A Revision of Solanum Section Gonatotrichum

Abstract

Solanum section Gonatotrichum (Solanaceae) includes eight species native to North, Central, and South America. Plants of this section are herbs to woody shrubs that lack spines, are pubescent with simple or stellate hairs, and have berries that swell due to increased turgor pressure and explosively dehisce to disperse the seeds. Section Gonatotrichum is closely related to section Brevantherum, from which it differs by the presence of explosive fruit dehiscence and simple hairs in all taxa except S. lignescens, which has stellate pubescence. The morphology, taxonomic history, nomenclature, ecology, distribution, and reproductive biology of Solanum section Gonatotrichum are reviewed. A dichotomous key is provided for the species of the section.

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A Revision of Solanum Section Gonatotrichum
Author(s): Stephen Stern , Lynn Bohs , Leandro Giacomin , João Stehmann and Sandra Knapp
Source: Systematic Botany, 38(2):471-496. 2013.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364413X666624
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Systematic Botany (2013), 38(2): pp. 471–496
©Copyright 2013 by the American Society of Plant Taxonomists
DOI 10.1600/036364413X666624
A Revision of Solanum Section Gonatotrichum
Stephen Stern,
1,4,5
Lynn Bohs,
1
Leandro Giacomin,
2
Joa
˜o Stehmann,
2
and Sandra Knapp
3
1
Department of Biology, University of Utah, 257 South 1400 East, Salt Lake City, Utah, 84112-0840, U. S. A.
2
Departamento de Bota
ˆnica, Instituto de Cie
ˆncias Biolo
´gicas, Universidade Federal de Minas Gerais,
Avenida Anto
ˆnio Carlos, 6627, 31270-901, Belo Horizonte, MG, Brazil
3
Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD, U. K.
4
Department of Biology, Colorado Mesa University, 1100 North Avenue, Grand Junction, CO, 81501, U. S. A.
5
Author for correspondence (sstern@coloradomesa.edu)
Communicating Editor: Carol Anne Wilson
Abstract—Solanum section Gonatotrichum (Solanaceae) includes eight species native to North, Central, and South America. Plants of this
section are herbs to woody shrubs that lack spines, are pubescent with simple or stellate hairs, and have berries that swell due to increased
turgor pressure and explosively dehisce to disperse the seeds. Section Gonatotrichum is closely related to section Brevantherum, from which
it differs by the presence of explosive fruit dehiscence and simple hairs in all taxa except S. lignescens, which has stellate pubescence.
The morphology, taxonomic history, nomenclature, ecology, distribution, and reproductive biology of Solanum section Gonatotrichum are
reviewed. A dichotomous key is provided for the species of the section.
Keywords—explosive fruit dehiscence, Neotropics, taxonomy.
Solanum (Solanaceae) contains approximately 1500 species
and is one of the 10 largest genera of flowering plants (Frodin
2004; Bohs 2005). Due to the large size and the morphological
complexity of the genus, many infrageneric groups within
Solanum are not well defined. Recent species level taxonomy
(e.g. Bennett 2008; Knapp 2008; Peralta et al. 2008; Tepe and
Bohs 2011; www.solanaceaesource.org) has led to a better
understanding of Solanum systematics and numerous molecu-
lar phylogenetic studies have helped elucidate clades within
the genus (Bohs 2005; Weese and Bohs 2007). One clade
that has been recognized in molecular studies is the
Gonatotrichum clade, which corresponds to the formally
named Solanum section Gonatotrichum Bitter. This contribu-
tion is a revision of that section.
Section Gonatotrichum includes eight species native to
North, Central, and South America. They are all herbaceous
or small, woody shrubs with simple hairs (except S. lignescens
Fernald which has stellate pubescence). The fruits are unique
within the genus and are a key diagnostic character for the
section. They have a thin pericarp with a watery mesocarp
that is held under pressure until the fruits explosively dehisce
(Nee 1989). While it is unusual to witness the explosive
dehiscence, the syndrome is apparent, as the fruits are white,
yellow, or green and nearly transparent when mature. After
dehiscence, the fruits appear deflated and shriveled. Section
Gonatotrichum has previously been taxonomically challenging
because while some species are relatively widespread in
their distribution (S. deflexum Greenm., S. olympicum Hassl.,
and S. turneroides Chodat), others are narrowly distributed
and relatively inconspicuous, making them among the least
collected species of Solanum.
Materials and Methods
The taxonomic conclusions presented are a result of extensive field and
herbarium work. We examined specimens from the following herbaria:
A, BH, BHCB, BM, BR, CEPEC, CESJ, CORD, CTES, ESA, FUEL, G, GH,
HAS, HB, HNUP, HUEFS, IAC, IAN, IBGE, ICN, INB, INPA, JPB, K,
LPB, M, MBM, MBML, MEXU, MO, NY, P, PACA, PEL, QCA, QCNE, R,
RB, SI, SP, SPF, SPSF, TEX, UC, UEC, UPCB, US, USZ, UT, VIC, WIS,
W, WU. We also field-collected five of the eight recognized spe-
cies, including two recently described species (Stern and Bohs 2009;
Giacomin and Stehmann 2011). Details of exsiccatae are cited here
and are also available on the Solanaceae Source webpage (www
.solanaceaesource.org). Specimens with sheet numbers are cited with
the herbarium acronym followed by a dash and the sheet number (i.e.
MO– 1781232); barcodes are preceded by the herbarium acronym
followed by a dash, then the barcode number (i.e. G –G00104280).
We have followed the morphological species concept or “morphologi-
cal cluster” concept in delimiting species of sect. Gonatotrichum (Mallet
1995). Taxa are recognized as distinct if they possess a unique suite of
characters and are separated from similar entities by morphological gaps.
In nearly all cases, taxa also occupy geographically circumscribed ranges.
Measurements were made from dried herbarium material supple-
mented by measurements from living material. Colors of corollas, fruits,
etc., are described from living material or from herbarium label data.
Specimens with latitude and longitude data on the labels were
mapped directly. Some species had few or no georeferenced collec-
tions; in these cases we retrospectively georeferenced the collections
using available locality data. Conservation status was assessed follow-
ing IUCN Red List Categories and Criteria (I. U. C. N. Standards and
Petitions Subcommittee 2010).
For seed SEM studies, seed coats were digested using a 1% cellulase
solution for 24 hr to remove the periclinal cell walls and expose the
anticlinal cell walls. For hair SEM studies, fresh stems, when available,
were mounted on a stub with double-sided tape and coated with gold-
palladium. For those species where fresh material was not available,
dried material from herbarium specimens was used. The material was
rehydrated and postfixed in osmium tetroxide and then dehydrated in
an ethanol series before being coated with gold-palladium.
Plants used in breeding studies were grown from seed in the green-
houses at the University of Utah. Voucher information and original
locality data of materials used are given in Appendix 1. For plants that
were not autogamous, crosses were made by shaking pollen onto a glass
slide that was rubbed gently across the stigma of the pollen-receptor
flower. Success or failure of the cross was monitored, as well as fruit
size, shape, color, and number of seeds in successful crosses.
Taxonomic History
The earliest descriptions of species that would later be
assigned to sect. Gonatotrichum were Sendtner’s (1846)
publication of S. adscendens Sendtn. and S. hoffmanseggii
Sendtn. in his treatment of the Solanaceae in Martius’ Flora
Brasiliensis. Asa Gray described Salpichroa wrightii A. Gray
in 1886, which would later be recognized as a synonym
of S. deflexum.In1897,FernalddescribedtheMexicanspe-
cies S. lignescens.Alsoin1897,GreenmandescribedaCen-
tral American species, S. deflexum. Chodat (1902) described
the South American species S. turneroides based on mate-
rial from E
´mile Hassler’s herbarium in Geneva. In 1911,
471

Hassler described S. olympicum Hassl. from material col-
lected in Paraguay.
In 1912, Bitter described two new species from Bolivia and
Paraguay, S. gonatotrichum Bitter and S. geniculatistrigosum
Bitter, both treated in this revision as synonyms of
S. turneroides. He also formally established sect. Gonatotrichum,
including these two species as well as S. adscendens and
S. deflexum. In his concept, plants of the section were
small annuals or perennials with few-celled, unbranched,
often geniculate hairs, few-flowered, nearly sessile inflo-
rescences, and glabrous filaments and styles.
One year later, Bitter (1913) described two other new spe-
cies from Paraguay, S. flavistrigosum Bitter (here considered
as another synonym of S. turneroides) and S. parcistrigosum
(here considered a synonym of S. olympicum), and assigned
S. hoffmanseggii to the section. Later, Bitter (1922) trans-
ferred Bassovia setosa Brandegee (Brandegee 1917) to Solanum
and placed it in sect. Gonatotrichum (Bitter 1922). Solanum
setosum (Brandegee) Bitter is now considered a synonym
of S. deflexum.
Solanum sect. Gonatotrichum received virtually no taxo-
nomic attention until Nee (1989) reduced the number of
species in the section to two, recognizing only S. adscendens
and S. turneroides;heleftS. hoffmanseggii as a name of
doubtful affinity. He reiterated the importance of hair charac-
ters in defining the section and also emphasized the explo-
sively dehiscent fruits as a distinctive character.
Recent herbarium and field studies have resulted in the
description of S. manabiense S.Stern from Ecuador (Stern
and Bohs 2009) and S. evolvuloides Giacomin & Stehmann
from Bahia, Brazil (Giacomin and Stehmann 2011). Con-
current phylogenetic and taxonomic work has led to the
present revision as well as a published molecular phylogeny
of the section (Stern and Bohs 2012; see below).
Morphology
Habit—Members of sect. Gonatotrichum are small herbs,
single- to few- branched from a lignescent base, with only
two species, S. lignescens and S. hoffmanseggii, becoming
subshrubs. Species range in height from mature individuals
of S. deflexum that reach only 5–10 cm to the shrubby
S. lignescens that can grow to 1 m. Some species (S. manabiense
and S. turneroides and rarely S. deflexum) are known to pro-
duce many upright stems from underground rhizomes. The
herbaceous habit of sect. Gonatotrichum most resembles
members of sect. Solanum (Morelloid clade sensu Weese and
Bohs 2007) but various other morphological and molecular
characteristics distinguish these sections (see Bohs 2005).
Trichomes—As in some other solanums (Seithe and
Anderson 1982), trichome morphology has proven to be
an excellent characteristic in delimiting species within the
section. Both branched and unbranched trichomes are
found in sect. Gonatotrichum (Fig. 1). Branched, stellate
hairs are found only in S. lignescens, which also has simple,
unbranched hairs. The unbranched hairs range from one-
to five-celled and can either be straight (in S. adscendens,
S. deflexum,andS. manabiense,) or bent at a 90!angle between
the first and second cells (S. olympicum and S. hoffmanseggii).
The trichomes of S. turneroides fall between these catego-
ries and vary from straight to bent at nearly a 90!angle.
Glandular hairs in sect. Gonatotrichum appear to be confined
to S. evolvuloides (Fig. 1F); however, SEM work has shown that
very small glandular hairs not visible with light microscopy
may be present in some other species (e.g. S. turneroides, Fig. 1B).
Among the species of sect. Gonatotrichum,trichomedensity
can vary from sparse to dense. Variability of trichome den-
sity changes both among and within species depending on
position on the plant. For instance, the abaxial surface of
the leaves is generally more pubescent than the adaxial
Fig. 1. Scanning electron micrographs of hairs in section Gonatotrichum.A.Solanum turneroides stem; note the hairs oriented downward at an
approximately 45!angle (Bohs 2715, UT). B. Higher magnification of S. turneroides; note that the base of the hair is swollen with what appear to be
reduced ray cells. Also note the small glandular hairs below the long, simple hairs (Bohs 2715, UT). C. Stem of S. deflexum; note the hairs are simple
and held perpendicular to the stem (Nee 51716, UT). D. Geniculate hairs of S. olympicum; note the sharp 90!angle between the first and second cells
(Bohs 3194, UT). E. Stellate hairs on abaxial leaf surface of of S. lignescens (Hampshire et al. 1151, BM). F. Solanum evolvuloides glandular calyx hairs
(Mattos Silva, s.n.,CEPEC).Scalebars:A,C=500mm. B = 50 mm. D, E = 200 mm. F = 100 mm.
472 SYSTEMATIC BOTANY [Volume 38

