Bohs, 2001 - Revision of Solanum Section Cyphomandropsis

Revision of Solanum Section Cyphomandropsis (Solanaceae)
Author(s): Lynn Bohs

Source: Systematic Botany Monographs, Vol. 61, Revision of Solanum Section Cyphomandropsis
(Solanaceae) (Aug. 30, 2001), pp. 1-85
Published by: American Society of Plant Taxonomists
Stable URL: http://www.jstor.org/stable/25027891
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REVISION OF SOLANUM SECTION

CYPHOMANDROPSIS (SOLANACEAE)
Lynn Bohs
Departmentof Biology
of Utah
University
Salt Lake City, Utah 84112

ABSTRACT.
ica. Plants
Solanum
of this section
Cyphomandropsis
it differs
somes
shrubs
branched
trichomes,
related
to Solanum
or dendritically
branched
section Cyphomandropsis
are woody
is closely
by lacking discrete,
characterize
trees that lack spines,
and have
on the abaxial
the Cyphomandropsis/Pachyphylla
13 species
(formerly
small
to South Amer
native
are glabrous
anthers with
tapered
sect. Pachyphylla
connectives
enlarged
includes
(Solanaceae)
to small
to pubescent
terminal
clade. The morphology,
large chromo
Exceptionally
taxonomic
un
Section
from which
genus Cyphomandra),
anther surfaces.
with
pores.
history,
nomencla
ture, ecology, distribution,economic value, reproductivebiology, and evolutionary relationshipsof Solanum

sect. Cyphomandropsis
name
RESUMEN.
Solanum
de esta secci6n
plantas
tricomas
secci6n
simples
are reviewed.
are proposed.
(S. pelagicum)
secci6n
son arbustos
o dendriticamente
esta estrechamente
genero Cyphomandra),
grandes.
ci6n,
la biologia
Aquf
nov.) y S. pelagicum
de espinas,
con
Cyphomandropsis
reproductiva,
ci6n Cyphomandropsis.
incluye
que carecen
la morfologia,
las relaciones
Se proponen
y tienen anteras atenuadas
ramificados,
la Solanum
secci6n
la historia
filogeneticas,
cuatro grupos
(nom. nov.). Se proporciona
con pequenos
y dos nombres
una clave dicot6mica
Esta
incluida
en el
la ecologia,
de las especies
nuevos,
y otra sin6ptica
con
terminales.
por sus cromosomas
y el valor econ6mico
de especies
poros
en la superficie
la nomenclatura,
Las
o pubescentes
(anteriormente
engrosado
se caracterizan
taxon6mica,
of the section.
sudamericanas.
son glabros
Pachyphylla
por la falta de un conectivo

y Pachyphylla
for the species
13 especies
(Solanaceae)
and one new
(S. hutchisonii),
keys are provided
leniosos o arbolitos
relacionada
se revisa
one new combination
groups,
and synoptic
Cyphomandropsis
del que se distingue
anteras. Las dos secciones

mente
Four species
Dichotomous
dorsal de las
excepcional
la distribu
de Solanum
S. hutchisonii
para las especies
sec
(comb.
de la
secci6n.
INTRODUCTION
Solanum, with approximately 1000 to 1400 species, represents one of the largest gen
era of angiosperms. Because of its large size and morphological
complexity, many infra
generic groups within Solanum are not well defined and their component species are
poorly known. Recent taxonomic work using morphological
and molecular approaches
has led to a better understanding of broad-scale relationships within the genus and more
precise knowledge of species limits in numerous sections, but a comprehensive
view of
the systematics of the entire genus is not yet within reach. This contribution aims to clar
ify the taxonomy of a poorly understood clade within Solanum by elucidating species re
lationships and generic placement of a formerly obscure section of the genus, Solanum
Bitter. This section includes 13 species native to South America.
The section is here defined
in the traditional sense of Bitter (1913), Seithe (1962), Danert

(1970), Morton (1976), and Child (1984). The goal of this treatment is to delimit and de
fine the species of the section, clarify nomenclatural
issues, and propose infrasectional
species groups that can be used as hypotheses of relationships for further study.
SYSTEMATICBOTANYMONOGRAPHS
VOLUME 61
MATERIALSAND METHODS
I have followed the morphological
species concept in delimiting species of sect.
Taxa are recognized as distinct if they possess a unique suite of
Cyphomandropsis.
characters and are separated from similar entities by morphological
gaps. In nearly all
taxa also occupy geographically
circumscribed
ranges. No formal infraspecific
taxa are recognized, and in fact many infraspecific names have been relegated to syn
onymy because they could not be defined by consistent morphological
and/or geographic
cases,
distinctions.
In general, measurements
have been made
from dried herbarium material with floral

possible, living and/or liquid-preserved material
has been used to supplement measurements made on dried specimens. In species with cor
date leaf bases, the length of the blade is given from the tip to the basalmost insertion point
parts rehydrated in boiling water. Where
of the blade on the petiole. The petiole
from this point. Colors of

or from herbarium label data. Con
ventions used inmeasuring and describing various organs follow Bohs (1994).
For SEM studies of pollen grains, dried pollen from herbarium specimens was
mounted on a stub with double-stick tape and coated with gold-palladium. Sizes of grains
were estimated from SEM photographs. Measurements
in polar and equatorial view are
corollas,
fruits, etc., are described
length is also measured
from living material
in Table 1. Herbarium vouchers for pollen SEM are listed in Appendix

1.
Plants used in the crossing studies were grown from seed in the greenhouses at the
University of Utah. Voucher information and original provenance data are given in Ap
pendix 2. Pollinations were effected by shaking pollen onto a clean glass slide that was
rubbed against the stigma of the female parent. All plants known or suspected to be self
compatible were emasculated in the bud before pollination. Success or failure of the cross
combined
was monitored,
as well as fruit size, shape, color, and number of seeds in successful
crosses. Seeds were initially judged to be viable or inviable based on their appearance, and
full-sized seeds of successful combinations were germinated in the greenhouse to deter
mine their viability. The number of accessions used, number of pollinations attempted,
and outcome
Pollen
of crosses is given in Appendix

tube growth was observed using
4.
the aniline-blue fluorescence technique de

At
least
(1991).
three flowers were examined per crossing combination.
In ambiguous cases, numerous flowers were examined for a given cross until a consistent
pattern was observed.
scribed in Bohs
Pollen viability was estimated in the parental plants using the aniline-blue-lactophe
nol technique of Hauser and Morrison
(1964) as described in Bohs (1991). At least five
flowers were observed per individual plant, and two to seven individual plants per acces
sion were monitored. Pollen was allowed to stain for at least one hour before observation.
first 300 grains encountered were scored as either viable or inviable. Unshriveled
grains staining deep blue were presumed to be viable. Pollen fertility is reported in Ap
The
pendix 4.
SECTIONALDELIMITATIONAND RELATIONSHIPS
All species of Solanum sect. Cyphomandropsis
share the following combination of
characters: 1) they are woody shrubs or small trees lacking spines; 2) trichomes, if pre
sent, are simple to dendritically branched; 3) the anthers are relatively narrow and distally
SOLANUM
2001
small terminal pores that usually do not enlarge into longitudinal slits; 4) a
region is lacking on the abaxial anther surface. At least eight
of the 13 species of the section have thick, angled seeds and relatively few seeds per fruit.
is closely related to Solanum sect. Pachyphylla
Solanum sect. Cyphomandropsis
tapered, with
discrete enlarged connective
formerly recognized as the segregate genus Cyphomandra

(Bohs 1994). Plants of the two sections can be very similar morpholog
are distinguished by having an enlarged connec
ically, but all taxa of sect. Pachyphylla
tive region on the dorsal anther surface that may function as a floral osmophore (Sendtner
which was
(Dunal) Dunal,
Mart.
ex Sendtn.
1994). All taxa of the two sections that have been inves
tigated cytologically have very large chromosomes and/or large amounts of nuclear DNA
(Roe 1967; Pringle & Murray 1991; Moscone
1992; Bohs 1994; see section "Chromo
somes"). Previous authors (D'Arcy 1972; Child 1984; Moscone
1992; Bohs 1994) ac
1845; Sazima
et al. 1993; Bohs
and Pachyphylla,
the close relationship between sections Cyphomandropsis
knowledged
and molecular data confirm that the sampled members of the two sections belong to the
same clade (Olmstead & Palmer 1992, 1997; Bohs & Olmstead 1997, in press; Olmstead
et al. 1999; Fig.
sect. Pachyphylla
1). Other characters
that may distinguish
include plant architecture
(Leeuwenberg's
from
to Chamberlain's model in
sect. Cyphomandropsis,
Prevost's model in sect. Pachyphylla),

fruit texture (mesocarp
in sect. Cyphomandropsis,
abundant and juicy in sect. Pachyphylla),
and habitat preferences (drier and cooler, often highland areas in sect. Cyphomandropsis,
wetter and warmer regions at lower elevations in sect. Pachyphylla). These characters are
scant and gummy
described
in more
detail
in the sections
"Morphology"
and "Habitats and Distribution"
below.
data place the Cyphomandropsis/Pachyphylla
of members
of Solanum subg. Leptostemonum
solanums),
some
a large clade
group within
Molecular
composed
(Dunal) Bitter
(the spiny
of subg. Minon Raf. [sections Brevantherum
Seithe,
(Moench) Bitter, Extensum D'Arcy, and Holophylla
(G. Don) Walp., pro
representatives
Pseudocapsicum
parte], sect. Geminata
and two representatives
(G. Don) Walp.,
of problematical
groups,
S. allophyllum
(Miers) Standl. [sect. Allophyllum
(A. Child) Bohs] and S. wendlandii
Hook.f. (sect. Aculeigerum
Seithe; Fig. 1). This clade, which includes nearly half of the
species in Solanum, is morphologically
diverse, biogeographically
the sister group to the Cyphomandrop
and poorly understood. Likewise,
clade has not been identified with certainty. Trees resulting from com
sis/Pachyphylla
bined nuclear and chloroplast sequence data sets place members of sections Aculeigerum,
currently recognized
widespread,
Allophyllum, Brevantherum, and Extensum as sister to the Cyphomandropsis/Pachyphylla

clade (Bohs & Olmstead, in press; Fig. 1), but this clade has very low bootstrap support,
and many members of potentially related groups have not been sampled.
TAXONOMICHISTORY
of sect. Cyphomandropsis
Cyphomandra. All species described
Species
have been placed
in the past in either Solanum or
established the
genus Cyphomandra, were assigned to Solanum. Bitter set up the sect. Cyphomandropsis
in 1913, which included two newly described species (S. stuckertii and S. semicoalitum)
and one previously known species (S. clavatum), and transferred two species previously
described as Cyphomandra
to Solanum (S. johannae and S. luridifuscescens). Subsequent
authors assigned
the section
before
to Solanum
1845, when Otto Sendtner
(Seithe
1962; Gilli
1970; Danert
1970; Morton
VOLUME 61
100
96
100
L
100
I
89
1
9
3
12
72 |
99
84
32
100
93
100
85
91
L 100
93
|
17
99
37
100
r1
100
100
1. Strict consensus
FIG.
above
are bootstrap
branches
tails). Subgeneric
generic
has been
of 16 most
ndhF and nuclear

values
and sectional
taxonomy
most
frequently
placed
Minon
Minon
Minon
Leptostemonum
None
Solanum cordovense
Solanumwendlandii
Solanum allophyllum
Solanum betaceum
Solanum luteoalbum
Solanum glaucophyllum
Solanum ptychanthum
Solanum villosum
Solanum physalifolium
Solanum tripartitum
Solanum palitans
Solanumwallacei
Solanumdulcamara
Solanum nitidum
Solanum appendiculatum
Solanum tycopersicum
Extensum
Aculeigerum
Allophyltum
Pachyphylla
Cyphomandropsis
Cyphomandropsis
Solanum
Solanum
Solanum
Parasolanum
Parasolanum
Califomisolanum
Dulcamara
Holophylla
Anarrichomenum
Lycopsersicum
Minon
Potatoe
Potatoe
Solanum tnzygum
Solanum trisectum
Trigueraosbeckii
Pteroidea
Normania
None
Bassovia
Potatoe
None
Solanum aviculare
Solanum laciniatum
Archaesolanum
Archaesolanum
Archaesolanum
Archaesolanum
parsimonious
trees of 1444 steps from parsimony
ITS sequences
(modified
see Bohs
(500 replicates;
assignments
and subgeneric
Leptostemonum
Leptostemonum
Leptostemonum
Leptostemonum
Leptostemonum
Leptostemonum
Leptostemonum
Leptostemonum
Minon
Solanum
Genus Cyphomandra
Debated
Debated
Solanum
Solanum
Solanum
Solanum
Solanum
Potatoe
Potatoe
JaltQmataprocumbens
Capsicum baccatum
Lycianthesheteroclita
Physalis alkekengi
Witheringiasolanacea
66
data from chloroplast
Subgenus
Leptostemonum
Leptostemonum
Nyctenum
Solanum vespertilio
Micracantha
Solanum adhaerens
Solanum jamaicense
Eriophylla
Leprophora
Solanum elaeagnifolium
Solanum campechiense
cf. Cryptocarpum
Torva
Solanum torvum
Acanthophora
Solanummammosum
Lasiocarpa
Solanum candidum
Solanum argentinum
Holophylta
Geminata
Solanum arboreum
Solanum pseudocapsicum Pseudocapsicum
Solanum abutiloides
Brevantherum
~g
17
Section
Melongena
Melongena
Solanummacrocarpon
Solanummelongena
follow D'Arcy
of sect. Cyphomandropsis
in either
Solanum
from Bohs
and Olmstead,
(1972,
in press,
1991), Child
or
in press). Numbers
for methodological
(1998),
and Nee
Traditionally,
has been controversial.
subg. Leptostemonum
of combined
analysis
and Olmstead,
in the genus
de
(1999).
The
the section
Cyphomandra
(see text).
1976) or to Cyphomandra
transferred to Solanum
is S. pubescens
1972; Child
1984). All species of Cyphomandra
were
in 1995 (Bohs 1995).
The earliest description

dropsis
(D'Arcy
of a species
that would
later be assigned
from Ruiz and Pavon's Flora Peruviana
renamed as S. luteoalbum by Persoon
(1805). Desfontaines
a new species of the section, S. glaucophyllum,

ber of the group. His description was
later
and most widespread mem
taken from a plant cultivated
records indicate that this showy species was cultivated
(1799),
(1829) was the next to describe
the best-known
in Paris. It is not known how or when S. glaucophyllum
to sect. Cyphoman
et Chilensis
at the Botanic Garden
arrived in France, but herbarium

in several European botanical gar
dens in the early 1800's, probably introduced from Argentina. Whereas most of the species
now
included within
have been at least loosely associated with
SOLANUM
2001
each other in the past, S. glaucophyllum has been an exception owing to its aberrant mor
and striking convergence with S. amygdalifolium Steud., an unrelated non-spiny
Solanum of the dulcamaroid group. Many authors since Sendtner (1846) and Dunal (1852)
than to
have considered S. glaucophyllum
to be more closely related to S. amygdalifolium
phology
other species of sect. Cyphomandropsis.

Dunal (1852) added to the confusion when he
placed S. glaucophyllum and S. malacoxylon,
here considered synonyms, into two separate
groups within
considered
Solanum
("Anthoresis"
S. glaucophyllum
and "Subdulcamara," respectively). Child (1986)

to sect. Cyphomandropsis
and removed it to
to be unrelated
its own section within Solanum subg. Solanum. Morphological,

cytological, and molecular
work (Morton 1976; Moscone
1992; Bohs & Olmstead, in press; Fig. 1) has confirmed the
placement of S. glaucophyllum within sect. Cyphomandropsis,
and has discounted its rela
tionship with the superficially similar S. amygdalifolium.
Nearly concurrently with Desfontaines's
(1829) publication of S. glaucophyllum, Vel
lozo (1829) featured two new species, S. ellipticum and S. cylindricum, in his account of
plants of the Rio de Janeiro region. Unfortunately,
the plates and descriptions from this
work are barely suitable for identification of many taxa, and no authentic herbarium ma
terial is known to exist. Solanum ellipticum is considered to be a synonym of S. cylin
dricum in the present work.
In 1846, Sendtner published his landmark treatment of Brazilian Solanaceae forMar
tius's Flora Brasilensis. He had established the genus Cyphomandra
in the previous year
and transferred to it a number of Solanum species, including S. ellipticum and S. cylin
dricum. In the same work, he also described two new species that would later be assigned
to sect. Cyphomandropsis,
one as a Cyphomandra
(C. velutina, a synonym of S. luridi
and one as a Solanum (S. malacoxylon,
fuscescens)
a synonym of S. glaucophyllum).
Dunal's
(1852) treatment of Solanaceae
for Candolle's Prodromus basically followed
Sendtner's scheme, but Dunal added the new taxa C. cornigera (a synonym of S. pelag
icum), C. betacea var. velutina (a synonym of S. fallax), and S. glaucum (a synonym of S.
He considered S. glaucum and S. malacoxylon
to represent two distinct
they are treated as conspecific
in the present work. Miers (1855) transferred C.
cylindrica, C. elliptica, C. cornigera, and C. velutina to his genus Pionandra. Later au
thors (e.g., Smith & Downs 1966; D'Arcy 1973; Child 1984; Bohs 1994) did not recog
glaucophyllum).
species;
nize the distinction between Cyphomandra and Pionandra, and Pionandra was relegated
to synonymy under Cyphomandra. Cyphomandra
is currently treated as a synonym of
Solanum (Bohs 1995).
The prolific taxonomist Georg Bitter published his works on Solanaceae during the
early part of the 20th century. Bitter (1913) erected Solanum sect. Cyphomandropsis
to in
clude two new species (S. stuckertii and S. semicoalitum) and two species transferred from
Cyphomandra
(C. velutina and C. elliptica). The next year he described S. narcoticum
(Bitter 1914), which he did not assign to sect. Cyphomandropsis,
but noted that it occu
pied an isolated position among the non-spiny taxa of Solanum. The identity of S. nar
coticum is in question, because Bitter's type specimens were destroyed in the Second
World War, and his description is not adequate to assign the name with certainty. In 1918,
Bitter transferred Cyphomandra
lauterbachii H. Winkler
to Solanum and assigned it to
sect. Cyphomandropsis.
Its oblong rather than tapered anthers exclude this species from
the section, and it actually belongs to Solanum sect. Geminata.
A few taxa were described during the next 40 years, but the more noteworthy devel
opments did not occur until the 1960's and 1970's with the publication of floras by
Macbride
(1962), Smith and Downs (1964, 1966), and Morton (1976). Solanum nitidum
var. hutchisonii
(now recognized
as S. hutchisonii)
VOLUME 61
and S. luteoalbum var. tunya (a syn
onym of S. luteoalbum) were first published inMacbride's

(1962) Flora of Peru. Smith
and Downs
(1964, 1966), in their works cataloging the flora of Santa Catarina state in
Brazil, proposed the novelties S. fusiforme, S. matadori, C. maritima (a synonym of S.
pelagicum), and S. catanduvae, S. iraniense, and S. subhastatum (the latter three here con
sidered synonyms of S. cylindricum). They maintained Cyphomandra as a separate genus,
based on the enlarged anther connective. They assigned S. fusiforme and S. matadori to
two large subgenera within Solanum, subg.
sect. Cyphomandropsis,
and recognized
to belong
Solanum and subg. Leptostemonum. They considered sect. Cyphomandropsis
within Solanum subg. Leptostemonum, which otherwise includes the spiny solanums. In
the
doing so, they emphasized the narrow tapered anthers with small pores and minimized
have neither stellate hairs nor spines. In real
fact that species of sect. Cyphomandropsis
ity, it appears that there is no clear-cut division of Solanum into two large subgenera based
on anther shape, trichome characters, and presence of spines (Bohs & Olmstead 1997).
contributions to our understanding of the Solanaceae
Conrad V. Morton's
included
much herbarium work on Solanum as well as several publications (Morton 1944, 1976).
Most
noteworthy is his treatment of the genus Solanum inArgentina (Morton 1976). Mor
ton also included sect. Cyphomandropsis within subg. Leptostemonum on the basis of its
tapered anthers with small pores. He described S. confusum and S. adelphum, which are
considered conspecific in the present treatment. He clarified the nomenclature of S. con

fusum and S. clavatum, and resolved some difficult problems of synonymy and typifica
tion in S. glaucophyllum.
The taxonomy of sect. Cyphomandropsis
has received little attention until relatively
recently. Many of the species occur largely outside the area of major published floras, and
no revisionary or monographic
or
studies have been available for sect. Cyphomandropsis
its close relative sect. Pachyphylla.
In the last 15 years, Child (1984) and Bohs (1994)
have published
that examined the Pachyphyllal

taxonomic conspecti or monographs
clade, and molecular data (Olmstead & Palmer 1992, 1997; Bohs &
Olmstead 1997, in press) have elucidated the position of this clade within the larger pic
ture of Solanum. Child (1984) included sections Cyphomandropsis
in the
and Pachyphylla
Cyphomandropsis
genus Cyphomandra,
and made a number of implicit transfers of Solanum taxa to
Cyphomandra; however, most of these transfers are not validly published under Art. 33.2
of the ICBN (Greuter et al. 2000), because no reference was made to the basionyms.
These combinations were validated in Bohs (1994). Child's concept of Cyphomandra was
quite broad, and he included within it several elements that are now thought to belong to
distinct clades of Solanum [e.g., S. graveolens Bunbury, amember of Solanum subg. Pota
toe (D'Arcy 1972; Bohs 1994), and S. allophyllum, of Solanum sect. Allophyllum
(Bohs
1990), which apparently belongs to an isolated clade along with S. wendlandii of Solanum
sect. Aculeigerum
(Bohs & Olmstead 1997)]. With these anomalous elements included,
Child recognized six sections within Cyphomandra. Though his work is insightful, it did
not examine species limits and nomenclature in the taxa under consideration.
In 1994, Bohs published amonograph of the genus Cyphomandra. She recognized 32
species and two insufficiently known taxa. She combined the sections Cyphomandra and
Ceratostemon Miers recognized by Child, and defined five informal species groups. She
excluded four of Child's
six sections (Allophylla, Rhynchantherum,
and
Cornigera,
from inclusion in the genus.
Cyphomandropsis)
At about the same time, molecular
data clearly established
that the genus
is nested deeply within the genus Solanum (Olmstead & Palmer 1992,
Cyphomandra
SOLANUM
2001
1997; Bohs & Olmstead 1997, 1999), and thus recognition of Cyphomandra as a distinct
untenable. Accordingly,
all species of Cyphomandra were trans
genus is phylogenetically
ferred to Solanum (Bohs 1995). The infrageneric name of this group is now Solanum sect.
Pachyphylla
(Dunal) Dunal.
The present treatment aims to clarify the species limits of the remaining taxa in the
clade. Included are members
PachyphyllalCyphomandropsis
and Cornigera, which are not recognized

Cyphomandropsis
species formerly considered to belong to sect. Pachyphylla,
Floral morphology
the sections. Monophyly
S. fallax,
and architectural
hypothesis
sections,
of relationships within
is now included in
characters
of each section has not been definitively
and an explicit phylogenetic

tailed molecular
of two of Child's
here as distinct sections. One

serve to delimit
established,
however,
the entire clade awaits de
work.
MORPHOLOGY
are gener
HABIT, STEMS, AND ARCHITECTURE.Members of sect. Cyphomandropsis
can
a small
is
m
tall.
An
S.
which
be
shrubs
less
than
3
fallax,
ally small woody
exception
a
narrow
in
contrast
to
tree up to 5 m tall. The upright stems are usually slender with
pith,
the thick fleshy stems of species of Solanum sect. Pachyphylla. Obvious
scars are present on young stems. Older stems generally have smooth white
elevated
leaf
to dark brown
longitudinal fissures and raised lenticels. Some species of the section

S. hibernum, S. luteoalbum, S. stuckertii) produce
(e.g., S. confusum, S. glaucophyllum,
several tomany upright stems from an underground rhizome. In S. confusum and S. glau
clumps composed of hundreds of
cophyllum, this habit may result in large monospecific
bark marked with
stems.
the architecture of Cyphomandropsis
species is similar to that of many
(Fig. 2). Initially a single stem is produced with spirally arranged leaves.
After reaching ca. 0.5 to 2 m in height, this upright stem ends in a terminal inflorescence.
Two to three lateral shoots then develop from axillary buds just proximal to the inflores
Where
known,
other solanums
cence. These
lateral shoots ascend at an acute angle, in contrast to the plagiotropic lateral

in the same morphological
(see
position in species of sect. Pachyphylla
shoots produced
Bohs,
1989). Growth of the lateral shoots is sympodial,
eral the sympodial units are plurifoliate

from the orthotropic
axis. Thus,
as in sect. Pachyphylla,
and not clearly differentiated
the trunk and branches
are more
but in gen
in leaf arrangement
or less equivalent
in
the branches differ from

species of sect. Cyphomandropsis, whereas in sect. Pachyphylla
the trunk in both orientation and leaf arrangement. The architecture exhibited by most
conforms to Prevost's model in the scheme of Halle et al.
species of sect. Pachyphylla
(1978). This is an uncommon model in the Solanaceae, and may be a synapomorphy that
defines sect. Pachyphylla. Furthermore, species of sect. Pachyphylla
that exhibit Prevost's
model
sympodial modules on the plagiotropic (crown) branches,

usually have 3-4-leaved
and early orders of branching on the plagiotropic shoots are generally dichasial; that is,
two replacement shoots grow out from buds beneath the inflorescence so that the inflo
rescence
appears to be situated in a branch fork. Sympodial
modules
in species of sect.
Dichasial branching
vary from 3- (in S. fallax) to 4- to many-foliate.
Cyphomandropsis
can also occur in sect. Cyphomandropsis,
but monochasial
branching ismuch more com
mon, especially on later orders of branching. Thus, architecture in species of sect.
Cyphomandropsis
lies along a continuum
between
Leeuwenberg's
and Chamberlain's
VOLUME 61
ste showin
... l .
bpatr;..........
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Roe,
Rof,
2001
SOLANUM
[In the former hair type, all lateral rays emerge from one point on
the axis. In the latter two types, the lateral rays are numerous, very short, and emerge at
various points along the central axis (Roe 1971).] Likewise, prickles are absent in species
of this section.
have entire, petiolate, el
of species of sect. Cyphomandropsis
species have truncate, cuneate, or decurrent leaf
blades, but in S. amotapense, S. hibermum, and S. hutchisonii the leaf base can be subcor
LEAVES. The majority
liptic to elliptic-ovate
leaves. Most
date, and in S. fallax

leaves
the leaf base is usually deeply cordate. Solanumfallax

has the largest
in the section, with blades often reaching 20 cm or more in length and width.
Leaves of the trunk and crown portion of the plants are more or less similar in shape and
size, and do not exhibit the degree of morphological
heterogeneity
commonly seen in
species of sect. Pachyphylla.
Solanum fusiforme and S. pelagicum frequently have both pinnately compound and
leaves on a single plant, with the compound leaves especially common on the
trunk. A few specimens of S. cylindricum have small hastate lateral lobes near the base of
simple
the blade.
Leaf texture in the group is generally chartaceous; subcoriaceous or fleshy leaves are
found only in S. hutchisonii and S. glaucophyllum. The latter species is notable because
most plants are covered with a dense glaucous
bloom.
INFLORESCENCES.Inflorescences
late, scorpioid
cymes. As
in sect. Cyphomandropsis
are ebracteate, peduncu
in all other species of Solanum, the inflorescence is develop
mentally
terminal, and shoots distal to the first inflorescence
modules.
Most
species
have unbranched
to forked
are composed
inflorescences.
of sympodial
Branched
inflores
in S. confusum, S. glaucophyllum,
and S. matadori; S. luteoalbum and S.
stuckertii rarely have branched inflorescences. In S. fusiforme, the inflorescence axis is no
cences
occur
ticeably zigzag between
adjacent pedicels.
The
inflorescences
are generally
nodding
to
pendent.
In all species,
except S. fallax, the pedicels are articulated at the base and after ab
leave scars or short pegs up to 1mm long on the rachis. In S. fallax, the pedicels
can have an articulation some distance above the point at which the pedicel joins the
rachis, and upon abscission can leave remnants up to 6 mm long (see Fig. 16). This pat
scission
tern is also seen in some species of sect. Pachyphylla

(Bohs 1994).
In some specimens of S. cylindricum, the flowers are borne in subumbellate clusters
with the pedicels inserted on a swollen platform. Pedicel platforms are a synapomorphy
found in some groups of Solanum sect. Holophylla
(Knapp 1989). Because the groups that
possess this character have no close phylogenetic connection with sect. Cyphomandrop
sis (cf. the position of Solanum nitidum Ruiz & Pav. with
in Fig. 1), the pedicel platform
Cyphomandropsis
ample of convergence.
FLOWERS. The
actinomorphic,
flowers
and 5-merous.
five lobes up to 3 mm
of
species
The calyx
long (up to 6 mm
of
the two representatives
in S. cylindricum
of sect.
appears to be an ex
are all perfect,
is green or greenish white
and cupulate, with
long in S. cylindricum).
In S. amotapense and
S. fallax the calyx tube is somewhat inflated, then constricted at the point where the
lobes emerge. This morphology
is not common
in members
of the Pachyphyllal
10
VOLUME 61
Cyphomandropsis
clade, but it occurs in at least a few species of Solanum sect. Brevan

therum (e.g., S. hazenii Britton) and is apparently convergent in the two groups.
are chartaceous in texture and stellate, rotate-stel
Corollas in sect. Cyphomandropsis
in outline. Corolla color is most commonly white or purple,
late, or stellate-campanulate
with both color morphs appearing in several species. Solanum glaucophyllum and S. con
fusum frequently have pinkish corollas, often with a darker or lighter central star. In gen
in sect. Cyphoman
eral, corolla shape, texture, and color are much more homogeneous
dropsis than in sect. Pachyphylla, which exhibits a remarkable diversity in floral color and
morphology.
are narrow and distally tapered, with small
The stamens of sect. Cyphomandropsis
terminal pores that do not open into longitudinal slits with age. Solanum cylindricum is an
exception, for older anthers in this species often have pores that extend into slits. This
character is also seen in several groups thought to be related to the Pachyphyllal
and Holophylla
Cyphomandropsis
clade, such as sections Geminata, Pseudocapsicum,
pro parte (Knapp, in press). Whether this indicates that S. cylindricum might be relatively
basal in the PachyphyllalCyphomandropsis
clade remains to be investigated.
such as Bitter (1913) and Morton (1976), attached much signifi
to the degree of anther connation in distinguishing
species. This is not a reliable
character at the specific level, for both connivent and free anthers can occur within a sin
gle taxon or within a single plant. Connivent anthers in Solanum occur due to interlock
ing papillae or hairs on the lateral anther surfaces (cf. Bonner & Dickinson
1989). Envi
Previous workers,
cance
ronmental factors or developmental

stage may thus affect the degree of anther connation
in a particular flower.
In contrast to members of sect. Pachyphylla,
lack
species of sect. Cyphomandropsis
a discrete enlarged connective region on the dorsal surface of the anthers (Bohs 1994;
Fig. 3). The presence
Pachyphylla
of this connective
and serves to differentiate
is the primary defining
characteristic
it from sect. Cyphomandropsis
of sect.
regardless of vari
ation in other characters. However, some species of sect. Pachyphylla can have small or
inconspicuous connectives
[e.g., S. corymbiflorum (Sendtn.) Bohs, S. pinetorum (L. B.
Sm. & Downs) Bohs], and some taxa of sect. Cyphomandropsis may have anthers that are
dorsally thickened or elaborated (e.g., S. fallax, S. luridifuscescens). The key to distin
guishing between the two situations is whether the connective
is discrete and clearly
from the thecal tissue (sect. Pachyphylla)
differentiated
or whether
it covers nearly
all of the dorsal anther surface and is not clearly demarcated
from the thecae
(sect.
Cyphomandropsis).
The gynoecium of species of sect. Cyphomandropsis
consists of a conical bicarpel
late ovary, a straight, slender, cylindrical style, and a truncate to subcapitate stigma more
or less the same diameter as the style. This conforms to gynoecia inmany other groups of
and contrasts with species of sect. Pachyphylla, which may have greatly ex
stigmas or distinctive stylar morphology
(see Bohs, 1994, for examples). Most
sect.
of
species
Cyphomandropsis
have glabrous ovaries and glabrous to sparsely puberu
lent styles; exceptions include S. fallax and some collections of S. amotapense, S. cylin
dricum, and S. stuckertii.
Solanum,
panded
POLLEN. Pollen grains were examined with SEM in the species listed in Table 1;
herbarium vouchers for the pollen studies are given inAppendix 1. Pollen of these species
is tricolporate, with the colpi not fused at the poles (Fig. 4). The grains are spheroidal to
in equatorial view; in polar view the apertures often protrude so that the
prolate-spheroidal
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12
TABLE 1. Pollen
Herbarium
vouchers
of Solanum
are listed in Appendix
examined
VOLUME 61
with
SEM.
1.
Species
Sculpturing
S. amotapense
S. confusum
S. cylindricum
S. fusiforne
S. glaucophyllum
S. pelagicum
Granulate
Granulate
Granulate
Granulate
Psilate
Granulate
Size (pm)
15-16.25
13.75-16.25
11.25-11.75
11.25-12.5
12.5-13.75
11.25-12.5
is semiangular. Grain diameter is approximately 10-20 Pm. Inmost of the species

examined, the pollen grains are finely ornamented with small bumps or granules less than
1 pm in diameter. The only exception is S. glaucophyllum,
in which the grains are nearly
outline
smooth (psilate).
Passarelli
(1999) examined pollen grains of three Argentine

species of sect.
S. glaucophyllum, S. stuckertii, and S. confusum (as S. adelphum). Her
Cyphomandropsis,
results were similar to those reported here, except she found the surface sculpturing of
grains of S. glaucophyllum
to be granulate rather than psilate. The diameter of acteolyzed
grains was 16-20 pm in S. glaucophyllum,
17-20 pm in S. stuckertii, and 17-24.5 Pm in
S. confusum. The number of pollen grains per flower estimated using a hemacytometer
ranged from about 1.1 to 1.3 x 106 in the three species.
Pollen
size and surface sculpturing in sect. Cyphomandropsis

are very similar to that
of sect. Pachyphylla
and to many other species of Solanum (Anderson &
Gensel 1976; Edmonds 1984; Bohs 1989, 1994; Passarelli 1999; Knapp, in press) and do
not appear to provide useful taxonomic characters at the sectional or specific level.
of species
FRUITS. The fruits in sect. Cyphomandropsis

are usually globose berries ca. 1-2
in diameter. The fruit pulp in most species is sparse and much drier in texture than in
succulent, juicy berries common in sect. Pachyphylla. The surface of the berry can
shiny (e.g., S. amotapense, S. hibernum, S. stuckertii) or dull (e.g., S. glaucophyllum,
cm
the
be
S.
luteoalbum). Ripe fruits are most commonly yellow or orange, but S. amotapense has
bright red fruits. Solanum glaucophyllum
is distinctive in producing dark purple or black
ish fruits usually covered with a glaucous bloom. The yellowish or pale green mesocarp
is juicier than in other members of the section and is extremely bitter. In one collection of
this species from lowland Bolivia (Nee 37532) the fruits are described as green when they
abscise. I have only seen blue-black mature fruits on plants of S. glaucophyllum.
is also very unusual in its fruit morphology. As the name implies,
Solanumfusiforme
the fruits are elongated and distally pointed. They turn yellow when ripe (Dottori 1995).
Solanum fallax is distinctive
in having fruits that are densely pubescent with un
branched eglandular hairs. Several authors (Smith & Downs 1966; Child 1984) have de
scribed the fruits of S. pelagicum as being dendritically pubescent, but I have only seen
glabrous fruits in herbarium material of this species.
Stone cell aggregates are commonly found in the fruits of species of sect. Pachy
are macroscopic
sclerified structures that often occur in the distal and me
dial parts of the fruit. Large stone cell aggregates occur in just a few species of sect.
Cyphomandropsis
(e.g., S. fusiforme, S. pelagicum). Dottori (1995) reports that the fruits
phylla. These
rains
Plar
C view
13
SOLANUM
2001
....~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
j.j.j.
:~~~~~~~~~~~~~~~~~~~~~~
:;:
:i...
.:.::
|~~~~~~~~~~~~~~~
..,.:..i?'
....
..........:.....
...
.......
.....j .
of
S.lanum
x350of
served
sc
C.hom
w
observed
?itht dm
S... E
... ........
,D.
pm).
Exm
surfac
(sale
bars=5
. ........0......
. .p
...5
S c.nfuum
and
S.!
:ckerii
them
:in
these
species
also
The
S.
. C,D
.
...
contain
aggregations,of
*
sciereids
but
..... . , .i.,X,SU .,, ;
have ......not ob-...

