entire,
Internodes 1.5-7 cm. Inflorescences 4-10 x 2-3 cm, slender, extra-axillary, unbranched with 17-58 flowers
cm
peduncle 2-4.5
flowers apparently perfect, the axes glabrous, very slender; long; rachis ca.
(scars), all
mm
mm
8-10 15-20 spaced
cm; enlarged glabrous, nearly
6 pedicels in flower, slender, in fruit, apically,
mm
mm
contiguously to 12 apart, articulated at base. Flowers with the calyx 1-1.2 long, glabrous to minutely
mm
mm,
and sparsely along margins, the tube 0.5-0.7 long, the lobes 0.5-0.6 x 0.8-1 rounded,
ciliate
mm.
somewhat
rounded weakly acuminate calyx accrescent, the lobes 0.6-0.8 x
fruiting
to at tips; ca. 1
mm
cm
Corolla 0.8-1 in diameter, 5 long, membranous, green to white near the margins of the
ca. stellate,
mm,
4-5 and
the lobes x 1.2-2.5 reflexed maturity, acute apices, glabrous adaxially abaxially, the
petals, at at
mm
margins Stamens subequal, with filaments 0.8 long, glabrous, anthers 1.5-2 x 0.7-1.2
ca. free;
ciliate.
mm,
opening with
oblong, not connivent, yellow, the pores directed distally, into short longitudinal slits
mm,
Ovary glabrous; 4-4.5 x 0.1-0.2 glabrous, slightly clavate; stigma truncate. Fruit (immature)
age. style
unknown.
0.9 x 0.6 cm, ovoid, pointed apex, green, glabrous. Seeds
ca. at
Solanum with deep purplish-green above with whitish and
striking species leaves that are veins,
pacificum a
is
and been
weakly intensely purple below. This species apparently quite rare has not collected extensively,
to
is
members by
but from other of Herpystichum. recognizable climbing habit,
it is clearly distinct sect. It is its
and weakly herbaceous
completely glabrous vegetative parts, large, very thin leaves slender, stems.
its
Solanum specimens have been but has wiry
identified as dolichorhachis, dolichorhachis
pacificum S. 5.
woody and
with spindle-shaped
stems, chartaceous coriaceous leaves distinctly oblique leaf bases, fruits
to
membranaceous more
2-3 times longer than wide the weak-herbaceous stems, leaves or less symmetrical
vs.
and somewhat than twice long wide)
at the base, globose to conical (but less as as fruits of 5. pacificum. It
and but distinguished from these
is also related to 5. crassinervium, S. evolvulifolium, S. loxophyllum, is easily
sym-
species by the texture, shape, and size of the leaves. Solanum pacificum also bears a resemblance to the
and
patic Geminata) due simple leaves, long, slender inflorescences, small,
5. leptorhachis Bitter (sect. to its
new
Tepe and Bohs, Three species of Solanum Herpystichum)
(sect.
514
Journal of the Botanical Research Institute of Texas
3(2)
m
greenish-white However, free-standing shrub 1-2 with
stellate, flowers. S. leptorhachis is a tall unifoliate
sympodial units on flowering stems vine with sympodial units in
vs. a plurifoliate pacificum.
5.
The lowlands Ecuador between Ocean and Andes under
of the Pacific the are large-scale cultivation
of cacao, papayas, bananas, African palm, and very few areas of undisturbed habitat remain
intact.
oil etc.,
Most collections of pacificum are from the small, but well-preserved Centro Cientifico Rio Palenque reserve.
S.
There are no large reserves in western Ecuador (like Yasuni in the east) and likely that pacificum and
it is 5.
other plant species endemic to western Ecuador are surviving in small, isolated reserves. As a result,
5.
made
and
pacificum in critical danger of extinction due to habitat loss every effort should be to maintain
is
the reserves in this region.
—
named
lowlands Ecuador where endemic, and
Etymology. Solarium pacificum after the Pacific of
is is
it
and companion Maria Paz Moreno.
as a tribute to ET's wife frequent field
—
Habitat and Distribution. Solarium pacificum occurs in rainforest habitats in the Pacific lowlands of
m
50-300
Ecuador,
in elevation.
—
Phenology. Flowering specimens have been collected from Feb-Aug; the type collection, collected in
Feb, the only fruiting specimen seen. likely that fruiting more frequent than the collection record
is It is is
indicates.
May
19 1987 H.H. van der Werff9496 (holotype: QCNE!; MO!, NY!).
isotypes:
(fl, fr),
m
Vine or scandent shrub, climbing understory trees as high as 4 via adventitious roots at the nodes; leafy
branches spreading pendulous. Stems thickly herbaceous weakly woody, somewhat glabrous
to to fleshy, to
very sparsely pubescent and soon glabrescent. Sympodial units plurifoliate to rarely unifoliate, not geminate.