surface. Similarly, young growth is commonly densely
pubescent while older growth on the same plant may be
nearly glabrous.
Leaves—The leaves of sect. Gonatotrichum are all entire,
petiolate (or nearly sessile in S. hoffmanseggii), and elliptic
or elliptic-ovoid (except in S. adscendens with cordiform
leaves and S. hoffmanseggii which can have linear leaves).
The base of the leaf is somewhat decurrent into the petiole
in all species except in the cordate leaves of S. adscendens.
Leaf size is relatively uniform within the section, with the
smallest leaves ca. 1 cm long and the longest ca. 7 cm long,
with greater intraspecific than interspecific variation. Leaf
texture is chartaceous to membranaceous. Venation is typi-
cally pinnate with one major vein from the base, except in
S. adscendens, which has 3–5 major veins extending from
the leaf base.
Inflorescences—The inflorescences in sect. Gonatotrichum
are all sessile or nearly so, unbranched, and extra-axillary
or subopposite the leaves. The inflorescences are all few-
flowered, with 1–6 (12) flowers. They are typically very
congested with a short to almost absent peduncle and a
condensed rachis. The pedicels are all articulated at the
base. The sessile inflorescence is a shared characteristic of
all species of sect. Gonatotrichum.
Flowers—The flowers of sect. Gonatotrichum are all per-
fect, 5-merous, and white or sometimes light pink to purple
in color. The flowers are small compared to many other
Solanum species, with corollas typically about 1.5 cm in
diameter (to 1.8 cm in S. lignescens, 2.5 cm in S. turneroides,
and 3 cm in S. evolvuloides). The flowers of S. adscendens,
S. deflexum,S. lignescens, and S. manabiense are actinomor-
phic. The flowers of S. olympicum and S. hoffmanseggii are
very slightly heterantherous, with the lowermost anther
slightly projected due to an elongated filament, although
this is often so inconspicuous as to be very difficult to see on
herbarium specimens. Solanum turneroides and S. evolvuloides
both have strongly heterantherous flowers with the lower-
most filament elongating to twice the length of the others.
These species also have a style that is deflected upwards at
the apex. Solanum turneroides has flowers that are strongly
fragrant, a unique trait in the section and uncommon in
Solanum in general.
Fruits—The fruits in sect. Gonatotrichum are unique in
the genus and are an excellent diagnostic character for the
section. They are glabrous, green, white, or yellow when
immature and green or green-purplish and transparent
when mature. The fruits have a thin pericarp with a watery
mesocarp that is held under pressure until the fruits explo-
sively dehisce. This dehiscence can propel seeds over 15 m
from a 30 cm plant in S. turneroides.
Seeds—The seeds of sect. Gonatotrichum are flattened and
reniform like those of many other Solanum species (Fig. 2).
In overall shape, S. deflexum is unique in that it has
a swollen margin and a large notch where it connects
Fig. 2. Scanning electron micrographs of seeds in section Gonatotrichum. All seeds were treated with 1% cellulase for 24 hr and dried prior
to imaging. A. Solanum olympicum; note the raised ridges radiating from the center to the margins (Bohs 3194, UT). B. S. adscendens; note the raised
ridges (Leite 658, NY) C. S. deflexum; note the swollen margin and large notch (Nee 51716, UT). D. Solanum turneroides (Bohs 2715, UT). Scale bars = 2 mm.
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 473

to the placenta (Fig. 2C). In S. turneroides the seeds are
twisted and bent, in contrast to the flat, reniform seeds
of other species in the section. The arrangement of cells
on the seed surface can be diagnostic. The individual cells
can be seen when the periclinal cell wall is enzymatically
removed and range from raised fibrils that radiate from
the seed center to the margins in S. olympicum (Fig. 2A) to
the netlike cell walls of S. turneroides (Fig. 2D).
Breeding Systems
Plants of S. deflexum, S. olympicum, S. manabiense,and
S. turneroides were studied in the greenhouses at the Uni-
versity of Utah, while those of S. adscendens and S. evolvuloides
were studied at the Jardim Bota
ˆnico da Fundac¸a
˜oZoo-
Bota
ˆnica, Belo Horizonte, Brazil. All species were her-
maphroditic and self-compatible, and all but S. evolvuloides
and S. turneroides were autogamous. Among the autoga-
mous species, S. adscendens, S. deflexum, and S. olympicum
were strongly autogamous, while a smaller proportion of
flowers of S. manabiense set fruit without manipulation.
Habitats and Geographic Distribution
Species of sect. Gonatotrichum are native to the south-
western United States, Mexico and Central America as far
south as Costa Rica, and to coastal Ecuador, Brazil, Paraguay,
Bolivia, and Argentina in South America. Species can be
found from sea level to 2,700 m. Although they occupy
various habitats, all seem to prefer dry sites, including dry
deciduous forests, roadsides, grazed areas, and forest edges.
Some of the taxa in the section are widespread: S. deflexum
ranges from the southwestern United States to Costa Rica,
S. lignescens ranges from Mexico to Nicaragua, and
S. turneroides and S. olympicum range throughout Bolivia
and Paraguay and into southeastern Brazil and northern
Argentina. Other members are restricted to much smaller
regions: S. manabiense is only found in central coastal Ecuador
in the Manabı
´and Guayas Provinces, S. adscendens is only
found in Rio Grande do Sul State in Brazil and adjacent
Corrientes and Misiones Provinces in Argentina, S. hoffmanseggii
is only known from collections in Para
´and Tocantins
States in northern Brazil, and S. evolvuloides is only known
from southeastern Bahia State, Brazil.
A notable aspect of the distribution of members of sect.
Gonatotrichum is their clustering in four regions (North and
Central America, coastal Ecuador, northern Brazil, and
Bolivia to southeastern Brazil/northeastern Argentina) with
large disjuctions between them. Bitter’s (1912) description of
the section noted the large geographic disjunction between
the southern South American and North to Central American
members of the section and predicted that a close relative
of these species might be expected in the territories lying
between these disjunct areas. In fact, S. manabiense of coastal
Ecuador was found in the intervening area (Stern and Bohs
2009). Solanum olympicum is the only species known to
span a large disjunction, being found in both the northern
and southern Brazilian centers of distribution. However,
specimens from the northern part of the range are some-
what aberrant in hair structure, and further collections and
study may show that they are distinct from the southern
populations (see taxonomic treatment below). Part of the
explanation for the large disjunctions in this group may be
due to their preference for dry habitats, but even so, they
are not found in all of the drier areas of the Neotropics.
Phylogenetic Relationships
Previous authors (Sendtner 1846; Nee 1999) allied species
of sect. Gonatotrichum with sect. Solanum of subg. Solanum.
Both groups are superficially very similar in their herba-
ceous, often weedy habit, pubescence of unbranched hairs,
usually short, unbranched, often extra-axillary inflorescences,
and small white flowers. However, molecular data (Bohs
2005; Weese and Bohs 2007) showed that sect. Gonatotrichum
is unrelated to sect. Solanum and instead belongs to the
Brevantherum clade of Solanum. This clade consists of about
60 Neotropical species lacking prickles and with short,
broad anthers. Most species of the Brevantherum clade have
stellate or lepidote pubescence, often large, branched inflores-
cences, and fleshy, non-dehiscent fruits. Section Gonatotrichum
is unusual in the Brevantherum clade because of its herba-
ceous habit, reduced inflorescences, simple hairs in all species
except S. lignescens, and its explosively dehiscent fruits.
Phylogenetic relationships of Solanum sect. Gonatotrichum
were further examined by Stern and Bohs (2012). Molecular
data indicate that sect. Gonatotrichum forms a monophyletic
group including S. lignescens that is sister to all other sam-
pled members of the Brevantherum clade (Fig. 3; Stern and
Bohs 2012). Within sect. Gonatotrichum, phylogenetic results
closely follow the geographic distribution of species in the
section, with the exception of S. adscendens, a species from
southern Brazil that does not form a clade with the other
South American species, but is sister to the rest of the spe-
cies in the section (Fig. 3); this relationship, however, is
not well supported. The two Mexican and Central American
species, S. lignescens and S. deflexum, are strongly supported
as sister species. Solanum evolvuloides, S. turneroides, and
S. olympicum, all have heterantherous flowers (although
those of S. olympicum are weakly so), and form a monophy-
letic group with S. evolvuloides sister to the other two species
(see Fig. 3). Multiple accessions of S. olympicum are grouped
together, with those from the northeasternmost extent of
the range in Bahia, Brazil, sister to the other accessions.
While these populations have some slight morphological
differences, they are treated here as the same species. Solanum
hoffmanseggii was not included in the phylogenetic study
due to a lack of high quality genomic DNA.
Taxonomic Treatment
SOLANUM section GONATOTRICHUM Bitter, Repert. Spec.
Nov. Regni Veg. 11: 230. 1912.—TYPE species: Solanum
gonatotrichum Bitter (= Solanum turneroides Chodat).
Herbs or shrubs, sometimes rhizomatous, lacking prickles.
Stems nearly glabrous to densely pubescent with straight
or geniculate, unbranched and/or stellate hairs. Leaves
simple, the blades chartaceous to membranaceous, elliptic
to elliptic-ovoid to cordiform, glabrous to densely pubes-
cent with straight or geniculate, unbranched and/or stellate
hairs, the base rounded to obtuse to truncate or cordate,
often decurrent into petiole, the margins entire and often
ciliate, the apex acute; petioles sparsely to densely pubes-
cent. Inflorescence nearly sessile, extra-axillary or sub-
opposite the leaves, unbranched, with 1– 6 (12) flowers, the
474 SYSTEMATIC BOTANY [Volume 38