,6, iS*,;;.~~~~~~~~~~.
ss~~~~~~~~~~~~~~
.:j:.;e
|ii'
S
g.
:.:
.:
..:&.x
.:.
.. ... ...::.....:'.:'
::.:
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
taxonomic
utility
and
functional
signific;ce
of;
thi
....
-:'~~
,.,>X8
.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
~ ......
when they have been identifiedwith certaintyto determine if thischaractr is primitiveor

............
...a.'..^
~ ..~
~ ~...
'..t .. t ...
i' 'ti5i.
,'M
-B~~~~~~~~~~~~~~~~~~~~~~~~~....
....
.. ....^.
........
~[~~~~~~~~~~~~~~~~~~~~.......
...
denved
inthemgnrus
There are only two known reports of fruit dispersal in the group, both for S. glauco
of fruit and seed dispersal of Cyphomandropsis
species are
phyllum (q.v.). Observations
urgently needed. This dearth of information regarding dispersal agents also extends to
14
species of Solanum in general, where

1979, for an exception).
SEEDS. Two basic
species have relatively
flattened, and rounded
long and 3-5 mm wide,
few well-documented
VOLUME 61
studies exist
(see Symon,
seed morphologies
occur in sect. Cyphomandropsis.
At least four
small seeds (ca. 2-3 mm in diameter) that are lenticular, strongly
in outline. The rest of the species have large seeds, ca. 4-5 mm
which are rather thick, not very flattened, and angled in outline.
Whether
these two seed types define monophyletic
lineages remains to be determined.
Taxa known to have each of these seed types are listed in the synoptic key. Large-seeded
species usually have fruits with very sparse mesocarp, and seeds occupy most of the vol
taxa also contain only on average about 10-50 seeds per fruit.
taxa have more abundant mesocarp and may contain hundreds of seeds.
Small-seeded
As is common in many species of Solanum, the outer walls of the cells of the seed
coat are very thin and easily rub off. The anticlinal walls of these cells are thickened in
comblike ridges that remain after the outer wall has disappeared (Soueges 1907; Edmonds
ume of the fruit. These
1983; Lester & Durrands 1984; Bohs 1994; Dottori 1995). Thus, many Solanum seeds ap
pear to be "pubescent," but the apparent hairs (termed "pseudohairs") are not composed
of cells but rather of the remnants of the thickened anticlinal walls.
CHROMOSOMES
All species of sect. Cyphomandropsis
that have been investigated cytologically have
a chromosome number of n = 12 (2n = 24; Appendix 3). The base chromosome number
in Solanum and in most other genera of the subfamily Solanoideae
is x = 12 (Moscone,
1992, and references cited therein). All species of sect. Pachyphylla
investigated thus far
also have n = 12 and 2n = 24 chromosomes
(Bohs 1994). Polyploidy does not appear to
be a factor in the evolution of the PachyphyllalCyphomandropsis
clade, as it has been in
some groups, such as the potatoes (Solanum sect. Petota Dumort.; Hawkes 1990) and the
nightshades (Solanum sect. Solanum; Edmonds 1972, 1977).
Although chromosome number is not unique in the PachyphyllalCyphomandropsis
clade, the group is cytologically
distinctive due to its large chromosome
size and high
amounts of nuclear DNA. All species from this clade that have been investigated have
chromosomes
that range in length from about 3 to 14 ,um,making them on the order of 2
to 5 times larger than those of other species of Solanum (Roe 1967; Pringle & Murray
1991; Moscone
1992; Bohs 1994). Pringle and Murray (1991), using flow cytometry, in
cluded S. luteoalbum from sect. Cyphomandropsis
in their investigation of DNA content
in the PachyphyllalCyphomandropsis
group. DNA content in this species was measured
at 15.2 picograms per 2C nucleus. This value falls within the range reported for species
(Pringle & Murray 1991) and is substantially greater than that re
ported for other species of Solanum (including formerly recognized genus Lycopersicon
Mill.; range = 1.7-4.2 pg per 2C nucleus; Bennett & Smith 1976; Bennett et al. 1982; Aru
muganathan & Earle 1991). The functional significance of this remarkable difference in
chromosome size and DNA content is unclear, but itmay represent a synapomorphy that
defines the Pachyphylla/Cyphomandropsis
clade. Efforts should be made to examine
chromosome number, size, and morphology
in all the species of this clade with respect to
its sister group(s) in order to establish a hypothesis of chromosomal evolution in sect.
and its relatives, and to gain insight into the possible adaptive signifi
Cyphomandropsis
cance of increased chromosome size and DNA content in flowering plants.
2001
SOLANUM
15
BREEDING SYSTEMSAND CROSSING STUDIES

Limited
terspecific
in sect. Cyphomandropsis.
in floral form. Appendix 4 documents
monomorphic
the distribution of self-incompatibil

in species of the section as determined by fruit and seed set and
ity and self-compatibility

tube growth
pollen
have been done to determine breeding systems and in

All species are hermaphroditic and
crossing experiments
compatibility
in selfs vs. sib- or outcrosses.
style in incompatible pollinations,
Pollen tube growth was arrested in the

to the pattern seen in other Solanaceae with
conforming
self-incompatibility
(Nettancourt 1977). Self-compatibility
may be more
in sect. Cyphomandropsis
than in sect. Pachyphylla:
two out of five species
investigated in sect. Cyphomandropsis were self-compatible, whereas only two out of ten
gametophytic
widespread
species were
in sect. Pachyphylla
self-compatible
were set without manipulation,
which
lination. Solanum glaucophyllum
(Appendix 4; Bohs 1994). No fruits

indicates that biotic vectors must be present for pol
and S. hibernum occasionally
set some fruits with seeds
after selfing in greenhouse
indicates pseudocompatibility
from environmental
(cf. Pandey
Passarelli
crosses. This probably

conditions in the greenhouse
(1999) calculated
the pollen/ovule
resulting
1959).
(P/O) ratio for three Argentine
species
of sect. Cyphomandropsis (S. confusum, S. glaucophyllum, and S. stuckertii). The

ratio is thought to reflect efficiency
pollen/ovule
dicator of breeding
were
very high
of pollen
transfer and thus be a rough in
system (Cruden 1977). The P/O ratios obtained
(37,875
in S. confusum,
25,309
for the three species
in S. glaucophyllum,
and 23,220
in S.
stuckertii)and are consistentwith xenogamy (Cruden1977).Passarelli (1999) reportsthat

the three Cyphomandropsis
greater than the highest
species are self-incompatible.

ratios calculated
by Mione
These P/O ratios are equal to or
and Anderson
(1992) for species of
Solanum sect.BasarthrumBitter known to be self-incompatible.

Artificial interspecific intrasectionalcrosses were attemptedusing four species of
(S. glaucophyllum,
S. hibernum, S. luteoalbum, and S. stuckertii).
Appendix 4 summarizesthe resultsof thesecrosses. Some combinations(e.g., S. luteoal

bum x glaucophyllum,S. stuckertiix hibernum)producedhybridplants thatwere inter
inmorphology
mediate
between
the parental species;
in the case of S. luteoalbum x glau
cophyllum, the cross was successful only in one direction.Pollen tube growth into the
ovary was observed
which
in several crossing combinations
that did not produce fruits or seeds,

crossing baffiers may be present. Both pre- and post-zy
implies that post-zygotic
gotic baffiers appear to be effecting

they are in sect. Pachyphylla
isolation among species of sect. Cyphomandropsis,
as
(Bohs 1991, 1994).
Artificial intersectional crosses were attempted in both directions between four

species of sect. Cyphomandropsis
S. stuckertii)
and seven
species
(S. glaucophyllum,
S. hibernum,
(S. betaceum,
S. luteoalbum,
S. circinatum,
and
S.
corymbiflorum,S. diploconos, S. diversifolium, S. roseum,and S. unilobum;Appendix

4). Small
seedless
fruits developed
in many
of these crosses,
but none produced
viable
seeds. Pollen tubegrowth in intersectionalcrosses ranged from very good (many tubes
around ovules)
to poor (abnormal
tube growth,
few to no tubes in ovary), depending
on
the crossing combination.Reproductive isolation appears to be complete or nearly so

between sections Cyphomandropsisand Pachyphylla. This is perhaps to be expected,
given the strongcrossing barriersbetweenmany specieswithin sect. Pachyphylla (Bohs
1991).
16
VOLUME 61
RELATIONSHIPS
SPECIESGROUPS AND INFRASECTIONAL
Phylogenetic
been established
will confound
have not
relationships among the species of sect. Cyphomandropsis
using rigorous analytical methods. It is likely that extensive convergence
cladistic analyses based on morphological
characters, as is evident in other
Solanum groups (Bohs & Olmstead 1997, 1999). Molecular

studies are in progress to ex
amine evolutionary
relationships among species in the PachyphyllalCyphomandropsis
clade (Bohs, unpubl.).
In the absence of cladistic
analyses, the following species groups are proposed based

on my intuitive interpretation of overall morphological
similarity. Though not intended to
and rigorously analyzed statements of relationship, they will
represent well-supported
serve as hypotheses for further testing.
I. S. luteoalbum group (S. hibermum, S. luteoalbum, S. stuckertii). These three species
share 1) stellate corollas, 2) long narrow anthers unelaborated on the dorsal surface, 3)
usually unbranched inflorescences, 4) globose, yellow to orange fruits, and 5) very large
branched hairs are present in S. hibemnum and in some popu
angled seeds. Dendritically
lations of S. luteoalbum. Corollas are white in S. stuckertii and deep purple (rarely white)
in the other two species.
II. S. amotapense group (S. amotapense, S. fallax). This group is probably most
closely related to the S. luteoalbum group with which it shares large angled seeds. The two
species are similar in their cordate leaf bases and in having the calyx tube swollen at the
distinctive in sect. Cyphomandrop
base of the lobes. Solanum fallax is morphologically
sis due to its large stature, long pedicel remnants, large leaves, and pubescent fruits. Few
leaved sympodial units may also distinguish it from other species in the section.
S. luridifuscescens,
III. S. glaucophyllum group (S. confusum, S. glaucophyllum,
S.
matadori). The taxa of this group all have elliptic and often rather narrow leaves with ta
pered bases, stellate to stellate-campanulate
corollas, and relatively broad anthers, often
with the dorsal surface roughened by scaly papillae. Branched inflorescences occur in all
except S. luridifuscescens. The seeds of S. confusum and S. glaucophyllum
are
large and angled, as in the species groups described above. Fruits and seeds are unknown
in S. matadori, and poorly known in S. luridifuscescens;
the latter appears to have small
lenticular seeds. Solanum confusum and S. glaucophyllum
range from central Bolivia to
the mouth of the Rfo de La Plata; S. luridifuscescens and S. matadori are found in south
eastern Brazil. Further investigation may reveal this group to be non-monophyletic.
species,
IV. S. cylindricum group (S. cylindricum, S. pelagicum). These two species from
southeastern Brazil are distinctive due to their usually abundant pubescence of dendriti
cally branched hairs. Although S. cylindricum is morphologically
variable, at least some
dendritic hairs are present on most specimens. Other species in sect. Cyphomandropsis,
such as S. luteoalbum and S. amotapense,
can have populations with dendritically
branched pubescence, but they differ from S. cylindricum and S. pelagicum in other mor
phological characters. Lobed or compound leaf blades may also link the two species; S.
pelagicum has pinnately lobed trunk leaves, and some specimens of S. cylindricum have
small hastate lobes at the base of the leaf blades.
V. Species not placed in a group (S. fusiforne, S. hutchisonii). Solanum fusiforme and
S. hutchisonii are each morphologically
distinctive, and their affinities to other species in
the section are not obvious. Both species are nearly glabrous, but otherwise they are dis
similar. Solanumfusiforme
is found inmesic areas of southeastern Brazil and adjacent Ar
and Paraguay, whereas S. hutchisonii occurs in dry valleys of northern Peru.
gentina
SOLANUM
2001
17
can be recognized by its pinnately compound leaf blades and fusiform

in the section has fruits with this morphology. Although S. pelag
icum also has pinnately compound leaves, it differs from S. fusiforme inmany characters.
Solanum hutchisonii has distinctive fleshy leaf blades, often with subcordate bases. It
may be most closely related to species of the S. amotapense group, which also have cor
date leaf bases and large angled seeds, and are found in nearby areas of Ecuador and
Solanumfusiforme
fruits. No other species
northernPeru.
HABITATSAND DISTRIBUTION
are found at a range of elevations from sea level
Species of sect. Cyphomandropsis
to snow line (ca. 4000 m). Although they occupy several distinct habitat types, in general
they prefer sunny areas, such as light gaps and clearings in primary forest, and disturbed
sites, such as roadsides or ravines in secondary vegetation. Approximately
half the species
in the group are typically found in seasonally arid inter-Andean valleys, often on rocky
slopes or in canyons. Some of them may lose their leaves during dry periods. These
species include S. amotapense, S. fallax, S. hibernum, S. hutchisonii, S. luteoalbum, and
S. stuckertii. Solanum confusum, in contrast, is generally found in wetter cloud forest
areas, especially in groves of aliso (Alnus acuminata). In the southern part of its range, S.
confusum is found in the bosque tucumano-boliviano
floristic province (Cabrera 1976;
Killeen et al. 1993). Solanum confusum occupies the highest elevations of any species of
the group (up to ca. 4000 m), tolerating at least short periods of frost and snow.
Some species of Cyphomandropsis,
such as S. cylindricum, S. fusiforme, and S. mata
dori, typically occur in clearings inAraucaria forests in southeastern Brazil and adjacent
areas at mid-elevations
(300-1200 m). These regions are warmer and more humid than
those described above, but can experience cold or frost in winter. A unique Brazilian en
is found at lower elevations and typically occupies restinga (coastal
demic, S. pelagicum,
scrub or dune habitats) along the coast of the state of Santa Catarina. Some populations of
this species also occur inland in forest clearings and margins. Another Brazilian endemic,
S. luridifuscescens, grows in higher elevation rain forest (ca. 1100-2600 m) and prefers
or swampy ground.
the more unusual habitats exploited
waterlogged
Among
the seasonally
inundated depressions
by species of sect. Cyphomandropsis

are
in clay or sandy soil frequented by S. glaucophyl
lum. These are found throughout the Chaco floristic province in south-central Bolivia,
Paraguay, north-central Argentina and adjacent areas, and in the Pampas floristic province
of Argentina (Cabrera 1976). Solanum glaucophyllum can form dense monospecific
thick
ets in isolated low-lying areas (called "curiches" in Bolivia and "duraznillares" in the Ar
gentine pampas) and on flooded or waterlogged
river banks and islands (Okada et al.
1977). This species is apparently deciduous during the winter dry season and can tolerate
frost in at least some parts of its range.
In contrast to species of sect. Pachyphylla, which range from Mexico
to Argentina,
all species of sect. Cyphomandropsis
are restricted to South America (Fig. 5). Only the or
namental species S. glaucophyllum has been taken to the Old World; cultivated specimens
have been recorded from various European botanic gardens as well as from two sites in
Asia. Solanum glaucophyllum was collected in Florida around 1900. It is thought that
these plants were adventive from seeds carried in ships' ballast (D'Arcy 1974). Whether
the Asian collections of S. glaucophyllum
also represent adventive plants is not known.
61
MONOGRAPHSVOLUME
BOTANY
SYSTEMATIC
18
Solanum section Cyphomandropsis
~~~~~~~~~~0~~~~~~~~
10
0~~~~~~~~~~~~~~~
0~~~~~~~~~~~~~~~~~~~~~~~~~1
0~~~~~~~~~~~~~~~~~~~~~~~~~~~2
~~~~~~~~b00~~~~~~~~~~~~~~~~~3
0
80
FIG.
included
70
5. Distribution
of Solanum
in the systematic
treatment.
*
60
50
This map
200 400 s00 600 1000k,
,00200 300 400 500S600mdOiS

0~~~~~~~~~~~~~~~~~~
40
is a composite
species
In comparison to the members of sect. Pachyphylla,
sis exhibit a pronounced tolerance of drier and cooler conditions.
this section extend further south than those of sect. Pachyphylla,
pass higher elevations. They exclude the more tropical parts of
of all the species maps
of sect. Cyphomandrop
The ranges of species of
and, in general, encom

South America, such as
the Amazon and Orinoco basins, Guayana highlands, Planalto de Mato Grosso in Brazil,
and the northern and western coasts of Colombia. Solanum glaucophyllum and, to a lesser
extent, S. stuckertii, are found in the seasonally dry Chaco and pampas regions from
is absent. The distribution of species by country is given in Ap

sect. Pachyphylla
the
Brazil
has
largest number of endemics (3), followed by Peru and Bolivia
pendix 5.
one
with
each.
which
SOLANUM
2001
In terms of the species groups outlined above,

distribution,
ranging
from southern Ecuador
19
the S. luteoalbum group is Andean
to approximately
32?S
in Argentina.
in
The
three species constituting this group are more or less allopatric and replace each other as
one moves from north to south. Hybrid plants have resulted from artificial crosses be
tween S. hibernum and S. stuckertii (see section "Breeding Systems and Crossing Stud
ies"). The ranges of these two species partially overlap in south-central Bolivia, but in
general they occupy different habitat types and occur at different elevations; thus, eco
differen
factors may prevent them from exchanging genes. Morphologically
tiated geographical variants are found in S. luteoalbum. Species and even populations
within this group are likely isolated by distance and by ecological factors, such as unfa
vorably hot and dry conditions in intervening Andean valleys.
Solanum amotapense and S. fallax both occur west of the Andes from Ecuador to
geographic
northern Peru; an isolated collection

Colombia.
Both
Jauneche
mesic
of S. fallax
is known
from the Cauca Valley
of
to seasonally arid sites, but S. fallax is found in the more

forest of northern and central Ecuador. Solanum amotapense has a dis
are adapted
junct distribution, and its range abuts that of S. fallax in southern Ecuador. It is possible,
however, that additional field work will expand the ranges of these little-known species.
Solanum hutchisonii is restricted to a small area of the Rio Marani6n valley in north
ern Peru, where
it is sympatric or nearly so with S. amotapense. Altitudinal differences

appear to separate the two species; S. hutchisonii occurs below 600 m and S. amotapense
above 700 m in elevation.
of the S. glaucophyllum
group are almost completely allopatric, and, where

their ranges approach each other, are separated by obvious ecological differences. This is
most pronounced in the case of S. glaucophyllum
and S. confusum, which occupy totally
Species
different habitats, even in areas where their ranges abut. Solanum glaucophyllum
is found
in seasonally inundated depressions in Chaco forest, whereas S. confusum occurs in the
cloud forest understory. Both S. matadori and S. luridifuscescens are native to southeast
ern Brazil, but the two species
differences. Solanum matadori
Catarina
are separated by geographic, ecological, and elevational

restricted to mid-elevation
Araucaria
forests of Santa
state, and S. luridifuscescens
found at higher elevations and more northerly lati
tudes in the Atlantic
coastal rain forest.
Likewise, S. cylindricum and S. pelagicum are species of southeastern Brazil and ad

jacent areas. Their known ranges barely overlap, however, and they are likely separated
by ecological preferences; S. pelagicum is restricted to low elevation coastal areas, and S.
cylindricum occurs inAraucaria groves inmore upland areas. The range of S. cylindricum
extends
into Prov. Misiones,
another species of Araucaria
Argentina, where
it overlaps with
the enigmatic S. fusiforme,
forest.
POLLINATION
The function of the anther connective
or dorsal anther surface in providing rewards

needs further study in sections Pachyphylla and Cyphomandropsis.
Inmost
members of sect. Pachyphylla
that have been investigated, the swollen anther connective
acts as an osmophore to secrete volatile compounds that are gathered by male euglossine
for pollinators
bees
(Gracie 1993; Sazima
Pachyphylla
et al. 1993). Although
may primarily be accomplished
this indicates
by passive
gathering male bees, the only taxa that have been well
that pollination
pollen deposition
in sect.
on fragrance
studied are those with
large anther
20
It remains
connectives.
VOLUME 61
to be determined what
role female buzz-pollinating

bees play in
At least one member of sect. Pachyphylla,
S. pinetorum, is not visited by male euglossines but is buzzed by several genera of female
bees (Sazima et al. 1993). Similarly, it is not known whether species of sect. Cyphoman
the reproductive biology
of sect. Pachyphylla.
those with elaborated anther surfaces, also produce volatile compounds

has been studied in only two species of sect. Cyphoman
S.
S.
dropsis,
glaucophyllum and
stuckertii, and they were found to be buzz-pollinated by
dropsis, especially
that attract male bees. Pollination
are needed to document

(L. Passarelli, pers. comm.). Field observations
flower visitors and to distinguish between two rather different pollination syndromes that
might occur in the section (buzz-pollination by female pollen-collecting
bees vs. fragrance
gathering by male euglossine bees).
female bees
HERBIVORY
Little is known about herbivory in sect. Cyphomandropsis.
Okada et al. (1977) note
that the caterpillars of Antarctia fusca Eslker burrow in the pith of S. glaucophyllum and
cause damage to the aerial stems. They have also been seen on the leaves, but it is not
known if they feed on them. Aside
of herbivores
on species
of
from this account,
the section;
there are only two published
both are ithomiine
butterflies
reports
in the genus
Mechanitis
[M. polymnia casabranca Haensch and M. lysimnia lysimnia (Fabricius)] that

use S. luridifuscescens as a larval host plant in Brazil (Drummond & Brown 1987; species
cited as Cyphomandra
of Solanaceae, where
array of phytochemicals
velutina Sendtn.). These butterflies are specialists

they may discriminate
1986,
1991). Although
to locate and identify potential
it is hoped that the present taxonomic

stimulate observations
herbivores.
they may
solanaceous
kaloid sources, such as species of Boraginaceae

tasteful to predators. This extremely interesting
will
oviposition
produced by the plants (see Brown
1987; Vasconcellos-Neto
pounds
in choosing
on many
1987; Drummond
use
host plants,
and Asteraceae,
species
sites among a rich

& Brown
these secondary
com
they rely upon other al

to render the adults dis
interrelationship needs further study, and

treatment, by facilitating identification of the plants,
of use of Cyphomandropsis
species by ithomiines and other
in turn provide insight into the phytochemistry
and evolu
Such studies may
tionary ecology
of the PachyphyllalCyphomandropsis
group.
USES
Unlike taxa in the related sect. Pachyphylla,
few uses have been reported for species
of sect. Cyphomandropsis.
The fruits are small and unpalatable to humans, and there are
no instances of their being used as food. Solanum glaucophyllum
is the only species in the
group with any recorded uses by people: it has been employed medicinally
as a purgative,
and the stems have been used as firewood and building materials (see notes under species
treatments
weigh
in section
its usefulness,
stock poisoning
in South America
increased calcification
investigated
"Taxonomy").
Yet,
for S. glaucophyllum
the deleterious
effects of this species
has been responsible
(Okada et al. 1977). Because
for many
cases of live
its toxicity
of soft tissues after ingestion, S. glaucophyllum

as a source of bone growth stimulants useful in medicine
far out
is due to
is currently being
(Morris 1977; B.
Barr, pers. comm.).
21
SOLANUM
2001
TAXONOMY
Solanum L., Sp. pl. 1: 184. 1753.-TYPE:
See D'Arcy (1972, 1973) for generic
Solanum nigrum L.
synonymy.
Herbs, shrubs, trees, or vines, with or without spines, glabrous or pubescent with un
branched or branched, often glandular hairs. Leaves alternate or paired and frequently un
equal in size, simple to pinnately lobed or compound, petiolate or sessile, without stipules.
Inflorescences cymose, branched or unbranched. Flowers usually perfect, (4-) 5-merous,
actinomorphic or zygomorphic; calyx campanulate, sometimes accrescent in fruit; corolla
rotate, campanulate, stellate, or urceolate, white, green, yellow, pink, or purple; stamens
equal or unequal, the filaments generally short and inserted at the corolla base, the anthers
basifixed, blunt or tapered toward apex, opening by terminal pores (sometimes expanding
into longitudinal slits); ovary 2-carpellate; ovules many; style articulated at base, usually
slender; stigma capitate. Fruit a berry, usually fleshy but occasionally dry, usually many
seeded, the seeds often flattened; embryo curved, embedded in abundant endosperm.
Chromosome number: n = 12, 23.
Bitter, Repert. Spec. Nov. Regni Veg. 12: 461. 1913.
(Bitter) D'Arcy, Ann. Missouri Bot.
Cyphomandra
Gard. 59: 277. 1972.-LECTOTYPE, designated by Seithe, 1962: Solanum stuck
ertii Bitter.
section Cornigera Child, Repert. Spec. Nov. Regni Veg. 95: 293.
Cyphomandra
Solanum
1984.-TYPE:
Solanum
Cyphomandra
section Glaucophyllum
cornigera Dunal
Child,
Repert.
[=Solanum pelagicum Bohs].

Spec. Nov. Regni Veg. 97: 144.
1986.-TYPE: SolanumglaucophyllumDesfontaines.
Herbs, shrubs, or small trees, sometimes rhizomatous, lacking spines. Stems glabrous
to densely pubescent with unbranched glandular, unbranched eglandular, and/or dendriti
cally branched eglandular hairs. Leaves 3 to many per sympodial unit, the blades charta
ceous to subcoriaceous or succulent, simple and elliptic to ovate or pinnately compound,
acute to acuminate at apex, cuneate to decurrent to deeply cordate at base, glabrous to
densely pubescent, the petioles glabrous to densely pubescent. Inflorescence unbranched
or forked to highly branched; pedicels articulated at or above the base, leaving scars or
pedicellar remnants; inflorescence glabrous to densely pubescent. Flowers perfect, actin
omorphic,
5-merous. Calyx
sometimes
bescent,
tuse to acuminate
campanulate,
inflated below
tips. Corolla
not accrescent
in fruit, glabrous
to densely pu
lobes, the lobes deltate to narrowly triangular with ob

to
white, pink, purple, or bluish, membranaceous
the lobes narrowly trian

stellate, rotate-stellate, or stellate-campanulate,
subcoriaceous,
gular to ovate, subacute to apiculate at apex, glabrous to densely pubescent abaxially with
pubescent adaxially. Stamens
branched or unbranched hairs, glabrous to moderately
equal, filaments very short, glabrous, inserted in corolla tube near its base; anthers con
nivent or free, yellow to greenish, reddish, or purplish, ovate to narrowly triangular, ta
adaxially, not (or rarely)
pered toward apex, the pores directed distally and occasionally
opening into longitudinal slits. Ovary glabrous to densely puberulent; style glabrous to
moderately puberulent, cylindrical to subclavate; stigma truncate to subcapitate, the same
diameter as the style. Fruits globose, ellipsoidal, ovoid, or fusiform, obtuse, acute, or apic
ulate at apex, glabrous to densely puberulent, yellow, orange, red, brownish, or blue-black
22
when
ripe; stone cell aggregates
present or absent. Seeds
strongly flattened, smooth to felty-pubescent.
Chromosome
VOLUME 61
thick and angled to lenticular or

number: n = 12.
SYNOPTIc LIST OF CHARACTERS OF SOLANUM SECTION CYPHOMANDROPSIS

The following
is a list of distinctive
Numbers
Cyphomandropsis.
ment. Numbers
character states found among species of section

to the species number in the taxonomic treat
indicate that the character state is uncommon or insuffi
correspond
in parentheses
ciently studied in that species.
Inmost cases, diagnostic character states are listed, but the

are not. This is not intended to be an exhaustive list of characters, but an aid
to identification of distinctive taxa.
alternatives
Plants found in seaside habitats of southeastern Brazil:
12
Plants glabrous or nearly so: 2, 3, 5, 6, 8, 9, 11

Plants pubescent: 1, 2, 3, 4, (6), 7, 9, 10, 12, 13
Plants with branched hairs: 1, 3, 7, 10, 12
Plants glaucous:
Leaf surfaces strongly discolorous:

Pinnately
Leaf bases cordate or subcordate:
than two rachises: 2, 4, 6, (10), 11, (13)

than 1mm long: 4
articulated above the base, leaving remnants more

at base of lobes: 1, 4
2
tube swollen
Calyx
1, 4, 7, 8, 12
branched, with more
Inflorescences
Pedicels
leaves present: 5, 12
compound
Corolla
stellate-campanulate:
Corolla
rotate-stellate:
1, 6
4, (12)
Fruits pubescent:
Fruits fusiform, much
longer than wide:
Fruits dark blue or black when

Seeds very large and angled:
ripe: 6
1, 2, 4, 6, 7, 8, 10, 13
KEY TO THE SPECIES OF SOLANUM SECTION CYPHOMANDROPSIS

1. Plants
or nearly
glabrous
2. Plants
so; if pubescent,
the hairs sparse and/or minute
and only visible
under high
(lOx or more).
magnification
often with
some pinnately
compound
leaves only;
fruits globose.
as well
as simple
leaves;
fruits elongate-fusiform.
5. S. fusiforme.
2. Plants with
simple
3. Leaf bases
rounded
to subcordate;
3. Leaf
cuneate,
truncate, or decurrent;
leaf blades
somewhat
thick and fleshy; northern Peru.