Leaves simple, 3.5-14 x 1.5-8 cm, ca. 1-2 times as long as wide, gradually reduced in size toward the inflo-
rescence, ovate to elliptic, somewhat fleshy, glabrous adaxially and abaxially, the secondary veins 5-7 per
conspicuous and prominent abaxially, the base rounded truncate cordate, sometimes oblique, the
side, to to
margin entire, the apex shortly acuminate; petioles (0.2-)l-1.5 cm, glabrous. Internodes 1.5-5.5 cm. Inflo-
cm cm
rescences 1-3 x 1-4 to ca. 6 x ca. 5.5 in fruit, stem-terminal to axillary to extra-axillary, unbranched
2-16
to branched, with flowers flowers apparently perfect, the axes glabrous; peduncle 0.1-0.5
(scars), all
mm mm
cm
4-12 9-18
long; rachis 0.1-2 cm; pedicels in flower, in only slightly enlarged apically,
fruit,
glabrous to rarely very sparsely pubescent, spaced nearly contiguously, articulated base. Flowers with the
at
mm mm
calyx 2.5-3.5 long, glabrous, the margins thickened, the tube 2.5-3 long, the lobes 1.5-2.5 x 1.5-2
mm,
somewhat
deltoid, acute to acuminate at tips, white to pale pink; fruiting calyx accrescent, the lobes
mm
mm. cm
5-8
somewhat
1.5-2 x .5-2.5 Corolla 1.5-2 in diameter, long, white, the lobes
1 stellate, fleshy,
mm,
5-8
x 2.5-3 planar at anthesis, acute to acuminate at apices, glabrous adaxially, sparsely pubescent
mm
near the apex abaxially, the margins densely Stamens equal, with filaments 1-1.5 long, glabrous,
ciliate.
mm,
3-4
anthers x 1.2-1.5 oblong, not connivent, yellow, the pores directed opening into
free; distally,
mm,
4-6
Ovary
short longitudinal slits with age. glabrous; style x ca. 0.3 glabrous, cylindrical, sometimes
deflected one side of flower ID); stigma Fruit (immature) 0.7-1 x 0.7-0.9 cm, ovoid
to (Fig. capitate. to
somewhat brownish
nearly globose, slightly flattened, pointed at apex, green to pale orangish to at maturity,
mm,
glabrous. Seeds 2-2.2 x 1.8-2 flattened-reniform, tan, the surface minutely reticulate-rugulose.
Nambi, margen derecha del Rio Nambi, 18'N, 78°08'W, 1325 m, Dec 1993 Franco 4707 (NY); Mpio. Tumaco, Resguardo
l 1 (fl), P. et a\.
ECUADOR.
Resguardo Indigena de Saunde, 1°30'N, 78°20'W, 350 m, 21 Jan 1996 B.R. Ramirez 9699 (NY). Carchi: Canton
(fr), et a\.
km
1813 (BM, NY, QCNE); border area between Prov. Carchi and Esmeraldas, 20 past Lita on road Lita-Alto Tambo, 550 m, 25
et
al.
Jun 1991 H. van der Werjfet al 11972 (MO, NY, Q
(fl),
new
Solanum
Tepe and Bohs, Three species of (sect. Herpystichum)
Fig. 2. Solanum crassinervium Tepe. A. Habit. B. Inflorescence. C. Bud. D. Flower. E. Flower, longitudinal section. F. Stamens. G. Infructescence {fruits
immature). drawn from van derWerff11972 [MO] and EJ. Tepe2729 [photo]; B drawn from EJ. Tepe2729 [photo]; D drawn from Stem
(A, C, E-F H. 5.
400 [photo]; G drawn from Tipaz 1813 [MO] and EJ. Tepe 2729 [photo].
G.
(QCNE); Canton Quinine
31
0-600
m, 26 Sep 1994
(fr),
Journal of the Botanical Research Institute of Texas 3(2)
&
QCA, QCNE,
0°20 49"N 79°4241"W, 540 m, 13 Feb 2009 Tepe Stern 2729 (BM, NY, UT).