Fig. 3. Parsimony strict consensus tree of Solanum section Gonatotrichum and outgroups based on plastid trnT-trnF and nuclear waxy and ITS
(from Stern and Bohs 2012).
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 475

axes sparsely to densely pubescent; pedicels articulated
at the base. Flowers 5-merous, perfect, actinomorphic or
zygomorphic due to heteranthery. Calyx campanulate to
spreading, sparsely to densely pubescent, the lobes linear-
lanceolate to broadly triangular, erect to reflexed at anthesis,
not to slightly accrescent in fruit. Corolla white to pale pink
or purple, membranaceous, stellate to rotate with abundant
interpetalar tissue, glabrous adaxially, glabrous to sparsely
pubescent abaxially, mostly on apex and main veins. Sta-
mens equal or one stamen borne on an elongated filament
to twice the length of the others, the filaments short to
greatly elongated, glabrous, inserted in corolla tube near
its base; anthers connivent, yellow, sagittate at the base,
poricidal at the tips, the pores directed distally, not (or
rarely) opening into longitudinal slits. Ovary glabrous
to sparsely pubescent; style equal to or exserted beyond
stamens, cylindrical, glabrous, the stigma capitate. Fruit
a globose to ovoid berry, obtuse at apex, glabrous, green
to white or purplish-black when ripe, the pericarp thin,
the mesocarp watery and held under pressure, dehiscing
explosively at maturity. Seeds somewhat flattened, often
with raised ridges on outer margin.
A Key to the Species of SOLANUM Section GONATOTRICHUM
1. Plants usually woody (occasionally herbaceous), 0.5– 1.5 m tall; hairs predominantly stellate; sympodia 3-foliate
to plurifoliate; peduncle 3– 8 mm . ............................................................................ 5.S. lignescens Fernald
10. Plants herbaceous from a slightly woody base or woody in few cases (S. hoffmanseggii), 0.3– 0.6 (0.8) m tall;
hairs simple; sympodia 2-foliate; peduncle absent or nearly so ...................................................................... 2
2. Cauline hairs exclusively geniculate (bent at a 90!angle between first and second cells), at least on the older
parts of the stems, and lying flat along stems . . . .............................................................................. 3
3. New growth densely pubescent with simple hairs, these lying flat along stem but lacking a strong 90!bend
(see Figs. 1A, B); corollas 1– 2.5 cm in diameter, white to purple; flowers heterantherous with one filament
longer than the rest, 2– 5 mm long; mature fruits 10 –20 mm in diameter, the fruit wall thickened at apex
resulting in dehiscence at proximal end near calyx ...................................................... 8.S. turneroides Chodat
30. Hairs strictly geniculate with a strong 90!bend; corollas 0.5–1.3 ( –1.5) cm in diameter, white; flowers
homantherous or very weakly heterantherous (only faintly noticeable in live plants and barely visible
in dried specimens), all filaments equal or nearly so, 1– 2 mm long; mature fruits up to 12 mm in diameter,
the fruit wall not thickened at apex ...................................................................................... 4
4. Plants subshrubs, 50– 80 cm tall; leaves lanceolate, more than three times longer than wide; plants
of Amazonian Brazil (Para
´and Tocantins) . . . ..................................................... 4.S. hoffmanseggii Sendtn.
40. Plants herbaceous, rarely woody, often rhizomatous, under 40 cm tall; leaves elliptic to elliptic-ovate,
less than three times longer than wide; widespread species of Paraguay, Argentina, Bolivia and
the xeric axis of Brazil (Bahia, Goia
´s and Mato Grosso do Sul) .......................................... 7.S. olympicum Hassl.
20. Cauline hairs straight, not geniculate, and not lying flat along stems . . . ............................................................ 5
5. Corollas 1– 2.5 cm in diameter; flowers heterantherous with one filament longer than the rest, 2 –5 mm long;
mature fruits 10– 20 mm in diameter ..................................................................................... 6
6. Leaves typically 1.5– 3.5 (4) cm long; abaxial surface of calyx and pedicels with long
multicellular glandular hairs (ca. 0.5 mm long); plants of east-central Bahia, Brazil . . . ..... 3.S. evolvuloides Giacomin & Stehmann
60. Leaves typically 3.5 –8 cm long; abaxial surface of calyx and pedicels with exclusively eglandular
hairs; plants of Argentina, Bolivia, Paraguay, and Brazil (Mato Grosso do Sul and Sa
˜o Paulo) . . . .......... 8.S. turneroides Chodat
50. Corollas 1 –1.7 cm in diameter; flowers homantherous, all filaments equal, 1– 2 mm long; mature fruits 5–12 mm in diameter ........... 7
7. Plants much-branched and spreading from a slightly woody base; leaves cordate, with small glandular
hairs on both sides; plants of northeastern Argentina (Provs. Misiones and Corrientes) and southern
Brazil (Rio Grande do Sul) . . ..................................................................... 1.S. adscendens Sendtn.
70. Plants not or few-branched, the base typically not woody; leaves elliptic to elliptic-ovoid, the base rounded to obtuse,
with exclusively eglandular hairs on both sides; plants of North and Central America and coastal Ecuador . . ................... 8
8. Plants arising from a single base, rarely rhizomatous; abaxial surface of leaves densely pubescent;
seeds with swollen margins and a pronounced notch where connected to placenta; plants
of southwestern USA and widely distributed from Mexico through Costa Rica . . ..................... 2.S. deflexum Greenm.
80. Plants always rhizomatous; abaxial surface of leaves nearly glabrous to sparsely pubescent;
seeds uniformly flattened throughout without a prominent notch where connected to placenta;
plants of coastal Ecuador . .................................................................... 6.S. manabiense S.Stern
1. SOLANUM ADSCENDENS Sendtn., Fl. Bras. (Martius) 10: 17,
Table 1, Figs. 9–12. 1846.—TYPE: BRAZIL. “Brasilia
australiore,” (fl), F. Sellow s.n. (lectotype, here desig-
nated: P-P00319345!; isolectotype: B (destroyed), photos
of isolectotype [F neg. 2798]: F!, G!, GH!).
S. amarantoides Dunal var. hirtellum Dunal, in DC., Prodr.
13 (1): 56. 1852.—TYPE: BRAZIL. “Province de Rio-
Grande”, [“de Minas Geraes” handwritten on label],
1833 (fl), C. Gaudichaud 1745 (holotype: P- P00319346!).
Herb, sometimes woody at base, much-branched, the
branches decumbent with apices upright, 1–3 dm tall. Stems
sparsely to densely pubescent with 2–5-celled unbranched,
straight hairs. Sympodia 2-foliate, the leaves solitary or gemi-
nate. Leaf blades 1.5–4
+
1– 3.5 cm, cordiform, chartaceous
to membranaceous, nearly glabrous to sparsely pubescent
adaxially and abaxially with 1– 2 (4)-celled unbranched
eglandular hairs, these 0.5–1.5 mm, erect or lying flat along
blade, denser along veins, mixed with small, glandular hairs,
these 0.1–0.2 mm, barely visible in dried material; base trun-
cate to cordate, often asymmetrical, slightly decurrent into
petiole; main veins 3– 5, palmately leaving the leaf base;
apex acuminate to acute; petioles 0.5– 1.5 cm, moderately
pubescent with unbranched hairs like those of stems.
Inflorescences with 1–3 flowers, the axes sparsely to mod-
erately pubescent with unbranched hairs; peduncle absent
or nearly so; rachis absent; pedicels 5–15 mm in flower,
10– 20 mm in fruit, pendent. Flowers with the calyx 3– 10 mm
long, the tube 1–3 mm, the lobes 2–7
+
0.5– 1.5 mm, linear-
lanceolate, moderately to densely pubescent. Corolla
0.5–1.5 cm in diameter, rotate with abundant interpetalar
tissue, chartaceous to membranaceous, white, the tube
476 SYSTEMATIC BOTANY [Volume 38

3– 6 mm long, the lobes very short, 1–2
+
0.5–1 mm, trian-
gular, acute at apices, glabrous abaxially and adaxially.
Stamens 2–4 mm long; filaments up to 1 mm long; anthers
1.5– 3
+
0.5–1.5 mm, oblong, the base cordate, the apex
emarginate. Ovary glabrous; style 4– 6
+
0.5–1 mm, equal
to or exserted beyond stamens; stigma to 1 mm wide.
Berries 5–12 mm in diameter, globose, white to yellow
when immature, maturing semitransparent, drying brown,
glabrous, the mesocarp probably watery and held under
pressure until dehiscing explosively at maturity. Seeds
10– 35 per fruit, ca. 2.5
+
1.5 mm, with a small notch where
connected to placenta, the margin not swollen, surface with
fine raised ridges radiating from center to edges and shal-
low ridges running parallel to margin. Figure 4.
Habitat and Distribution—The majority of collections of
S. adscendens are from Rio Grande do Sul State in Brazil
Fig. 4. Solanum adscendens (Stehmann 6005, BHCB). A. Habit; note the cordate leaf bases. B. SEM of the abaxial leaf surface; note the long, straight
hairs and the minute glandular hairs. C. Fruit. D. Flower. E. Calyx and leaves. Scale bars: A = 2 cm. B = 200 mm. C, E = 1 cm. D = 5 mm.
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 477