8. S. hutchisonii.
bases
gentina,
4.
Paraguay,
Leaves
Uruguay,
usually
4.
Leaves
2-6 mm
green,
and southeastern
corollas
glaucous;
lobes 5-10 mm wide;
not glaucous;
rotate-stellate
corollas
surface of anthers
smooth
to fleshy; Bolivia,
stellate
and plicate,
the tube 5-8 mm
and glaucous.
or stellate-campanulate,
ripe fruits yellow

to roughened,
long,
the
6. S. glaucophyllum.
not plicate,
the tube
or orange.
but not obviously
papillate;
and northwesternArgentina.
5. Abaxial
Ar
Brazil.
ripe fruits dark blue-black
long, the lobes 2-5 mm wide;
5. Abaxial
leaf blades membranaceous
Bolivia
2. S. confusum.
surface of anthers with band of scaly papillae
(these most
obvious
in dried ma
terial);southeasternBrazil.
6. Adaxial
ormore.
corolla
surfaces
pubescent;
petioles
glabrous;
inflorescence
branches
three
11. S.matadori.
SOLANUM
2001
23
6. Adaxial corolla surfacesglabrous or nearly so; petioles pubescent; inflorescences

unbranched
or at most
7. Abaxial
surface of anthers with
forked.
a dense band of scaly papillae;
7. Abaxial
1. Plants obviously
8. Pedicels
bases
deeply
8. Pedicels
hairs frequently
above
cordate
the base,
leaving
at or very near the base;
articulated
fine or inconspicuous
very
papillae;
cm wide.
0.5-3.5
3. S. cylindricum.
magnification.
remnants more
truncate
(rarely shallowly
with
9. S. luridifuscescens.
leaf blades
present;
the hairs visible without
pubescent,
articulated
at most
of anthers
surface
branched
if present,
hairs,
unbranched;leaf blades 2-7 (-9) cmwide.
than 1mm
long; fruits pubescent;
leaf
to subcordate).
4. S. fallax.
leaf bases
cuneate
or at most
ratio generally
more
than 4.5:1.
fruits glabrous;
shal
lowly cordate.
9. Vegetative
pubescence
of unbranched
10. Leaf blades
glaucous,
10. Leaf blades
green, not glaucous,
hairs only.
the length:width
very narrow,
6. S. glaucophyllum.
11. Calyx
length:width
less than 4.5:1.
ratio usually
tube inflated at base of lobes; corollas
the tube 5-7 mm
rotate-stellate,
long; leaf
bases often subcordate,rarelycuneate.

11. Calyx
tube not noticably
(-6) mm
swollen;
long; leaf bases
corollas
cuneate,
or decurrent.
rounded,
an obvious
the tube 2-4
to stellate-campanulate,
stellate
truncate,
surface of anthers with
12. Abaxial
1. S. amotapense.
band of scaly papillae;
southeastern
Brazil.
9. S. luridifuscescens.
surface of anthers
12. Abaxial
or roughened,
smooth
but not obviously
scaly-papillose;
Andean Ecuador,Peru,Bolivia, and northwesternArgentina.

13. Pubescence
to dense, with many
sparse
1mm
of the hairs
long or more;
las stellate to stellate-campanulate.

13. Pubescence
to dense,
moderate
corol
2. S. confusum.
less than 1mm
the hairs generally
long; corol
las stellate.
14.Corollaswhite; south-centralBolivia to northwesternArgentina.

13. S. stuckertii.
14.Corollas purple (rarelywhite); southernEcuador to southernPeru.
10. S. luteoalbum.
9. Vegetative
pubescence
at least partly of branched
15. Calyx
tube inflated at base of lobes; corollas
15. Calyx
tube not inflated; corollas
hairs.
the tube 5-7 mm
rotate-stellate,
long.
1. S. amotapense.
stellate,
the tube 2-4 mm
long.
16.Leaves strongly discolorous, the adaxial surface green with sparse pubescence,
the abaxial surfacedenselywhite-pubescentwith branchedhairs; centralBolivia.
7. S. hibe mum.
16.Leaves not stronglydiscolorous, green on both sides or with pubescence evenly dis
tributedthroughout;Ecuador,Peru,Argentina, and southeasternBrazil.
17. Leaves
often pinnately
date; seaside
17. Leaves
compound
habitats
simple or with
and upland Ecuador,
or lobed; if simple,
in Santa Catarina,
1-2
small basal
Peru, Argentina,
the bases
truncate
Brazil.
lobes,
to subcor
12. S. pelagicum.
the bases
cuneate
and southeastern
to decurrent;
inland
Brazil
18. Peduncles0.5-3.5 cm long; inflorescenceunbranchedor forked,1-15-flowered;

southeasternBrazil and adjacentArgentina.
3. S. cylindricum.
18. Peduncles 3.5-6.5 cm long; inflorescence unbranched, forked, or further
branched,
1. Solanum
amotapense
tapensis
(Svenson)
(5-)
10-25-flowered;
Svenson,
Amer.
A. Child
TYPE: PERU. Piura: Amotape

4045'S,
2300
ft, 28-30 Mar
Andean
J. Bot.
ex Bohs,
Hills,
Ecuador
and Peru.
33: 483.
1946. Cyphomandra
Fl. Neotrop. Monogr.

near summit
1941, Haught
10. S. luteoalbum.
of Cerro Prieto,
& Svenson
11634
amo
63: 153. 1994.

81?15'W,
(holotype: BKL!;
isotype: US! #1832594).
24
VOLUME 61
villosa Steyermark, Phytologia 9: 348. 1964.-TYPE:

ECUADOR. Loja:
Las Chinchas, 2250 m, 12 Apr 1944, Acosta Soli's 7743 (holotype: F!; photo of
holotype [F neg #51366]: F!).
Cyphomandra
Shrub 0.5-2 m tall. Stems densely pubescent with unbranched eglandular hairs and
often also some short-stalked glands (dendritically branched hairs occasionally present).
Leaves 7-many per sympodial unit, the blades 4-23 cm long, 2-14 cm wide, length:width
ratio ca. 1.5-2.5:1, simple, ovate to elliptic-ovate, acuminate at apex, cuneate to subcordate
at base, chartaceous, nearly glabrous to densely pubescent adaxially with hairs like those
of the stem, moderately pubescent to densely so on veins and abaxial surface, the petioles
1-8 cm long, densely pubescent.
2.5-14
Inflorescence unbranched or forked, ca. 6-30-flowered,

cm long; rachis 1.5-9 cm long; pedicels 5-15 mm long,
long in fruit, spaced 1-25 mm apart, articulated at the base; inflorescence axes
to densely pubescent. Calyx moderately pubescent with unbranched eglandu
cm long; peduncle
10-15 mm
moderately
1-6.5
lar hairs and stalked glands, constricted at apex of inflated tube, the radius 2.5-5 mm, the
lobes 1-3 mm long, 1-1.5 mm wide, deltate to triangular-subulate, acute at tips. Corolla
white or purple, chartaceous tomembranaceous,
rotate-stellate, the radius 10-25 mm, the
tube 5-7 mm long, the lobes 6-13 mm long, 4-12 mm wide at base, triangular-ovate, acute
at apex, sparsely tomoderately puberulent abaxially, especially near tips of lobes, glabrous
adaxially. Anthers connivent or not, yellow or greenish yellow, ovate, 4-6 mm long, 2-2.5
mm wide, abaxial surface smooth to roughened but not obviously papillate, the pores di
rected distally. Ovary glabrous; style glabrous (moderately pubescent in Acosta Solis
7743), cylindrical, 6-10 mm long, 0.5 mm in diameter; stigma truncate. Fruits 1.5-2 cm
long, 1.5-2 cm in diameter, globose, obtuse at apex, glabrous, pale orange, red, or brown
ish when ripe; stone cell aggregates absent. Seeds 4-5 mm long, 4 mm wide, angled,
smooth to rugose. Chromosome number: 2n = 24. Figs. 6, 7.
and fruiting in January through April and in November.
(Fig. 8). Southwestern Ecuador and northwestern Peru; cliff edges, dry
and rocky stream beds, and under deciduous vegetation, primarily in seasonally and areas;
Phenology.
Flowering
Distribution
600-2300 m.
Local name. Ecuador: Sabalucu
(Acosta Solis 7743).
ADDITIONAL SPECIMENS EXAMINED. Ecuador.

Harling
& Andersson
Stdihl 26473
Peru.
(NY).
127 E of Olmos,
Sdnchez
9614
(MO, NY);
hwy between
Lbpez
Olmos
Prov. Contumaza,
between
Prov. Tumbes,
at Royal
Gardens,
Botanic
Empalme,
Km
(on the Rio Huancabamba),

7894
Chilete
mountains
Edinburgh,
ca. 12 km from Celica,
Toldo-Chaco,
and Jaen, Hutchison
& Sagdstegui
(NY).-TuMBES:
Cult.
LOJA: road Celica-El
road Cariamanga-Yambaca-El
CAJAMARCA: Prov. Jaen, Pucara
near San Benito,
Vega 4220
Cultivated.
(GB, NY);
Mesones-Muro
Prov. Contumazd,
Sagdstegui
22510
& Wright
3548
(MO); Prov. Cajamarca,

and El Rupe, W
E of Hacienda
Scotland,
10-20,
vicinity
Harling
&
of town, Km
(F,MO,
NY, US);
Chilete-Magdalena,
of road Chilete-Contumaza,
Chicama,
Child C9469
Weberbauer
7634
(F).
(E, G).
Solanum amotapense can be distinguished from the other species in the section by its
inflated calyx tube, rotate-stellate corolla, long slender filaments, and subcordate leaf
bases. The only other species of sect. Cyphomandropsis
occurring in arid areas of north
ern Peru is S. hutchisonii. Solanum amotapense differs from it in its abundant pubescence
and chartaceous rather than succulent leaf blades.
All
specimens of S. amotapense have unbranched eglandular hairs on the stems,

leaves, and inflorescences, except the type collections of S. amotapense and C. villosa,
which have a few dendritically branched hairs, especially on the leaf margins. Corolla
2001
SOLANUM
FIG. 6. Solanum
amotapense.
flower. D. Gynoecium.
E. Stamen
Child C9469;
B-E,
greenhouse
A. Habit.
B. Flower,
(left to right: abaxial,
material
of Bohs
25
seen from above. C. Lateral

lateral, and adaxial
views).
view of partially
F. Fruit.
(Based
2479.)
sectioned
on: A, F,
26
VOLUME 61
k;~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..
.. ...
'l
......... I...L
FIG. 7. Flowers
of S. amotapense.
Scale
bar = 1 cm.
color varies from white to light blue or purple in this species,

rather than glabrous in the type of C. villosa.
and the style is puberulent
2. Solanum confusum C. V. Morton, Contr. U.S. Natl. Herb. 29: 70. 1944.-TYPE: Bo
LIVIA. Cochabamba: Mount Tunari, near snow line, 1891, Bang 1118 (holotype:
isotypes: C! F! G! L! LD! M! NY! WU! Z!).
US! #1177847;
Solanum adelphum C. V. Morton, A Revision of the Argentine Species of Solanum,
185. 1976. Cyphomandra adelpha (C. V. Morton) A. Child ex Bohs, Fl. Neotrop.
ARGENTINA. Tucuma6n: Depto. Burruyacu,
Monogr.
63: 153. 1994.-TYPE:
Cerro del Campo, 1800 m, 14 Dec 1928, Venturi 7732 (holotype: US! #1441095;
photos of holotype: GH! NY!; isotypes: GH! S! SI! US).
with glandular and
Shrub 1-3.5 m tall. Stems glabrous to densely pubescent-pilose
eglandular hairs. Leaves 4-5 per sympodial unit, the blades 5-19 cm long, 1.5-6.5 (-8)
cm wide, length:width ratio ca. 2-4.5: 1, simple, elliptic, acute or acuminate at apex,
cuneate or decurrent at base, chartaceous to subcoriaceous, nearly glabrous tomoderately
adaxially and abaxially, the petioles 0.5-4 cm long, glabrous tomoder
pubescent-pilose
denser in adaxial channel. Inflorescence un
with pubescence
ately pubescent-pilose,
3-20 cm
branched to twice forked or further branched, 5-30 (-40 or more)-flowered,
long; peduncle 1.5-9 cm long; rachis 1-10 cm long; pedicels 5-30 mm long, 15-35 mm
long in fruit, expanded distally below calyx, spaced 1-14 (-40) mm apart, articulated at
or near base, leaving scars or short pegs up to 1mm long; inflorescence axes glabrous to
with glandular and eglandular hairs. Calyx nearly glabrous to
densely pubescent-pilose
densely pubescent, the radius 2-8 mm, the lobes 1-7 mm long, 1-2 mm wide, triangular
deltate, often narrowed into acuminate tips. Corolla white to pink or violet, chartaceous to
the radius 8-15 mm, the tube 2-4 mm
stellate or stellate-campanulate,
membranaceous,
long, the lobes 6-12 mm long, 2-5 mm wide at base, narrowly triangular to ovate, acute
27
SOLANUM
2001
A S. amotapense
7S. fallax
10
*S.
0
200
100 200 300 400 500 600 miles l\
FIG.
400
hutchisonii
8. Distribution
600
800
1OO0km
of S. amotapense,
S. fallax,
and S. hutchisonii.
or subacute at apex, nearly glabrous to densely pubescent abaxially, nearly glabrous adax
ially except on midribs and tips of lobes. Anthers connivent or not, greenish white to pur
plish, yellowish, or reddish brown, ovate to lanceolate, 5-9 mm long, 1.5-3.5 mm wide,
glabrous, abaxial surface smooth to roughened but not obviously papillate, the pores di
rected distally. Ovary glabrous; style glabrous or sparsely puberulent, cylindrical, 6-10
mm long, ca. 0.5-1 mm in diameter; stigma truncate. Fruits ca. 0.8-2 cm long, 0.8-2 cm
in diameter, globose, obtuse at apex, glabrous, yellow to orange when ripe; stone cell ag
gregates present or absent. Seeds 4-6 mm long, 3-5 mm wide, angled, glabrate and ru
gose or pubescent with dense white pseudohairs, especially along margin. Chromosome
number: n = 12. Figs. 9, 10.
Phenology. Flowering in all months except July and August; fruiting in all months ex
cept July, August, and September.
Distribution
(Fig. 11). Bolivia and northwestern Argentina; cloud forest, open areas,
or secondary vegetation, often on slopes and in groves of aliso (Alnus acuminata); ca.
900-4000 m.
28
VOLUME 61
)~~~~~~~~~~~
FIG. 9. Solanum
15442;
B-F,
Sleumer
confusum.
E. Stamen
A. Habit.
B. Flower,
seen from above.

lateral, adaxial
C. Lateral
views).
view
F. Fruit.
of partially
(Based
3056.)
sectioned
on: A, Meyer
2001
SOLANUM
FIG.
10. Inflorescence
of S. confusum
29
(Yungas de Mairana,
Nov.
Florida, Dept.
Santa Cruz, Bolivia).
Scale
I cm.
bar
Local name. Bolivia:
Tanta sacha (Beck & Seidel
REPREsENTATIVE SPECIMEiNs. Bolivia.

prox. 200 m
al E del campamento
rov. Sud Cinti,
CHUQUISACA: Nrov. Sud Cinti,
Rinconada
entre campamento
camino
14478).
del Bufete,
Rinconada
del Bufete
base oriental
64o22P28*W
20_4949PS,
y la cumbre
del Cerro Bufete,

et al. 931
Arroyo
ap
(USZ);
en al lado N del
del Cerro Bufete,
Serrano et al. 1299 (USZ); Nrov. Sud Cinti, approx. 500 m al N del cam
64022'28"W,
transecto 4-B-13,
del Bufete,
et al. 1385 (USZ).
Serrano
20'49149S,
pamento Rinconada
64022'28"W,
ca. 10 km al NW de Independencia,
COCHABAMBA: Nrov. Apopaya,
Beck & Seidel 14478 (LPB); Tako Tako,
farm near Miske,
18'000#S, 65'15'W, Brooke 5947 (BM, F, NY); Nrov. Chapare,
Incachaca, small power station
Cerro Bufete,
20o49~49"S,
about 80 mi NE
3260
denas
Corani,
of Cochabamba,
Cdrdenas
(US); Siberia,
5936
Km
(K, US);
l7o00'S,
5741
65'30'W,
Brooke
6806
(US); above Pojo, Cdrdenas
Hensen
Corn. Lachujmayu,
Nrov. Carrasco,
61.4 Cochabamba-Chapare
road, Kessler
(K, US); Nrov. Carrasco,

936
& Kelschebach
canyon
of Rio Monte
& Solomon
Steinbach
36628
8660
romarca, Beck
Puncu,
5 km NE of Monte
(NY); Nov.
fall above Circuata

km S of Choquetanga,
down
Cant6n
from and NE
Plowman
on old road, 16019'5,
16'47'S,
1605415, 67'17'W,
67053'W,
Lewis
16016'S,
1603515,
& Davis
et al. 36494
Kehuifial,
of Comarapa,
(by air) ESE
17049PS,
of Siberia,
8.5 km
(LPB, UT); Nrov. Carrasco,
narrow
of Epizana,
Sarav(a
5150
Solomon
67'17'W,
38954
67044'W,
8694
Dorr
Lewis
17'33'S,
& L6pez
Inquisivi,
(LPB); Nrov.
et al. 6932
38693
65'16'W,
Nee
11 71 (USZ); Pocona,
Pizarro
91
unos 8 km de Quime
Inquisivi,
30226
9km
Inquisivi,
Pata, 7
by road (ca. 4km
by air)
Sud Yungas,
road from Unduavi
to
3.1 km SE of Unduavi
(LPB, NY); Nrov. Nor Yungas,
Rio
Tambo
Inquisivi,
(LPB, NY); Nrov.
(LPB); Nrov. Nor Yungas,
Nrov. Sud Yungas,
hacia
first large water
(LPB, UT); Nrov.
(LPB); Nrov.
(LPB); Nrov. Nor Yungas,

Nee & Solomon
67047'W,
(GH, K, MO);
(LPB); Nrov.
Beck 24352
on road to Inquisivi,
67'15'W)
of Chuspipata,
Beck 21830
16058#S, 67'12'W,
of Choquetanga,
Lamnbate, Chillkni,
Sacramento,
16'11I'S, 67043'W,
(16.385,
7 km NNE
7 km
Nor Yungas,
arriba del pueblo,
Inquisivi, Camillaya,
(by road) NW
"Churro,"
PAZ: Nrov. Sud Yungas, debajo de Unduavi,

subiendo al valle de Cer
ca. 4.5 km al S de Coroico, Beck 17219 (NY);
rov. Nor Yun
(BM, GH, K, NY).-LA

14665
gas, ca. 2 km al S de Coroico,
Pavimani,
comunidad
Centr. Hidroel.
Chapare,
canmino K'uri
(NY); Nrov. Mizque,
on road from Comarapa
to Cochabamba,
4
10kmu (by air) NW
Puncu,
(LPB, USZ, UT); Nrov. Mizque,
(NY); Nrov.
233
230S, L6pez & Sarav(a 427 (USZ); Nrov. Carrasco,

K'asa-Lagunas,
km W of border with Depto.
Santa Cruz, 20 kmn (by air) and 28 km
(LPB, NY); Nrov. Carrasco,
64041'W, Nee & Solomon 34038
SW of limit with Depto. Santa Cruz, 17'50'S, 64042'W Nee
Sailapata, Cdrdenas
Jattin Pino, Cdr
(BM); Nrov. Ayopaya,
5773
bridge
7.8 km SE (above) Yolosa
(below)
on road
VOLUME 61
30
20~~~~~~~
"S
.~~~~~~~~~~~........
..........
10
0030
40
50.0
'
mls
30
...
1~~~~~~~~~~~~~
FS.confusumc2fu
100
600
400
200
400
300
200
800
1000Okm
6100 miles
500
80
7060
1 1. Distribution
FIG.
to San Juan de La Miel,
16016'S,
just S of Parque Nacional
ballero,
64032'W, Abbott & Jardim 17232

bra 2619
Cerro Bravo
(USZ); Prov. Caballero,
(LPB); Prov. Florida, Valle de "El Fuerte,"
Prov. Vallegrande,
air ESE of Vallegrande,

"Mataralcito"
18032'30"S,
18?32'30"S,
and "El Palmar"

63057'30"W,
reaches of Quebrada
Nee
63?57'30"W,
et al. 37409
Agua Blanca,
18?01'S,
Los
Sitanos,
Sitanos,
38546
UT);
Nee
38389
(LPB, USZ, UT);
18?47'S,
64?02'W,
Nee
1 km N of turnoff to Sitanos
(LPB, NY, UT);
Parque Nacional
Prov. Florida,
Ambor6,
SW
38427
map
ca. 65?50'W,
km N of Comarapa,
(NY); Prov. Florida,
(LPB, USZ,
Aguadita,
UT);
7 km (by air) NE of Mairana,

region of park, N of Comarapa,
(USZ);
17 km by
Prov. Vallegrande,
between
17 km by air ESE of Vallegrande,

top of ridge at upper
(LPB, USZ, UT); Prov. Vallegrande,

and 5 km N of Khasa Monte,
8 km (by air) S of Khasa Monte
Prov. Vallegrande,
on road from Abra Tabla
ca. 17?51'S,
to Tierras Nuevas,
5 km SW of Yerba Buena,
64?03'W, Nee & Vargas 38292

ca. 1km N of Lagunillas
of
de Tablar, Coim
Ibisch et al. xx.efl
18?12'S,
(LPB, NY, UT);
to Tierras Nuevas,
10 km NE
CRUZ: Prov. Ca
San Juan del Potrero, Yungas
et al. 37398
Prov. Vallegrande,
Altos
area, 4-10
Prov. Nor Yungas,
(LPB, NY).-SANTA
on road from Vallegrande
on road from Vallegrande
ca. 6 km S of Abra Tablas on road to Los Sitanos,

64?02'W,
Nee
12566
Cant6n
Samaipata,
and El Palmar
between Mataralcito
(K, LPB, NY);
Solomon
67?47'W,
16?16'S,
Ambor6,
9347
Solomon
67043'W,
on road to Yolosa,
(below) Chuspipata
of S. con.fusum.
5 km
to Los Sitanos,
18?4'S,
63055'W,
(by air) NW
18'50'30"'S,
Nee
18?39'S,
on road to
40633
in tree plot of I. Vargas
C.,
of Los
6400'W, Nee
(LPB, USZ,
17?49'45"S,
SOLANUM
2001
& Nee
MO, NY);
17969
17?52'S,
Ambor6,
La Yunga,
I. Vargas
trayecto
18?04'S,
entre Chape
del Cerro Bravo
proximadades
y Peniones,
10 km al E de la ciudad de Vallegrande,
I. Vargas
y el Corral
Chape
I. Vargas
3714,
Prov. Arce,
Rancho
km al NE
on trail between
2618
Sidras
(NY, US).
Esquina
tenna, Ahumada
5316
(SI); Depto.
to Valle Grande,
4 km before Abra
contra
el valle
(CTES).-SALTA:
Burkart
13219
brada de San Lorenzo,

del pueblo, Marmol
8773
(MCNS);
Depto.
Chaile,
19508
13658
15442
Depto.
Olea
& Hjerting
565
(C, MO,
los Sosa, Km 39 to Tucuman

galito,
Slanis
W); Depto.
del Valle,
SI); Depto.
et al. S124
NY);
Depto.
3067
Chicligasta,
3056
11433
Quebrada
Las Pavas,
del Clav
Depto.
Chicligasta,
LIL);
(C, MO);
Depto.
Chicli
(F, GH, NY); Quebrada
(CTES); Depto.
de Cainzo,
Burruyacu,
Sleumer
Valle
Venturi 4589
between
70 (F, GH,
Taft, Parque Aconquija,
618
35090
24871
Santa Rosa, Meyer
O'Donell
& Hjerting
& O'Donell
Schulz
de los
s.n. (W);
of Cuesta
Puesto
(L); Depto.
Cochuna,
s.n. (BM, LIL, NY);

Petersen
Guachipas,
& Vervoorst
before beginning
16374
Meyer
Depto.
36 (Ruta prov. 307), Hunziker
(G, LIL); Depto. Monteros,
Estancia
Grassi
et al.
Vaqueros,
road to Comiza,
Caldera,
Chicligasta,
Chicligasta,
Taft, Parque Aconquija,
Venturi 5688
slightly
(GH), Schreiter
on Ruta 307 to Taft del Valle,
(G); Depto.
Taft, La Hoyada,
Taft, Raco,
10393
Taft, Puerta de San Javier, Sleumer

Sleumer
to Nogalar,
La Caldera,
269 (LIL); Quebrada
Bailetti
road
Que
3-4 km alW
Chaile,
155 (MCNS);
de Cainzo,
(A, LIL, NY); Depto.
Rio Loro, Olea
Schreiter
(LIL); Depto.
Burruyacu,
Taft, Quebrada
(W); Depto.
18911
89 (A, NY);
gasta, bridge over Rio Cochuna,
Depto.
road to Tafi del Valle, Km
San Pablo, Lillo 4543

Jaya, Meyer
(G, NY); Depto.

Varela
Capital,
a Porongal, Marmol
(CTES); Depto.
Tilian,
station of Rio Cochuna,
(B); road from Tafi del Valle
and Aser.
Taft, Siamb6n,
Petersen
LP); Depto.
(UT); Depto. Monteros,
15445
(B, W),
(CTES,
et al. 5567
Chicoana,
944
La Hoyada,
Santa Victoria,
(CTES); Depto.
de Baritu
de San
Rotman
Moldes,
del Arroyo
cuenca
camino
et al. 20370
of town, Novara
(LIL); Depto.
(A, BM, GH, MO).-TucuMAN:
(UT); Sierra San Javier, Alto

Pte. Santa Rosa
km W
3-4
Vaqueros,
9938
road
Ledesma,
de Perico
to Centinela,
36 (CORD); Depto.
& Cuezzo
Santa Victoria,
5 km alW
(CTES); Coronel
SI);
road to the an
a la antena, Cabr
camino
ascent
below
(GH, MO,
(SI); Depto.
(LIL); Ruta
Cocucci
Legname
Los Toldos, Meyer
in front of the gauging
Chicligasta,
y Solazati,
1472
Cerro Zapla,
et al. 8295
1571
entre
63?53'W,
18?04'S,
(LPB, NY).-Without
Jbrgensen
Santa Barbara,
Depto.
& Malmierca
La Caldera,
(MCNS); Depto.
et al. 11849C
Cabrera
illo, Hunziker
5567
Santa Victoria,
Schiavone
Sosa, Alisales,
Depto.
Lotti 86 (CTES); Depto.
Venturi 9837
Alemania,
(MCNS);
18032.5'S,
trayecto
(LPB, USZ).-TARIJA:
11274
Capital,
El Palmar
of Rio Chillaguatas,
9 de Octubre,
8220
& Cuezzo
ca. 15 km from Los Toldos,
& Niifiez
of Arroyo
streambed
El Lapachar,
247
El Rey, Brown
Oran, entre Rio Pescado
to Lipeo,
Grande,
Depto.
cerca de El Fuerte, Kiesling
de Canias, Legname
del rio, Novara
(SI); Depto.
(G).-JUJUY:
8672
valley
Solomon
64?25'W,
Sierra de Zapla, Mina
Capital,
Santa Barbara,
Parque Nacional
from Los Toldos
10788
Jorgensen
(CTES); Depto.
22?5'S,
CATAMARCA: Esquina
Argentina.
Grande,
era et al. 32059

Antonio,
and Tariquia,
San Rafael),
Park), Wood
(Ambor6
17?49.5'S,
Nuevas,
La Yunga,
el
Ambor6,
permanente,
Ambor6,
(LPB); Prov. Arce,
Beck 14092
hacia
Parque Nacional
tramo entre la comunidad
hacia el Rio
del Rio
Parque Nacional
senda de la Yunga
de la parcela
Parque Nacional
above Mairana
Huancanqui,
y cabeceras
el camino Vallegrande-Tierras
senda de la Yunga
64?24'W,
(A, BM, F, GH, K,
Prov. Florida,
de Mairana,
alrededores
siguiendo
Prov. Florida,
de Mairana,
entre Yunguillas
Prov. Vallegrande,
(USZ, UT);
(USZ, UT); La Yunga
3718
Andalgala,
3017
(USZ, UT);
43 km hacia Padcaya,
Bang
locality:
et al. 3043
(8-10
Nogalar
Depto.
& Jardim 3001,
18?04'S,
8384
Prov. Caballero,
(USZ);
a 10 km al N de Comarapa,
I. Vargas
Isidro,
NY),
(NY, USZ);
km al NE
(8-10
et al. 2514
64?32.5'W,
63?57.5'W,
1363
(NY, USZ),
y El Corral
I. Vargas
5.7 km al SE de San
(BM, F, GH, K, MO,
San Juan del Potrero,
Amboro,
et al. 1362
63?53'W,
8383
Steinbach
Parque Nacional
64025'W,
San Rafael),
Prov. Caballero,
(USZ);
(LPB); Comarapa,
Prov. Caballero,
Zapallar,
Rio
et al. 15, 30
Skinner
64033'05"W,
Solomon
31
2094
de los Sosas,
de
Sa. El No
(G, NY, P,
road to Taft
(A, F, GH, GOET,
LP, NY,
(A).
Solanum confusum is characterized by its unbranched and often sparse hairs, elliptic
leaves, long peduncles and pedicels, and usually campanulate-stellate
corollas. It is most
similar to S. glaucophyllum and to S. hibernum, S. luteoalbum, and S. stuckertii in the S.
luteoalbum species group. Solanum glaucophyllum
differs from S. confusum in its pur
plish black fruits, narrower glaucous leaves, and rotate-stellate corollas. The collection
Schreiter 11214 mentioned by Morton (1976) as similar to S. confusum actually belongs
to S. glaucophyllum.
Solanum confusum differs from species of the S. luteoalbum group
in its more campanulate corollas and in its often sparse and shaggy pubescence.
Morton (1976) distinguished S. confusum and S. adelphum in part on the basis of con
nate vs. free anthers, respectively, but this character is difficult to evaluate critically in
herbarium material.
I can find no consistent morphological
distinctions
between
the two
32
VOLUME 61
taxa. Collections
from southern Bolivia (Depto. Tarija) and Argentina are morphologi
cally homogeneous, with sparse pubescence and usually branched inflorescences. These
from elsewhere in Bolivia
populations have been recognized as S. adelphum. Collections
can vary widely in degree of pubescence,
inflorescence branching, and corolla size and
shape. Plants that are glabrous or nearly so were assigned to S. confusum; however,
glabrous and densely pubescent individuals are found growing side by side in several Bo
livian localities and apparently represent morphological
variants of the same species.
Therefore, the name S. confusum is particularly appropriate for this taxon.
Solanum confusum is confined to cloud forest areas above 1700 m inwestern and cen
tral Bolivia, whereas it is often found at lower elevations in southern Bolivia and north
western Argentina. In these regions, it occurs in the forest type known as the "selva sub
tropical boliviano-tucumana"
or "bosque tucumano-boliviano"
(Cabrera 1983; Killeen
et
al. 1993).
The type collection
of S. confusum, Bang 1118, is amixture of two species separated

sheets. Rusby based his description of Solanum clavatum on one of the ele
ments, which has obtuse anthers with large terminal pores; this element is S. aligerum
Schltdl. of sect. Holophylla
(S. Knapp, pers. comm.). The other element has tapered an
on different
thers with small terminal pores and belongs to sect. Cyphomandropsis.