, (fl, fr), E.J. S.
somewhat
and by
Solarium crassinervium the most robust species in sect. Herpystichum is recognizable the
is
and with conspicuous secondary
fleshy texture of the leaves, stems, flowers, ovate to elliptical leaves
its
veins in fresh and dried material, and occasionally branched inflorescences. Its distribution is restricted to
NW
SW
the Choco region of Colombia and Ecuador, a biodiversity hotspot with one of the highest species
common
some
per area (Myers 2000), but apparently quite in parts of range.
diversities et is its
al. it
and but from both
species
Solarium crassinervium closely related to evolvulijolium loxophyllum, differs
is S. S.
may
and and be
in robust habit, ovate leaves mostly oblong), fleshy calyx corolla, inflorescences that
its (vs.
many
The and than
simple and branched within an individual. leaves, petioles, internodes are times larger
than but crassinervium
those of evolvulijolium. Solarium loxophyllum is larger in stature 5. evolvulijolium, 5.
5.
The
on and
easily distinguished from the former based leaf inflorescence characters. leaves of S. crass-
is
more
somewhat but crassinervium has secondary veins
inervium, those of loxophyllum, are fleshy,
like 5.
S.
(5-7 3-4 and these are prominent abaxially, whereas those of loxophyllum are obscure within
vs. pairs) S.
and The
These characters are apparent in both fresh dried material. often stout
the fleshy leaf blade. leaf
which and
from apparently always simple
inflorescences of crassinervium differ those of 5. loxophyllum, are
S.
borne on
very and Furthermore, inflorescences of loxophyllum are typically older, leafless
slender
delicate. 5.
parts of the stem, as compared to those of 5. crassinervium, which appear to always be produced in the leafy
member
sometimes been
parts of the stem. Solanum crassinervium has identified as siphonohasis Bitter, a of
5.
Solanum Anarrhichomenum but from species in having > 2 flowers per inflorescence
differs this
sect. Bitter,
and
and
has 1-2 flowers per inflorescence), in the fleshy texture of the leaves, stems, flow-
siphonohasis
(S.
from
Anarrhichomenum node-rooting but they Herpystichum
Species of are also vines, differ sect.
ers. sect.
primarily possessing pseudostipules the base of the petioles (Correll 1962).
in at
—
The prominent secondary veins help distinguish
Etymology. epithet crassinervium describes the that
from
this species closest relatives.
its
—
and premontane
lowland
and Solanum crassinervium occurs in rainforest habitats in
Habitat
Distribution.
the Choco region of Colombia and Ecuador, including the Mache-Chindul mountain range in northwestern
m
150-1800
Ecuador, in elevation.
—
specimens have been from
Flowering apparently occurs year-round; fruiting collected
Phenology.
and Sep-Dec.
Jan-Feb,
Solanum
limoncochaense ECUADOR.
Tepe, sp. nov. (Fig. 3). SuciMBfos: Limoncocha, Reserva Biologica Limoncocha,
Type.
Sympodial
Terrestrial herb, trailing, rooting at the nodes. Stems slender, glabrous. units apparently plurifoli-
somewhat
not geminate. Leaves simple, the blades 3.5-7 x 3.5-8 cm, slightly wider than long, rounded,
ate,
margins
glabrous; venation palmate with 5 (-7) veins, the base cordate, the entire, slightly revolute
fleshy,
3-16
on some leaves, the apex rounded to obtuse to shortly acuminate; petioles cm, glabrous. Internodes
new
Tepe and Bohs, Three species of Solanum Herpystichum)
(sect.
(immature).
Fruit dr
F. (All
4-8 2-3
2.5-20 cm. x 3 cm, unbranched, with
Inflorescences extra-axillary, flowers, flowei
ca. all
ently the axes glabrous; peduncle 1.3-4.5 cm, slender; rachis 0.9-1.5 cm; pedicels 15-2:
perfect,
mm mm
25-30
flower, in slender, glabrous, spaced ca. 15 apart, articulated at the base. Flowers
fruit,
mm mm
mm,
calyx 2.5-4.2 long, the tube 1-1.5 long, the lobes 1.5-2 x ca. 1.2 rounded, acuminat
glabrous sparsely pubescent abaxially, densely pubescent adaxially, purplish; fruiting calyx ver)
to
mm, cm
5-8
accrescent, the lobes 1-1.2 x 1.5-2 truncate-acuminate. Corolla 1-1.6 in diameter, n
mm,
mm,
6-10
membranous, white, the tube 1.5-2 the lobes x 1.5-3 lanceolate, narrow
stellate,
518 Journal of the Botanical Research Institute of Texas
3(2)
apex and margins Stamens
at tips, the papillose adaxially abaxially, the ciliate apically. equal, the filaments
mm, mm,
glabrous; anthers 2-2.5 x tapered, not connivent, yellow, the
ca. 1 ca. 1 slightly sagittate basally,
and opening with Ovary
pores large directed distally, into longitudinal slits age. sparsely papillose; style
mm,
somewhat
2-2.5 x 0.3 straight, cylindrical, stout, sparsely papillose in lower half; stigma capitate,
when
1-3 x 0.6-3.2 cm, ovoid-rhomboid, greenish-brown purplish near apex
2-lobed. Fruits flattened, to
mm
when when
immature, bronze-brown mature, sparsely pubescent with hairs < 0.1 to glabrous mature,
the apex truncate to emarginate, strongly fragrant with a sweet, heavy scent, very juicy, the flavor sweet.
mm
Seeds 2-2.5 in diameter, lenticular, light reddish-brown, the surface minutely rugose.
