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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
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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
sect. Cyphomandropsis
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.

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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

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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

sect. Cyphomandropsis
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

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VOLUME 61

SYSTEMATICBOTANYMONOGRAPHS

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

sect. Cyphomandropsis

have been at least loosely associated with

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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

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SYSTEMATICBOTANYMONOGRAPHS

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

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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

sect. Cyphomandropsis.
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

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SYSTEMATICBOTANYMONOGRAPHS

VOLUME 61

ste showin
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All use subject to JSTOR Terms and Conditions

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
sect. Cyphomandropsis.

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

sect. Cyphomandropsis

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

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10

SYSTEMATICBOTANYMONOGRAPHS

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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2001

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12

SYSTEMATICBOTANYMONOGRAPHS

TABLE 1. Pollen
Herbarium

vouchers

of Solanum

section Cyphomandropsis

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

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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.
:.:
.:
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.:.

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::.:
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

taxonomic

utility

and

functional

signific;ce

of;

thi

....
-:'~~
,.,>X8
.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
~ ......

when they have been identifiedwith certaintyto determine if thischaractr is primitiveor


............
...a.'..^
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'..t .. t ...
i' 'ti5i.

,'M

-B~~~~~~~~~~~~~~~~~~~~~~~~~....
....
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........
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...

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

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SYSTEMATICBOTANYMONOGRAPHS

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
of sect. Pachyphylla
(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.

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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
sect. Cyphomandropsis

(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,
of sect. Pachyphylla

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).

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16

SYSTEMATICBOTANYMONOGRAPHS

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

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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.

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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

sect. Cyphomandropsis.

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

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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

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SYSTEMATICBOTANYMONOGRAPHS

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.).

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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.
section Cyphomandropsis
Bitter, Repert. Spec. Nov. Regni Veg. 12: 461. 1913.
section Cyphomandropsis
(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

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22

SYSTEMATICBOTANYMONOGRAPHS

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.

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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).

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SYSTEMATICBOTANYMONOGRAPHS

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

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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.)

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sectioned
on: A, F,

SYSTEMATICBOTANYMONOGRAPHS

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

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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.

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SYSTEMATICBOTANYMONOGRAPHS

28

VOLUME 61

)~~~~~~~~~~~

FIG. 9. Solanum
flower. D. Gynoecium.
15442;

B-F,

Sleumer

confusum.
E. Stamen

A. Habit.

B. Flower,

(left to right: abaxial,

seen from above.


lateral, adaxial

C. Lateral
views).

view

F. Fruit.

of partially
(Based

3056.)

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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

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(below)
on road

VOLUME 61

SYSTEMATICBOTANYMONOGRAPHS

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.,

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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

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the two

SYSTEMATICBOTANYMONOGRAPHS

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,

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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

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384

34

SYSTEMATICBOTANYMONOGRAPHS

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).

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2001

SOLANUM

35

-V~~~c

/~~~~~~~~~~~~~~~~~~~~~~~~~~~

ttlitEX

d!~~~~~~~~~~~~~~~

B~~~~~

FIG. 13. Solanum


of partially

sectioned

(Based on Hatschbach

cylindricum

(element

flower. D. Gynoecium.

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.)

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36

VOLUME 61

SYSTEMATICBOTANYMONOGRAPHS

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

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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

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SYSTEMATICBOTANYMONOGRAPHS

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,

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and its

2001

SOLANUM

FIG.

15. Solanum

flower. D. Gynoecium.
ster & Hebert 27710;

fallax.

A. Habit.

E. Stamen

B. Flower,

seen

(left to right: abaxial,


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

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40

SYSTEMATICBOTANYMONOGRAPHS

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)

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[=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,

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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

SYSTEMATICBOTANYMONOGRAPHS

42

VOLUME 61

ol

FIG.

17. Solanumfusiforme.

partially

sectioned

(Based

on: A, C-G,

A. Habit.

flower. E. Gynoecium.

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.)

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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).