Bitter (1913) re
described what he thought was Rusby's S. clavatum, remarking that Rusby's description
contained many inaccuracies; however, Bitter had the Cyphomandropsis
element rather
than the Holophylla
element. Morton (1944) described the Cyphomandropsis
element as
a new species, Solanum confusum. Sleumer (1957) provided a description of what he
called "Solanum
clavatum," but his description refers to S. confusum. He provided un

information from the Bolivian botanist M. Cardenas that the snow line on Mt.
Tunari is at approximately 4000 m in elevation, which is the source for the maximum el
evation reached by S. confusum given above under "Distribution."
published
3. Solanum cylindricum Vellozo, Fl. flumin. 2: 87. 1829 (text); 2: t. 119. 1831 (icones).
Cyphomandra cylindrica (Vellozo) Sendtner inMartius, Fl. bras. 10: 121. 1846.
Pionandra
cylindrica (Vellozo) Miers, Ann. Mag. Nat. Hist. 15, ser. 2: 199.
1855.-TYPE:
BRAZIL. Locality unknown, Vellozo s.n. (holotype: unknown).
Solanum ellipticum Vellozo, Fl. flumin. 2: 84. 1829 (text); 2: t. 100. 1831 (icones),
non Solanum ellipticum R. Brown, 1810. Cyphomandra
elliptica
(Vellozo)
Sendtner inMartius FH.bras. 10: 121. 1846. Pionandra elliptica (Vellozo) Miers,
Ann. Mag. Nat. Hist. 15, ser. 2: 199. Solanum johannae Bitter, Repert. Spec.
Nov. Regni Veg.
of Pharmacopolis
12: 465. 1913.-TYPE:
BRAZIL. Rio de Janeiro: seaside forest

[Parati], Vellozo s.n. (holotype: unknown).
Solanum subhastatum Smith & Downs, Phytologia 10: 432. 1964. Cyphomandra sub
hastata (Smith & Downs) A. Child ex Bohs, Fl. Neotrop. Monogr. 63: 154.
1994.-TYPE:
BRAZIL. Santa Catarina: Lauro Muiller-Urussanga,
Pinhal da
Companhia, 300 m, 23 Aug 1958, Reitz & Klein 7053 (holotype: US! #2323379;
isotypes: HBR, L!).
Solanum catanduvae Smith & Downs, Fl. Illustr. Catar., Solanaceas, 1: 135, fig. 14a,
a-c. 1966.-TYPE: BRAZIL. Santa Catarina: Mpio. Catanduvas, E of Catanduvas,
ruderal, ca. 27?3'S, 51?45'W, 700-800 m, 7 Nov 1964, Smith & Klein 12980
(holotype: US! #2568831;
isotypes: B! F! HBR, R, US!).
Solanum iraniense Smith & Downs, Fl. Illustr. Catar., Solanaceas,
1: 137, fig. 14a,
d-f. 1966.-TYPE:
BRAZIL. Santa Catarina: Irani, Campo de Irani, ca. 26?57'S,
SOLANUM
2001
51P50'W,
#2568834;
700-900
m,
13 Oct
33
1964, Smith & Reitz
12457
US!
(holotype:
isotypes: HBR, NY! R, US!).
Shrub ca. 1-2 (-5) m tall. Stems glabrous to densely pubescent with unbranched or
branched hairs, the branched hairs often long-stalked and with three or more
dendritically
terminal rays. Leaves 4-many per sympodial unit, the blades 3-12.5 cm long, 0.5-3.5 cm
to narrowly elliptic or
wide, length:width ratio ca. 2-10:1 or more, simple, elliptic-ovate
lanceolate, acute to acuminate at apex, cuneate to decurrent and occasionally
subhastate
at base with 1-2 small lobes, chartaceous to subcoriaceous, glabrous to moderately pu
bescent
to densely pubescent abaxially with simple or branched hairs,

denser on margins and veins, petioles 0.3-2.5 cm long, sparsely to densely
pubescent. Inflorescence unbranched or forked, 1-15-flowered,
0.5-5 cm long; peduncle
0.5-3.5 cm long; rachis up to 3 cm long; pedicels 5-15 (-20) mm long, nearly contiguous
adaxially, glabrous
the pubescence
or spaced
ca. 1-15 mm apart, articulated at the base; inflorescence axes glabrous to

pubescent with unbranched, forked, or dendritically branched hairs and some
times also with small stalked glands. Calyx sparsely to densely pubescent, the radius 2-9
mm, the lobes 1-6 mm long, 1-2 mm wide, deltate to narrowly triangular and often
abruptly narrowed distally, acute to acuminate at tips. Corolla purple or white with a
lighter or darker star at base, chartaceous, stellate, the radius 8-13 mm, the tube 2-3 mm
densely
long, the lobes 5-10 mm long, 2-4 mm wide at base, triangular to narrowly triangular,
acute at apex, sparsely to densely puberulent abaxially with unbranched or branched hairs,
nearly glabrous adaxially except for a few hairs on midveins. Anthers usually
yellow, lanceolate to narrowly oblong, 4-6 mm long, 1.5-2 mm wide, abaxial
most with very fine or inconspicuous scaly papillae, the pores directed distally
ially. Ovary glabrous; style glabrous tomoderately puberulent, cylindrical, 5-6
connivent,
surface at
and adax
mm
long,
ca. 0.5 mm in diameter; stigma truncate to subcapitate. Fruits 1.5-2.5 cm long, 0.8-2 cm
in diameter, globose, ellipsoidal, or ovoid-fusiform,
obtuse or acute at apex, glabrous,
color unknown; stone cell aggregates absent. Seeds 2-2.5 mm long, 2 mm wide, lenticu
lar, puberulent on margin but otherwise
12, 13.
nearly glabrous. Chromosome
number unknown.
Figs.
Phenology. Collected in flower in January, February, May, June, and August through
December; collected in fruit in January, February, September, November, and December.
Distribution
(Fig. 14). Southeastern Brazil (Parana', Rio Grande do Sul, Santa Cata
rina), and perhaps Rio de Janeiro and in adjacent areas of Misiones, Argentina; clearings
inAraucaria
forest; 300-1100 m.
Local names. Brazil: Jua, joa'manso, joaimanso de f6lha comprida (all from Smith &
Downs, 1966).
REPRESENTATIVE SPECIMENS. Brazil.
7081
(F, GH,
Hatschbach
NY);
9668
19384
15436
(F, MBM,
Sao Mateus
Z); Mpio.
19391
(F); Mpio.
Laranjeiras
Ortiguera,
do Sul, Vargem
Hatschbach
43491
Branco
do Sul, Rod.
PR-092,
do Sul, Klein & Eskuche

of Rio Tigre Preto,
2839
PARANA: Marichal
17508
(GH,
Grande,
do Sul, Rio
Hatschbach
(F, MBM);
Hatschbach
19-50
(Z); Mpio.
Mpio.
Rambo
23255
53509
Igua,u,
prox.
Hatschbach
(MBM,
(B).-RIo
Itaperussii,
7924
22933
Jonsson
Hatschbach
45519
1009a
Kummrow
1355
Palmas,
Pal
a barra, Hatschbach
Hatschbach
(C, CTES, MBM,
Z); Planalto,
Dusen
do Carumbe,
(US); Mpio.
prox.
a barra do Desengano,
Hatschbach
& de Haas
(S, US);
do Sul, Varginha
Rio Chopim,
do Sul, Bromado,
Lindeman
3056
10538 & Pereira
(UT); Itaperussu,
Rio Branco
Dusen
Bocaiuva
Vizinhos,
Irati, Riozinho,
48833
ca. 25 km S of Marmaleiro,
(NY); Cascavel,
Dois
Serra dos Mulatos,
Mallet,
S); Mpio.
do Sul, Hatschbach
(F, L, LIL, NY, US); Mpio.
Palmas,
& de Haas
Dusen
(MBM, US); Laranjeiras
mas, Hatschbach
Guimardes
Tres Barras,
26472
(C, MBM,
Rio
Laranjeiras
(UT); near source
(NY); N of Campo Novo,
GRANDE DO SUL: Neu-Wurttemberg,
(UT); near
Z); Mpio.
(F, GH, K, NY);

et al. 3067
&
NY, Z); Mpio.
Lindeman
Bornmiiller
384
34
VOLUME 61
A~~~~~~~
ciii
branched
(element 4; see text). A. Habit. B. Dendritically
trichome. C.
view of partially sectioned
F. Stamen (left to right:
flower. E. Gynoecium.
G. Fruit. (Based on: A, Hatschbach
15436; B-G, Smith & Klein 12982.)
Flower,
12. Solanum
cylindricum
seen from above. D. Lateral
abaxial,
lateral, adaxial
FIG.
views).
2001
SOLANUM
35
-V~~~c
/~~~~~~~~~~~~~~~~~~~~~~~~~~~
ttlitEX
d!~~~~~~~~~~~~~~~
B~~~~~
FIG. 13. Solanum

of partially
sectioned
(Based on Hatschbach
cylindricum
(element
5; see text). A. Habit.