MT. MO, QCA,
environs of Limoncocha, 240 m, 16 Jun 1978 Madison 5327 (AAU, K, NY, SEL); near northwest corner of
(fl, fr), et al. F,
Solarium limoncochaense belongs to a distinctive group of species within Solanum sect. Herpystichum character-
ized by herbaceous, ground- trailing stems that root at the nodes and bizarre, flattened, arrowhead-shaped
group and Solanum
this includes pentaphyllum, phaseoloides, dalibardiforme, limon-
fruits; 5. S. 5. 5. trifolium.
can be from group by simple and
cochaense distinguished other species in this leaves stellate flowers.
its
most from Colombia, and most specimens have been
Vegetatively, similar to dalibardiforme central
it is S.
annotated as this species. Both species have simple, cordate leaves, but S. limoncochaense is distinguish-
because and has than Furthermore,
able vegetative parts are glabrous, rather rotate corollas.
stellate
all S.
found low whereas found > 2000 m. Solanum
limoncochaense elevations, dalibardiforme elevations
is at S. is at
on
grows presumably
limoncochaense in dense, clonal patches th
weak
and were
apparently a climber living plants in the see
field
&
ground Tepe
the (E Stern, pers. obs).
S.
J.
Mature fruits are bronze-brown in color, very juicy, and have a strongly sweet aroma and flavor. The
when
on the ground stems are or are pendulous on climbing stems. The dispersal agents
fruits trailing,
lie
unknown, on and
of these fruits are but based the strong scent, color, presentation, is likely that they are
it
W.
eaten by rainforest rodents such as the guatusa or agouti (genus Dasyprocta; Haber, pers. comm.).
known
Solanum endemic Ecuador and from
limoncochaense is to all collections are the 'terra firme' for-
Limoncocha Limoncocha,
ests at the northern side of the lake within the Reserva Biologica a small reserve
Much made
4613 which 250 up
of hectares (including the lake hectares). of the rest of the reserve
is ca. is
of swamps and seasonally flooded and limoncochaense was not encountered in any of the flooded
forests,
S.
forest habitats. The reserve is surrounded by cultivated land and oil platforms, whose effects on the habitats
unknown.
within are If this area is truly the only locality for this species, then should be considered
it it
much
and enormous
endangered; however, Reserva de Yasuni nearby,
the Biosfera of this re-
critically la is
serve has not yet been explored botanically. hoped that this species more widespread than appears
It is is it
on
be based
the collections currently available.
to
—
name
Laguna Limoncocha
Etymology. Solanum limoncochaense takes from the de in western Ecuador,
its
known
the geographic locality of collections of the species.
all
—
Napo
endemic and Sucumbios
Habitat and Distribution. This species apparently to Provinces,
is
m
240-300
Ecuador, near the Laguna de Limoncocha, where grows in primary forests and clearings, in
it
elevation.
—
and
Flowering specimens have been Oct; specimens
Phenology. collected in Jan, Jun, Sep, fruiting
have been and
in Jan
collected Jun.
ACKNOWLEDGMENTS
We
thank Stephen Stern and Freddy Villao for assistance in the the herbaria AAU, BM, K, MO, NY,
field; F,
UT
Ann
and SEL for loans of specimens; Kelsey from for assistance with loans; David Neill and the staff at
QCNE
and Hendry Moya UISEK Amazonica
for assistance with logistics in Ecuador; Katy Coral of the Estacion
and and
Limoncocha; Freddy Villao and the the Centro Cientifico Rio Palenque; Carlos Aulestia
at staff at
.
new
Tepe and Bohs, Three Solanum Herpystichum) 519
species of
(sect.
the Fundacion Jatun Satcha for access to the Bilsa Biological Station; Bobbi Angell for the illustrations; and
two anonymous whose and comments improved manuscript. This work was
the reviewers questions
this
NSF
by DEB-0316614,
supported through Solanum
the the PBI: grant, to LB.
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