(Fig. 5). A few collections exist from Misiones and Corrientes
Provinces in Argentina, but these are found near the border
between the two countries. Solanum adscendens is a weedy
species of interior forests and river banks as well as fields
and roadsides (commonly ruderal) in seasonal deciduous
forests at elevations from 0–600 (900) m.
Phenology—S. adscendens has been collected in flower and
fruit in all months except January and May.
Conservation Status—While S. adscendens status is Least
Concern because it does not meet IUCN Red List qualifica-
tions for a Threatened or even Near Threatened species, it is
still of some concern due to the limited geographic distribu-
tion and rapid conversion of wildlands to grazing and
farming in Rio Grande do Sul (IBGE 2010). It is advised
that this species be monitored in the future.
Etymology—The epithet adscendens refers to the species’
growth form, which begins by spreading horizontally with
subsequent ascending, erect flowering stems.
Additional Specimens Examined—BRAZIL.Rio Grande do Sul: Balnea
´rio
Iraı
´, 27 Oct 1976 (fl, fr), Arzivenco s.n. (ICN); Derrubadas, Parque Estadual
do Turvo, 31 Jan 1997 (fl, fr), Brack 1714 (ICN); same loc., 31 Jan 1997
(fl, fr), Brack 1807 (ICN); Montenegro, Polo Petroquı
´mico, 28 Jun 1977
(fl), Bueno 344 (HAS); Santo Amaro, 6 Jun 1996 (fl, fr), Carneiro 443
(ICN); General Ca
ˆmara, Santo Amaro, na quadra da igreja, 8 Oct 1995
(fl, fr), Carneiro s.n. (ICN); same loc., 15 Mar 1996 (fl, fr), Carneiro s.n.
(ICN); Trindade do Sul, assentamento Trinidade, 28 Feb 2008 (fl, fr),
Grings 340 (ICN); Marcelino Ramos, barranca do Rio Uruguai,
23 Sep 1987 (fl, fr), Jarenkow 720 (ICN, MBM, PEL); Cambara
´do
Sul, Itaimbezinho, 27 Dec 1988 (fl, fr), Jarenkow & Bueno 1171 (ESA);
Dois Irma
˜os, Cascata de Sa
˜o Miguel, 1 Nov 1984 (fl), Jeisen s.n. (ICN);
Vicinity of S. Leopoldo, Oct 1941 (fl, fr), Leite 658 (NY, RB, SP, UEC);
Sa
˜o Leopoldo, 1941 (fl, fr), Leite 1864 (RB, SP, UEC); Nonoai, Cascata
do Legeado Tigre, 2 Nov 1993 (fl, fr), Matzenbacher s.n. (ICN); Vena
ˆncio
Aires, Vol. da Pa
´tria, 5 Aug 1984 (fl), Pilz s.n. (ICN); Santa Clara, p.
Lageado, 18 Nov 1940 (fl), Rambo s.n. (PACA); Nonoai, ad. fl. Uruguai,
Mar 1954 (fl, fr), Rambo s.n. (PACA); Harmonia, 6 Oct 1945 (fl, fr),
Sehnem 1546 (PACA, US); Verano
´polis, pro
´ximo ao Rio das Antas,
2 Nov 1989 (fl, fr), Silveira 1699 (HAS); Tenente, Parque Estadual do
Turvo, na estrada para Salto de Yucuma
˜, 11 Sep 1990 (fl, fr), Silveira
8734 (HAS); Triunfo, Estrada para Taquari, 24 Sep 1987 (fl), Silveira
9634 (HAS); Derrubadas, Parque Estadual do Turvo, na Estrada para
Porto Garcia, 20 Jul 1995 (fl), Sobral & Almeida 7911 (ICN); Bage
´,junto
ao Rio Camaqua
˜, 26 Sep 1984 (fl, fr), Stehmann 473 (BHCB, ICN, RB);
General Ca
ˆmara, Santo Amaro, Estac¸a
˜oFerrovia
´ria de Amaro
´polis,
28 Mar 2009 (fl), Stehmann et al. 6001 (BHCB); General Ca
ˆmara, Santo
Amaro, Eclusa, 29!56034.8800S, 51!53030.5100 W, 18 m, 28 Mar 2009 (fr),
Stehmann et al. 6002 (BHCB); same loc., same date (fr), Stehmann et al.
6003 (BHCB); same loc., same date (fl), Stehmann et al. 6004 (BHCB);
same loc., same date (fr), Stehmann et al. 6005 (BHCB); Montenegro,
Arroio Bom Jardim, 30 Aug 1977 (fl, fr), Ungaretti 549 (HAS);
Triunfo, 30 Dec 1996 (fl, fr), Ungaretti 595 (HAS); Montenegro, Polo
Petroquı
´mico, 13 Sep 1977 (fl, fr), Ungaretti 646 (HAS); same loc.,
19 Oct 1977 (fr), Ungaretti 730 (HAS); Montenegro, 24 Jul 1979 (fl, fr),
Waechter & Zanette s.n. (HAS).
ARGENTINA.Corrientes: Dept. Santo Tome
´,Garruchos,Estancia
San Juan Batista, 28!100S, 55!3805100W, 100 m, 17 Apr 2005 (fl, fr),
Barboza et al. 1494 (CORD); Garruchos, Estancia San Juan Batista,
costa del Rı
´o Uruguay, 20 Sep 1974 (fl, fr), Krapovickas et al. 25819
(CTES, MBM). Misiones: Concepcio
´ndelaSierra,entreAzaray
Ciudad de la Sierra, 24 Aug 1978 (fl, fr), Cabrera et al. 29445
(CTES, SI); San Pedro, Parque Provincial Mocona
´, 24 Oct 2006 (fl, fr),
Keller 3746 (CTES).
Fig. 5. Distribution of S. adscendens,S. evolvuloides,S. hoffmanseggii,S. olympicum, and S. turneroides.
478 SYSTEMATIC BOTANY [Volume 38