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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.

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SOLANUM

2001

45

.:

B
FIG.

19. Solanum

glaucophyllum.

tioned flower. D. Gynoecium.


C, F, greenhouse

material

E. Stamen

of Bohs

A. Habit.

B. Flower,

(left to right: abaxial,

seen from above. C. Lateral


adaxial,

lateral views).

E
~~~~~~~~~~~~~~~~
2530; D, E, Crist6bal

~~~~D
view

F. Fruit.

of partially
(Based

et al. 1437.)

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sec

on: A, B,

AA:
ON

..........

7,11

. .........

FIG. 20. Solanum

glaucophyllum.

A. Flowering

branch;

scale bar

2 cm. B. Flowers

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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).
REPRESENTATIVE SPECIMENS. Brazil.
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,

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48

SYSTEMATICBOTANYMONOGRAPHS

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);

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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.).
Solanum glaucophyllum
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.).

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SYSTEMATICBOTANYMONOGRAPHS

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
sect. Cyphomandropsis

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.

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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


Solanum glaucophyllum

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).

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52

SYSTEMATICBOTANYMONOGRAPHS

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.)

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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

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54

SYSTEMATICBOTANYMONOGRAPHS

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

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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

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in
and

SYSTEMATICBOTANYMONOGRAPHS

56

FIG. 24. Solanum


flower. D. Gynoecium.

hutchisonii.

A. Habit.

E. Stamen

(left to right: abaxial,

B. Flower,

VOLUME 61

seen from above. C. Lateral


lateral, adaxial

view).

view of partially sectioned


(Based on Hutchison

F. Fruit.

1490.)

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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

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SYSTEMATICBOTANYMONOGRAPHS

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.
REPRESENTATIVE SPECIMENS. Brazil.
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).

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SOLANUM

2001

59

ci~j

m
FIG. 25. Solanum
tioned

; C

B
luridifuscescens.

flower. D. Gynoecium.

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.)

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SYSTEMATICBOTANYMONOGRAPHS

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);

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2001

SOLANUM

61

NI~

II

-4.

CN

FIG. 26. Solanum

luteoalbum.

flower. D. Gynoecium.
E. Stamen
material of Bohs 2336.)

A. Habit.

B. Rlower,

(left to right: abaxial,

seen from above. C. Lateral


lateral, adaxial

views).

view of partially sectioned


(Based on greenhouse

F. Fruit.

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WIN
......
.....

JI:

-er
MW
'T:
.............
.............
. .........

. ... . . ......
....... ..........
..... . .
.... .......

oRX
. .........
..
.........
.............
......

....... ....
.........
...

............
.
..........
i...X

............
..
..........

FIG. 27. Solanum

luteoalbum.

A. Habit;

scale bar

2 cm. B. Fruits,

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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.

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SYSTEMATICBOTANYMONOGRAPHS

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

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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;

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stone

cell

SYSTEMATICBOTANYMONOGRAPHS

66

VOLUME 61

D
FIG. 29. Solanum
of partially

sectioned

(Based on: A, Occhioni

pelagicum.

A. Habit.

flower. E. Gynoecium.
5307; B-F, Hatschbach

B. Compound
F. Stamen
& Forero

leaf. C. Flower,

(left to right: abaxial,


40366;

seen from above. D. Lateral


lateral, adaxial

G, Reitz & Klein

views).

6770.)

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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).

REPRESENTATIVE SPECIMENS. Brazil.


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]).

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SYSTEMATICBOTANYMONOGRAPHS

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

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2001

SOLANUM

69

A~~~~~~~~~~~~~~~~~~~~~~~~'

FIG. 30. Solanum


flower. D. Gynoecium.
material

of Bohs

stuckertii.
E. Stamen

A. Habit.

B. Flower,

(left to right: abaxial,

seen from above. C. Lateral


lateral, adaxial

views).

view

F. Fruit.

of partially
(Based

sectioned

on greenhouse

2522.)