E. Stamen
B. Flower, seen from above. C. Lateral view

(left to right: abaxial, lateral, adaxial views). F. Fruit.
22933.)
36
VOLUME 61
10
0~~~~~~~~~~~~~~~~~~~~~~~~2
A~~~~~~~~~~~~~~A
-I
1..
::-
1-----
'
AS.cylindricum
S. glaucophyllum
20
30
200 400 600 800 1000km

~~~~~~~~0
0 100 200 300 400 500 600miles
40
0
70
80
FIG.
(GH); Farroupilha,
Camargo
60
14. Distribution
2101
50
of S. cylindricum
(B); Farroupilha,
Rambo
and S. glaucophyllum.
40302
(LIL); Santa Rita
p. Farroupilha,
Rambo
45760 (B);Montenegro, Sehnem 3859 (B);Gramado,Sehnem4162 (US).-SANTA CATARINA:

Mpio. Caqador,
8 km N of Cagador,
Smith & Reitz 8956
Ere, Smith & Klein

12982
(NY, US).
11572
Argentina.
Iguazu, Ao. Yacuf, Klein

San Antonio,
C.E.B.S.,
MIsIoNES:
& Eskuche
Klein
(US); Mpio.
(US); Mpio.
1-38
& Eskuche
Catanduvas,
Chapec6,
Fazenda
E of Catanduvas,
Campo
Sao Vicente,
ca. 27?03'S,
24 km W
51045'W,
ca. 5 km de Bernardo
de Irigoyen, Bernardi
18835
(US); San Antonio-S.,
Ao. Guayuvira,
& Eskuche
9058
(US);
Iguazu, Osten
& Rojas
Klein
8262
of Campo
Smith & Klein
(BM); Parque Nac.

7-27
(US, Z);
(K).
Solanum cylindricum differs from other species of sect. Cyphomandropsis

from
southeastern Brazil in its relatively narrow, simple leaf blades and frequent occurrence of
branched hairs. Solanum pelagicum also has stems covered with dense dendritic pubes
cence, but it usually has pinnately compound leaves and is restricted to coastal restinga
vegetation. Solanum luridifuscescens
has generally larger, wider simple leaves, lacks
branched hairs, and has a dense band of scaly papillae on the abaxial surface of the an
thers. Solanum matadori
is glabrous, and has subcoriaceous leaf blades and branched in
florescences with very long peduncles.
My concept of S. cylindricum encompasses a great deal of variation in pubescence,
leaf size and shape, and inflorescence and fruit morphology. Specimens range from nearly
to densely
pubescent
with dendritic
to substellate
hairs. Leaf blades
range from long and narrow to elliptic-ovate,
and inflorescences vary from raceme-like
with an extended rachis to contracted and umbel-like with pedicels clustered close to
gether at the distal end of the axis. Fruits are obtuse to acute at the apices. Six morpho
glabrous
types can be discerned in the material of S. cylindricum available tome, but all intergrade
inmorphological
features and geographical distribution, which argues against recognition
of separate taxa. Most specimens from Rio Grande do Sul have very narrow coriaceous
leaves (length:width ratio ca. 3-10:1) with the blades nearly glabrous above and moder
ate to dense dendritic pubescence on the axes and abaxial leaf surfaces. The inflorescence
2001
SOLANUM
is raceme-like
37
at the bases of the pedicels. Many collections

from
also have very narrow leaf blades with sparse adaxial pubes
cence, but differ in inflorescence structure: the flower pedicels emerge at nearly the same
and lacks swellings
Parana and Santa Catarina
point on the axis (umbel-like)

represented
by Smith & Klein
and are subtended by a swelling or sleeve. Another element,

11572, 12980, 12982, and Hatschbach & Pereira 10538
from Parana and Santa Catarina, has rather broad leaf blades with a length:width
ca. 2-3.5:1.
Pubescence
raceme-like, without
ratio of
is relatively
sparse in these collections, and the inflorescences are

swellings. This element was described as Solanum catandu
pedicel
it on the basis of its supposedly

(1966), who differentiated
habit and relatively broad leaf blades with obvious lateral veins. The fourth mor
photype also has rather broad leaves, but is notable in its extremely dense dendritic pu
bescence
that nearly obscures the stems and abaxial leaf surfaces. This element has
vae by Smith
and Downs
woody
raceme-like
inflorescences without
an obvious pedicel
swelling or sleeve. Specimens
with
sparse to moderate
branched pubescence, mostly from western parts of Parana and Santa

Catarina, were described as S. subhastatum Smith & Downs and constitute a fifth mor
photype (Fig. 13). The sixth morphotype consists of nearly glabrous plants, usually with
narrow leaves, ellipsoidal
elements
to ovoid and sometimes
it was
segregated
is somewhat
distinctive
inflorescences;
or geographical
morphological
pointed fruits, and short, few-flowered
as S. iraniense Smith & Downs.

when
in isolation,
viewed
disjunctions
Though
each of these
I cannot discern consistent
that separate them, and therefore I have sub
sumed all these variants under one taxon.

Solanum cylindricum may be relatively basal in the Cyphomandropsis
clade, for it has
a number of characters thatmay link it to putative sister groups in Solanum subg. Minon.
These include anther pores that eventually open into longitudinal slits, pedicels often in
serted in a basal sleeve, collar, or platform, and branched pubescence that resembles the
stellate hairs seen in some groups within subg. Minon. Although most of the hairs of S.
are dendritically
cylindricum
branched, some specimens have the lateral branches
in several
arranged
tiers ("substellate";
Fig. 12). Seithe
(1979) postulated
that dendriti
cally branched and stellate hairs are derived from different developmental pathways, and
thus discounted an ontogenetic relationship between the two types of branched hairs seen
in Solanum species. This question should be re-examined in light of phylogenetic
evi
dence that shows complex
patterns of hair types throughout the genus (Bohs & Olmstead
1999).
Though
morphic
highly
species,
stylized, Vellozo's
plates evidently
lipticum depicts a plant with narrowly elliptic

obtuse
soidal,
Solanum
fruits. He
cylindricum
describes
pointed fruits and 3-4-flowered,

dricum as described
is Vellozo's
cylindricum
leaves, racemose
this species
is very similar, with
of this species as glabrous

mens
represent variants of this poly
and thus his names have been adopted. Vellozo's
subumbellate
as pubescent
narrowly
elliptic
inflorescences.
("laevi"). Itmay conform
plate of Solanum
inflorescences,
("sericeis").
Vellozo's
leaves but with

Vellozo
somewhat
describes
to the sixth morphotype
el
and ellip
the stems
of S. cylin
above. The one point that is difficult
collecting
to reconcile with recent speci

locality in Rio de Janeiro. No other herbarium material of S.
has been seen from this state, nor from intervening areas in Sao Paulo.
4. Solanum
fallax Bohs, Taxon 44: 585. 1995. Cyphomandra hypomalaca Bitter, Repert.
Spec. Nov. Regni Veg. 17: 346. 1921, non Solanum hypomalacum (Bitter) C. V.
Morton, 1944.-TYPE:
ECUADOR. Gualea, subtropical woods, May 1886, Sodiro
38
114/60
photo of holotype,
(holotype: B, destroyed;
VOLUME 61
F neg. 2934: F! G! GH!;
iso
type:P!).
Cyphomandra
var. velutina Dunal
betacea
PERU. Pavon
in DC., Prodr. 13(1): 394. 1852.-TYPE:
s.n. (holotype: G!).
to densely pubescent with un

Shrub or small tree 3-5 m tall. Stems moderately
branched glandular and eglandular hairs. Leaves 3 per sympodial unit, the blades 6-37 cm
long, 4-25 cm wide, length:width ratio 1-2:1, simple, ovate, acute at apex, truncate to
deeply cordate at base, chartaceous, sparsely to moderately pubescent adaxially, more
to densely pubescent with
densely so abaxially, the petioles 2-15 cm long, moderately
hairs like those of the stem. Inflorescence usually forked or further branched, 20-50-flow
ered or more, 5-20 cm long; peduncle 2-8 cm long; rachis 2-12 cm long; pedicels 10-20
mm
long in fruit, spaced 2-9 mm apart, articulated above the base, leav
long, 15-30 mm
remnants 1-6 mm
ing pedicellar
distally, moderately
long; inflorescence moderately pubescent. Calyx inflated

the radius 2-3 mm, the lobes 0.5-1 mm long, 2 mm wide,
pubescent,
truncate, with acuminate
purple, chartaceous to subcoriaceous,

stellate, the
long, the lobes 8-13 mm long, 1.5-3 mm wide at base,
tips. Corolla
the tube 1-2 mm
radius 10-15 mm,
triangular, acute at apex, sparsely pubescent abaxially, nearly glabrous adaxially.

not connivent, yellow or purplish, narrowly triangular, 4-5 mm long, 1.5-3 mm
abaxial surface with thickened discrete papillate connective, the pores directed dis
narrowly
Anthers
wide,
tally. Ovary densely puberulent; style sparsely puberulent, cylindrical, 7-9 mm long,
0.5-1 mm in diameter; stigma truncate. Fruits 1-1.5 cm long, 1-1.5 cm in diameter, glo
bose, obtuse at apex, densely puberulent, color unknown; stone cell aggregates absent.
Seeds 4-5 mm
long, 3-4 mm wide,
angled, whitish
number unknown. Figs. 15, 16.

Phenology. Collected in flower in January, April
reticulate on margin.
puberulent,
Chromosome
ber, and November;
collected
through June, and August,
in fruit in January, February, May,
June, August
Septem
through Oc
tober, and December.

Distribution
(Fig. 8). Colombia
and western
or scrub, Jauneche forest (tropical moist

REPRESENTATIVE SPECIMENS. Colombia.
and Zarzal,
El Medio,
Hacienda
tanga, Sodiro
BABURA: Collapi,
F).-MANABI:
Acosta
12847
Solis
Cerro Montecristo,
S, US).-EL
ORO: ravine above Pifias

Quito,
airline
SE of Nanegal,
tween Mocachi
tween Babahoyo
Sparre
on W
00?7.5'N,
and Palenque
and Montalve,
on Estero
Sparre
(S); El Recreo,
trail from Hacienda
78?38'W,
Webster
& Hebert
Pefnafiel, Dodson
17919
Panamericana
Ecuador.
on the San Lorenzo
19484
slope of Andes,
Maquipucuna,
(NY).
03?09'S,
27710
& Valverde
Sparre
R.R., Madison
30'S, Eggers
79008'W,
El Carmen
Knight
to Hacienda
(TEX).-Los
6952
between
La Paila
CHIMBORAZO: Valle
ca. 25 km S of Esmeraldas,
Timbre,
in dry savanna
m.
Zarzal, Carretera
et al. 2606
(F); Tercer Paso
S. Manta,
CHA: Cant6n
Reserva
VALLE: Mpio.
Hacienda
forest pockets
forest); 20-1300
Silverstone-Sopkin
s.n. (P).-ESMERALDAS:
Ecuador;
15248
et al. 4972
15069
673
(AAU,
(C, F, GH, K, M,
(WIS).-PICHIN
Esparragos,
RIoS: Cant6n
(MO); Hacienda
Palla
(S).-IM
ca. 5 km
Vinces,
Clementina,
be
be
(S).
The affinities of S. fallax are problematical. This species was originally described as
of the genus Cyphomandra
(now Solanum sect. Pachyphylla)
and was treated
as such by Bohs (1994). However,
the anther connective
is not greatly enlarged in this
a member
species, and its flower and fruit structure, especially its distally swollen calyx and thick
angled seeds as well as its tolerance of drier sites, are more suggestive of species of sect.
It is anomalous in the latter section due to its very large cordate leaves,
Cyphomandropsis.
its pedicels
articulated
above
the base
leaving prominent
pedicellar
remnants,
and its
2001
SOLANUM
FIG.
15. Solanum
ster & Hebert 27710;
fallax.
A. Habit.
E. Stamen
B. Flower,
seen

et al. 2606.)
F, Silverstone-Sopkin
from above.
lateral, adaxial
39
C. Lateral
views).
view
F. Fruit.
of partially sectioned
on: A-E, Web
(Based
40
VOLUME 61
Nam;
ir~~~~~111
FIG.
fallax].
Note
16. Photo
of holotype
prominent
pedicellar
of Cyphomandra
hypomalaca
remnants on inflorescence.
Bitt.
(Sodiro 114/60,
B; F neg. 2934)
[=Solanum
2001
SOLANUM
fruits. If it indeed belongs
pubescent
41
to sect. Cyphomandropsis,
closely
related to S. amotapense
species
share include cordate leaf bases, swollen calyces,
5. Solanum
from coastal Ecuador
S. fallax may be most
and Peru. Characters

and Ecuadorian
that the two
distributions.
Smith & Downs,

10: 431. 1964. Cyphomandra
Phytologia
(Smith & Downs) A. Child ex Bohs, Fl. Neotrop. Monogr. 63: 154.
1994.-TYPE:
BRAZIL. Santa Catarina: Mpio. Dionisio Cerqueira, near Dionisio
fusiforme
fusiformis
Cerqueira,
800-850
30 Dec
m,
1956, Smith & Reitz
9658
US!
(holotype:
#2423799; isotype:HBR).
Shrub 0.5-2 m
tall. Stems glabrous
4 per sympodial
Leaves
tic-ovate
to pinnately
acute to acuminate
to sparsely puberulent with unbranched

the simple
7-11-compound,
at apex, truncate to cuneate
upper lateral leaflets often basiscopically

(rarely moderately)
puberulent
oles 1-7 cm long, glabrous

branched,
or nearly so. Calyx
purple, chartaceous,
mm
adaxially. Anthers
usually
the peti
puberulent. Inflorescence un
cm long; rachis 1-11 cm long,
3-7
(-30) mm apart, articulated at or

remnants up to 1mm long; inflorescence axes
the radius 3-5 mm,
often unequal, obtuse
at base, narrowly
long, 1.5-2.5 mm wide,
to sparsely
glabrous
along midribs,
long, spaced 1-10
stellate, the radius 10-15 mm,
long, 3-4 mm wide
ratio 2-3:1,
leaves with the
the compound
subcoriaceous,
(rarely moderately)
glabrous,
triangular-deltate,
hairs.
simple and ellip
length:width
and abaxially, especially
slightly above the base, leaving pedicellar

glabrous
at base,
cm long; peduncle
6-18
10-25 mm
cm wide,
leaves with
decurrent,
adaxially
to sparsely
6-12-flowered,
often zigzag; pedicels
mm wide,
cm long, 3-17
unit, the blades 7-20
the lobes 1-2 mm
to acute at tips. Corolla

the tube 2-3 mm
long, the lobes 8-13
triangular, acute at apex, glabrous abaxially and
the color unknown,
connivent,
long, 1-3
pink to dark
triangular, 5-7 mm
narrowly
abaxial surface with an obvious band of scaly papillae,
the pores
directed distally. Ovary glabrous; style glabrous to sparsely puberulent, cylindrical, 7-9
mm long, 0.5-1 mm in diameter; stigma truncate. Fruits 3-5 cm long, 0.5-1.5 cm in di
ameter, elliptic-fusiform,
acute at apex, glabrous, yellow when ripe; stone cell aggregates
several per fruit, small. Seeds 2.5-3 mm long, 2 mm wide,
lose. Chromosome number: 2n = 24. Fig. 17.
Collected
Phenology.
in flower in January, February, April,
cember, with a peak inDecember
and January; collected
Distribution
(Fig. 18). Argentina,
Paraguay,
the rivers Parana and Uruguay; Araucaria

zone (fide Smith & Downs,
Local names. Brazil:
Rond6n,
5373
(US).-Rio
RINA: Liso, Guaraciaba,

(SP, US).
Antonio,
and October
through De
in fruit in January through April,
Paraguay.
C.E.B.S.,
Caraguatay
Bom
jua' (Smith & Downs
de San Pedro,
camino
16873
ALTO PARANA: Rio

& Eskuche
1573
10404
a Tobuna,
in drainages
Seco
(C, CTES,
(NW),
(L, NY, US).-SAO
Itab6, Caballero
3498
(Z); Posadas,
263
Sr. Leib.,
Morrone
Sehnem
10884
281 (CTES).
Bonpland,
Km
Ekman
m.
(G, LD,
9500
CATA
4421
Loefgren
MIsIONEs:
S, US);
Ruta Nac.
Gral. M. Belgrano,
San
San Pedro,
(BH, G); Depto.
Cruce Caballero,
et al. 799 (SI); Depto.
(Z); Mpio.
do Iguacu,
(US).-SANTA
Argentina.
845
17, Montes
San Pedro, Parque Provincial
1'S, 53059W,
18384
UT, Z); Parque Nacional
PAULO: Vale do Paranapanema,
(W); Iguazii, Puerto Segundo,
(LP); Depto.
do Sul, Hatschbach
HBG,
Marmori
of
and thickets in "mata branca"

1966).
PARANA: Laranjeiras
40573
GRANDE DO SUL: Herval
(Centro), Montes
Puerto Wanda, Montes
Jardim, Hatschbach
Reitz & Klein
Ahumada
and southeastern Brazil
forests, clearings,
1966), riparian forests, and disturbed areas; 500-850

Joaimanso,
ADDITIONAL SPECIMENS EXAMINED. Brazil.

Candido
Pereira
flattened, tomentu
and December.
October,
Mal.
strongly
Iguazui,
14, 14 km
8 km de
42
VOLUME 61
ol
FIG.
17. Solanumfusiforme.
partially
sectioned
(Based
on: A, C-G,
A. Habit.
B. Compound
F. Stamen
Smith & Reitz 9658; B, Hatschbach
(left
1~~~~~~~~~~~~~~~~~~~~
leaf. C. Flower, seen from above. D. Lateral view of

to right: abaxial, adaxial,
lateral views). G. Fruit.
18384.)
2001
43
SOLANUM
20
A S.
fusiforme
0 S. luridifuscescens
*S. matadori
US. pelagicum
0
FIG.
San Antonio,
18. Distribution
26?01'S,
53047W,
Parque Provincial
Pedro,
Morrone
Cruce Caballero,
This distinctive
S. luridifuscescens,
of S. fusiforme,
et al. 2073
26?31'S,
and S. pelagicum.
S. matadori,
745
(SI); Santa Ana, Rodriguez

53?59'W,
600 800 1OOOkm
0 100 200 300 400 500 600 miles

40
50
60
70
200 400
30
Zuloaga
et al. 5538
species has several unusual morphological
(A, BM,
SI); Depto.
San
(SI).
characters not commonly
in the other
species of sect. Cyphomandropsis,

including pinnately compound
leaves, fusiform fruits, and nearly complete lack of pubescence. Some species of Solanum
found
sect. Pachyphylla
exhibit these characters, and S. fusiforme may have a closer relationship
with sect.Pachyphylla, especially theBrazilian species S.melissarumBohs, S. diploconos

(Mart.) Bohs,
S. pinetorum
(Smith & Downs)
Bohs,
and S. sciadostylis
(Sendtn.) Bohs,
thanwith members of sect. Cyphomandropsis.Solanumfusiforme, however, completely

lacks the swollen anther connective
cal of sect. Cyphomandropsis.
Solanumfusiforme
jacent Argentina
serve populations
is uncommon
that defines
sect. Pachyphylla
and has stamens
typi
and restricted to southeastern Brazil and areas of ad
(Misiones) and Paraguay. Efforts

of this interesting species.
6. Solanum
Desfontaines,
glaucophyllum
based on living plants cultivated
should be made
to relocate and pre
Cat. P1. Hort. Paris, ed. 3: 396. 1829.-TYPE:

in the Botanical Garden at Paris (holotype: un
known).
Solanum malacoxylon
Sendtner inMartius, Fl. bras. 10: 51. 1846. Solanum mala
var.
coxylon
genuinum Hassler, Repert. Spec. Nov. Regni Veg. 15: 120. 1918.
TYPE: BRAZIL. Sellow s.n. (holotype: B, destroyed; fragment of holotype: F!;
photos of holotype,
F neg. 2837: F! G!; lectotype, designated
by Morton,
1976:
K!).
Solanum glaucum Bertoloni, Mem. Accad. Sci. Ist. Bologna 3: 188, t. 13. 1851, nec
Solanum glaucum Dunal, 1852, nec Solanum glaucum Rojas, 1897.-TYPE:
based on living plants cultivated at the Botanical Garden of the University of
Bologna
(holotype: unknown).
44
SYSTEMATIC
BOTANY
MONOGRAPHS
Solanum
glaucum
Dunal
in DC.,
Prodr. 13(1):
VOLUME61
100. 1852, nec Solanum
glaucum
Bertoloni, 1851, nec Solanum glaucum Rojas, 1897.-TYPE: ARGENTINA.

Buenos Aires, Bacle 43 (lectotype, designated by Morton, 1976: G!; photo of lec
totype, Morton neg. 8586: F! GH!; isolectotype: G-DC!; photo of isolectotype: F
neg. 6796 F! GH!).
var. angustissimum Kuntze, Rev. gen. pl.
Solanum malacoxylon
("melanoxylon")
var. subvirescens
f. vulgare subf. an
3(2): 227. 1898. Solanum malacoxylon
gustissimum (Kuntze) Hassler, Repert. Spec. Nov. Regni Veg. 15: 120. 1918.
TYPE: PARAGUAY.Rio Tebicuary, Kuntze s.n. (lectotype, here designated: NY!).
("melanoxylon") var. latifolium Kuntze, Rev. gen. pl. 3(2):
PARAGUAY.Sep 1982 (fr), Kuntze s.n. (lectotype, here desig
Solanum malacoxylon
227. 1898.-TYPE:
nated:NY!; isolectotype:US!).
var. albo-marginatum Chodat, Bull. Soc. Bot. Geneve, ser. 2,
Solanum malacoxylon
8: 153. 1916. Solanum malacoxylon
var. subvirescens f. albo-marginatum
(Cho
PARAGUAY.
dat) Hassler, Repert. Spec. Nov. Regni Veg. 15: 121. 1918.-TYPE:
Lago Ypacarai, Tuilerie, San Bernardino, Chodat & Vischer 36 (holotype: G!).
var. subvirescens Hassler, Repert. Spec. Nov. Regni Veg. 15:
Solanum malacoxylon
120. 1918. Solanum malacoxylon
var. subvirescens f. vulgare Hassler, Repert.
PARAGUAY. Margin of Lake
Spec. Nov. Regni Veg. 15: 120. 1918.-TYPE:
Ypacarai, Hassler 3201 (lectotype, here designated: NY!; isolectotypes: BM!
W!).
Rhizomatous
shrub or slender treelet ca. 0.5-4 m tall. Stems glabrous (rarely moder
ately to densely puberulent-pubescent),
usually light-colored and smooth. Leaves many
per sympodial unit, the blades 6-18 cm long, 0.6-3.5
(-5) cm wide, length:width ratio
4.5-10 (-15): 1, simple, narrowly elliptic, acute at apex, tapered to decurrent at base, suc
culent or fleshy with the midrib and margin often thickened and whitish, glabrous adaxi
ally and abaxially (rarely moderately
to densely puberulent-pubescent),
surfaces glaucous
in fresh material, the petioles 1.5 cm long or less, glabrous, often slightly winged. Inflo
rescence branched, sometimes highly so, ca. 20-50-flowered
or more, 3.5-9 cm long; pe
duncle 1-3.5 cm long; rachis 2-7 cm long; pedicels 12-15 mm long, ca. 15-20 mm long
in fruit, spaced 1-15 mm apart, articulated at the base; inflorescence axes glabrous (rarely
puberulent-pubescent).
Calyx glabrous except for some sparse puberulence at
margin, the radius 2-3 mm, the lobes 0.5-1.5 mm long, 1.5-2.5 mm wide, deltate, acute.
moderately
to pink or violet, often with a white central star, chartaceous, rotate-stel

late and plicate, the radius 10-30 mm, the tube 5-8 mm long, the lobes 4-10 mm long,
5-10 mm wide at base, broadly triangular, apiculate at apex, moderately
to densely pu
berulent abaxially, especially on distal parts of lobes and plicae, glabrous adaxially except
for a few sparse hairs at tips of lobes. Anthers usually connivent, yellow to orange-yellow,
ovate, 5-7 mm long, 2 mm wide, abaxial surface smooth to roughened but not obviously
Corolla whitish
papillate, the pores directed distally. Ovary glabrous; style glabrous, cylindrical, 5-7 mm
long, 0.25-0.5 mm in diameter; stigma truncate. Fruits 0.75-2 cm long, 0.75-2 cm in di
ameter, globose, sometimes apiculate at apex when young, obtuse at apex when mature,
glabrous, dark purple or blue-black and glaucous when ripe; stone cell aggregates absent.
Seeds ca. 4-6 mm long, 3.5-4 mm wide, angled, smooth or with minute scalloped ridges.
Chromosome
number: n = 12. Figs. 19, 20.
Phenology.
Flowering
and fruiting throughout
the year, with
a peak
in November
through March.
SOLANUM
2001
45
.:
B
FIG.
19. Solanum
glaucophyllum.
tioned flower. D. Gynoecium.

C, F, greenhouse
material
E. Stamen
of Bohs
A. Habit.
B. Flower,

adaxial,
lateral views).
E
~~~~~~~~~~~~~~~~
2530; D, E, Crist6bal
~~~~D
view
F. Fruit.
of partially
(Based
et al. 1437.)
sec
on: A, B,
AA:
ON
..........
7,11
. .........
FIG. 20. Solanum
glaucophyllum.
A. Flowering
branch;
scale bar
2 cm. B. Flowers
with
rotate
2001
SOLANUM
47
Distribution
(Fig. 14). Bolivia, southern Brazil, Paraguay, northern Argentina, and
Uruguay; low, swampy ground at margins of marshes and ponds in seasonally inundated
areas; ca. 0-600 m.
Local names and uses. Argentina: Corcho del agua (Torres 9), duraznillo (Boffa 133;
Gibson s.n.; Huidobro 1452; Schulz 2053; Tweedie s.n.), duraznillo blanco (Gibson s.n.;
Leftbre s.n.), duraznillo de baniado (Schinini 9033), duraznillo hediondo (Hunziker 310),
duraznillo negro (Cordini 61), na'kyet (Arenas 2000), palo hediondo (Boelcke 1396),
palo-ne or palone (Boelcke 1396; Irigoyen 221), varilla (Burkart 8727, Hunziker 25304),
yaa'tuk or ye:'tuk (Maranta & Arenas 153). Bolivia: Bobo (Murquia 502). Paraguay: sin
hejuik (Arenas 714). Uruguay: duraznillo blanco (Gallinal et al. A831; Gibert 140), du
raznillo negro (Osten 21883). Brazil: espichadeira (Prance et al. 26173), espixadeiro
(Macedo et al. 1209).-The
dried stems are used for firewood and for wattling (Burkart
as a purgative (Morton, 1976; Gibert 140;
s.n.). Plants are used medicinally
Tweedie s.n.). Foliage is very toxic for cattle and other animals (Prance et al. 26173;
8727; Gibson
Schulz 2053).
Tokarnia
699
(US); Transpantaneira
Transpantaneira
way,
Dobereiner
Mpio.
& Tokarnia
Dobereiner
Guimardes
Brasil
800
& Tokarnia
Fazenda
Aquidauana,
Fronteira,
Dobereiner
(F, US);
Mpio.
(F); Mpio.
Miranda,
Rio Miranda,
& da Silva 49222
(NY, US);
Pantanal,
Fazenda Miranda,
Silva 80 (SP).-Rio
et al. 26173
Fazenda
Fazenda
Hatschbach
Schaller
(NY);
794 (F, US);
Hatschbach
Cruz,
38640
(F, US);
Porto Murtinho,
301
(NY); Mpio.
de Miranda,
GRANDE DO SUL: I.A.S, Pelotas, margem
Tu
Fazenda
Aquidauana,
Santa
High
Sao Sebastiao,
& Tokarnia
797 (US); Mpio.
Aquidauana,
et al. 1209
(NY); Transpantanal
Corumbd,
&
Dobereiner
Cassange,
Tabaco, Dobereiner
& Tokarnia
21941
60 km N de Guaicurus,
Prance
56?30'W,
GROSSO DO SUL: Mpio.
(NY).-MATO
717 (L, US); Mpio.
Fazenda
Fazenda
Pacone,
Santa Rosa, Km 40, Macedo
Fazenda
Pocone,
Jofre, 17?10-17'S,
258
Jofre, Schaller
Corumbd,
pacireta,
Fazendo
Highway,
Fazenda
MATO GROSSO: Mpio.
Highway,
&
Hatschbach
Marimbondo,
do Canal
do I.A.S.,
Sao
Bolivia.
BENI: Prov. Cercado, 7 km SW of Trinidad, vic. Puerto Almacen,
along
Gonqalo, Sacco 816 (F, NY).
the Rio Ibare, 14?52'S, 64?57'W, Nee 37532 (G, LPB, NY, USZ).-CHUQUISACA:
Prov. Luis Calvo, margenes
del lugar "Arbol Solo," Murquia
orillas
US);
de represa, Penseiro
Prov. Andres
17047'S,
Puesto
Izozog, Navarro
2767
4 km NW
Corrales,
& Vargas
346
SE of Comunidad
17050'S,
Don
Nee
62049'W,
40174
de Estancia
Arenas
(MO, NY);
Zinfunke,
Cachari,
1407
guna Ypacarai,
Ascunci6n,
13344
del Salado,
querenza,
cauce
del Rfo Timane,

Nueva Misi6n,
Morong
163
Jukyty, cercanfas
del Cruce
I. Vargas
Schinini
3919
704
Mereles
181
(CTES, NY);
(G); Trinidad,
Mereles
20?0'S,
60?45'W,
Schinini
Schinini
& Bordas
17881
et al. 1935
35 (G); Laguna
& Bordas
camino
14871
F, G, MO,
17?51'S,
Don
Lorenzo,
a la localidad
de Izozog,
Guarayos-Santa
Casas
Puerto Casado
5 km de
(CTES);
Puerto
4061
Simpio
4434
Pedersen
ltd Enramada,
(CTES);
463
Sparre & Vervoorst
(CTES); Mayor
& Molero
and vicinity,
NY, US, WIS);
de
500 m al N de las
(USZ); Missiones
Ipacarary, Fiebrig
et al. 496
P. Caballero,
2692
Ibaniez, 12 km
[Rfo Guapail,
Estigarribia,
de
del Rio
12 km E of center
Prov. Andres
costa del Lago Ypacarai, Mereles
Aregud,
inundado
bosque
de Santa Cruz,
(Jukyty), Bordas
trail
Bafiados
ALTO PARAGUAY: Puerto Diana,
(CTES);
Piquete-cue
& Ramella
(CTES,
Alto,
Baniados
200 km NE
(NY, USZ);
the Rio Grande
(BM, F, GH, K, MO,
Bahia
75300
Ibaniez, 7 km SE of comunidad
Paraguay.
del arroyo ka'a niave, Perez
seco del Rfo Zimane,
of
cerca y al S de Mariscal
(CTES, G, MO).-CENTRAL:
s.n. (K,
(USZ); Curuyuqui,
Ibaniez, 10 km S de Cotoca,
S).
33 (G); Laguna Ypoa, Chodat
Chodat
0.5 km W
62025'W,
(K, LPB, MO,
Quemado,
Arnold
San Miguelito,
586
(JBSC, LPB, NY);
(USZ); Prov. Cordillera,
del Rio Pilcomayo,
las del Lago Ypacaraf, Mereles

Italia, Compafiia
18055'S,
2625
35186
Prov. Andres
518
& Foster
Gentry
Prov. Andres
(CTES).-BOQuERoN:
orillas
(BR, C).-CAPIrAL:
& Bordas
I. Vargas
Estancia
Fuentes
61?43'W,
Ibaniez, Jardin Botdnico

Nee
Seca, Campo
San Miguelito,
de Izozog, Monte
Banados
Caracore,
(JBSC, LPB);
de Chavez,
1709'S,
62?20'W,
63004'W,
(LPB, USZ);
62?57'05"W,
Bahia Negra,
4198
Estancia
39996
de la Sierra, Werdermann
Schinini
17047'S,
Lorenzo,
& Coimbra
17?49'0"S,
instalaciones
Cruz
del puesto,
(LPB); Prov. Andres
of Santa Cruz on road to Cotoca,
Paurito,
Prov. Nuflo
18050'S,
Laguna
12 km E of center of Santa Cruz on road to Cotoca,
de Santa Cruz,
(USZ, UT);
El Salvador,
CRUZ: Prov. Velasco,
(SI).-SANTA
et al. 19 (LPB); Prov. Cordillera,
Parapeti, Navarro
Nee
4423
ca. 8 km SW of airstrip,
along Rio Parapeti
62047'W,
(LPB); Prov. Luis Calvo,
Ibafiez, Jardin Botdnico

Bohs & Nee
63004'W,
de Santa Cruz,
502
& Marino
Salado,
(E, F, G, GH);
1017
La
oril
(G); Nueva
a Emboscada,
67 (S).-CHACO:
cerca
La
Pedro Lagerenza,
cauce
(CTES); Cerro Le6n,
20026'S,
en Laguna
NY).-CONCEPCION:
Chaco
60015'W,
seco
y prope Concepci6n,
48
34 (G); near Concepci6n,
Chodat
CORDILLERA: Altos,
Schinini
57?22'W,
MO,
NY);
& Mereles
Dist.
inundable
del Rio Negro,
937
La Plata, Berisso,
BAB 52328
pana, Campana,
de Monte,
(K); Laguna
& Beetle
Eyerdam
of the Rio de La Plata, Gibson

1452
Huidobro
ramar, Partido Gral. Alvarado,
Belen,
s.n. (K); Buenos

1212
Aguilar
& Eskuche
Charpin
(BR, MO);
Benitez,
Ruta
(CTES, MO);
Esquina,
25, 29?48'S,
(desvio
a Pto. Gonzalez),
4 km E de Paso de la Patria
11 km NE de Chavarria,
tancia Caaguazd,
riente, Arbo
et al. 6675
et al. 1437
Cristobal
(LPB); Goya,
4 leguas al E de El Pollo,
Rio
Ibarrola
a Bella Vista,
empalme
Depto.
(CTES); Depto.
La Blanca,
30?20'S,
Santa Lucia, Lourteig

Santa Maria,
2595
Pedersen
(CTES); Depto.
383
Depto. Mercedes,
9659
(CTES,
Schinini
10 km S de Corrientes,
G, MO);
& Cristobal
km N de Mercedes,
al. 11829
12666
Martin,
9870
Laguna
(CTES); Depto.
por Ruta
et al. 531
Ibera, Tressens
Cuatia,
10094
Patifno, Pozo Navagdn,

a la Esmeralda,
a orillas
Saladas,
27,
10 km S de Bella
Saladas,
Culantrillar,
Schinini
sobre
de Perugorria,
(CTES); Depto.
Laguna
brazo
Ruta
61 (SI); Depto.
Formosa,
Goya,
23 y Rio
Plowman
Curuzd
24?15'S,
Estancia
2724
Ruta
Picada,
Burkart
8727
25304
& Walter
Eyerdam
del Rio
Cap
et al.
Parana,
75
Trin, Schinini
Schinini
13956
(CTES); Depto.
Depto.
et
& Ahumada
Goya,
San
36 km S
12, 8 km E de Paso L6pez,
Reserva
60?0'W, Arenas
Depto.
Schinini
orilla
(F,
et al.
(CTES); Depto. Mercedes,
(CTES, MO);
Hunziker
Formosa,
Corrientes,
Santa Lucia,
Cuatia,
Ibera, Paso
(CTES, WIS);
de la Laguna
12, Schinini
largo del Rio Parand, Walter

Pilagds,
Ruta
Vista,
Toropi,
inundable
et al. 16868
RIOS: Islas de Victoria,
de Indigenas
Vista,
et al. 11059
la orilla
13 km NW
Mercedes,
Bella
1044
12,
26 km SE de Libertador,
Capital,
de la ciudad, Rio
Schinini
Depto.
(CTES, G, MO);
(A); Depto. Mburucuya,
& Schinini
(CTES); Depto.
del Rio Parand, Bajada Grande,
(B); Constanza,
Reducci6n
Quarin
del
et al. 26742
25 km E de San Luis de Palmar, Quarin
Riachuelito,
orillas
(CTES).-ENTRE
Cordini
Ruta
6 km del
Ruta Nacional
road toMburucuya,
Saladas,
San Roque,
et al. 18929
(CTES, G); Depto.
S, US);
et al. 6915
sobre el Rio Uruguay,
12, Schinini
al S de la ciudad,
Parand), Meyer
comayo
Trin, Estancia
Curuzd
et al. 3580
Vista,
(CTES); Depto.
6 km SW de La Cruz,
de Goya
Bella
San Roque,
(CTES); Depto.
Tressens
un poco
Depto.
6458
Empedrado,
a San Roque,
Krapovickas
Esquina,
(CTES); Depto.
Capital, Meyer
Cnia. Pellegrini,
Schinini
et al. 26861
et al. 27503
Es
bajos del Santa Lucia,
Rio Empedrado,
Depto.
San Roque,
orrilas del Rio Cor
(NY, S); Depto.
costa del Rio Corriente,
Krapovickas
2021
San Cosme,
Paso de La Patria, orillas
San Cosme,
(CTES, G, MO);
(BR, C, G, GH, K, MO,
890
de Bella Vista
Empedrado,
Esquina,
Capital,
San Luis del Palmar, Arroyo
(CTES, US, WIS);
Depto.
(CTES); Depto.
GH, L, S, US); Depto.

ital, Riachuelo,
G, WIS);
(CTES, LP);
13, Ahumada
Depto.
San Roque,
Ibarrola
camino
Bella Vista,
Krapovickas
59?20'W,
et al. 2678
San Roque,
Cercania,
Ahu
Rzepecki,
(CTES); Depto.
approx. 4 km al S del casco,
(SI); Depto.
et al. 27317
et al. 3458
Ruta
et al. 721 (CTES, MO, WIS);
Arbo
islas frente a Esquina,
Esquina,
12, 40 km S de Goya, Krapovickas
Estancia
Ahumada
Esquina,
a casi 20
117, 7 km E
12 km. S de Caa-Catf,
San Miguel,
(CTES); Depto.
Itati, Pueblo
12964
Aldao,
et al. 1449
126, Ahumada
59?15'W,
(AAU, CTES,
Colonia
12, 60 km E de Itati, Arrocera

Ruta
9033
(CTES);
Vista, Ruta
Bella
Paz, 29 km S de Caa-Cati,
(F, L); Depto.
& Crist6bal
et al. 19897
(CTES, G, MO);
Ruta
221
1653
1396
Depto.
3205
Irigoyen
Parana, Krapovickas
Krapovickas
Boelcke
(AAU, G, CTES);
22746
Cerrito,
Zaparinqui,
Schinini
Benitez,
Schinini
Aires,
Margarita
Antequera
Juarez, unos 8 km al SE de Camilo
a Tacuaritas,
camino
of Buenos
Gral. Giiemes,
General
Depto.
sobre Ruta
de Mayo,
Depto.
s.n. (BR); Mi
Resistencia,
(CTES); Colonia
Tranqueras,
de Sauce,
Ahumada
(CTES, MO, WIS);
Primero
(UT).-CORRIENTES:
47 km W
mouth
111 (A, SI); 2 km
outskirts
Depto.
US); Depto.
4230
Dos
Itatf, Ruta Na_.
Depto.
Esquina,
28 km E de Corrientes,
1762
5, Ahumada
(CTES); Depto.
309
& Hilfer
Cam
(NY); Depto.
Rodriguez
(CTES, MO);
(GH, MO,
Estancia
18957
Platanos,
(LP); Depto.
2204
663
12613
(NY); Tigre,
near Cape San Antonio,
76 (K, S).-CHACO:
Depto. Marcos
(CTES).-CORDOBA:
(CTES, F); Depto.
San Cosme,
Depto.
13 km N de La Verde,
del limite con Santa Fe, Hunziker
et al. 858
63
Piccinini
60?33'W,
4026
& Solomon
Jorgensen
et al. BAA
2795
(NY); La Plata, Lefebre
S, US); Quilmes,
Biraben
island
Ruta 74 a 7 km
Plata, Go'mez Sosa 61 (CTES); Azul,
310
Hunziker
al Sud, Venturi
Makal1e,
26?04'S,
Donovan,
de Ruta 27, Ahumada

mada
(C, BR,
10, Solomon
(G); Las Palmas,
20145
2053
Schulz
km al W
Otamendi,
1045
Barracas
Aires,
13 km S de J. J. Castelli,
(CTES); Depto.
Pedersen
on Ruta Provincial
SW of Ensenada
Tweedie
de Campana,
campo
PEDRO: Puerto Rosario,
Dawson
Aj6,
La Plata, Punta Lara-La
s.n. (BM); Depto.
(NY, S); Partido
23?40'S,
La Golond
Estancia
Indio, Boeleke
La Adela,
Los Yngleses,
(BH, G, GH, MO);
(BM, F,
Pyta,
(CTES); Chacoi,
de La Plata, Los Talas, Cabrera
6 (SI); Laguna
Cordini
23064
Punta
Pdo. Magdalena,
(CTES);
Loma
2592
25?13'S,
18517
BUENOS AIRES: Pdo. Maipu,
Argentina.
NY, S, W).
Salado,
Bernardi
(CTES); Villa Hayes,
(CTES, NY).-SAN
133 (F); alrededores
Boffa
Humaita,
HAYES: Estancia
1490
26770
Schinini
et al. BAA 5888
Boelcke
(CTES); Depto.
(CTES); Rio
120, Ruta Trans Chaco, Mereles
(C, K, S, SP, U, US).
de Las Armas,
6721
Curupayty,
Paratodo, Arenas
57038'W,
(BM, G, GH, K, LIL, MO,
(BM).-PRESIDENTE
(BH, NY); Km
25?12'S,
in Rio Paraguay, Woolston
Castro
93
Walter
714 (NY); Colonia Menno,

671
7483
Itagaza, Schinini
(CTES, G).-NEEMBUCU:
Albera,
57?40'W, Hahn
del arroyo
Cud, orillas
24560
Cambacu,
59?35'W, Arenas
rina, 24?55'S,
shores of Rio Paraguay, Hassler
Cnia. Bernal
VOLUME 61
Natural
(F); Depto.
Provincial
(UT); Isla Puentes

216
2000
& Beetle
de
Parand, Parand,
(frente a
(B).-FORMOSA:
Depto.
(CTES);
de Pil
22974
camino
(BH, G, GH, K);
2001
SOLANUM
Depto.
3 km al N de Las Lomitas,
Patinio,
alrededores
de Laguna
de Puerto Velaz,
Morel
305
Depto.
US);
of Riacho
Ruta
3592
(BR, G);
Ing. Judrez, Torres 9 (CTES).-MISIONES:

del Palmar, Hieronymus
SALTA: Ordn, Laguna

Schinini
30853
nas 153
(CTES, NY);
(CTES); Depto.
Depto.
10 km al E de Hickmann,
Hickmann,
Schreiter
US).-SANTA
3419
al. 24857
11214
Gral. L6pez,
Barboza
113 (UT); Depto.

Arroyo
(US); Depto.
Gral. Obligado,
Spegazzini
Barra Arroyo,
Osten
Sierra del Acegud,

video,
Gibert
140
HBG, MO, NY,
616
21883
18512
(K, W);
Carrasco,
(BM).
Japan.
U.S.A.
Fe, Km
590
(CTES, WIS);
camino
Ragonese
El Puerto,
Terribile
LARGO: Rio Negro,
estancia
5281
Aerodromo,
Herter
(GOET, US).-SAN
unos
a
58, Vespucio
Rafaela,
a Coronda,
(A, SI,
Huidobro
et
Hunziker
34, a 1 km al E de la ruta, Moscone
La Capital,
Vieja,
& Are
Venturi 5158
Castellanos,
115, cerca del desvio
e Ibarlucea, Ruta
Quarin
Ordn, Km
orilla del Rio Bermejo,
&
Krapovickas
81 y el Rio Bermejo,
Depto.
762 (CTES); Depto.
Franceschi
(K).-COLONIA:
Osten
(MCNS);
Ordn, Embarcaci6n,
Capital,
(GH, S).-CERRO
Herter
500
(F,NY).
62?53'W, Maranta
23?28'S,
de los Ilanos" entre Ruta
(CTES); Depto.
(K); Depto.
NY, S,
599
12 km de Hickmann,
San Martin,
Novara
entre San Juan y Adelaida
64608
S,SI, U, US, WIS,
Introductions.
bremez
BAB
(C, K, MO,
Schwarz
Barrio Mataco,
(C);
(F, K);
3546
Candelaria,
5087
4085
Pierotti
J. Page, Renvoize
Mocovi,
3058
Quarfn
Sastre, Morel
Las Lomitas,
1346
(BR, MO); Depto.
Puerto Santa Ana,
entre La Salada
Gral. Obligado,
Las Garzas,
Puente
Pilcomayo,
Bermejo,
of Pirane,
11, Km 6, Morel
3739
Isla Pe, Morel
11, 10 km N
3 km W
Pirane,
Ruta
5 km E of Capitdn
Fca. "Marianito
Chapuy,
Depto.
Pilcomayo,
Pilcomayo,
Ruta
Laishi,
Depto.
autopista Rosario-Santa
San Lorenzo,
Reserva,
Mascias,
(B); Depto.
(C); Depto.
s.n. (Z); Depto.
(F, GH, SI); Depto.
San Jeronimo,
(UT); Depto.
Pilcomayo,
antes de liegar a Dragones,
FE: Depto.
(G); Depto.
(B); Depto.
J. Sold (Morillo),
Rivadavia,
San Martin,
(G); Depto.
5174
(SI); Depto.
US, WIS);
655
4072
Morel
Parque Nacional
Pilcomayo,
et al. 2041
(CTES, MO,
Morel
50, Morel
4157
Pierotti
Pirane, Chacras,
Negro,
86 al Km
(SI); Depto.
Fortunato
13196
& Crist6bal
Puente Ceibo, Morel
2 km NW
Pilcomayo,
110
Filipov
58?06'W,
Pirane, Los Matacos,
Depto.
Pilcomayo,
Pilcomayo,
Depto.
25?10'S,
Krapovickas
(L, MO);
(F); Depto.
Blanca,
49
3255
563
a Guadalupe,
Obligado,
Garay,
Uruguay.
Gallinal
Colonia
CANELONES:
et al. A831
(US, Z).-MONTEVIDEO:
JOSE: Barra, Herter
659
La
2139
Ragonese
(US); Depto.
(L).
Palleros,
659a
General
Depto.
&
(US);
Monte
(B, F, G, GH,
Z).
Koyasan,
Wilson
& Suzuki
FLORIDA: Pensacola,
Curtiss
s.n. (A).
6862
Nepal.
Kathmandu,
(BH, G, HBG,
NY, WU,
27?42'N,
85?19'E,
Do
Z).
Solanumglaucophyllumcan be distinguished fromother species in sect.Cyphoman

dropsis by its usually glabrous, glaucous, lanceolate to elliptic leaves with decurrent bases
and short, slightly winged petioles, its rotate-stellate corolla with a relatively long tube
and broad lobes, and its globose, purple-black, glaucous fruits. The stems are smooth and
light yellowish or whitish, and the leaf midribs and margins are often thickened,
inrolled, and whitish or cream-colored. Throughout its range, S. glaucophyllum
exhibits
great morphological
in leaf shape, which ranges from elliptic to
variability, especially
usually
nearly linear, and in corolla and fruit size. Narrow-leaved

plants generally have smaller
flowers and fruits than those with broader leaves, and narrow-leaved
forms seem more
common in the northern part of the geographical range of this species.
This species grows commonly
known
as "varillales"
in flooded or swampy ground, where
or "duraznillales,"
of virgate
stems
it forms thickets,
from spreading
rhizomes
(Okada et al. 1977; Cabrera & Zardini 1978). It is reportedly deciduous in winter (Okada
et al. 1977). The fruits apparently float readily and may be dispersed by water (Nee
37532); they are also eaten by birds (Gibson s.n.).
is of economic importance mainly because it causes a dis
ease, "enteque seco" or "espichamento," of grazing animals (D'Arcy 1974; Wasserman
1974; Morris 1977; Okada et al. 1977). The disease is characterized by calcification of soft
tissues, frequently leading to death, and has caused losses of millions of dollars annually
to livestock ranchers in Argentina (Cabrera 1983). The active principle of S. glaucophyl
lum has been shown to be a vitamin D-like substance that increases calcium and phos
phorous absorption (Wasserman 1974; Morris 1977). Extracts of S. glaucophyllum
are
currently being tested for activity as bone growth factors useful
medicine
in human and veterinary
(Morris 1977; B. Barr, pers. comm.).
50
Solanum
and by having
pubescent,
tubes with narrower corolla
is glabrous,
like most
rather than glaucous
green
corollas,
of southeastern Brazil. Solanum confusum differs
in being commonly
shorter corolla
Solanum matadori
similar to S. confusum of northwestern Argentina
and to S. matadori
from S. glaucophyllum
petioles,
is most
glaucophyllum
and adjacent Bolivia,
VOLUME 61
leaves with
longer
to orange
fruits.
lobes, and yellow
but it has stellate

plants of S. glaucophyllum,
and is known only from Santa Catarina,
leaves,
Brazil.
characters, Child (1986) considered S. glaucophyllum
He removed it to its own section, Solanum sect.
On the basis of morphological

to be unrelated
to sect. Cyphomandropsis.
Glaucophyllum
Child, included within Solanum subg. Solanum. Dottori (1995) investi
of this species and concluded that there were significant
gated fruit and seed morphology
differences in fruit and seed characters between S. glaucophyllum and the other species of
and S. stuckertii) she investigated.

These differences included epicuticular wax covering the fruit exocarp, ventilation cracks
rather than stomata in the fruit surface, and seed coat cells with a distinct shape and pat
exhibits some unique characters and
tern of wall thickening. Although S. glaucophyllum
may
S. fusiforme,
(S. confusum,
represent an isolated clade within the section, its tapered anthers, large angled seeds,
establish it as a member of sect. Cyphomandropsis.
and large chromosomes

A few collections
& Guimardes
21941,
(Arenas 1407, Dobereiner & Tokarnia 794, 797, 800, Hatschbach

301) from ca. 20?S latitude in the Rfo Paraguay drainage
Schaller
to densely puberulent-pubescent
axes and leaves.
in having moderately
the
in
the
but
never
herbarium
segregated
pubescent plants
published the varietal
are anomalous
Morton
name he assigned
phyllum
them. Because
they conform
in all other respects and occur within
to typical representatives
the range of the glabrous
of S. glauco
forms, I do not
them to be taxonomically distinct.

Solanum glaucophyllum has often been confused with S. amygdalifolium Steud. (also
known under the synonyms S. angustifolium Lam., S. persicifolium Mart., S. handelianum
consider
Morong,
amygdalifolium,
stems and ovate-lanceolate
gitudinal
which is sympatric and is found in similar habitats.

however, is a twining or scandent vine with strongly ridged
leaves. The anthers dehisce by broad pores that open into lon
S. brittonianum Morong),
Solanum
slits, and the seeds are much
amygdalifolium
belongs
Jasminosolanum
(Moench) Dumort.,
smaller than those of S. glaucophyllum.
to the dulcamaroid
group of Solanum
Solanum
(sections Dulcamara
Seithe, and relatives) and, though strikingly conver
gent in severalmorphological features,is not closely allied to sect.Cyphomandropsis.

Desfontaines
cultivation
(1829) published
the name S. glaucophyllum
in the Paris Botanic Garden.
name, and a search of the collections

candidates. D'Arcy
Paris Cat. Ann.
I was unable
at P by Sandra Knapp
(1974) cited a specimen
1829, p. 396" and speculated
found this specimen
atMPU,
but with
ing that itwas grown at theMontpellier
atMPU
in reference
to plants
to locate a type specimen
in 1998 failed to turn up any
labelled as "S. glaucophyllum
that itmay be a type. Sandra Knapp
the additional annotation

Botanic Garden
in
for this
Hort.
in 1998
"h.m. Jul 1840," indicat
in 1840 and thus cannot be a type.
Specimens from BR, Fl, and G attest to the cultivation of this species in the Paris Botanic
Garden in the early nineteenth century. Bertoloni (1851) and Dunal (1852) indicate that
both of the names S. glaucophyllum
and S. glaucum were in use to refer to this species.
Because S. glaucophyllum was described from living material, it is likely that no type
specimen exists; however, Desfontaines's
original description
is adequate to fix the appli
cation of the name.
SOLANUM
2001
Kuntze
(1898), Chodat
51
(1918) proposed a number of infraspecific
(1916), and Hassler
taxa based on leaf color and size. All of these taxa are subsumed within
the range of vari
inmy concept of the species.
ation that is encompassed

was cultivated
in French botanical
gardens at Angers, Hy
and early twentieth century, and specimens are also
eres, and Dijon in the mid-nineteenth
known from Japan, Nepal, and the United States (Pensacola, Florida). It is not known
whether
the Asian plants were cultivated or adventive, but probably at least the Nepal col
introduction. D'Arcy (1974) speculates that S. glaucophyllum was
lection was a deliberate

introduced
to Florida
in ship's ballast. It has not been collected
7. Solanum hibernum Bohs, Novon 4: 203. 1994.-TYPE:

Florida, 3 km S of Mataral, valley of Rio Cienega,
Feb 1987, Nee & Coimbra
33970
from Florida
since 1901.
BOLIVIA. Santa Cruz: Prov.

1400 m, 1
18?08'S, 64013'W,
isotypes: K! LPB,
(holotype: US! #3146788;
NY! TEX!).
Small
shrub 0.5-1.5
eglandular
hairs mixed
occasionally
present.
with
Leaves
tall. Stems
some
densely
short-stalked
pubescent
unbranched
branched
dendritically
unit, the blades
per sympodial
6-many
with mostly
glands,
4-11
hairs
cm
long,
ratio ca. 1.5-3.3:1, simple, ovate or elliptic, acute at apex,
cuneate to subcordate at base, chartaceous to subcoriaceous, sparsely pubescent adaxially
with curled, white, usually dendritically branched hairs, these more abundant on midvein
1.5-5.5
cm wide,
length:width
and margin, densely white-pubescent

abaxially with dendritically-branched
hairs, the peti
oles 1-2.5 cm long, densely pubescent with hairs like those of the stem. Inflorescence un
branched (rarely forked), ca. 10-flowered,
1.5-3.5 cm long; peduncle 0.5-1 cm long;
cm long; pedicels
rachis 0.5-2.5
at or near the base,

sparsely
to moderately
erately pubescent,
10-15 mm
long, spaced 1-5 (-10) mm apart, articulated

less than 1mm long; inflorescence axes
leaving scars or short pegs

pubescent with mostly
the radius 3-5 mm,
unbranched
the lobes 1-3 mm
hairs. Calyx
sparsely
long, ca. 2 mm wide,
to mod
often un
equally divided, deltate, with apiculate tips. Corolla violet, chartaceous, stellate, the radius
the tube 2 mm long, the lobes 7 mm long, 2.5 mm wide at base, narrowly triangu
to densely puberulent abaxially, sparsely puberulent adaxi
lar, acute at apex, moderately
9 mm,
ally. Anthers
usually connivent, yellow, lanceolate, 5-7 mm long, 1.5-2 mm wide, abax

to roughened but not obviously papillate, the pores directed distally.
ial surface smooth
style glabrous, cylindrical to subclavate, 7-8 mm long, 0.5 mm in diam

eter; stigma truncate to subcapitate. Fruits 1.5-2 cm long, 1.5-2 cm in diameter, globose,
obtuse or slightly apiculate at apex, glabrous, yellow to orange when ripe; stone cell ag
Ovary glabrous;
gregates very small or absent. Seeds 4-5 mm

with white pseudohairs.
Phenology.
Chromosome
Collected
in fruit in February, March,

Distribution
long, 3-4 mm wide, angled, felty-pubescent

number: 2n = 24. Figs. 21, 22.
in flower in January, February, May,

June, August,
September,
(Fig. 23). Central Bolivia
Santa Cruz; sandy or rocky soil in semiarid
in Deptos.
and December;
collected
and December.
Chuquisaca,
inter-Andean
valleys,
Cochabamba,
often
and
in thorn scrub
communities; 1250-2600 m.
Local names and uses. Bolivia:
Andres huaylla (Spanish; I. Garcfa 71B), bolo bolo

(Nee & Coimbra 33970), kita (Quechua; I. Garcfa 30).-Used
as an antidote for snake
bites in cattle (Nee 46611). Fruits said to not be eaten due to their bitter taste (Nee & Coim
bra 33970).
52
VOLUME 61
E~~~~
FIG. 21. Solanum hibernum.A. Habit. B. Dendritically branched trichome.C. Flower, seen from above.
D. Lateral view of partially sectioned flower.E. Gynoecium. F. Stamen (left to right:abaxial, lateral,adaxial
views). G. Fruit. (Based on greenhousematerial of Bohs 2443.)
SOLANUM
2001
FIG. 22. Flowers
SPEcIMENs ExAmINED
ADDITONAL
km E de Zudaiiez, Muihlbauer
71B (LPB).-SANTA
of central
square
385
inMairana,
Bolivia.
(LPB, USZ);
18007'S,
Saipina,
Nrov. Florida,
63056'W,
scale bar
I cm.
CHUQUISACA: Nrov. Zudafiez,
s.n. (LPB).-COCHABAiMBA:
CRUZ: Nrov. Caballero,
Balcazar
64039'44"W,
of S. hibernum;
53
Bohs
Rodeo
Nrov. Camparo, Rumicancha

Estancia Buena Vista, 6 km NW
Monte Espinoso,
20
(Aiquile), L Garc('a 30,
del pueblo, 18'03'18"S,
to Yunga de Mairana,
2 km NE
along road from Mairana
et al. 2770 (USZ, UJT); Nrov. Vallegrande,
10 km by air
NNW
of Vallegrande,
18'23'S, 6408'W, Nee & Coimbra 33945 (NY); Nrov. Vallegrande,
Quebrada
14 km by air SSE of Mataral,
1801 4'S, 64011IW, Nee & Coimbra 33950 (LPB, NY); Nrov. Florida,
ca. 18o00S, 64'05'W, Nee 35544 (LPB, NY);
to Comarapa,
of Los Negros, along road from Mairana
18'26'S, 6407'W, Nee & Solomon 36559 (LPB, NY);
legrande, 6.5 kmnNNW of center of Vallegrande,
legrande, Lagunillas,
UT); Nrov. Caballero,
Liullucha,
7 km NNE
Nrov. Val
Nrov. Val
5 km N of El Trigal on road toMataral,

18015'S, 64'09"W, Nee 38328 (LPB, NY, USZ,
6.5 km (by road) SW of Mataral on highway
to San Isidro, narrow gorge at confluence
of
and Quebrada Seca, 18008'S, 64'16'W, Nee 46524 (USZ); Nrov. Caballero,
10 km (by road) SE of
on highway
to San Isidro, just NW of turnoff to Pulquina Arriba, along Quebrada Pujio (a dry wash),
tramo entre Santa Rosita, Blanquiscal
17059PS, 64029'W Nee 46611 (USZ); Nrov. Vallegrande,
y San Antonio
18030'32P"S, 64006'12"W,
(2-3 km al S de Vallegrande),
ILVargas et al. 1999 (USZ).
Rio Karikari
Comarapa
This species is most similar to S. lutecalbum of Peru and S. stuckertii of south

central Bolivia and Argentina. Solanum hibernum is distinctive in having strongly discol
orous leaf surfaces with the lower surface densely covered with dendritically branched
hairs. Many specimens of S. lutecalbum also have dendritically branched pubescence,
but the hairs are more evenly distributed on both leaf surfaces. Solanum hibemnum and
54
80
VOLUME 61
60
8
10
~~~
10
F0
23 Drt
FIG
hibesrbnumofS
2 00 400 6 00 800 1000km