Notes—Solanum adscendens is similar to S. olympicum and
has been considered to be conspecific with it (Nee 1989).
Mentz and de Oliveira (2004) also placed specimens of
S. adscendens and S. olympicum together in S. adscendens. This
placement was largely due to the absence of type material,
as the presumed holotype was destroyed in Berlin and the
photographs of it were not sufficient to distinguish the
species. Mentz and de Oliveira’s (2004) photographs of a
duplicate of the type at Paris enabled us to readily distin-
guish S. adscendens from the more common and widespread
S. olympicum. The cordate leaf bases, straight (not genicu-
late) hairs, glandular hairs on the abaxial leaf surface,
and sprawling growth form with many branches from
a single base characterize S. adscendens and differentiate
it from S. olympicum,asdomoleculardata.
Solanum amarantoides var. hirtellum has not previously been
considered a synonym of S. adscendens, but both Bitter
and C.V. Morton annotated the holotype at P as “affine
S. adscendens”and“probably=S. adscendens,” respectively.
The original printed label indicates the specimen was col-
lected in “Province de Rio-Grande” but unidentified hand-
writing above this label states “de Minas Geraes.” Solanum
adscendens has its northern limit at the border of Rio
Grande do Sul and Santa Catarina States, so it is unlikely
that the Gaudichaud 1745 specimen was collected in Minas
Gerais State.
2. SOLANUM DEFLEXUM Greenm., Proc. Amer. Acad. Arts 32:
301. 1897.—TYPE: MEXICO. Oaxaca: Cuicatla
´n, 15 Jul
1895 (fl, fr), L.C. Smith 403 (lectotype, here designated:
GH-GH00295516!; possible isolectotype: MEXU-00540158).
Solanum setosum (Brandegee) Bitter, Repert. Spec. Nov. Regni
Veg. 18: 307. 1922. Bassovia setosa Brandegee, Univ. Cal.
Publ. Bot. 6: 373. 1917.—TYPE: MEXICO. Veracruz:
Zacuapan, Aug 1915 (fl, fr), C.A. Purpus 7509 (holotype:
UC-178623!; isotypes: G-G00016530!, GH-GH00217435!,
MO-825706!, NY-NY00138551!, US-884541!).
Salpichroa wrightii A. Gray, Syn. Fl. of N. Amer., ed. 2(1).
1: 231. 1886.—TYPE: UNITED STATES. Arizona: on
the Sonoita, towards the San Pedro, 17 Sep 1851 (fr),
C. Wright 1592 (erroneously as “1692” in protologue;
lectotype, here designated: GH-GH00217433!; isolecto-
types: G-G00016521!, GH-GH00217434!, K! [2 collections,
both labeled as Wright 1592,mountedononesheet],
MO-3830682!, NY-NY00138909!).
Herb, sometimes slightly woody at base, single- to few-
branched, 1– 3 (4) dm tall. Stems sparsely to densely pubes-
cent with straight, one- or two-celled unbranched hairs,
these denser on new growth. Sympodia 2-foliate, usually
geminate. Leaf blades 1–4.5
+
0.5–2.5 cm, elliptic to elliptic-
ovoid, chartaceous to membranaceous, sparsely to moder-
ately pubescent adaxially and abaxially with 1 or 2 celled
unbranched hairs, these lying flat along blade, denser along
veins; base rounded to obtuse, often decurrent into petiole;
apex acute; petioles 0.5–2 cm, moderately pubescent with
unbranched hairs. Inflorescences with 1– 5 flowers, the
axes sparsely to moderately pubescent with unbranched
hairs; peduncle absent or nearly so; rachis absent; pedicels
5–12 mm in flower, 10–20 mm in fruit. Flowers with the
calyx 3– 9 mm long, the tube 1 – 3 mm, the lobes 2 – 6
+
0.5– 1.5 mm, linear-lanceolate, moderately to densely pubes-
cent. Corolla 0.5– 1 cm in diameter, rotate with abundant
interpetalar tissue, chartaceous to membranaceous, white, the
tube 3– 6 mm long, the lobes very short, 1–2
+
0.5–1 mm,
triangular, acute at apices, glabrous abaxially and adaxially.
Stamens 2– 4 mm long; filaments up to 1 mm long; anthers
1.5–3
+
0.5–1.5 mm, oblong, the base cordate, the apex
emarginate. Ovary glabrous; style 4– 6
+
0.5– 1 mm, equal
to or exserted beyond stamens; stigma to 1 mm wide.
Berries 5– 12 mm in diameter, globose, white to yellow
when immature, maturing semitransparent, drying brown,
glabrous, the mesocarp watery and held under pressure
until dehiscing explosively at maturity. Seeds 5–20 per
fruit, ca. 2.5
+
1.5 mm, significantly notched where con-
nected to placenta, the outer margin of seeds swollen with
large raised ridges running perpendicular to margin and
connected by smaller cell walls running parallel to margin,
the center of seed smoother. Figure 6.
Habitat and Distribution—Weedy in grazed areas, along
roadsides, and disturbed areas in dry forests from southern
Arizona, U. S. A. through Mexico to Guatemala, Honduras,
Nicaragua, and Costa Rica at elevations from 0– 1,550 m
(Fig. 7).
Phenology—Flowering specimens have been collected
in all months with a peak from May through September.
Fruiting specimens have been collected in all months of the
year with a peak from June through October.
Conservation Status—The widespread distribution and
abundant populations of S. deflexum give it an IUCN Red List
Status of Least Concern.
Etymology—The epithet deflexum indicates a structure
bent sharply downward but it is somewhat unclear what
structure this refers to. In the description, Greenman notes
that the species is well characterized by the inflorescence,
calyx, and pubescence. Since only the inflorescence, specifi-
cally the pedicels, could be characterized as deflexed, it is
likely that the epithet refers to this structure.
Additional Specimens Examined—UNITED STATES.Arizona: Fresnal
Canyon, Baboquivari Mts., Pima Co., 2 Sep 1931 (fr), Gilman 80 (WIS);
Baboquivari Mts., 26 Sep 1927 (fr), Harrison 4777 (US); Tumacacori,
Santa Cruz Co., 29 Aug 1931 (fr), Harrison 8147 (US); Baboquivari
Mts., 30 Sep 1934 (fr), Kearney & Peebles 10387 (US); near Patagonia,
18 Aug 1928 (fl, fr), Peebles et al. 5625 (US).
MEXICO.Baja California: Cape Region, Oct 12 1899 (fr), Brandegee
s.n. (F, NY); Corral Piedra, 10 Sep 1893 (fl), Brandegee s.n. (NY, US);
Corral Piedra, 9 Sep 1893 (fl, fr), Brandegee s.n. (GH); Sierra San
Lazano, 11 Sep 1893 (fl, fr), Brandegee s.n. (UC); San Jose
´del Cabo,
17 Sep 1890 (fl, fr), Brandegee 412 (UC). Baja California Sur: Sierra
de la Giganta, Mesa del Potrero de San Javier (northeast of Misio
´n
San Javier), ca. 25!520N, 111!32.50W, 800– 850 m, 20 Sep 1965 (fr),
Carter 4991 (NY, UC, US); Sierra de la Giganta, Can
˜on de Arroyo
Hondo, N side of Cerro Giganta, ca. 26!8.50N, 111!350W, ca. 750 m,
29 Aug 1971 (fl), Carter 5621 (UC); Sierra de la Giganta, Can
˜ada del
Encinal, S side of Valle de Los Encinos (S side of Cerro Giganta),
ca. 26!3.5–40N, 111!340W, ca. 750 m, 1 Oct 1967 (fr), Carter & Moran
5376 (MO, TEX, UC). Chiapas: Mun. Cintalapa, 23 km west of Las
Cruces along road to La Mina Microwave Station, 870 m, 14 Sep 1981
(fr), Breedlove 52762 (NY); Mun. Jiquipilas, a 7.5 kn al ONO de
Francisco Villa, 646 m, 16!2404800N, 93!4300000 W, 11 Jul 2004 (fl, fr),
Martı
´nez & A
´guilar 36935 (MO); between Topana, Oaxaca & Tonala,
200–500 ft, 1–3 Aug 1895 (fl, fr), Nelson 2876a (GH, US). Chihuahua:
near Rio Urique, near Urique, ca. 1700 ft, 31 Jul 1971 (fl, fr), Bye Jr.
1712 (GH). Colima: near km 288, ca. 20 miles ESE of Manzanillo,
ca. 50 m, 23 Jul 1957 (fl, fr), McVaugh 15658 (US); along Rı
´oCihuatla
´n
in gorge near bridge 13 miles north of Santiago, 175 m, 27 Jul 1957
(fl, fr), McVaugh 15796 (BM, G, NY, TEX, US). Guerrero: Dist.
Coyuca, Pungarabato, 22 Aug 1934 (fl, fr), Hinton et al. 6481 (BM,
G, GH, K, K, MO, US); Dist. Mina, Placeres, 350 m, 23 Jul 1936
(fl, fr), Hinton et al. 9143 (GH, K, NY, US); Distr. Montes de Oca,
Vallecitos, 7 Mar 1937 (fl), Hinton et al. 10557 (F, GH, K, NY, UC, US);
along highway, 8 miles S of Mexcala, 23 Jul 1939 (fl, fr), Langman 2145
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 479

(PH); Rio Balsas, 26 Aug 1910 (fl, fr), Orcutt 4210 (F, GH, K, MO,
US); Rio Balsas, 26 Aug 1910 (fl, fr), Orcutt 4389 (F); Flora de la cuenca
del Rı
´o Balsas, Mun. Tepecoacuilco de Trujano, Oapan, between
Oapan/Ahuelica
´n, 17!5805300N, 99!2605000W 1846 ft, 2 Aug 2003 (fl, fr),
Smith & Rojas 348 (NY); Flora de la cuenca del Rı
´o Balsas, Mun.
Tepecoacuilco de Trujano, Oapan, between Oapan/Ahuelica
´n, 17!5903800N,
99!2605700W, 2018 ft, 30 Aug 2003 (fl, fr), Smith & Rojas 561 (NY). Jalisco:
Mun. Huejuquilla, Rancho Los Arroyos del Agua, 15 km al NW de
Huejuquilla, 1550 m, 4 Aug 1990 (fl, fr), Flores 2025 (WIS); near
Guadalajara, 24 Aug 1901 (fl, fr), Rose & Hay 6288 (US). Mexico:
Dist. Temascaltepec, Tenayac, 1450 m, 14 Jul 1933 (fl, fr), Hinton
et al. 4219 (BM, F, G, K, NY, US); Dist. Temascaltepec, Ixtapan, 1000 m,
2Aug1933(fl),Hinton et al. 4470 (BM, F, G, GH, K, K, NY, US).
Michoaca
´n: Dist. Apatzingan, Mun. Apatzingan, 300 m, 19 Aug 1938
(fl, fr), Hinton et al. 12057 (K, NY, US); Mun. Apatzingan, bank of Rio
Apatzingan 2 mi S of Apatzingan, 1200 ft, 5 Aug 1940 (fl), Leavenworth
473 (F, MO, NY); Mun. Uruapan, Tancitaro Region, 2 miles west of
Uruapan, 6000 ft, 2 Aug 1941 (fl, fr), Leavenworth & Hoogstraal 1279 (F);
Mun. Apatzingan, Tancitaro Region, La Majada, 1200 ft, 7 Aug 1941
(fr), Leavenworth & Hoogstraal 1344 (F, GH, MO, NY); near Morelia,
ca. 2000 m, 14 Jul 1941 (fl, fr), Schery 107 (MO); cuenca media del Rio
Balsas, en La Cuesta del Mango, 30 km al N de Huetamo, carr.
AZita
´cuaro, 650 m, 6 Sep 1978 (fr), Soto Nu
´n
˜ez 952 (MO). Morelos:
hillside near Cuernavaca, 5000 ft, 26 Jul 1896 (fr), Pringle 6400 (BR,
E, G, GH, GOET, MEXU, PH, W, WU); near Yautepec, 13– 13 Jul 1905
(fl, fr), Rose et al. 8587 (US). Nayarit: Cerro de la Cruz, east of Tepic,
1000 m, 17 Sep 1926 (fr), Mexia 673 (UC). Oaxaca: Dist. Tehuantepec,
Mun. Santiago Astata, Barra de la Cruz, 2 km W, por la vereda hacia
Zimata
´n, 15!5002200N, 95!590400 W, 105 m, 3 Sep 1998 (fr), Elorsa 700 (NY);
Dist. Tehuantepec, Mun. Santiago Astata, Las Tres Peladas, 3 km al
SE de Barra de la Cruz cerca de la playa, 15!4905100N, 95!5701300 W,
10 m, 15 Jun 2000 (fl, fr), Elsora 3117 (NY); 6 km NE of Juchita
´n,
along the Pan-American highway (route 19), elevation less then
50 m, 9 Jul 1958 (fl, fr), King 446 (TEX, WIS); 11 km NW of the village
of La Ventosa, along the Trans-Isthmian highway (route 185), eleva-
tion 50 m or less, 14 Jul 1958 (fl, fr), King 555 (TEX); 2 km NW of
Juchitan, 29 Jun 1981 (fl, fr), LaSalle et al. 810629–2 (TEX); Mun. Salina
Cruz, Dto. Tehuantepec, Carnero, 10 km al NO de Rincon Bamba,
16!050N, 95!300W, 8 Jul 1988 (fl, fr), Martinez 1461 (F, MO); near
Oaxaca, 5000 ft, 5 Jul 1897 (fl, fr), Pringle 6729 (BH, BR, F, G, GH, K,
M, MO, NY, PH, UC, US, WU); Jomillin Can
˜on, 6 Jul 1897 (fl, fr),
Pringle 7508 (MO); Santa Catarina, 14 Jul 1910 (fl, fr), Rusby 85
(NY, US); Mun. Cuicatla
´n, Dist. Cuicatla
´n, 4 km al E de Cuicatla
´n,
brecha a Concepcio
´npapalo,17
!4103100N, 96!5704400W, 925 m, 19 Jun 1993
(fl), Salinas et al. 7242 (NY); Almoloya, 100–250 m, Jul 1937 (fl, fr),
Williams 9819 (F); 80 miles S of Oaxaca on Rt 190, km mark 670,
22 Jun 1966 (fl), Windler & Snider 994 (MO). Puebla: near Oaxaca,
vicinity of San Luis Tultitlanapa, Puebla, Jul 1908 (fr), Purpus 3563
(F, GH, MO, NY, UC, US). Quere
´taro: Mun. de Arroyo Seco, 1 km al
NNW de Vegas Cuatas, 760 m, 19 Aug 1991 (fl, fr), Carranza 3413
(TEX); Mun. Jalpan, 4.7 mi W of Jalpan, 1260 m, 7 Jul 1985 (fr), Cowan
et al. 5455 (MO, NY, TEX). San Luis Potosı
´:edge of hill and corn field
beside Tamasopo Falls, 12 Sep 1978 (fr), D’Arcy 11896 (MO, TEX).
Sinaloa: vicinity of Culiacan, Culiacan, 18 Oct 1904 (fl, fr), Brandegee
Fig. 6. Solanum deflexum (Bohs 2715, UT). A. Habit. B. Closed flower in the late afternoon; note straight hairs on calyx and pedicel. C. Open flower
in the day; note straight hairs on stem. Scale bars: A = 1 cm. B, C = 5 mm.
480 SYSTEMATIC BOTANY [Volume 38