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VOLUME 61

SYSTEMATICBOTANYMONOGRAPHS

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

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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).

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5808

72

SYSTEMATICBOTANYMONOGRAPHS

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

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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.

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74

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 61

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51: 748-752.

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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.

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in Bo

in

VOLUME 61

SYSTEMATICBOTANYMONOGRAPHS

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, +]

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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, -]

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79

SYSTEMATICBOTANYMONOGRAPHS

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).

This content downloaded from 192.188.53.215 on Fri, 17 Apr 2015 02:02:40 UTC
All use subject to JSTOR Terms and Conditions

(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

Gutierrez, J. 333 (13).

5936 (2).

R., et al. 5817

Knapp,

S., & J.Mallet

Knight,

D. H. 673

Krapovickas,

A.

(12).
6465

(10).

(4).
1711

(13).

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(12), 8834

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MONOGRAPHSVOLUME
61
SYSTEMATIC

82
Krapovickas,

A., & C. Crist6bal

17367

(13), 44389

Krapovickas,

A., & J. Irigoyen

Krapovickas,

& A.

A.,

(6), 13196

12964

(6),

(6).
17844

Schinini

(6).

30853

(6),

38643

0.

Navarro,

G., &

Navarro,

G., et al. 19 (6).

Nee, M.

25505

(13).

38328

Krapovickas,

et al. 19897

A.,

(6), 26742

M.

Kummrow,

R., et al. 3067

Lillo, M.

4543

(13), 8220

7454

(2), 7284

J. C., et al. ICN # 21246


(5), 5880

L6pez,

A., & A. Sagdstegui

L6pez,

A., & E. Saravia

Lopez,

A.,

et al. 9089

P. G.

Lossen,
Lotti,

7894

427

(1).

(2).

15 (13), 29 (13),
(6), 1018

101 (13),

lOlb

(13),

360 (13), 568

(6).

Luti, R. 4628
Macbride,

Madison,

M.,

(10).

& Arenas

1038

(4).
(13).

153 (6).
(13).

P., et al. 7040

Nuiiez,

L., & V. Nduiez

Marmol,

L., et al. 8773

Martinez

Crovetto,

V. 155 (2).

(2).
(2), 37409

(6), 8171

(2).

(2).

(2).
(10).

8667

(10).

(10).

P. 5307

O'Donell,

C. A. 70 (2), 4163

(12).

F. I., & J. Jaramillo

Ortfz,

510

(13).

140 (10).

(6).

5567

(2).

R. 1230

8262

(13).

F. 163 (6), 704

(6), 1017

F., & L. Ramella

2692

Mexia,

Y. 4039

(9), 4043

Meyer,

T. 4543

(13), 6458

T., et al. 20370

(6), 2592

(6).

T. M.

J., & G. Marino

1045

(6), 10094

(2), 18911

(6), 15442

(2),

(2).

E. 5373

E., & J. P. Hjerting

Piccinini,

B., & J. Hilfer


S. 4063

(5).

T. 2724

Plowman,

T., & E. W. Davis

J. 3455

P. C. 772
C. 590

Quarfn,

C., & A. Schinini

1044

(6).

Quarfn,

C., & S. Tressens

1436

(6).

(6), 3058

2139

Ragonese
Rambo,

B. 40302

(6).
(6).

(3), 45760

3650

(6), 3739

(6), 4072

(6), 4910

(6), 5087

(6),

Reitz, R., & R. Klein

5174

(6), 5268

(6), 6566

(6).

R. 1732

(6).

(6), 3255

Reitz,

(13).

(2).

(6).

Quarfn,

(6),

(12), 5081

(3).

(3), 53509

(12).

4 (12), 6770

(12), 7053

(3), 7254

(11), 16873 (5).

(6).

R. R278

Renolfi,
(5).

(5), 218

Rizzini
(5).

(13), 414

S. A. 3546

Renvoize,

113 (6).
217

(10), 5150

(9).

(6),

E. A., & G. Barboza

4913

(9).