jfL/e.Q
.............................................................
of S
ibru
adS.Itoabm
SOLANUM
2001
S. luteoalbum are geographically
55
disjunct, and no hybrids resulted from greenhouse
cross
ing attempts.
In general, S. hibernum can be distinguished
from S. stuckertii by hair morphology
and corolla color. Solanum stuckertii invariably has unbranched hairs and whitish corol
las. All specimens of S. hibernum have dendritically branched hairs (at least on the abax
in Nee 46524 the collector
ial leaf surfaces) and most have purplish corollas. However,
described the corollas as white with a yellow central vein. Polymorphism
for blue and
white flowers, even within a single population, has been noted in species of Solanum sec
tions Petota and Solanum (Correll 1962; Henderson
1974; Edmonds 1977), but the fre
quency of white corollas in S. hibernum needs further investigation. Furthermore, hybrids
have been produced between S. hibernum and S. stuckertii in greenhouse crosses. The F1
S. hibemnum in pubescence characters, but had whitish or very
light lavender corolla lobes with a yellowish central star.
Many individuals of S. hibernum become leafless during the dry season, but remain
conspicuous because of their persistent bright orange fruits that hang like tiny Halloween
pumpkins from the bare branches (M. Nee, pers. obs.).
hybrid plants resembled
8. Solanum hutchisonii
J. F. Macbride,
(Macbride) Bohs, comb. nov. Solanum nitidum var. hutchisonii

Field Mus. Publ. Bot. 13, pt. V-B, no. 1: 209. 1962.-TYPE:
PERU. Amazonas: Prov. Bagua, "St. Julian" hill, on the Rio Utcubamba, Ha
cienda Marerilla near Bagua Grande, 600 m, 1Oct 1957, Hutchison 1490 (lecto
type, here designated: F! #1559946; photo of lectotype, F neg #51398: F!; isolec
totypes, G! K! NY! US!). [The type was cited erroneously in the protologue as
Hutchison 1243.]
Herb or shrub up to 3 m tall. Stems glabrous and white-punctate with cells contain
ing crystal sand. Leaves ca. 4-5 per sympodial unit, the blades 2.5-9 cm long, 1-3 cm
wide, length:width ratio 1.75-3.5:1,
simple, elliptic-ovate, acute at apex, rounded to sub
cordate at base, subcoriaceous or somewhat succulent, glabrous and white-punctate
adax
ially and abaxially, the petioles 1-2.5 cm long, glabrous. Inflorescence unbranched or
rarely forked, 5-25-flowered,
1.5-6 cm long; peduncle 1-4 cm long; rachis 0.2-2 cm
long; pedicels 5-15 mm long, 15-20 mm long in fruit, unevenly spaced 1-5 mm apart, ar
ticulated at base, leaving scars on the axis; inflorescence glabrous. Calyx glabrous abaxi
for a few sparse hairs at tips of lobes, veiny and sand-punctate, irregularly
splitting, the radius 3-6 mm, the lobes 2-5 mm long, 1.5-3 mm wide, deltate, with apic
ulate tips. Corolla purple, chartaceous, stellate, the radius 8-15 mm, the tube 2-3 mm
long, the lobes 6-12 mm long, 2-4 mm wide at base, triangular-ovate, acute at apex,
glabrous abaxially and adaxially except for the ciliolate margins. Anthers not connivent,
color unknown, narrowly triangular, 5-8 mm long, 1.5-2.5 mm wide, abaxial surface
smooth to roughened but not obviously papillate, the pores directed distally. Ovary
ally except
glabrous; style glabrous, cylindrical, 6-11 mm long, 0.5-1 mm in diameter; stigma trun
cate. Fruits 1-2.5 cm long, 1-2.5 cm in diameter, globose, obtuse at apex, glabrous, dark
colored when ripe; stone cell aggregates absent. Seeds 4 mm long, 3 mm wide, angled,
whitish
pubescent and reticulate. Chromosome number unknown. Fig. 24.

Phenology. Collected
in flower in February, October, and November;
fruit in February.
Distribution
Cajamarca;
(Fig. 8). Northern Peru, Rio Marafioin valley

m.
in Deptos.
collected
Amazonas
dry slopes; 450-600
in
and
56
FIG. 24. Solanum

hutchisonii.
A. Habit.
E. Stamen
B. Flower,
VOLUME 61

lateral, adaxial
view).

(Based on Hutchison
F. Fruit.
1490.)
SOLANUM
2001
ADDITIONAL SPECIMENS EXAMINED. Peru.

al Rfo Marafn6n,
convergen
con Marafion,
Sagdstegui
Sdnchez
5862
Vega 3970
57
AMAZONAS: Prov. Bagua,

(F, NY).-CAJAMARCA:
(US); Prov. Chota, Huambos,
9 Nov
cerca a Corral Quemado,

Prov. Jaen, confluencia
1956, Soukup
4531
laderas que
del Chamaya
(US).
Solanum hutchisonii can be distinguished from the other species in sect. Cyphoman
dropsis by its fleshy subcordate leaves and lack of pubescence. Some other species of the
and S. fusiforme, are glabrous or nearly so, but they
section, such as S. glaucophyllum
occur far to the south in Brazil, Argentina, and adjacent areas.
Macbride
(1962) cited the type of S. nitidum var. hutchisonii as Hutchison 1243. It is
obvious from herbarium notations
that the correct type number of S. nitidum var.
1490, and that the citation inMacbride's
protologue is an error.
S. nitidum var. hutchisonii should not be considered as a synonym of S. ni
tidum, contrary to Knapp's (1989) treatment. The collection Hutchison 1243 is S. nitidum
hutchisonii
is Hutchison
Consequently,
Ruiz & Pav., a species of the S. nitidum group of Solanum sect. Holophylla
(Knapp 1989).
tomolecular data, S. nitidum belongs to a clade consisting of S. dulcamara L.,
S. wallacei
(A. Gray) Parish, and relatives, and is unrelated to sect. Cyphomandropsis
According
(Bohs & Olmstead,

9. Solanum
in press; Fig. 1).
luridifuscescens
Bitter, Repert. Spec. Nov. Regni Veg. 12: 466. 1913.

Cyphomandra velutina Sendtner inMartius, Fl. bras. 10: 120, t. 17. 1846, non
Solanum velutinum Dunal, 1814. Pionandra velutina (Sendtner) Miers, Ann.
Mag. Nat. Hist. 15, ser. 2: 199. 1855.-TYPE:
BRAZIL. Goias: ad Fazendam S.
Cruz da Donna Tereza, Pohl 3455 (lectotype, here designated: W!; photos of lec
totype, US neg. 8531: F! GH! LL! NY!, US neg. 8758: GH! LL! NY!; isolecto
types: BR! F! G, M! W!).
Cyphomandra glaberrima Dusen, Ark. Bot. 9(5): 19. 1909.-TYPE:

BRAZIL. Rio de
Janeiro: Serra do Itatiaia, ca. 1200 m, Oct 1903, Dusen 2057 (lectotype, here des
ignated: S!).
Perennial
herb or shrub up to 2 m tall. Stems nearly glabrous to densely puberulent

stalked glandular and unbranched eglandular hairs. Leaves 4-5 per sym
podial unit, the blades 4.5-23 cm long, 2-7 (-9) cm wide, length:width ratio 1.5-3 (-4): 1,
simple, elliptic to elliptic-ovate,
acute to acuminate at apex, cuneate to decurrent at base,
pubescent with
chartaceous, nearly glabrous to moderately puberulent-pubescent

adaxially and abaxially
with unbranched eglandular hairs, the pubescence denser on veins, the petioles 0.3-2.5 cm
long, sparsely to densely puberulent-pubescent.
Inflorescence unbranched, 5-12 (-25)
flowered, 4-20 cm long; peduncle 2-6 cm long; rachis 2-18 cm long; pedicels (10-)
mm long, 20-30 mm long and thickened distally in fruit, spaced 3-14 mm apart,
articulated at or slightly above the base, leaving pedicellar remnants up to 1mm long; in
florescence axes sparsely to densely puberulent-pubescent
with hairs like those of the
stem. Calyx sparsely tomoderately puberulent-pubescent,
the pubescence denser on mar
20-30
gins and tips of lobes, the radius 4-6 mm, the lobes 1.5-3 mm long, 1.5-2.5 mm wide,
deltate, narrowed to acute or acuminate tips. Corolla white to purple, subcoriaceous, stel
late, the radius 13-18 mm, the tube 3-4 mm long, the lobes 9-15 mm long, 3-5 mm wide
at base, narrowly triangular, acute at apex, sometimes with a small subapical projection,
glabrous to moderately puberulent-pubescent
abaxially, with the pubescence denser dis
tally, glabrous adaxially. Anthers connivent or free, yellow, lanceolate, 7-9 mm long,
1.5-2 mm wide, abaxial surface with an obvious band of scaly papillae, the pores directed
58
VOLUME 61
distally. Ovary glabrous; style glabrous, cylindrical, 5-8 mm long, 0.5-1 mm in diameter;
stigma truncate. Fruits 1-2 cm long, 1-2 cm in diameter, globose, obtuse at apex,
glabrous, the color when ripe unknown; stone cell aggregates absent. Seeds 3-4 mm long,
2-3 mm wide, lenticular, densely puberulent with whitish pseudohairs. Chromosome
number unknown. Fig. 25.
Phenology. Collected in flower in February and June through November; collected in
fruit in October, November, and February.
Distribution
(Fig. 18). Eastern to southeastern Brazil (Espirito Santo, Goias, Minas
Gerais, Parana, Rio de Janeiro, and Sao Paulo); moist forest, often in wet or swampy
areas; 1100-2650 m.
Guimardes
Mpio.
Serra, perto de Vit6ria,
rani et al. 164 (SP).-MINAS

Kuhlmann
Maurfcio
8302
2597
ESPIRITO SANTO:
(UT); SE slopes of Serra da Capara6, Mexia
46879
Esta,co
Serra do Gongo,
(SP); Ervdlia,
Ferreira Fernandes,
(L, US); Mpio.
JANEIRO: Teres6polis,
Biol6gicado
V;eira 630
Campina
Grande
Serra dos Orgaos,
Serra do Coelho
et Nova
Tiete, Rio Claro,
Loefgren
Mestre
GERAIS: Serra do Capara6,
Friburgo
5880
arredores
4039
16926
7 km do Carego
Mpio.
do Sul, Serra Capivari
Grande,
Pedra do Frade, Brade
an Conego,
Glaziou
(US); Serra dos Orgaos,
13084
fazenda
16604
(RB, UT);
Alvaro,
Pi
propriedade
de
Grande, Hatschbach
20327
Hatschbach
&
(G, S, US);
Serraria Boa Vista,
das Olivieras,
Cerro Azul, Morro
(BR, C, F [fragment],
Ule 4314
4043
do Mestre
(RB); Sapucai-Mirim,
(arraial),
(GH, VIC).-PARANA:
Hatschbach
Freire,
NY, US),
subida para o Morro
Alvaro,
Brade
de Muniz
(BM, GH, MO,
environs
(CTES, Z).-RIO
of Rio
DE
de Janeiro,
PAULO: Alto
G, K).-SAO
(HBG).
This species is distinctive in having rather large elliptic leaves with a tapered base and
a dense obvious band of scaly papillae on the abaxial anther surfaces. Vegetative pubes
cence is quite variable in S. luridifuscescens; plants range from nearly glabrous to densely
pubescent on axes and leaf surfaces. Solanum luridifuscescens
is superficially similar to
and has been confused with S. melissarum of Solanum sect. Pachyphylla.
It can be dis
tinguished from the latter species by its stellate corollas with very short tubes and by its
papillose
anther surfaces.
The epithet velutina
cannot be used when Cyphomandra
velutina
is transferred to
Solanum, because the name is already occupied by S. velutinum Dunal in Poiret (1814).
Bitter provided a new name in Solanum, S. luridifuscescens, which refers to the dark
brown tobacco color of the dried plants. He pointed out several discrepancies
in Sendt
ner's original description (Sendtner 1846) and provided a new diagnosis based solely on
the collection Ule 4314 from the Organ Mountains near Rio de Janeiro.
Dusen (1909) did not specify a holotype for C. glaberrima;
the protologue, has been chosen as the lectotype.
his specimen
at S, which
matches
10. Solanum luteoalbum Persoon, Syn. 1: 221. 1805. Solanum pubescens Ruiz & Pavon,
Fl. peruv. 2: 36, pl. 169, fig. b. 1799, non Solanum pubescens Willdenow,
1794.
Cyphomandra
luteoalba (Persoon) A. Child ex Bohs, Fl. Neotrop. Monogr. 63:
154. 1994.-TYPE:
PERU. "In Peruviae nemoribus ad Cuchero tractus, floret Jan
uario et Februario," Ruiz & Pavon s.n. (lectotype, here designated: MA; frag
ment of lectotype: F!; photo of lectotype, F neg. 29723: F! GH!).
Solanum semicoalitum Bitter, Repert. Spec. Nov. Regni Veg. 12: 463. 1913.-TYPE:
ECUADOR. Azuay: hills above Gualaceo, Sep 1864, Jameson s.n. (holotype: W!;
photo of holotype, F neg. 33110: F! G!).
Solanum luteoalbum var. tunya Macbride, Field Mus. Publ. Bot. 13, pt. V-B, no. 1:
206. 1962.-TYPE:
PERU. CUZCO:Ollantaytambo,
ca. 3000 m, 18 May 1915,
Cook & Gilbert 804 (lectotype, here designated: US! #603970).
SOLANUM
2001
59
ci~j
m
FIG. 25. Solanum
tioned
; C
B
luridifuscescens.
E. Stamen
,i:C D
B. Flower, seen from above. C. Lateral view of partially sec

(left to right: abaxial, lateral, adaxial views). F. Fruit. (Based on: A,
A. Habit.
Brade 16926;B-F, Hatschbach 20327.)
60
VOLUME 61
Shrub 1-3 m tall. Stems moderately to densely pubescent with eglandular unbranched
or dendritically branched hairs and also with short-stalked glands; tips of shoots some
times densely glandular. Leaves 6 tomany per sympodial unit, the blades 3.5-18 cm long,
1.5-7.5 cm wide, length:width ratio 2-3:1, simple, elliptic or ovate-elliptic,
acute to
acuminate at apex, cuneate to slightly decurrent at base, chartaceous, moderately
to
densely pubescent adaxially and abaxially with eglandular unbranched or dendritically
hairs and often also with short-stalked glands, the petioles 1-4.5 cm long,
densely pubescent with hairs like those of the stem. Inflorescence unbranched, forked, or
4-10 (-15) cm long; peduncle
sometimes further branched, ca. (5-) 10-25-flowered,
3.5-6.5 cm long; rachis 1-5 (-10) cm long; pedicels 10-25 mm long, 10-35 mm long in
fruit, spaced ca. 2-10 (-15) mm apart, articulated at base; inflorescence axes moderately
branched
to densely puberulent with eglandular unbranched and/or dendritically branched hairs and
often also densely beset with glands. Calyx moderately puberulent with glandular and eg
landular hairs, the radius 3-4 mm, the lobes 1.5-2 mm long, 1.5-2 mm wide, deltate, often
abruptly narrowed into acuminate tips. Corolla purple (rarely white), chartaceous, stellate,
the radius 11-20 mm, the tube 3-4 mm long, the lobes 8-17 mm long, 3A mm wide at
base, narrowly triangular, acute at apex, sparsely to densely puberulent abaxially with eg
landular unbranched and/or dendritically branched hairs and often also glandular hairs,
nearly glabrous adaxially except for some hairs on midrib and near apices of lobes. An
thers usually connivent, yellow, lanceolate, 5-8 mm long, 1.5-2 mm wide, abaxial surface
smooth to roughened but not obviously papillate, the pores directed distally. Ovary
glabrous; style glabrous or sometimes with a few hairs, cylindrical, 7-9 mm long, 0.5 mm
in diameter; stigma truncate. Fruits 1.5-2 cm long, 1.5-2 cm in diameter, globose, obtuse
at apex, glabrous, orange or red when ripe; stone cell aggregates absent. Seeds 5-6 mm
long, 4-5 mm wide, angled, smooth to rugose. Chromosome
number: 2n = 24. Figs. 26,
27, 28.
Phenology.
months
Collected
in flower
in August
through March;
collected
in fruit in all
except July.
Distribution
(Fig. 23). Andean slopes from southern Ecuador to southern Peru; grav
elly or rocky slopes and cliffs tomoist river valleys; 2200-3300 m.
Local names.Peru:Pajarito(Macbride& Featherstone 1038), tunya(Herrera3346),

tunya-tunya
(Marin 159, Cook & Gilbert
REPRESENTATIVE SPECIMENS. Ecuador.

tidero sitio denominado
02047'S,
(BM, F,WIS);
Caribamba,
WIS),
al. 601
&
Jaramillo
ca. 1 km above
of two rivers,
Iltis et al. 591
(MO); Prov. Andahuaylas,
mazd,
ca. 16 km S of Contumaza
6088
(NY, UT),
Gilbert
387a
Cuzco, Herrera
Urubamba
921 (W); valley
(BM, F, K, WIS);
valley
valley
of Pincos,
(MO),
valley
of Rio Urubamba,
just SE of Urcos
Urubamba,
near Ollantaytambo,
at Km
10711
Cachil,
et al. 15117
on road to Ollantaytambo,
of Rio Urubamba,
57
ca. 6 km W
4 km below
from Cuzco,
Rio Vilcanota
valley,
13?40'S,
13?l5'S,
of Rio
5 km NW
Aymaraes,
Colcachaca
Iltis et al. 589
(BM, F,
of Chalhuanca,
Iltis et
near Catholic
14?03'S,
Church,
(UT).-Cuzco:
3346
of Calca
(F,MO,
(F); Prov. Calca,
(6 km E of Yucay),
Urubamba,
71?40'W,
Prov. Contu
& Sagdstegui
Ollantaytambo,
et al. 19785
Iltis & Ugent

Iltis & Ugent
72?20'W, Knapp
& Mallet
Cook
NY);
bottom
&
near
of Rio
Iltis et al. 923
1141
1213
6465
Iltis
Nuiiez
73'15'W,
Dillon
78?46.5'W,
A. Gentry
at
Iltis et al. 528
(F, GH, K).-CAJAMARCA:

07?24'S,
de Cuenca,
on road to Caraibamba,
(WIS),
of Abancay
Herrera
Ollantaitambo,
Pachar,
ENE
(NY); par
sector NW
from Chalhuanca
Iltis et al. 584
(F); Challhuanca,
Bosque
Sagdstegui
valley
of Rfo Chalhuanca,
Stork & Horton
en route to Cascas,
56 on road Cuzco-Urubamba
Urubamba,
APURIMAC:
of Rfo Apurimac,
1872
Camp E-4675
de agua,
ca. 15 km (air) S of Chalhuanca,
(BM, F,WIS),
Curahuasi, Marin
13 km N of Pisaq
valley, Km
Peru.
de Oro,
Represa
km (air) S and SW
of Acobamba,
et al. 9089
Lbpez
(NY);
(NY).
5-15
Curahuasi,
et al. 736 (WIS); Prov. Abancay,

7137
140
Iltis et al. 582
(WIS); vicinity
(WIS); Prov. Abancay,
Guandum-La
junction with Rfo Chalhuanca,
of Rio Chalhuanca,
valley
junction
km N of Sevilla
AZUAY: 4-6
Llantera-Chiquintad-Saucay,
Ortiz
79008'W,
Cotarusi-Colca
804).
(WIS); Prov.
(WIS);
Prov.
(F, K, MO, NY);
2001
SOLANUM
61
NI~
II
-4.
CN
FIG. 26. Solanum
luteoalbum.
E. Stamen
material of Bohs 2336.)
A. Habit.
B. Rlower,

lateral, adaxial
views).