s.n. (GH, US); vicinity of Culiacan, Culiacan, 10 Sep 1904 (fl, fr),
Brandegee s.n. (UC); Cerros del Fuerte, 18– 24 miles north of Los
Mochis, 200– 1000 ft, 3 –5 Oct 1954 (fr), Gentry 14430 (LL, US); near
Rosario, on the road to Aeaponeta, 27 Jul 1897 (fl, fr), Rose 1852
(F, GH, K, US); E of Villa Union, on Hwy 40, near 106!080W, 23!140N,
200 m, 18 Aug 1988 (fl), Sanders et al. 8033 (NY). Sonora: 0.5 mi SE of
Alamos at junction of Gu
¨irocoba and El Fuerte Rds., 1400 ft, 1 Sep 1973
(fr), Fish 73 (UC); El Naranjo, ca. 23.5 mi NE Alamos, 9 Aug 1980 (fl),
Lehto 24716 (NY); Alamos, 4 miles N of A., 250 – 300 m, 3 Aug 1935 (fl),
Pennell 19496 (PH); Mun. de Soyopa, arroyo Las Tinajas below ruins
of Toledo smelter, near Loma Maderistra, 3.5 km S of To
´nichi, west
side of Rio Yaqui, 220 m, 28!3400300N, 109!3302500 W, 17 Sep 2006 (fl, fr),
Van Devender & Reina 2006–937 (MO); Mun. de Alamos, 1 km E of
Yocogigua on road to Capitahuasa, 26!4702500 N, 109!005500 W, 230 m,
25 Sep 1993 (fr), Van Devender et al. 93– 1114 (MO). Tamaulipas: 3miles
SofVilladeAldama,NofTampico,18Jul1971(fl),D. Spellman et al.
95 (MO). Veracruz: La Granja Vista Hermosa, entrada a la presa de
Temascal, km 25, 8 m, 20 Nov 1966 (fr), Caldero
´n 1169 (BM, MO);
Mun. Apazan, Ban
˜os de Carrizal, 7 km SE of Emiliano Zapata
[=Carrizal], 19!190N, 96!380W, 250 m, 27 Jun 1980 (fr), Hansen & Nee
7447 (F, MO); Pte. Nacional, 3 km al E del Crucero, camino de
terracerı
´a, 8 Jul 1986 (fl, fr), Ortiz 1029 (MO); Ban
˜os del Carrizal,
Aug 1912 (fr), Purpus 6100 (BM, F, GH, MO, NY, UC, US);
Zacuapan, Sep 1907 (fl), Purpus 7860 (MO, NY, UC, US); Zacuapan,
1917 (fl, fr), Purpus 8016 (GH, M, UC); Barranca de Panoaya, Sep 1919
(fr), Purpus 8498 (GH, M, MO, NY, UC, US); Rancho Tlacosinatla,
Aug. 1929 (fl), Purpus 10771 (M, NY, PH, US); Zacuapan, Aug 1929
(fl, fr), Purpus 12055 (NY); Zacuapan, Aug 1929 (fl, fr), Purpus 13000
(A); Mun. Xalapa, Chitares, antes Arteria, cerca de San Nicola
´s, 640 m,
9Jul1971(fl,fr),Ventura 3835 (CORD, F); Mun. Totutla, Encinal,
750 m, 21 Aug 1971 (fl, fr), Ventura 4123 (CORD, F); Mun. de Puente
Nacional, Mata de Can
˜a, 100 m, 20 Aug 1973 (fr), Ventura 8883 (F);
La Concepcio
´n, Mun. de Jilotepec, 900 m, 7 Aug 1978 (fr), Ventura
11750 (F). Zacatecas: 5 miles NE of Mesquitula near the Rio Juchilipa,
3500 ft, 11 Aug 1969 (fl, fr), Taylor &. Taylor 6073 (NY, US). Distrito
Federal: Vicinity of Rancho del Rosario, 10 miles N of Mexico City
near Atzcapotzolco, 7300– 7500 ft, 1 – 15 Jul 1937 (fr), Happ 101 (MO).
Without State: Mexico, (fl), Orcutt 5286 (MO).
GUATEMALA.Chiquimula: bei Chiquimula, 400 m, Jul 1881 (fl, fr),
Lehmann 1671 (BM, G, US); near divide on road from Zacapa
´to
Chiquimula, ca. 660 m, 9 Oct 1940 (fr), Standley 73719 (F); Chiquimula,
transecto La Hondonada, 300–400 m, 14!5100300N89
!3100800W, 8 Jul 2003
(fl), Ve
´liz & Ramirez 13712 (MO). El Progreso: San Agustı
´n, Aldea
El Rancho, 200–300 m, 14!5404200N, 90!0200800W 10 Jul 2003 (fr), Co
´bar &
Garcı
´a520(MO); El Jı
´caro, 200–300 m, 14!5501900N, 89!5202500 W,
9 Jun 2003 (fr), Garcı
´a et al. 715 (MO); Moraza
´n, Aldea Pasasagua,
300– 400 m, 14!5504100 N, 90!0304200 W, 10 Jul 2003 (fr), Ramirez & Veliz
948 (MO). Guatemala: 15 km SE of Granados, dry hillside above
Rio Motagua, 15 Jul 1970 (fl, fr), Harmon & Dwyer 3085 (MO, WIS).
Jutiapa: near Mongoy, 1800 m, 27 Jun 1949 (fr), Williams & Molina
16796 (F). Suchitepequez: EsideofSanAntonio,30Jul1970(fr),
Harmon & Dwyer 3399 (MO). Zacapa
´:Rı
´o Hondo, 170– 200 m,
15!0103200N, 89!3500600 W, 8 Jul 2003 (fl, fr), Co
´bar & Garcia
´385 (MO);
Caban
˜as, 200– 300 m, 14!5505400N, 89!4605500 W, 9 Jul 2003 (fr), Ramı
´rez &
Ve
´liz 813 (MO); Rı
´o Hondo, Transecto Rı
´o Hondo, 210 m, 15!0103200N,
89!3500600W, 8 Jul 2003 (fl, fr), Ve
´liz & Ramı
´rez 13681 (MO).
Fig. 7. Distribution of S. deflexum,S. lignescens, and S. manabiense.
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 481