(6), 3592

E. A., & J. R. Davifia

(6).

(6).

(6), 2602

Moscone,

(2).

L. 91 (2).

(6), 2286

Moscone,

(2), 618

(6).

(6), 4157

(6), 4085

Plowman,

(6), 1158 (6), 1346

(5), 2073

565

4230

J. R., et al. 164 (9).

(6), 1670

O., et al. 799

(6).

(6).

(6).

Petersen,

1624

181 (6), 1530

(6), 4198

4423

(5).

L., et al. 496

Quarfn, C., et al. 383


(5), 10404

(6), 898

Morong

(11).

(6), 2595

Prance, G., et al. 26173

(2).

Morrone,

6086

1 (6), 2 (6).

Pedersen,

Porto,

(6).

(9).

(5), 9500

I. 305 (6), 655

L. M.

Penseiro,

Pohl,

Mereles,

(2), 16374

(3).

82 (13), 106 (13).

Pizarro,

(6).

Mereles,

J. E. 1573

C., & Rojas

Pirani,

(2).

V., et al. 595

T. 565

(2), 36559

Occhioni,

Pierotti,

(10).

Marmol,

Morel,

(2), 34038

Peter, A. 54 (6).

N. E. de, et al. 10225C

Meyer,

Nuinez,

Perez,

(6).

159 (10), 1872

15445

P., & E. Bengoa

Pereira,

(13), 1013

Marin

Montes,

(10).

(6).

Marco,

Marufiak,

Nuiiez,

Passarelli,

M. R. 28 (13).

Malvarez,

30226

(10), 7361

Pabst, G., & E. Pereira


(10), 3795

et al. 4972
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Maranta

(2), 500

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Osten,

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(7), 33970

(2), 37398

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(13).

Macedo

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(13).

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(7), 33950

Osten, C. 5281 (6), 21883 (6).

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(6),
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(6).

194 (13).

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Lourteig,

Mdfioz,

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(3).

(9).

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(2), 2839

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L. 247

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(2).

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(2), 46524

(2),

(7), 36628 (2).

(2),

Nee, M., &


(2), 38954

Lindeman,

Loefgren,

(6), 40633

(7), 37532

38427

(2),

& J. Solomon

Nee, M.,

(3).

9938 (2).
38693

(6).

(6), 35544

(6), 35186

Nee, M., & G. Coimbra

(9).

P. R., & A. R. Cuezzo

Lewis, M.

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(7), 39996 (6).

Kuhlmann,

2597

(6).
I. Vargas

(7), 38389

40174

(6), 26861

(6), 27317 (6), 27503 (6), 27517 (6).

Legname,

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Murquia,

268

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(6).

(9).

A. P. 395

(13), 486

111 (6), 464

(13), 994

(3), 543

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Rodriguez,D. 1087 (13).


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T. 318

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J. da Costa

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Solomon, J. 3030 (10), 8694 (2), 9347 (2), 11274 (2),


12566 (2).

816

A. 5862

Sagastegui,

A.,

et al. 15117

Saint-Yves,

A.

156 (6).

(8), 9614

Soukup,

(8), 4220

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Schaller

258

Schiavone,

(6), 301

1171

(2).

F. 162

A.

3919

12611

(6),

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(6), 5350
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(13),

(6), 6721

(6), 22746

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(13).
64608

(6), 9033
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(13).

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(2), 1385

C. 200

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(13), 3336

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93 (6).

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(2), 76 (6), 312

H., & E. Walter

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Smith, L. B., & R. Reitz

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(13), 1451

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et al. 15 (2), 30 (2).

Sleumer,

(2), 3718

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(4).

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(10), 9325

Vargas,

(5).

(6).

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Vargas,

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et al. S124

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(13),

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(9).

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(3), 4162

et al. 1299

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(6),

VOLUME 61

SYSTEMATICBOTANYMONOGRAPHS

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

sect. Cyphomandropsis
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

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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

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