(Based on greenhouse
F. Fruit.
WIN
......
.....
JI:
-er
MW
'T:
.............
.............
. .........
. ... . . ......
....... ..........
..... . .
.... .......
oRX
. .........
..
.........
.............
......
....... ....
.........
...
............
.
..........
i...X
............
..
..........
FIG. 27. Solanum
luteoalbum.
A. Habit;
scale bar
2 cm. B. Fruits,
showing
rough du
2001
FIG. 28. Inflorescence of S. lu::oaGbum
Prov. Calca,
.~~~~~~_
Pisac,
Mann 159 (F); Huayoccari
63
showing
stellate
.X
!_#
*l~. .. ......
..._:,......
SOLANUM
to Yanacocha
.
.c
::o:l
. .cp
. :.. ......
.
Urubamba,
NW.fromCuzco;
3,,i,S
74....'W
~~~~~~~~,
.~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
.,...
...........I
Sw... ,
........
,: ... ... ... ......^X-^
FI.
2. Inlrscec
of_ 'S.ueolu
Naiihez et aL. 7040
(F, NY);
Yaurisque,
Prov. Urubamba,
67 km
from Cuzco,
72'16'W,
Naiitez & Bengoa
man & Davis
9325
(WIS);
Tarma, Macbride
Huamba,
Boeke
Cano
3173
from Rio
Calicanto
(NY); Nrov. Paucartambo,

Pisac, hillsides
scal
brIcm
on road from Cuzco

to Chacchapara,
near Paucartambo
to Paruro, Nunfez 7361 (MO);

Muris
and environs,
13'16'S,
on road to Abra Acanacu,
Plow
ruins of Ollantaytambo,
1.5 km above
& Featherstone
1557
around ruins, Solomon 3030 (F,MO); Nrov. Anta,

Urco, Vargas 686 (GH); Nrov. Urubamba, Yanahuara, Var
near HuAnuco, Kanehira
West 6476 (GH, MO).-HuANuco:
79
222 (BM, F, WIS);
(W of) Palca, Iltis et al. 221 (BM, F, NY, WIS),
(GH); Nrov. Calca, Hacienda
Nrov. Tarma,
(GH).-JUNIN:
8667
Paruro, SW of Cuzco
coolas
(F, K, GH); Rio Urubamba,
Vargas 200
Limatambo,
gas
4913
S~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..
.....................
shwintt
(NY).-PuNo:
(NY, TEX).-Department
1038
(F); Mito,
Nov.
Macbride
Carabaya
unknown:
3273
Ollachea,
Yanano, Macbride
(F, G).-PIURA
across
San Gaban
3795
Nrov. Ayabaca, Bosque

iver from town Boeke
de
&
(F).
lutecalbum is similar to S stuckertii and S. hibenum.

all have elliptic
stellate
and
Solanum
seeds.
luteoal
leaves,
corollas, orange globose fruits,
large angled
bum can be distinguished
from S. stuckertii by its purple rather than white corollas and
more northerly distribution. Hairs of S stuckertii are exclusively
unbranched whereas
those of S. luteoalbum can be either unbranched or dendritically branched. Solanum lu
teoalbum and S. hibernum are very similar morphologically.
Both species have purple
Solanum
corollas and are often covered with dendritically branched trichomes, but in S. hibernum
the leaf surfaces are strongly discolorous due to the dense whitish covening of dendriti
cally branched hairs on the lower surface, whereas the pubescence of S. luteoalbum is
more or less evenly distributed.
Solanum luteoalbum is also similar to pubescent
forms of S. confusum, but can be dis

tinguished from the latter by its stellate and usually deep purple corollas shorter and often
branched trichomes, and its more northerly geographical range.
64
Peruvian
specimens
have an obvious,
of S. luteoalbum
often dense
VOLUME 61
from the Departments
of Apurimac
indumentum of forked or dendritically
and Cuzco
branched
trichomes
especially on young parts. Macbride (1962) differentiated

these collections
as S. luteoalbum var. tunya. Plants from further north in Deptos.
Huanuco and Junin usually have a few branched hairs on the axes and leaf midribs, but
on the axes and leaf undersides,
as in the collections
these are not nearly as noticeable
from southern Peru. Collections
Cajamarca and Piura in Peru, as well as the types of S. pubes

have exclusively
cens and S. semicoalitum,
unbranched hairs and numerous stalked
from Ecuador and Deptos.
these variants merit
glands. Whether
is debatable. Herbarium
taxonomic recognition
imens from these areas do not have any obvious morphological
spec
except for in
differences
dumentumcharacters.
and Pavon's
Ruiz
dropsis. A specimen
and plate of S. pubescens
description
atMA
the lectotype. The only discrepancy is their

the plants listed here have purple
and Pavon's Flora (1799) and here designated
color as "albo-lutescens";
of the corolla
description
it in sect. Cyphoman
place
is very similar to plate 169 in Ruiz
annotated as S. pubescens
corollas.
A single specimen
of Cook & Gilbert
the type collection of S. lu

teoalbum var. tunya Macbride was examined from US. This specimen is not annotated by
Macbride, and he gives no indication of the type location in his protologue. Nonetheless,
since this is the only specimen
804 representing
that has been located of this number, it has been designated
as the lectotype.
11. Solanum matadori Smith & Downs, Phytologia 10: 432. 1964.-TYPE: BRAZIL.
Santa Catarina: Rio do Sul, Alto Matador, Araucaria
forest, 800 m, 16 Oct 1958,
7254 (holotype: US! #2323440;
isotypes: HBR, L! NY!).
Reitz & Klein
Shrub up to 2 m tall. Stems glabrous. Leaves many per sympodial unit, the blades
6-16 cm long, 1-2.5 cm wide, length:width ratio 4-8:1, simple, narrowly elliptic, acute at
apex, tapered at base, somewhat fleshy, glabrous adaxially and abaxially, the margins cil
thickened and inrolled,
iolate, often
branched,
cm long; pedicels
leaving
or more,
15-30-flowered
5-25 mm
the petioles
8-15
forked hairs at base of pedicels.

forked hairs, the pubescence
Calyx
4-6
Inflorescence
cm long; rachises 4-7
axes nearly glabrous, with

glabrate
denser on margins
long, 2-3 mm wide,
violet, chartaceous,
cm long, glabrous.
long in flower, spaced 2-10 mm apart, articulated at the base,
scars on the rachis; inflorescence
lobes 2-3 mm
0.5-2
cm long; peduncle
stellate,
a few simple or
to sparsely pubescent
with
simple or
and tips of lobes, the radius 3-5 mm,
deltate, narrowed
the radius 11-17 mm,
to acute or acuminate
the tube 3-6 mm
the
tips. Corolla
long, the lobes 7-11
mm
long, 3-4 mm wide at base, triangular, acute at apex, sparsely to moderately pubes
cent abaxially and adaxially with simple and forked hairs. Anthers connivent or free, color
unknown,
lanceolate, 7-9 mm
long, 2-3 mm wide,
abaxial surface with an obvious band
of scaly papillae, the pores directed distally. Ovary glabrous; style glabrous, cylindrical,
7-8 mm long, 0.5-1 mm in diameter; stigma truncate. Fruits and seeds unknown. Chro
mosome number unknown.
Phenology.
Collected
Distribution
(Fig.
in flower in October.
18). Southeastern
Brazil
(Santa Catarina);
Araucaria
forest;
800-1200 m.
Local names. Brazil:
Joa manso
de f6lhas compridas,
jua (Smith & Downs
1966).
SOLANUM
2001
ADDITIONAL SPECIMENS EXAMINED. Brazil.

pos do Areao,
Pabst
6086 & Pereira
6259
65
SANTA CATARINA: Alto
da Serra do Espigao,
pr. Vale Cam
(F, RB).
is known from only two collections from Santa Catarina, Brazil.

Solanum matadori
similar to two other southeastern Brazilian species, S. fusiforme and S. luridi
fuscescens. All three species have the abaxial anther surface scaly or papillose; these
It is most
papillae
are most
similar
in having
Solanumfusifonne
in S. luridifuscescens. Solanum matadori and S. fusiforme are

and in being nearly glabrous vegetatively.
inflorescences
has distinctive elongated ovaries and fruits. Fruits of S. matadori are
obvious
branched
but the ovaries appear to be elongated. Solanum matadori differs from S.

in its narrower and strictly simple leaves, pubescent corollas, and glabrous peti
oles. Solanum matadori, unlike S. luridifuscescens, has very narrow leaves, branched in
florescences, and corolla lobes that are pubescent adaxially as well as abaxially.
unknown,
fusiforme
12. Solanum
Bohs, nom. nov. Cyphomandra cornigera Dunal in DC., Prodr.

13(1): 401. 1852, non Solanum cornigerum Dunal, 1852. Pionandra cornigera
(Dunal) Miers, Ann. Mag. Nat. Hist., ser. 2, 15: 199. 1855.-TYPE:
BRAzIL.
Santa Catarina, Gaudichaud 160 (lectotype, here designated: P!; photo of lecto
pelagicum
type, F neg. 39255: WIS!;
isolectotype:
P!; fragment of lecto- or isolectotype:
F!).
Smith & Downs, Phytologia 10: 436, pl. 9, fig. 7. 1964, non
ex Nees, 1843.-TYPE:
BRAZIL. Santa Catarina:
Porto Belo, Bombas, strand, 1-5 m, 31 Mar 1957, Smith et al. 12322 (lec
Cyphomandra
maritima
Solanum maritimum Meyen

Mpio.
totype, here designated: US! #2423787;
isolectotypes:
HBR, R, US!).
Herb or shrub 0.5-1 m tall. Stems moderately

to densely pubescent with forked or
dendritically branched eglandular hairs and some scattered long-stalked glands. Leaves
4-6 (-many?) per sympodial unit, the blades 2-10 cm long, 1.5-10 cm wide, simple and
to pinnately (2-) 3-9 (-11) compound, the simple leaves with length:width
elliptic-ovate
ratio ca. 1.5-2.5:1,
the compound leaves with the upper lateral leaflets often basiscopi
cally decurrent, acute at apex, truncate to subcordate at base, chartaceous to subcoria
ceous, moderately pubescent adaxially with unbranched or dendritically branched hairs,
to densely pubescent abaxially with dendritic hairs and often with scattered
moderately
stalked glands, the petioles 1-4 cm long, densely dendritic-pubescent.

Inflorescence un
branched or forked, ca. 7-40-flowered,
2-12 cm long; peduncle 1-5.5 cm long; rachis
1-10 cm long; pedicels ca. 10-15 mm long, spaced 1-15 mm apart, articulated at or
slightly above base, leaving pedicellar scars up to 1mm long; inflorescence axes moder
ately to densely puberulent with dendritic hairs and sparser stalked glands. Calyx moder
ately puberulent with glandular and eglandular hairs, the radius 2-5 mm, the lobes 1-3
mm long, 1-3 mm wide, deltate, acute at tips. Corolla purple, lilac, or bluish, chartaceous
tomembranaceous,
stellate, the radius 8-13 mm, the tube 3 mm long, the lobes 5-10 mm
long, 2.5-4 mm wide at base, triangular, acute at apex, sparsely tomoderately puberulent
abaxially, glabrous to sparsely puberulent adaxially. Anthers often connivent, yellow,
lanceolate to narrowly oblong, 4.5-6 mm long, 1-2 mm wide, abaxial surface smooth to
roughened but not obviously papillate, the pores directed distally. Ovary glabrous; style
glabrous, cylindrical, 4-5 mm long, 0.5-1 mm in diameter; stigma truncate to subcapitate.
Fruits 1-2.5 cm long, 1-2 cm in diameter, ovoid or ellipsoidal, obtuse at apex, glabrous
(dendritically
pubescent,
fide Smith
& Downs,
1966),
color
unknown;
stone
cell
66
VOLUME 61
D
FIG. 29. Solanum
of partially
sectioned
(Based on: A, Occhioni
pelagicum.
A. Habit.
5307; B-F, Hatschbach
B. Compound
F. Stamen
& Forero
leaf. C. Flower,

40366;
seen from above. D. Lateral

lateral, adaxial
G, Reitz & Klein
views).
6770.)
view
G. Fruit.
2001
SOLANUM
aggregates
present. Seeds
ca. 3 mm
67
long, ca. 2 mm wide,
strongly
flattened,
rugulose.
Chromosome
number unknown. Fig. 29.

in flower in July and October through December; collected
Phenology. Collected
fruit inMarch, July, October, and December. A peak of flowering and fruiting occurs
in
in
October.
Distribution
(Fig. 18). Southeastern
coast (restinga vegetation)

Local
Brazil (Santa Catarina);

and inland in pastures; 0-300 m.
names. Brazil: Baga de urubui (Reitz & Klein
(Smith & Downs
& Kozicki
27529
(BH, C); Mpio.
LL, NY, Z); Praia de Laguna,
Hoehne
et al. 5817
Santa Catarina,
Klein
Bresolin
(US); Garopaba,
Laguna,
Retiro,
6386
6770); baga de bugre da praia
do Norte,
Klein
Reitz & Klein
Solanum pelagicum
24447
SANTA CATARINA: Garopaba,

Laguna,
do Sul, Klein
& Bresolin
Reitz
6770
8834
1732
Eskuche
Praia do Gi, Hatschbach
Itajai, Praia Braba, Hunt
(US);
(US); Pantano
4 (US); Orleaes,
Reitz & Klein

Braco
along
1966).

Hatschbach
sandy beaches
& Bresolin
1251-11
(Z); Laguna,
40366
(BH, C, CTES,
& Forero
6360
6289
(US); Pantano
(US); Morro
(US); Barra da Lag6a, Rio Vermelho,
(US); Campeche,
(NY, US); mainland
Ilha Santa Catarina,
opposite
Desterro,
Ule
do Sul, Ilha de
das Pedras, Klein

Occhioni
Reitz 5081
5307
&
(F);
(US); Bom
s.n. (HBG, US).
can be distinguished
by its often pinnately compound leaves with

subcordate bases, stalked glands on the stem and inflorescence axes, stone cell aggregates
in the fruits, and by its specialized habitat and restricted distribution. The dendritic hairs
of this species are similar to those of S. cylindricum.
Smith and Downs
(1966) distinguished
basis of leaf morphology

had pinnately
and ecological
between C. cornigera
characteristics.
According
and C. maritima
on the
to them, C. cornigera
compound
leaves, whereas their new species C. maritima had simple or

leaves. In reality, both variants occupy the same habitat and localities,
and some specimens have both simple and pinnately compound leaves on the same plant,
which argues against recognition as two separate taxa.
rarely few-lobed
Smith and Downs

was not obvious
(1966) describe the fruits as minutely

in the herbarium material I examined.
pubescent,
but this character
Three sheets of the type collection of C. cornigera Dunal (Gaudichaud 160) exist at
P. Two are annotated by Dunal, and one of these has been chosen as the lectotype of this
name.
Two sheets at US are annotated by Smith and Downs
them, #2423787,
as "C. maritima-Type."
One of
is here selected as the lectotype.
Solanum pelagicum occupies the unusual habitat of beach dune formations (restinga)
along the coast of southeastern Brazil. The herbarium label of Hunt 6360 indicates that
the plants were collected "above the high water mark." Several other collectors noted find
ing this species along forest margins and cleared pastures. A new name is required be
cause S. cornigerum and S. maritimum are already occupied. The epithet "pelagicum,"
from Greek 2wXocyo; (sea) has been chosen in reference to the seaside habitat of this
species.
13. Solanum
stuckertii
dra stuckertii
Bitter, Repert. Spec. Nov. Regni Veg.

(Bitter) D'Arcy,
Ann. Missouri
12: 461. 1913. Cyphoman
Bot. Garden
59: 277. 1972.

TYPE: ARGENTINA. Tucumain: Burruyacu', 20 Apr 1910, Stuckert 21589 (lecto
type, here designated: G!; photo of lectotype, F neg. # 23159: F!; isolectotypes:
CORD! GOET! [fragment]).
68
VOLUME 61
stuckertii var. angustifrons Bitter, Repert. Spec. Nov. Regni Veg. 12: 463.
1913.-TYPE: ARGENTINA. Cordoba: Los Cocos, Punilla, Stuckert 16234 (lecto
type, here designated: CORD!).
Solanum stuckertii var. atrichostylum Bitter, Repert. Spec. Nov. Regni Veg. 12: 463.
ARGENTINA. Tucuman: Burruyacui, 1 Apr 1900, Stuckert 9138
1913.-TYPE:
(lectotype, here designated: CORD!; isolectotype: GOET! [fragment]).
Solanum stuckertii var. trichostylum Bitter, Repert. Spec. Nov. Regni Veg. 12: 463.
Solanum
ARGENTINA. Cordoba: Salto, Rio Tercero, Stuckert 915a (lecto

type, here designated: CORD!; isolectotype: GOET! [fragment]).
Solanum stuckertii var. obrutum C. V. Morton, A Revision of the Argentine Species
of Solanum, p. 192. 1976.-TYPE: ARGENTINA. Salta: Depto. La Vinia, Coronel
Moldes, ca. 1200 m, 2 Feb 1941, Hunziker 1223 (holotype: US! #1804101).
1913.-TYPE:
stuckertii var. pilosistylum C. V. Morton, A Revision

of the Argentine
Species of Solanum, p. 193. 1976.-TYPE: ARGENTINA. Tucumain: Depto. Tran
cas, Vipos, 850 m, 9 Nov 1921, Venturi 1424 (holotype: US! #1548854;
isotype:
Solanum
GH!).
Shrub 0.3-3 (-7 fide Venturi 5613) m tall. Stems moderately
to densely pubescent
with unbranched eglandular hairs. Leaves 6-many per sympodial unit, the blades 3-15
(-25) cm long, (1-) 1.5-7 (-10) cm wide, length:width ratio ca. 1.7-3.5 (-6):1, simple,
ovate to elliptic, acute to acuminate at apex, truncate, cuneate, or rounded to slightly de
current at base, chartaceous, sparsely to moderately pubescent adaxially with curled un
branched uniseriate
to densely pubescent
hairs, more densely so on veins, moderately
abaxially, the petioles 0.5-4 cm long, densely pubescent with unbranched hairs. Inflores
cence unbranched or forked (rarely further branched), ca. 5-30-flowered,
2.5-10 cm long;
peduncle 1-5 cm long; rachis 1.5-8 cm long; pedicels ca. 6-15 mm long, spaced 1-7 mm
apart, articulated at the base; inflorescence axes sparsely to densely pubescent with eg
landular unbranched hairs and occasionally with a few stalked glands. Calyx moderately
to densely pubescent with eglandular hairs, the radius 3-6 mm, the lobes 1.5-3 mm long,
1.5-2 mm wide, often unequal, deltate, abruptly narrowed distally into acute or acuminate
tips. Corolla white, chartaceous, stellate, the radius 10-19 mm, the tube 3-6 mm long, the
lobes 7-15 mm long, 3-4 mm wide at base, narrowly triangular, acute at apex, moderately
to densely pubescent abaxially with mostly unbranched hairs, glabrous adaxially except
for a few hairs toward tips. Anthers usually connivent, yellow, lanceolate, 6-7 (-10) mm
long, 1-2.5 mm wide, abaxial surface smooth to roughened but not obviously papillate,
the pores directed distally. Ovary glabrous or sparsely puberulent at apex; style glabrous
or sparsely tomoderately pubescent, cylindrical to subclavate, 7-10 mm long, 0.5-1 mm
in diameter; stigma truncate to subcapitate. Fruits 1-3 cm long, 1-3 cm in diameter, glo
bose, obtuse at apex, glabrous, orange to reddish when ripe; stone cell aggregates present
or absent. Seeds 3-5 mm long, 34 mm wide, angled, rugose-pubescent. Chromosome
number: n = 12. Figs. 30, 31.
Phenology. Collected in flower in October through May; collected in fruit in October
through August.
Distribution
(Fig. 32). Southern Andes of Argentina (Catamarca, Cordoba, Jujuy, La

Rioja, Salta, San Luis, Santiago del Estero, and Tucumain); clearings, thickets, and open
woodland, often at the borders of streams, in relatively dry areas; 250-2000 m. Several
specimens have also been collected inBolivia in low-lying areas east of the Andean slopes
in Chaco forest, and populations of S. stuckertii are to be expected from suitable habitats
2001
SOLANUM
69
A~~~~~~~~~~~~~~~~~~~~~~~~'
FIG. 30. Solanum

material
of Bohs
stuckertii.
E. Stamen
A. Habit.
B. Flower,

lateral, adaxial
views).
view
F. Fruit.
of partially
(Based
sectioned
on greenhouse
2522.)
VOLUME 61
70
*~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
: .^
u..
; .... .. .. ...
.......
*:_~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...
_... .....,i
..;X-vv.......
=Il |- ..............................
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
_....o..cn
r.!.z.....~~~~~~~~~~~~~~~
A
FIG.
3 1. Mlowers
of S. stuckertii with white,
deeply
stellate
corollas;
scale bar =I
cm.
to Cabrera
in intervening areas of southern and perhaps southeastern Bolivia. According
of
Chaco
forests
and in the
(1983), this species is found in the phytogeographic
provinces
transition zone between the Chaco and Yungas provinces.
Local names. Bolivia: Malva
Argentina:
duraznillo
(Mdiifoz 565), pereremi grande (Vargas & Tapia 1055).

comida de vibora (Stuckert 9138),
del per-ro (Bartlett 20453),
hediondillo
negro (Schuel 36).
(Villafafie 361),
Bolillos
R.EPRE,sENTATIVE SPEcimENs.
565
(LPB).-SANTA
La Brecha,
(JBSC); Nrov. Cordillera,

Parapeti,
Potrero,
19035PS, 62035'W,
1113
Brizuela
Bolivia.
CHUQUISACA:
CRUZ: Nrov. Cordillera,

Bafiados
ILVargas & Tapia
(BH, G); Depto.
Cdrdenas
ov. Luis Calvo
2 km al NE de Prop. Inti, Mui'oz

336
Coimbra
(F) Nrov. Cordillera,
Charagua,
del hospital y 3 km E camino hacia Rio
alrededores
2731
de Izozog,
loss (NY, USZ).
Andalgald,
CATAMARCA: Depto. Ancasti, El

4 km N of Andalgald,
Cantino 669
0 Donell
4163 (B, G); Yacu
Capillitas,
Argentina.
along Rio Andalgali,
s.n. (F); Depto. Andalgahi,

& Hieronymus
162 (CORD); Canmzal de Augier, Villavil,
297 (CORD); Depto. Bel6n, Sierra
Schickendantz
de Bel6n, Las Faldas, Schreiter 35083 (GH); Depto. Bele'n, Quebrada de Belen, Sleumer & Vervoorst 2356 (G,
Cuesta de Tala Cafiada, cerca a Taninga, Burkart 20832 (SI); San Javier, slope of Cerro
US, W).-C6RDOBA:
(GH); Catamarca,
Lorentz
tula, Schickendantz
2001
SOLANUM
71
10
0~~~~~~~~~~~~~~~~~~~~~~~~2
,~~~~~~~~~~-
30
S. stuckertii
80
70
60
& Perez Moreau
Fabris
city of C6rdoba,
6769
ziker 639 (US); La Falda, Rio Grande,

W, WU),
194
Job 480
(BH, F, GH, M, Z); between
La Posta
and Mesillas,
Stuckert
1112 (CORD, G); Sierra Chica,
Stuckert
8246
10992
W).-JuJuY:
Fabris
cis 1830
(GH).-SALTA:
Del Costillo
et al. 3116
Oran, Eyerdam
Brown,
Legname
Depto.
Ledesma,
Coronel
1087
& Cuezzo
between
Anta,
F, GH, K, SI, US);
Depto.
Piedra Blanca,
US); Depto.
38643
7454
Gerth
s.n. (L).-SANTIAGO
Tigre, Ruta 5, entre Santa Rosa

Pozo Cavado, Maldonado
Taco Pozo, Peirano
nia, Ruta
Cristobal
17367
414
(CTES).-TucUMAN:
Stuckert
Venturi
(GH, US); Depto.
1603
19328
(CORD); Depto.
(SI); Depto.
Burruyacu,
Trancas,
Cerro del Campo,
& Cocucci
Rio Hondo-El
El Cevilar,
Leales,
Leales,
Vipos,
Santa Rosa,
Venturi
Venturi
7756
2366
(F, GOET,
Venturi 5613
US);
Bartlett
19855
(F, GH, NY, SI,
(CORD); Depto.
543
del
Ojo de Agua,
82 (GH); Depto.
Pellegrini,
Ruta Nac.
Puestos,
(A,
Luis: Merlo,
Sierra de San Luis, Quebrada
14877
(A, US, U); Tapia, Rodriguez
24?25'S,
Jose de San Martin,
F, GH, LP, US).-SAN
s.n. (A, US); Depto.
Los
a San Javier,
50 km NE of
(F); Oran, El Tabacal, Rodriguez
Alto, Ousset
Pierotti
360
Oran, Abra Grande,
(F, GH, NY, SI, US);
Depto.
7284
(A, BM,
(NE
and Almirante
Balboa
(CTES); Depto.
DEL ESTERO: S of Sumampa,

20453
Rio Hondo,
Guasayan,
Lillo
10001
de Cautana, Hunziker
981 (LP); Depto.
(CTES, LP); Vipos,
gre, Burruyacu,
754 C.N.,
y Bafiado
s.n. (GH); Depto.
3, Renolfi
Bartlett
Venturi
San Severo
between
(B,
Job de Fran
39 km NE of Las Lajitas,
4543
Varela 29 (CORD); Depto.
Alemania,
to Choya,
et al. 10225C
361
between
Rio de Las Piedras
Lorentz & Hieronymus
Juramento,
de Marco
Oran, El Tabacal, Meyer
(G); Depto.
Guachipas,
La Punta
Choya,
and Ledesma,
Santa Barbara,
border, S of Volcan,
de la Frontera,
Stuckert
Villafaiie
de Sanogasta,
camino
Anta,
Paz,
(G); Cruz de Eje,
S. Alberto,
Alnera,
Ledesma,
between
(CORD); General
Charbonier,
(SI); Alto
Anta, Rio Dorado,
Rosario
del Rio
(GH); Pasaje
et al. 595
Depto.
(CTES); Depto.
Joaquin V. Gonzalez,
12584
Oran, Rio Tarija on Bolivian
(BH, G, GH, MO);
Chalican
Cornejo, Maruinak
(NY); Depto.
Arroyo
22825
& Schinini
(LP, SI); Depto.
Burkart
843
(NY, US); Depto.
211 (NY); Depto.
Aguilar
Depto.
(MCNS);
Punilla,
Sierra Chica,
de Arhala,
Stuckert 5641
falda de Mina
et al. 22208
RIOJA: Huaco,
Stuckert
15 (BM, GOET,
(CORD);
(F); Pampa
prope C6rdoba,
(CORD);
et al. 23358
Cabrera
395
Rodrigo
(CORD, G); Depto.
near
(BH, G, W);
Lorentz
101
Lorentz
del Paraiso,
(CORD);
8884a
15196
Joaquin V. Gonzales,
& Beetle
64?W, Krapovickas
Stuckert
(LP, SI).-LA
& Varela 695, 699
Puesto
Capital,
Guti&rrez 333
banks of arroyo Alta Gracia, Hun
near C6rdoba,
de C6rdoba,
and Santa Maria,
El Fuerte, Cabrera
de los Morteros,
de Yuto),
Alta Gracia,
Germania
cercanias
Stuckert 2146
Stuckert
Santa Barbara,
and Abra
(F); Depto.
S. Miguel,
Sierra de C6rdoba,
Depto.
Santa Clara
Sierra
(CORD);
(CORD);
486
Rodrigo
Puerto Punilla,
Punilla,
(GH); Estancia
Cosquin
100 200 300 400 500 6f00miles

40
of S. stuckertii.
Santa Maria,
29 (BM, BR, G, GOET, M, W);
Lorentz
Lossen
(LP); Depto.
s.n. (BR, NY); Depto.
Hieronymus
200 400 600 800 1000km
\
50
FIG. 32. Distribution
Champaqui,
Pellegrini,
La Armo
9, Krapovickas
(A, CTES,
&
SI); Caniada Ale
Venturi 633
(A, US); Depto.
(US); Depto.
Trancas,
Cruz Alta, K.
Tapia, Venturi
(GH, US).
5808
72
VOLUME 61
similar to S. luteoalbum and S. hibermum. All three species

corollas and persistent showy orangish fruits with large angled seeds.
Solanum stuckertii can be distinguished from S. luteoalbum by its indumentum of exclu
sively unbranched hairs and its white corollas. Solanum stuckertii is found in southern Bo
livia to central Argentina, whereas S. luteoalbum occurs in Ecuador and Peru. Solanum hi
Solanum stuckertii ismost
have
stellate
bernum differs in having purple corollas and a dense covering of dendritically branched
hairs on the abaxial leaf surfaces. In Bolivia, S. hibemnum is generally found in the arid
vegetation of interandean valleys, whereas S. stuckertii occurs in lowland Chaco vegeta
tion, and thus the two species may be separated by ecogeographic differences.
Bitter (1913) and Morton
(1976) divided S. stuckertii into several varieties based
mainly on leaf and flower size, stylar pubescence,
and number of flowers per inflores
cence. All of these characters vary within
nizing
infraspecific
the species;
there is no justification
for recog
taxa.
D'Arcy (1972) designated Stuckert 21589 as the lectotype collection of S. stuckertii

from among the many syntypes cited by Bitter (1913) in the protologue. He gave the lo
cation of the lectotype as "B?, not seen; photo NY"; this specimen is no longer extant. I
have seen duplicates of Stuckert 21589 at CORD, G, and GOET (a fragment) and a photo
of the G specimen at F (neg. # 23159), but none at NY Therefore, I amend D'Arcy's
choice by designating the G specimen as the lectotype. Morton (1976) designated another
of Bitter's
syntypes, Stuckert 5641 (G), as the lectotype of S. stuckertii, but D'Arcy's

ear
lier choice must be followed under Art. 9.13 of the ICBN (Greuter et al. 2000). Bitter did
not specify the herbarium location of the specimens he examined for his description of S.
stuckertii and its varieties. Many
of these sheets were
likely at B and have been destroyed.
of some of these collections exist in other herbaria, such as GOET and CORD,
and have been used to designate as lectotypes.
Specimens of S. stuckertii have sometimes been misidentified
as S. sordidum Sendtn.
Duplicates
The
latter species belongs to Solanum subg. Leptostemonum

stellate hairs that usually occur inmembers of that subgenus.
and has the characteristic
DOUBTFUL AND EXCLUDED NAMES

clavatum Rusby, Mem. Torrey Bot. Club 6: 87. 1896. Cyphomandra
clavata
(Rusby) A. Child ex Bohs, FH.Neotrop. Monogr. 63: 154. 1994.-TYPE:
BOLIVIA.
Cochabamba: Mt. Tunari, 1891, Bang 1118 (E! NY! US!). = Solanum aligerum
1118 is a mixture of two different species separated on different
Schtldl.-Bang
herbarium sheets. One element, represented by sheets at C, F, G, L, LD, M, NY, US,
WU, and Z belongs to Solanum confusum of sect. Cyphomandropsis
(q.v.). The other
Solanum
element
is the basis of Rusby's Solanum clavatum. This name is a synonym of the

species S. aligerum Schltdl. of Solanum section Holophylla
(Knapp, in
widespread
press).
Ruiz & Pavon, Fl. peruv. 2: 39. 1799, non Solanumfoetidum
Solanumfoetidum
Rottb0ll,
1778. Solanum maleolens J. F.Macbride, Publ. Field Mus. Nat. Hist., Bot. ser. 8: 111.
1930.-TYPE:
PERU. "In Tarmae oppidi versuris et ruderatis, floret Julio et Augusto,"
Ruiz & Pavon s.n. (holotype: not located).-The
application of Ruiz and Pavon's
name is uncertain. The brief description is too vague to fix the name and not accom
panied by an illustration; no authentic material is known. Dunal (1852) placed S.
foetidum Ruiz & Pav. among members
of Solanum
sect. Geminata
and postulated
that
2001
SOLANUM
73
its affinities are with S. caavurana Vell. of that section; however, he included many
disparate elements in his circumscription of S. foetidum. Macbride (1930) accepted S.
foetidum Ruiz & Pav. as a species and, because the name is a later homonym, pro
vided the new name S. maleolens. Subsequently, he considered it conspecific with S.