HONDURAS.Comayagua: vicinity of Comayagua, ca. 600 m, 12–23 Mar
1947 (fl, fr), Standley & Chaco
´n5407(F); al Oeste de Comayagua cerca
de El Taladro, 650 m, 27 Jun 1964 (fl, fr), Standley & Molina 14278 (F, G,
NY, US). Copa
´n: Mun. Nueva Arcadia, 3 mi SW of Los Tangos along
Santa Rosa de Copa
´n-San Pedro Sula highway, 15!0602500N, 88!4105100W,
425 m, 26 Jun 1994 (fl, fr), Davidse et al. 35391 (MO, NY). Francisco
Moraza
´n: Tegucigalpa D.C., Col. Miraflores Sur y alrededores, 900 m,
26 Jun 1978 (fr), Dı
´az 63 (MO); drainage of the Rı
´o Yeguare, ca. 14!N,
87!W, 900 m, 2 Aug 1949 (fr), Molina 2535 (GH); El Zamorano, Dept.
of Agronomy, common in corn field of Monte Redondo drainage of
Yeguare river, 800 m, 11 Dec 1971 (fr), Molina 27185 (F, US); meadows
around Escuela [El Zamorano], Aug 1960 (fr), Pfeifer 1245 (US);
Zamorano, 800 m, 21 Sep 1943 (fr), Rodriguez 940 (F); same loc.,
24 Sep 1943 (fl, fr), Rodrı
´guez 1013 (F); same loc., May 1944 (fl, fr),
Rodrı
´guez 2126 (F); same loc., 800 m, 7 Mar 1945 (fl, fr), Rodrı
´guez
3800 (F, GH, MO, US); vicinity of El Zamorano, 780–900 m, 26 Nov
1946– 9 Jan 1947 (fr), Standley 1654 (F); same loc. and date? (fr),
Standley 1811 (F); same loc. (fr), Standley 3833 (F); same loc., 12 Oct 1948
(fl, fr), Standley 13024 (F); same loc. (fl), Standley 16093 (F); same loc.,
800– 850 m, 16 Jul 1949 (fl, fr), Standley 21306 (F); regio
´nofRı
´odela
Orilla, southeast of El Zamorano, 900–950 m, 5 Aug 1949 (fl, fr),
Standley 22210 (F). Olancho: matorrales al lado de la carretera 6 km a
Juticalpa, 430 m, 18 Nov 1963 (fr), Standley & Molina 13233 (F, NY, US).
Valle: Mun. Nacaome, 1 mi NE of Jı
´caro Gala
´n, along main highway
to Tegucigalpa, 13!3203700 N, 87!2503500 W, 80 m, 18 Jun 1994 (fl, fr),
Davidse et al. 35103 (MO).
ELSALVADOR.Ahuachapa
´n: San Francisco Mene
´ndez, El Corozo,
Mariposario, zona alta “Mariposario,” 13!490N, 89!590W, 250 m, 25 May
2000 (fl), Rosales 837 (BM, MO, NY). Caban
˜as: Cinquera, zona protegida,
quebrada calle antı
´gua, 500 m, 13!510N, 88!570W, 13 Aug 2002 (fr), Carballo
371 (MO); Cinquera, zona protegida, entrada cementerio, 400 m, 13!530N,
88!570W, 29 Aug 2002 (fr), Carballo & Carrillo 433 (MO). La Paz:
San Luis Talpa, Centro Experimental Universidad El Salvador, 29 Aug
1995 (fl, fr), Montalvo 6380 (MO, NY). Moraza
´n: adjacent to ditch lead-
ing to reservoir, Montecristo (ca. 15 km northeast of San Miguel),
13!360N, 88!040W, ca. 140 m, 9 Dec 1941 (fl, fr), Tucker 502 (BH, F,
G, GH, IAC, IAN, K, LL, NY, UC, US). San Salvador: vicinity of
San Salvador, 650– 850 m, 20 Dec 1921–4 Jan 1922 (fl, fr), Standley 19647
(G, GH, NY, US); vicinity of San Salvador, 650– 850 m, 30 Mar-24
Apr 1922 (fl, fr), Standley 22678 (US). San Vicente: vicinity of San Vicente,
400–500 m, 7– 14 Feb 1947 (fl, fr), Standley & Padilla 3808 (F). Sonsonate:
vicinity of Santa Emilia, ca. 135 m, 22– 25 Mar 1922 (fr), Standley
22063 (GH, S, US); vicinity of Santa Emilia, ca. 135 m, 22–25 Mar 1922
(fl, fr), Standley 22258 (GH, MO, NY, US).
NICARAGUA.Boaco: Km 57.5 on Hwy 7 (Carretera al Rama), ca. 5.0 km
NE of Managua line, ca. 12!220N, 85!520W, ca. 130 m, 16 Jul 1978 (fr),
Stevens 9363 (MO). Chinandega: E base of Coseguina [Cosegu
´ina]
Volcano, 6 Jul 1932 (fl, fr), Howell 10253 (F, US); Ameya, near sea level,
19– 21 Jun 1923 (fl, fr), Maxon et al. 7138a (US); Ameya, near sea
level, 19– 21 Jun 1923 (fl, fr), Maxon et al. 7191 (US); vicinity of
Chichigalpa, ca. 90 m, 12–18 Jul 1947 (fr), Standley 11148 (F).
Chontales: 0.9 km W of bridge at Lo
´vago, km 166, 95 m, ca 12!000N,
85!100W, 7 Jun 1981 (fl), Stevens & Henrich 20473 (MO). Esteli: near
El Dorado, 8 km N of Estelı
´,alongtheRı
´oLaSirena,13
!090N,
86!230W, 800 m, 30 May 1985 (fl, fr), Davidse et al. 30657 (MO, NY);
Mun. San Juan de Limay. Had. La Grecia “Cerro Quiniento,”
13!110N, 86!350W, 518 m, 3 Sep 1980 (fr), Moreno 2157 (MO); ca. 7 km
from Hwy 1 (at ca. km 193) on road to Pueblo Nuevo, from Quebrada
Jamaili to near summit of Cerro El Pedrero, ca. 13!240N, 86!270W,
600– 700 m, 3 Jul 1977 (fr), Stevens 2610 (BM, LL); Km 163 on Hwy
1, ca. 11.2 km N of entrance to Estelı
´,ca.13
!130N, 86!230W, ca. 920 m,
19 May 1981 (fl, fr), Stevens & Henrich 20185 (MO). Leo
´n: Mun. de
La Paz Centro, Miramar, nivel del mar, 12!100N, 86!450W, 27 Sep 1997
(fl, fr), Rueda 7420 (NY); Las Pen
˜itas, al SE de Poneloya, 12!210N,
87!010W, 10 m, 22 Jun 1982 (fl, fr), Sandino 3134 (MO); slope and
ridge immediately W of Quebrada Las Ruedas, N of road, NW of
El Transito, ca. 12!050N, 86!430W, ca. 15– 30 m, 16 Oct 1977 (fr),
Stevens 4708 (MO). Managua: Managua, Aug 1923 (fl, fr), Chaves 2 (US).
Matagalpa: 9 km south of Sebaco, collection from wet depression
along road (Nic. 1), 500 m, 7 Feb 1971 (fl, fr), Harmon & Fuentes 6016
(MO, WIS); just W of Puente de Rio Viejo, ca. 8 km SE of San Isidro,
12!530N, 86!090W, 460 m, 24 Jun 1982 (fl, fr), Kral 69080 (MO); Darı
´o,
“Cuajiniquilapa,” a 4 km de la Carretera Panamericana siguiendo
la carretera a Terrabona, 12!440N, 86!040W, 440– 480 m, 18 Jun 1981
(fl, fr), Moreno 9222 (MO); Darı
´o, “Las Joyas,” en la quebrada a 7 km
de la Carretera Panamericana, sobre el camino a Terrabona, 12!450N,
86!010W, 480– 500 m, 18 Jun 1981 (fr), Moreno 9280 (MO); Darı
´o,
“El Caracol,” a 14 km de la Carretera Panamericana, 12!450N,
85!590W, 560–580 m, 18 Jun 1981 (fl, fr), Moreno 9294 (MO); Entrada
Paso de Carreta, quebrada, 12!520N, 86!080W, 460– 480 m, 24 Jun 1982
(fl, fr), Moreno 16703 (MO); SW slopes of Cerro El Pilo
´nandadjacent
Laguna Tecomapa, 420–540 m, 12!370N, 86!020W, 20 Jul 1978 (fl, fr),
Stevens 9376 (MO). Nueva Segovia: above Rio Dipilto 6 km N of
Ocotal, 700 m, 15 Jun 1977 (fl, fr), Neill 2174 (MO). Rivas: Las Salinas,
sea level, 25 Aug 1977 (fl), Neill 2460 (MO).
COSTA RICA.Alajuela: Canto
´n de Orotina, Valle del Ta
´rcoles, de la
carretera Orotina-Caldera, ca. 1 km S rumbo a playa Bajamar, Guacalillo,
9!530000N, 84!3801000 W, 100 m, 21 July 1995 (fl, fr), Hammel & Grayum
19933 (INB); La Balsa de Rio Grande, 2 June 1911 (fl), Pittier 3646 (US).
Guanacaste: Canto
´ndeBagaces,P.N.PaloVerde,ValledeTempisque,
A.C.T. Sector Palo Verde, sitio la Carreta, 10!220000N, 85!170000W, 10–100 m,
30 July 1996 (fl, fr), Chavarrı
´a1503(INB, MO); Canto
´ndeBagaces,P.N.
Palo Verde, Cuenca del Tempisque, sector La Carreta, 14 m, 10!2104000N,
85!1902000W, 9 Oct 1997 (fr), Chavarrı
´a 1766 (MO); Comelco E, W of
Bagaces, 2 Aug 1971 (fl, fr), Heithaus 274 (MO); Rio Higuero
´n, near
Taboga, 0–100 m, 10!200N, 85!120W, 29– 30 Jun 1977 (fl, fr), Liesner et al.
2798 (MO); Finca La Pacı
´fica Can
˜as, Secondary Plot, 100 m, 13 Jun 1972
(fl, fr), Opler 860 (F, UC). Puntarenas: vicinity of Cascajal (25 km ESE of
Puntarenas), along road from Cascajal to Pigres, 30– 100 m, 6 Jul 1949
(fr), Holm & Iltis 295 (A, G, MO); entre Mata de Limo
´n y Cerro de las
Mesas, 50 m, 1 Jun 1963 (fl, fr), Jimenez 721 (F); Canto
´n de Montes
de Oro, cuenca del Aranjuez, San Isidro, Las Lomas, 2.5 km N de
Cuatro Cruces, 10 m, 10!0103300N, 84!4401200 W, 13 Aug 1998 (fr),
Rodrı
´guez et al. 3909 (F, MO); near junction of Hwy. 1 and road to
Miramar, 50 m, 28 Aug 1966 (fl, fr), Weston et al. 2060 (UC).
Notes—Solanum deflexum and S. lignescens are the only
two species in the section that occur in Central America.
The straight, multicelled, unbranched hairs of S. deflexum
are unlike the hairs of S. hoffmanseggii, S. olympicum, and
S. turneroides that bend up or down the stem, and, unlike
S. lignescens, they are not stellate. Solanum lignescens is sym-
patric with S. deflexum, but the latter is a diminutive non-
woody plant, reaching 30 to rarely 40 cm in height. Solanum
deflexum is frequently many-branched at the base, creating
a “flat” appearance that is distinct from the upright appear-
ance of other members of the clade. Solanum deflexum is also
unique in that the first inflorescence often appears with the
first pair of leaves after the cotyledons, giving the appear-
ance of flowering while still a seedling (Nee 1989). Unlike
other members of the clade, S. deflexum is rarely rhizomatous,
although it does appear in patches, likely due to the explo-
sive dehiscence of the fruits.
In the description of S. deflexum, Greenman lists three
collections, Nelson 2876a,Pringle 6400,andSmith 403,but
none was designated as the type. Smith 403 was chosen
as the lectotype from among the syntypes because of the
quality of the specimen, with both flowering and fruiting
material. A collection bearing a label for Smith 403 exists
at MEXU but this specimen also bears a label for Pringle
6400, a different syntype collection. The material is likely
that of Smith 403 as it is both flowering and fruiting while
all of the many Pringle 6400 collections at MEXU and
other herbaria only have flower buds or fruit. Nelson 2876a
is comprised of plants that are unusually large and are not
reflective of the species as a whole.
The type of Salpichroa wrightii exemplifies the problems
botanists have encountered with collections by Charles Wright
made along the boundary with Mexico. In the protologue,
Gray gives the collection locality as “Arizona on the Sonoita”
and the type as Wright 1692.ConsultationofShaw’s(1987)
book about Wright’s expeditions and W. T. Kittredge’s
label on the lectotype sheet, presumably with information
taken from Wright’s field notes, indicates that his field
number for this collection is 549 and it was made on Septem-
ber 17, 1851. The specimens, all apparently belonging to the
482 SYSTEMATIC BOTANY [Volume 38