luteoalbum but indicated in his discussion thatmore than one taxon may be included
(Macbride1962).
Solanum glaucescens Bacle ex Dunal in DC.,
Solanum glaucescens Zuccarini, 1835.
Solanum
100. 1852, pro syn., non
Prodr. 13(1):
lauterbachii
(H.Winkler) Bitter, Repert. Spec. Nov. Regni Veg. 15: 155. 1918.
lauterbachii H. Winkler, Repert. Spec. Nov. Regni Veg. 7: 247.
1909.-TYPE: BOLIVIA. La Paz: San Antonio bei Mapiri, 850 m, Dec 1907, Buchtien
1436 (holotype: WRSL;
isotype: US!). = Solanum corumbense S. Moore.-Bitter
Cyphomandra
(1918) assigns this species to Solanum sect. Cyphomandropsis,

but its oblong anthers
do not fit with the definition of this section. Instead, it belongs to sect. Geminata and
is a synonym of Solanum corumbense S. Moore (S. Knapp, pers. comm.). Knapp (in
press) notes that the flowers of S. corumbense
resemble those of
superficially
sect. Pachyphylla.
Solanum
narcoticum Bitter, Repert. Spec. Nov. Regni Veg. 13: 97. 1914.-TYPE:
Bo
LIVIA. Toldos prope Bermejo, ca. 1900 m, 26 Nov 1903 (fl), Fiebrig 2261 (holotype:
B, destroyed; photos of holotype, Morton neg. 2717: F! G! NY!).-The
description
is not sufficient to assign the name with certainty, and no isotypes have been found.
Solanum
this name applies
Possibly
to S. confusum,
but this cannot be ascertained
from the
photograph.
ACKNOWLEDGMENTS
I thank Solanaceae
R. N. Lester,
Knapp,
many
of Solanum
aspects
and invaluable
edge,
G. Anderson,
G. Bernardello,
A. Child, W. G. D'Arcy,
R. G. Olmstead,
and D. Spooner
for technical
specialists
E. Moscone,
biology.
assistance
Special
thanks are due toMichael
in the field. Financial
support was
Nee
for his unfailing
provided
A. Hunziker,
S.
and insight
into
help, patience,
by NSF
support, vast knowl
grants DEB-9207359
and
DEB-9726511, National Geographic Society Grant 6189-98, theUniversity of Utah Research Committee, the
of Utah Bioscience
University
Utah. Distribution
thank A. Purgue
information
Kelsey
of
seeds;
the Garrett
Herbarium
for their comments

S. Beck,
I thank the curators
HBG,
WIS,
A, AAU,
M. Moraes,
A. Hunziker,
and the Department
Base Map
for help with

the Botanic
loans;
I also acknowledge
Y. Roca,
E. Moscone,
M.
and staff of the following
herbaria
at Nijmegen,
for maintaining
and
I. Vargas
and A. Sersic
for providing
NY, RB,
University
at FH for
the Netherlands,
for pro
my
living collections;
G. Bemardello,
for facilitating
access
E, ECON,
botanists
field work
A.
and D.
of Bolivian
and Argentine
me with
of
I also
of Utrecht.
counts; C. Nepi
and C. Anderson,
B, BH, BKL, BM, BR, C, COL, CR, CTES, DUKE,

MO,
of Biology,
1, ?University
the help and companionship
Saldias,
L. Novara,
No.
the chromosome
Garden
greenhouses
herbarium
INB, K, L, LD, LIL, LL, LP, M, MA, MBM,

WU,
Program,
of Utah
for help with
on the manuscript.
F. Mamani,
A. Cocucci,
and/or facilities:
C. Christensen
the staff of the University
leagues
America.
Research
from Flora Neotropica
types of S. glaucophyllum;
Spooner
Bernardello,
prepared
for the line drawings;
on possible
Solanum
viding
Undergraduate
were
maps
col
G.
in South
to their specimens
F, G, GB, GH, GOET,
S, SI, SP, TEX, U, US, USM,
UT, VIC, W,
and Z.
74
VOLUME 61
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re
SOLANUM
2001
77
APPENDIX 1
HERBARIUM VOUCHERS FOR SEM STUDIES OF POLLEN GRAINS
S. amotapense:
Harling
S. confusum:
& Andersson
Petersen
15436, NY
Hatschbach
S. cylindricum:
Hatschbach
S. fusifonne:
et al. 721, MO.
Hatschbach
S. pelagicum:
UT.
40573,
Arbo
S. glaucophyllum:
NY.
22510,
618, MO.
& Hjerting
& Forero
LL.
40366,
APPENDIX 2
VOUCHERS FOR CHROMOSOMEAND CROSSING STUDIES
Solanum sect. Cyphomandropsis.
S. amotapense.
Voucher:
Prov. Cajabamba,
Bohs
Voucher: Moscone
S. fusiforne.
dor General
Voucher:
Bohs
S. hibermum. Voucher:
S. luteoalbum.
Bohs
Voucher:
Prov. Calca,
Pisac
S. stuckertii. Voucher:
2443
Bohs
(UT). From
Prov. Misiones,
and Pastoriza
seed accession
(UT). From
s.n., collected
seeds of Hawkes
(GH). From
2523
in Argentina,
seeds of Hunziker
seeds of Nee
(UT). From
2336
2522
Bohs
Collected
Cajamarca,
S.43).
Liberta
Dept.
in Ar
collected
25304,
Parana, Parana.
(Birmingham
Bohs
doba.-Voucher:
(CORD).
in Peru, Dept.
collected
2422,
seed accession
12, Tres de Mayo.
2530
Prov. Entre Rios, Dept.
gentina,
seeds of Hawkes
(Birmingham
218
& Davinia
Ruta Nac.
San Martin,
S. glaucophyllum.
(UT). From
2479
of Condebamba
valley
Santa Cruz, Mairana.
Dept.
collected
in Peru, Dept.
Cuzco,
S.1543).
seeds of Moscone
(UT). From
in Bolivia,
et al. 5416,
122, collected
seeds of Vargas
inArgentina,
collected
1055,
Prov. C6rdoba,
in Bolivia,
Dept.
C6r
Santa Cruz,
del Izozog.
Baniados
Solanum sectionPachyphylla.
S. betaceum
Cav. Voucher:
Bohs 2274
purchased
in La Vincentina
Colombia,
Putumayo,
in Bolivia,
S. circinatum
Bohs
bia, Dept.
1599
Bohs
in Cochabamba
purchased
Voucher:
(GH). From
Bohs
2442
(UT). From
(Sendtn.) Bohs. Voucher:
Bohs
2343
Tolima,
s.n., collected
(GH). From
in Ecuador,
seeds of Bohs
(UT). From
2468
Quito,
Pichincha,
1599,
seeds of Nee
collected
30359,
collected
market.
2301
daci6n Merenberg.-Voucher:
S. corymbiflorum
Bohs
valley of Sibundoy.-Voucher:
Cochabamba,
Bohs.
seeds of Sperling
(GH). From
market.-Voucher:
seeds of Buch
s.n., collected
seeds of Debouck
in Colombia,
et al. 2948,
Huila,
collected
Fun
in Colom
Chaparral.
(GH). From
seeds sent by G. Pringle, DSIR,
New Zealand
(NZ IP#27772), originally collected in southeasternBrazil, exact provenanceunknown.

S. diploconos
(Mart.) Bohs.
inally collected
S. diversifolium
Aragua,
Dunal.
Voucher:
Parque Nacional
S. roseum Bohs.
Voucher:
Dept. Nor Yungas,

S. unilobum
Voucher:
in Brazil,
Bohs
Henri
Bohs
2338
2335
(GH). From
Voucher:
La Paz, Larecaja,
seeds
sent by A. Child, Yorkshire,
England,
orig
city of Curitiba.
2341
(GH). From
seeds of Benftez
de Rojas
2744,
collected
in Venezuela,
Pittier.
(GH). From
8.7 km below Chuspipata
(Rusby) Bohs.
livia, Dept.
Bohs
Parana,
Bohs
between
2284
seeds of Solomon
& Escobar
12458,
collected
in Bolivia,
on road to Yolosa.
(GH). From
Consata
seeds of Sperling
& King
5500,
collected
and Mapiri.
in Bo
in
VOLUME 61
78
APPENDIX 3
CHROMOSOMENUMBERS IN SOLANUM SECT. CYPHOMANDROPSIS
to previously
References
reports are cited
published
in parentheses.
on vouchers
Information
is given
in
Appendix 2.
2n = 24, voucher:
S. amotapense:
Bohs
S. confusum:
n = 12 (Moscone
S. fusiforme:
2n = 24, voucher: Moscone
S. glaucophyllum:
Bohs
2n = 24, voucher:
S. luteoalbum:
pers. com.).
(Moscone,
1992).
2443.
Bohs
2n = 24 (Pringle & Murray
2336;
n = 12 (Ratera 1954; Moscone
S. stuckertii:
218
& Davinia
1974); n = 12 (Moscone
2n = 24 (Federov
2n = 24, voucher:
S. hibernum:
2479.
1992).
1991).
1992).
APPENDIX 4
AcCESSSIONS USED INCROSSING STUDIES
Collection
number
numbers
of plants
selfs/outcrosses,
pollen
used,
of selfs/outcrosses
tubes in self pollinations
pollen
of parents,
stainability
refer to vouchers
in parentheses
number
listed
growing
in Appendix
number
attempted,
into ovary
(SC = self-compatible,
compatibility
2. Numbers
in brackets
of fruit set containing
(+) or inhibited
in upper
viable
style
refer to
seeds
in
(-), average
nt = not tested.
SI = self-incompatible).
INTRASPECIFIC
CROSSES
S. amotapense
(2479)
S. glaucophyllum
S. hibernum
(2443)
S. luteoalbum
[2, 11/0, 11/0, +, 98.0%,
(2530)
[6, 31/28,
[4, 35/1,
(2336)
0/25,
1/0, -, 90.0%,
[4, 31/68,
SC]
-, 76.0%,
19/47, +, 73.0%,
S. stuckertii
(2522)
[7, 14/15 0/10,
S. stuckertii
(2523)
[2, 8/2, 0/2, nt, nt, SI]
SI]
SI]
-, 76.8%,
SC]
SI]
CROSSES BETWEEN SPECIES OF SECT. CYPHOMANDROPSIS

The
female
parent
fruits set, number

ovary
is listed
first. Numbers
of fruits produced
with
in brackets
indicate
number
seeds, F1s produced
full-sized
of crossing
attempts,
(y) or not (n), pollen
number
tube growth
of
into
(+) or not (-) or not tested (nt).
S. glaucophyllum
(2530)
x S. hibernum
S. glaucophyllum
(2530)
x S. luteoalbum
S. glaucophyllum
(2530)
x S. stuckertii
S. hibernum
x S. stuckertii
(2443)
[17, 9, 0, n, +]
(2443)
(2336)
(2522)
(2522)
[25, 0, 0, n, -]
[17, 7, 0, n, +]
[2, 2, 2, y, nt]
S. luteoalbum
(2336)
x S. glaucophyllum
S. luteoalbum
(2336)
x S. hibernum
(2443)
[22, 10, 0, n, +]
(2336)
x S. stuckertii
(2522)
[19, 11, 6, n, +]
S. luteoalbum
S. stuckertii
(2522)
x S. glaucophyllum
S. stuckertii
(2522)
x S. hibernum
S. stuckertii
(2522)
x S. luteoalbum
(2530)
(2530)
(2443)
[37, 29, 29, y, +]
[18, 8, 8, n, -]
[19, 10, 7, y, +]
(2336)
[18, 0, 0, n, -]
CROSSES BETWEEN SPECIES OF SECT. CYPHOMANDROPSIS AND SECT. PACHYPHYLLA

Number
into ovary
in brackets
indicate number
of crossing
(+) or not (-) or not tested (nt). No
seeds
S. betaceum
(2468)
x S. glaucophyllum
S. betaceum
(2468)
x S. hibernum
(2443)
[6, 0, -]
S. betaceum
(2274)
x S. hibernum
(2443)
[4, 0, +]
(2530)
attempts,
resulted
number
of fruits produced,
from any of the following
pollen
crosses.
[19, 5, +]
tube growth
SOLANUM
2001
[10, 0, nt]
S. betaceum
(1599)
x S. hibernum
S. betaceum
(2468)
x S. luteoalbum
(2443)
(2336)
S. betaceum
(2274)
x S. luteoalbum
(2336)
S. betaceum
(2274)
x S. stuckertii
[12, 0, -]
[9, 0, nt]
(2522)
[3, 0, +]
S. circinatum
(2301)
x S. glaucophyllum
(2530)
[20, 0, +]
S. circinatum
(2442)
x S. glaucophyllum
(2530)
[14, 0, +]
S. circinatum
(2301) x S. hibernum
(2443)
[10, 0, nt]
S. circinatum
(2442)
x S. hibernum
(2443)
[21, 0, +]
S. circinatum
(2301)
x S. luteoalbum
(2336)
S. circinatum
(2442)
x S. luteoalbum
(2336)
S. circinatum
(2301)
x S. stuckertii
(2522)
[15, 0, nt]
S. circinatum
(2442)
x S. stuckertii
(2522)
[18, 0, +]
[30, 0, -]
[21, 0, -]
S. corymbiflorum(2343) x S. glaucophyllum(2530) [20, 0, -]

S. corymbiflorum
(2343)
x S. hibernum
[18, 0, -]
(2443)
S. corymbiflorum(2343) x S. luteoalbum(2336) [20, 0, -]

S. corymbifiorum
(2343)
x S. stuckertii
S. diploconos
(2335) x S. glaucophyllum
S. diploconos
(2335) x S. luteoalbum
S. diploconos
(2335)
[20, 0, -]
(2522)
(2530)
x S. stuckertii
[0, 0, +]
(2522)
S. diversifolium
(2341)
x S. glaucophyllum
S. diversifolium
(2341)
x S. hibernum
S. diversifolium
(2341)
x S. luteoalbum
(2341)
x S. stuckertii
[10, 0, +]
(2350)
[20, 0, -]
(2443)
S. diversifolium
[19, 0, +]
[20, 0, -]
(2336)
[23, 0, -]
(2336)
[20, 0, +]
(2522)
S. glaucophyllum
(2530)
x S. betaceum
(2468)
[15, 7, +]
S. glaucophyllum
(2530)
x S. betaceum
(2274)
[5, 0, nt]
S. glaucophyllum
(2530)
x S. circinatum
(2442)
[16, 4, +]
S. glaucophyllum
(2530)
x S. circinatum
(2301)
[25, 5, +]
S. glaucophyllum (2530) x S. corymbiflorum(2343) [20, 2, +]

S. glaucophyllum
(2530)
x S. diploconos
S. glaucophyllum
(2530)
x S. diversifolium
S. glaucophyllum
(2530)
x S. roseum
(2530)
x S. unilobum
S. glaucophyllum
(2335)
[20, 7, +]
(2341)
[24, 9, +]
[4, 0, -]
(2338)
[5, 0, +]
(2284)
x S. betaceum
(2468)
[10, 0, +]
(2336)
x S. betaceum
(2274)
[10, 0, nt]
(2336)
x S. circinatum
(2442)
[24, 12, +]
S. luteoalbum
(2336)
x S. circinatum
(2301)
[20, 1, -]
S. luteoalbum
(2336)
S. luteoalbum
S. luteoalbum
(2336)
S. luteoalbum
S. luteoalbum
(2336)
x S. corymbiflorum
(2343) [22, 0, +]
x S. diploconos
(2335) [20, 4, +]
S. luteoalbum
(2336)
x S. diversifolium
S. luteoalbum
(2336)
x S. roseum
S. luteoalbum
(2336)
x S. unilobum
(2341)
S. roseum
(2338)
x S. glaucophyllum
S. roseum
(2338)
x S. luteoalbum
S. roseum
(2338)
x S. stuckertii
x S. betaceum
S. stuckertii
(2522)
x S. circinatum
[0, 0, +]
(2530)
(2522)
(2522)
[20, 0, +]
(2284)
(2336)
S. stuckertii
[21, 0, -]
[5, 0, +]
(2338)
[0, 0, -]
[0, 0, +]
(2468)
[0, 0, +]
(2442)
[0, 0, +]
S. stuckertii(2522) x S. corymbiflorum(2343) [0, 0, -]

S. stuckertii
(2522)
x S. diversifolium
S. unilobum
(2284)
x S. glaucophyllum
S. unilobum
(2284)
x S. luteoalbum
[0, 0, -]
(2341)
(2530)
(2336)
[8, 0, +]
[20, 0, -]
79
80
VOLUME 61
APPENDIX 5
DISTRIBUTION OF SPECIES OF SOLANUM SECT. CYPHOMANDROPSISBY COUNTRY
Asterisks
Colombia:
indicate
endemic
species
to the country
listed.
S. fallax
Ecuador:S. amotapense,S. fallax, S. luteoalbum

Peru: S. amotapense,S. hutchisonii*,S. luteoalbum
Brazil: S. cylindricum,S. fusiforme, S. glaucophyllum,S. luridifuscescens*,S.matadori*, S. pelagicum*
Bolivia: S. confusum,S. glaucophyllum,S. hibernum*,S. stuckertii
Paraguay:
S. fusifonne,
S. glaucophyllum
Argentina:S. confusum,S. cylindricum,S. fusiforme,S. glaucophyllum,S. stuckertii

Uruguay: S. glaucophyllum
NUMERICALLIST OF SPECIES
2. S. confusum
3. S. cylindricum
8. S. hutchisonii
9. S. luridifuscescens
10. S. luteoalbum
4. S. fallax
11. S. matadori
5. S. fusiforme
6. S. glaucophyllum
12. S. pelagicum
13. S. stuckertii
1. S. amotapense
7. S. hibernum
INDEXTO NUMBERED COLLECTIONS

The numbers in parenthesis refer to the correspondingspecies in the text and in theNumerical List of
Species presentedabove.
J. R., & A. Jardim 17232
Abbott,
Solfs, M.
Acosta
R. M.
Aguilar,
(1), 12847
211 (13), 280
0.
Ahumada,
7743
309
(2).
Bodenbender,W. SI 26573 (13).
(4).
Boeke,
(13), 1212
(6), 1653
(6), 1762
(6), 2021
(6),
2105 (6), 5316 (2).

Ahumada,
O., & U. Eskuche

O., et al. 858
E. 4450
3498
Arechavaleta,
J. 140 (6), 3057
Arenas,
P. 714
L., et al. 931
(6), 1407
Balansa,
B. 2105
Balcazar,
J. 385
Bang, M.
Bartlett,
Billiet,
(6), 1490
(6), 2000
(6).
M.
L., et al. 2770
Bordas,
E. 4061
Brown
2618
(13), 20453
(2), 14665
(2).
(10),
2443
(7), 2479
(1),
2767
(2), 2790
(2),
(6).
(7), 2776
(2), 2780
(6).
(2), 17219
14478
(6), 18835
(2).
(3).
(6), 3244
& Malmierca
Caballero
7737
(13),
Cabrera,
A.,
(2).
(2).
8727
(6), 12584
(6), 13658
et al. 22208
13219
(2).
(13), 23358
(13), 32059
J. 1198 (2).
2101
(13),
(2).
G. 281 (5).
Marmori,
A. 2795
Camargo
(9).
(2), 6806
1571
(13), 22103
Cabrera,
Cadifuerel,
(6).
A.
(2), 20832
(2), 21830
(9), 16926
J. 1113 (13).
Burkart,
(13).
(2).
63 (6).
2337
W. M. A. 5947
Brooke,
F., & B. Jadin 3036
Biraben, M.
L., & M. Nee
Bohs,
Brizuela,
(6).
18517
(6).
Bornmiiller,A. 384 (3).
(7).
14092
(6), BAA-12613
(13).
2836 (2), 2844 (2).
(2).
S. G., & R. Seidel
Bernardi,
(10),
Bohs,
(6).
1118 (2, S. aligerum),
(2), 24352
Beck,
(6).
(2).
H. H. 19855
S. G.
L. 2336
Brade, A. C. 16604
E. 269
(10).
2522 (13), 2523 (13), 2530 (6).
(6).
43 (6).
Bailetti,
Beck,
(6), 6675
3173
O., et al. BAA-5888

P. 133 (6), 1009
Bohs,
(6), 3458
(1).
et al. 721
Bacle
(5).
(6), 1449
Arbo, M. M.,
Arroyo,
Boelcke,
Boffa,
Ahumada,
Andre,
J. D., & S. Boeke
Boelcke, 0. 1396 (6).
(6).
(3), 2150
Camp, W. H. E-4675
(3).
(10).
(2).
81
SOLANUM
2001
Cano, A.
1557
Gibert,M. 140 (6).

Glaziou, A. 13084 (9).
(10).
Cantino, P. 669 (13).

M.
Cardenas,
(13), 3260
2731
(2), 5773
(2), 5741
(2),
Hahn, W. 671
Carter, G. F. 42 (6).
4434
Casas & Molero
(6).
Castro,R. BAB-52328 (6).

A., & U. Eskuche
Charpin,
20145
(6).
G., & L. Andersson
Harling,
G., & B. Stahl 26473
Hassler,
E. 317
Coimbra,G. 336 (13), 601 (13), 1006 (6), 2320 (6),

(2), 2618
2358
(2), 2619
O., & G. Gilbert
387a
(2), 2893
C. L., et al. 1437
Crist6bal,
A.,
Curtiss,
A. H. 6862
Dawson
663
(6).
A. de, & F. Varela
T. E. 473
e Otavio
695
(13), 699
(13).
237
6088
G., & E. Forero 40366

G., & 0.
717
(6), 794
687
(6), 690
(6), 800
(6), 797
#616
Dodson,
C. H., & F. M. Valverde
Dodson,
C. H., et al. 8731
L., et al. 6932
(6), 699
(6),
(6).
3206
Dusen,
P. 2057
(3), 7081
(9), 3056
(12).
(3), 17508
Herrera,
F. L. 921
(13), 22974
et al. 3116
(13), 5038
6769
(13).
Felippone, F. 5947 (6).
110 (6).
14 (6).
R., et al. 2041
(6).
T. 310
(6), 639
(6),
P., & J.Wright
et al. A-831
Garcfa,
I.H. 30 (7), 71B
(6).
Job, M. M.
Klein
(7).
C. 160 (12), 1833
Klein,
(13).
(13).
3548
(1).
(6).
(10), 222
(10).
(10), 528
(10), 582
(10), 589 (10), 591 (10), 601 (10), 736
480
(2).
(13), 737
1830
1009a
P. 1472
(6).
(13).
(3).
(2), 2204
(6), 10788
(2).
R. 79 (10).
M., & M. Kelschebach
R., et al. 8295
& U.
233
(2).
(2).
Eskuche
1-38
(3), 7-27
(3),
Klein,
75300
(10).
(13).
(6).
R., & A. Bresolin
6289
(12), 6386
(12).
(6).
(4).
et al. 19785
6290
19-50
(3),
9058 (3).
(6).
Gentry, A., & R. Foster
(13),
(10), 923 (10).

Irigoyen,J. 221 (6).
Kessler,
A. 586
14877
(6), 25325
1141 (10), 1213
et al. 221
(10), 584
(13).
Kiesling,
Fuentes,
1223
(2).
Iltis, H. H., & D. Ugent
Kanehira,
(6).
(6).
(13),
A. T., et al. 24857
Fries, R. E. 1640
M. L. 878
(6), 3419
A. T., & A. E. Cocucci
Friedrichs32880 (3).
Giardelli,
1452
Hunziker,
Jorgensen,
(6).
Gallinal
(6).
(12).
Hunziker,
Jonsson, G.
E. A. 762
Gentry, A.,
A.
Job de Francis
A.
Gentry, A. 30988
(10).
(6), 18512
(12).
A. R. M.
Huidrobo,
Juarez, F. 2023
(1).
Fiebrig,K. 463 (6), 1249 (6), 2261 (2).
Gaudichaud,
(10), 3346
(6), 659a
Ibisch, P., et al. xx.efl

22825
Fabris, H. A., & R. L. Perez Moreau
Fortunato,
(1).
(2).
F. C. 24447
Iltis, H. H.,
Flossdorf,
(6).
11634
Ibarrola, T. S. 890 (6), 3205
23064 (6).
Franceschi,
I. 936
Hutchison,
Eyerdam, W. J., & A. A. Beetle
A.
49222
O., & H. K. Svenson
Hensen,
(12).
(3).
Hutchison, P. 1490 (8).
(3).
(5).
R. 9141
27529
10538
(13), 7630 (13), 7980 (13), 18957 (6), 19508 (2),

24871 (2), 25304 (6).
Eskuche,U. 1251-11 (12).
Filipov,
G., & da Silva
(4).
Eggers, H. 15069 (4).
Ferreyra,
Hatschbach,
(4).
A. P., & J. Falcao
Fabris, H. A.,
G., & E. Pereira
Hunziker,
6952
(2).
E. L. 845
G., & C. Kozicki
Hatschbach,
Hunt, D. R. 6360
(6).
Duarte,
Ekman,
(6), 46879
Huber 111 (6).
J. F. DBR Nep
Dobremez,
(12).
21941
Guimaraes
Hatschbach,
Hoehne,
(9).
J., & Tokarnia
Dobereiner,
(9), 22240
Hieronymus,G. 3057 (6).
(10).
Dobereiner, J. 820 (6).
Doff,
Hatschbach,
Herter, W. 659
(13).
M. O., & A. Sagastegui
Dionisi
(3), 20327
(3), 19391
Hatschbach,
Haught,
(6).
(6).
Del Costillo,
DiFulvio,
(6).
et al. 11271C
Cuezzo,
(6).
(9).
N. E. 132 (13).
Crespo,
Dillon,
(2).
(10), 804 (10).
R. 6 (6), 61 (6).
Cordini,
(1).
(1).
(9), 22933 (3), 23255 (3), 26472 (3), 38640 (6),

40573 (5), 43491 (3), 45519 (3), 48833 (3).
A. 36 (2).
Cocucci,
22510
(6), 7483
(6), 3201
(3, 5), 19384
R. 32 (6), 33 (6), 34 (6), 35 (6), 36 (6).
Chodat,
(6).
Harling,
Hatschbach,G. 8302 (9), 9668 (3), 15436 (3), 18384
Child, A. C9469 (1).
Cook,
Sosa, E. 61 (6).
G6mez
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Krapovickas,
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6465
(10).
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1711
(13).
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BOTANY
MONOGRAPHSVOLUME
61
SYSTEMATIC
82
Krapovickas,
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(13), 44389
Krapovickas,
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J. C., et al. ICN # 21246

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VOLUME 61
84
INDEXTO SCIENTIFICNAMES
Accepted
names
are in Roman
type; the main
entry for each is in boldface.
Alnus
Solanum
acuminata
Antarctia
17, 27
subg. Bassovia
20
19, 41
Araucaria
Boraginaceae
subg. Minon
L. 4
sect. Allophylla
ex Sendtn.
A. Child
sect. Ceratostemon
sect. Cornigera
3, 4, 5, 6, 7
(Miers) A. Child
A. Child
sect. Cyphomandra
6, 7, 21
(Bitter) D'Arcy
(C. V. Morton)
amotapensis
Dunal
(Svenson)
Seithe
sect. Allophyllum
(A. Child)
6, 7, 21
sect. Brevantherum
sect. Californisolanum
clavata
(Rusby) Bohs
Dunal
elliptica
26
sect. Cryptocarpum
A. Child
ex Bohs
23
sect.CyphomandropsisBitter 21-22
72
sect. Extensum
D'Arcy
sect. Geminata
5, 32
sect. Holophylla
A. Child
ex Bohs
41
3, 4
(G. Don) Walp.
sect. Leprophora
Dunal
58
6, 65, 67
(L. B. Sm. & Downs)
A. Child
32
ex Bohs
sect. Melongena
(Mill.) Dunal
sect. Micracantha
Dunal
5, 20, 57
Steyerm.
sect. Normania
(Lowe) Bitter
(Vent.) Dunal
24, 26, 58
polymnia
(Cav.) J. L. Gentry
casabranca
Haensch
Physalis
alkekengi
L. 4
Miers
cornigera
cylindrica
20
(Vell.) Miers
32
elliptica
(Vell.) Miers
velutina
(Sendtn.) Miers
Solanaceae
5, 7, 20
Solanoideae
8, 14
65
32
(Moench)
sect. Pteroidea
Dunal
sect. Solanum
4, 14, 55
sect. Torva Nees
14, 55
sect. Pseudocapsicum
Bitter
3, 4, 10
[unranked]
AnthoresisDunal S
Dunal
[unranked]
Subdulcamara
abutiloides
(Griseb.) Bitter & Lillo

C. V. Morton
adhaerens
Roem.
aligerum
Schltdl.
allophyllum
amotapense
57
A. Child
sect. Petota Dumort.
adelphum
(Dunal) Miers
3, 4, 6, 7, 9, 10,
77, 78
sect. Parasolanum
Lycianthes
heteroclita (Sendtn.)Bitter 4
LycopersiconMill. 14
Mechanitis 20
lysimnia lysimnia (Fabricius)20
Pionandra
(Dunal) Dunal
12, 13, 14, 15, 16, 17, 18, 19, 20, 38, 43, 58, 73,
Jaltomata
procumbens
sect. Nycterium
sect. Pachyphylla
Sendtn.
sect.LycopersicumWettst. 4
73
stuckertii(Bitter)D'Arcy 67
velutina
50
37, 40
L. B. Sm. & Downs
subhastata
Seithe
(Dunal) D'Arcy
(Pers.) A. Child Bohs
maritima
3, 4, 9, 10, 32, 57,
72
sect. Lasiocarpa
H. Winkl.
lauterbachii
luteoalba
21, 50
57, 58
Bitter
hypomalaca
3, 4, 5, 10, 72, 73
A. Child
sect. Jasminosolanum
Dusen
4, 50
(G. Don) Walp.
sect. Glaucophyllum
(L. B. Sm. & Downs)
Dumort.
(Moench)
Dunal
21, 65, 67
Dunal
sect. Eriophylla
(Vell.) Sendtn.
glaberrima
ex Bohs
5, 38
3, 4, 10
A. Child
A. Child
cylindrica (Vell.)Sendtn. 32
fusifornis
3, 4, 6
15
Seithe
sect. Dulcamara
cornigera
Bohs
(Marzell) Danert
Bitter
betacea
var. velutina Dunal
4
3, 4, 6
sect. Aculeigerum
sect. Basarthrum
(Bitter) A. Child
4, 6
4, 5, 6, 50
sect. Archaesolanum
sect. Rhynchantherum
3, 4, 6, 72
sect.AnarrichomenumBitter 4
adelpha
(Dunal) Bitter
3, 4, 37
sect. Acanthophora
Mart.
4
4
(G. Don) D'Arcy
subg. Solanum
baccatum
villosa
Raf.
subg. Potatoe
20
Capsicum
Cyphomandra
Marzell
(Aubl.) Bitter
subg. Leptostemonum
20
are in italics.
L. 21
subg. Archaesolanum
fusca Eslker
Asteraceae
Synonyms
5
4
6, 12, 26, 31, 32
& Schult.
32, 72
(Miers) Standl.
Svenson
3, 4, 6
8, 9, 10, 12, 16, 17, 19, 23-26,
27, 41, 77, 78, 80

amygdalifolium
angustifolium
Steud.
Lam.
5, 50
50
2000
SOLANUM
appendiculatum
argentinum
Dunal
arboreum Dunal
var. tunya J. F. Macbr.
Bitter & Lillo
malacoxylon Sendtn. 5, 43
betaceum
Cav. 4, 5, 77, 78, 79
var. albo-marginatum
73
var. genuinum
campechienseL. 4
candidum
L. B. Sm. & Downs
circinatum
Bohs
6, 32, 37
Rusby
confusum
C. V. Morton
6, 7, 8, 9, 10, 12, 13, 15, 16,
Sess6 & Moc.
comigerum
Dunal
corumbense
S. Moore
cylindricum
Vell.
Bohs
10, 15, 77, 79
dulcamara
15, 42, 77, 79
15, 77, 79
R. Br. 32
ellipticum
Vell.
5, 32, 37
12, 16,
Rottb.
foetidum
Ruiz & Pav. 72, 73
17,
19, 23,
27,
72
17, 19, 22, 41-43,
6, 8, 9, 12, 13, 16,
50, 57, 65, 77, 78, 80
glaucescens
Bacle
ex Dunal
glaucescens
Zucc.
73
Desf.
glaucum
Dunal
5, 43, 44, 50
glaucum
Rojas
43, 44
77,
43, 44
Bohs
sciadostylis
Mill.
laciniatum Aiton
lauterbachii
43
3, 5, 58, 64
72
Sendtn.
3, 5, 7, 9, 10, 12, 13, 15, 16, 17, 18,
Bitter
68
var. obrutum C. V. Morton

var. pilosistylum
Bohs
6, 8, 9, 16, 17, 19,
L. B. Sm. & Downs
subhastatum
37
6, 32, 37
tripartitum Dunal
6, 32, 37
4
5, 73
3, 5, 10, 16, 17, 19, 20, 23,
26, 43, 57-58, 59, 65, 80

Pers. 4, 6, 7, 8, 9, 11, 12, 14, 16, 17, 19,
23,31,53,54,55,58,60-64,72,73,77,78,79,
80
trisectum Dunal
trizygum Bitter
Bitter
68
torvum Sw. 4
Bitter
68
C. V. Morton
var. trichostylumBitter 68
3, 32
(H. Winkl.)
luridifuscescens
Bitter
semicoalitum
7, 8, 9, 12, 15, 16, 17, 19, 23, 31,
(Bitter) C. V. Morton
Bitter
15, 77, 79
(Sendtn.) Bohs
10
L. B. Sm. & Downs
jamaicense
10, 20, 42
Ruiz & Pav. 4, 58, 64
var. angustifrons
(J. F. Macbr.)
Bohs
var. atrichostylumBitter 68
50
22, 24, 27, 55-57, 80

hypomalacum
Dunal
19,20,21,23,31,53,55,63,67-72,77,78,79,
80
51-55, 63, 72, 77, 78, 79, 80

hutchisonii
5, 6, 8, 9, 12, 13, 16, 17, 19, 21, 23,
stuckertii Bitter
graveolensBunbury6
Britton
6, 55, 57
(L. B. Sm. & Downs)
ptychanthum
sordidum
Bertoloni
Morong
J. F. Macbr.
Bohs
roseum Bohs
78, 80
hibernum
5, 73
pubescensWilld. 58
4, 5, 6, 7, 8, 9, 10, 12, 13, 15,
glaucum
handelianum
42, 58
36, 43, 65-67, 77, 80

persicifoliumMart. 50
physalifoliumRusby 4
pubescens
73
16, 17, 18, 19, 20, 21, 22, 23, 31, 36, 43-51,
luteoalbum
Bitter
pinetorum
L. B. Sm. & Downs
johannae
narcoticum
pelagicum
10,
3741, 80
iraniense
L. 4
var. hutchisonii
foetidum
hazenii
Bohs
melongena
palitans C. V. Morton
5, 7, 9,
glaucophyllum
melissarum
nitidum Ruiz & Pav. 4, 9, 57
ellipticum
fusiforme
6, 8, 9, 16, 17, 19, 22,
80
nigrum L. 21
Cav. 4
fallax Bohs
65, 67
melanoxylon 44
L. 4, 57
elaeagnifolium
ex Nees
L. B. Sm. & Downs
36, 43, 50, 64-65,
73
(Mart.) Bohs
72, 73
L. 4
Meyen
matadori
5, 6, 8, 9, 10, 12, 16, 17, 19, 23,
Dunal
J. F. Macbr.
maritimum
32-37, 67, 77, 80

diversifolium
maleolens
mammosum
5, 67
(Sendtn.)
44
44
subf. angustissimum(Kuntze)Hassl. 44
3, 6, 32, 72
cordovense
(Chodat) Hassl.
f. vulgare Hassl.
17, 19,22,23,26-32,50,63,72,73,77,78,80
diploconos
43
f. albo-marginatum
5, 77, 79
clavatum
corymbiflorum
Hassl.
var. latifoliumKuntze 44
var. subvirescensHassl. 44
Lindl.
catanduvae
44
Chodat
var. angustissimumKuntze 44
50
Morong
Vell.
L. 4
macrocarpon
G. Forst. 4
caavurana
6, 58, 64
lycopersicumL. 4
aviculare
brittonianum
85
unilobum
(Rusby) Bohs
velutinum
Dunal
57, 58
Aiton
vespertilio
villosum
wallacei
Triguera
(A. Gray)
wendlandii
Witheringia
Mill.
Hook.
osbeckii
15, 77, 79
Parish 4, 57
f. 3, 4, 6
(L.) Willk.
solanacea
L'Her. 4

Bohs, 2001 - Revision of Solanum Section Cyphomandropsis

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Bohs, 2001 - Revision of Solanum Section Cyphomandropsis

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Revision of Solanum Section Cyphomandropsis (Solanaceae)

Author(s): Lynn Bohs

REVISION OF SOLANUM SECTION

Salt Lake City, Utah 84112

trees that lack spines,

clade. The morphology,

ture, ecology, distribution,economic value, reproductivebiology, and evolutionary relationshipsof Solanum

y tienen anteras atenuadas

(nom. nov.). Se proporciona

una clave dicot6mica

por sus cromosomas

por la falta de un conectivo

for the species

and one new

keys are provided

one new combination

del que se distingue

anteras. Las dos secciones

in the traditional sense of Bitter (1913), Seithe (1962), Danert

have been made

from dried herbarium material with floral

parts rehydrated in boiling water. Where

of the blade on the petiole. The petiole

from this point. Colors of

fruits, etc., are described

length is also measured

from living material

in Table 1. Herbarium vouchers for pollen SEM are listed in Appendix

of crosses is given in Appendix

the aniline-blue fluorescence technique de

discrete enlarged connective

formerly recognized as the segregate genus Cyphomandra

et al. 1993; Bohs

1). Other characters

that may distinguish

include plant architecture

Prevost's model in sect. Pachyphylla),

and "Habitats and Distribution"

and two representatives

(G. Don) Walp.,

Allophyllum, Brevantherum, and Extensum as sister to the Cyphomandropsis/Pachyphylla

have been placed

in the past in either Solanum or

1845, when Otto Sendtner

ndhF and nuclear

trees of 1444 steps from parsimony

data from chloroplast

has been controversial.

1984). All species of Cyphomandra

in 1995 (Bohs 1995).

The earliest description

from Ruiz and Pavon's Flora Peruviana

renamed as S. luteoalbum by Persoon

a new species of the section, S. glaucophyllum,

and most widespread mem

taken from a plant cultivated

records indicate that this showy species was cultivated

(1829) was the next to describe

in Paris. It is not known how or when S. glaucophyllum

at the Botanic Garden

arrived in France, but herbarium

have been at least loosely associated with

other species of sect. Cyphomandropsis.

and "Subdulcamara," respectively). Child (1986)