same gathering, have been distributed with the numbers
1592 and 1692. In fact, on the lectotype sheet at GH the
handwritten “1592” appears to have been changed to
“1692”. Because the distribution number 1692 does not
exist for Wright’s Solanaceae collections according to Shaw
(1987), it is apparent that all the collections should bear
the distribution number 1592 and the protologue contains
a typographical error.
3. SOLANUM EVOLVULOIDES Giacomin & Stehmann, PhytoKeys
7: 1– 9. 2011.—TYPE: BRAZIL. Bahia: Mun. Jequie
´, Dist.
Cachoeirinhas, caatinga arbustiva em topo de morro, com
lajeados granı
´ticos, 13!54014.400 S, 40!01046.800 W, 299 m,
10 Jul 2009 (fr), L.L. Giacomin 974 (holotype, BHCB!;
isotypes to be distributed to BM, MBM, NY, RB).
Herb, slightly woody to woody at base, few- to many-
branched, 2–4 dm tall. Stems moderately to densely pubes-
cent with multicelled unbranched erect glandular hairs,
these mixed with less frequent slightly longer 1–3-celled
unbranched straight hairs. Sympodia 2-foliate, solitary or
more commonly geminate, unequal with the smaller leaves
up to half of the size of the larger ones. Leaf blades 1–4
+
1–3 cm, elliptic-ovoid to cordiform, chartaceous to mem-
branaceous, sparsely to moderately pubescent adaxially
and abaxially with 1 or 2-celled unbranched hairs, these
denser along veins; base attenuate to cordate, often decur-
rent into petiole; apex acute; petioles 0.5–2.2 cm, moder-
ately pubescent with hairs like those of the stem but with
fewer eglandular hairs. Inflorescences with 1–4 flowers,
the axes moderately pubescent with hairs like those of
the stem; peduncles absent; rachis nearly absent; pedicels
6–10 mm in flower, 7–14 mm in fruit. Flowers with the calyx
2–7 mm long, the tube 1–2 mm, the lobes 2–6
+
1–2.6 mm,
ovate-elliptic, the apex acuminate, moderately pubescent
abaxially with almost exclusively glandular unbranched
multicellular erect hairs; fruiting calyx accrescent, the lobes
up to 8 mm long, equal to or exceeding the berry at matu-
rity. Corolla 1–3 cm in diameter, rotate with abundant
interpetalar tissue, membranaceous, white, the tube 4–6 mm,
the lobes 2–4
+
1– 3 mm, triangular, acute at apices, with
a few eglandular hairs abaxially, mainly on the central part
of each lobe, glabrous adaxially. Stamens 4–9.5 mm long;
upper, shorter filaments 1–2 mm, the lowermost, longer fila-
ment 3–7 mm; anthers 4–6
+
1.3–2 mm, oblong, the base
cordate, with a small bulge dorsally, the apex emarginate,
the pores directed introrsely and subapically, not opening
into longitudinal slits. Ovary glabrous; style 7–9
+
0.5–1 mm,
longer than the smaller stamens, closely appressed to the
larger stamen, curved near apex: stigma to 1 mm wide.
Berries 8–15 mm in diameter, globose, greenish-white when
immature, maturing translucent, drying brown to blackish,
glabrous, the mesocarp watery and held under pressure
until dehiscing explosively at maturity, normally between
two calyx lobes. Seeds 10– 25 per fruit, 2.5–3.6
+
1.8–2.9 mm,
reniform, with a small hollow where connected to the
placenta, the margin flattened, the seed surface with
raised projections and grooves parallel to margin, giving
a netlike impression. Figure 8.
Habitat and Distribution—An herb of the transition
zone between deciduous forests and xeric formations of
shrubby caatinga. It is known only from a restricted area
in the southeastern part of Bahia State, Brazil at 0–275 m
in elevation (Fig. 5).
Phenology—Flowering and fruiting materials were col-
lected between February and August, with a flowering peak
from February to May; fruiting specimens were collected
from June to August.
Conservation Status—The IUCN Red List Status of
S. evolvuloides is Endangered based on an area of occu-
pancy <5000 km
2
,<5 collection localities and declines in
extent of occurrence, area of occupancy, quality of habitat,
and number of locations. The species is known from only
two localities where the landscape has been strongly modi-
fied in the last decades due to the expansion of urban
centers and extensive farming. The region has been the
focus of several surveys undertaken by the Centro de
Pesquisas do Cacau in association with the New York
Botanical Garden; despite this, just a few collections of
this species have been made. Although one collection was
made in a disturbed area (Jardim 1843), the most recent
collection is from a well-preserved forest fragment, and the
species was not found in surrounding areas. There are no
collections from within conservation units.
Etymology—The epithet is derived from the similar leaf
shape and habit shared with members of the genus Evolvulus
L. of the Convolvulaceae.
Additional Specimens Examined—BRAZIL.Bahia: Km 7 da estrada
Jequie
´/Ipiau
´, caatinga, 10 Feb 1983 (fl), Carvalho 1591 (CEPEC); Mun.
Itacare
´, Fazenda Monte Alegre, ca. 1 km a leste na rodovia para
Itacare
´,margemdoRiodeContas,10Aug1998(fl,fr),Jardim 1843
(CEPEC); Mun. Jequie
´, Rodovia Ipiau
´/Jequie
´, 12 May 1969 (fl, fr),
Jesus 367 (CEPEC); Mun. de Manoel Vitorino, Rod. Man. Vitorino/
Caatingal, Km 4, regia
˜odecaatinga,16February1979(fl,fr),Mattos
Silva s.n. (CEPEC).
Notes—The species is closely related to S. turneroides and
shares with it notable heteranthery, with the filament of one
stamen much longer than the other four. Like S. turneroides,
the flowers are only open in the morning and close by
midday. Solanum evolvuloides can be easily distinguished
from all other species in sect. Gonatotrichum by the glan-
dular, multicellular hairs on the stems, inflorescences,
and flowers that are not found in any other species of
the section.
Jardim 1843 from the banks of the Rio de Contas near the
city of Itacare
´might be an occasional case of water dispersal
by the Contas River, which originates in a xeric environment
near the center of Bahia in the caatinga biome.
4. SOLANUM HOFFMANSEGGII Sendtn., Mart. Fl. Bras. 10: 112.
1846.—TYPE: BRAZIL. “In provincia Paraensi a
Siber lectum comunicavit Com. de Hoffmansegg,” (fl),
F.W. Sieber [as Siber]s.n. (lectotype, here designated:
BR-BR000000699204!; isolectotype (but see note in com-
mentary): M-M0090348! [fragment in packet only]).
Small shrub from a thick woody taproot with the base
reaching 7 mm in diameter, not appearing rhizomatous,
much-branched from base, to 5–8 dm tall. Young stems
moderately pubescent with two-celled unbranched hairs,
these geniculate between first and second cells and point-
ing apically; older stems glabrescent. Sympodia appearing
unifoliate but the structure is difficult to evaluate. Leaf
blades 1.5 – 2.5
+
0.5– 1 cm, linear to elliptic, chartaceous
to membranaceous, nearly glabrous to moderately pubes-
cent adaxially and abaxially with 1– 2-celled unbranched
hairs, these lying flat along blade; base acute, often decur-
rent into petiole; apex acute; petioles absent to 4 mm,
moderately pubescent. Inflorescences with 1–3 flowers, the
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 483

axes sparsely to moderately pubescent with unbranched
hairs; peduncle absent or nearly so; rachis absent; pedicels
5–10 mm in flower, 10–20 mm in fruit. Flowers with the
calyx 2 – 4 mm long, the tube 1–2 mm, the lobes 2–3
+
0.5–1 mm, linear-lanceolate, moderately pubescent; fruiting
calyx 5– 8 mm in length. Corolla 8 – 12 mm in diameter,
rotate with abundant interpetalar tissue, membranaceous,
the tube 1.5–2.5 mm, the lobes 1.5 – 2.5
+
0.8–1.5 mm, triangu-
lar, acute at apices, moderately pubescent abaxially with
hairs like those of the leaves, glabrous adaxially. Stamens
1.5– 2 mm; filaments ca. 0.5 mm; anthers 1.5–2
+
0.5–1 mm,
oblong, the base cordate. Ovary glabrous; style 2.5–3
+
0.2–0.4 mm, exserted beyond stamens; stigma to 0.5 mm
wide. Berries 5– 12 mm in diameter, globose, white to
brown when dried, glabrous, appearing to be explosively
dehiscent due to the presence of small tears and the wrin-
kled, deflated appearance of the dried fruits. Seeds 5–15 per
fruit, ca. 2.5
+
1.5 mm, with a small notch where connected
to placenta, the margin not swollen, the surface with fine
raised ridges radiating from center to edges and shallow
ridges running parallel to margin. Figure 9.
Habitat and Distribution—Known from several localities
in Para
´State, Brazil and one collection from Tocantins State,
Brazil at elevations of 50–200 m (Fig. 5).
Phenology—Flowering materials were collected in June
and October to December; fruiting specimens were collected
from June to August and October through January.
Conservation Status—According to IUCN guidelines, the
status of S. hoffmanseggii is deemed of Least Concern due
to a sufficiently large number of collections over a rela-
tively large area of occupancy. Many areas of Amazonia
are under-collected, leading to difficulty in understanding
the species’ distribution (Milliken et al. 2011). Given this,
it is highly likely that S. hoffmanseggii is more widespread
and abundant than our data indicate.
Etymology—The epithet honors Johann Centurius
Hoffmann Graf von Hoffmannsegg, a German botanist,
entomologist, and ornithologist, who sent the sheet col-
lected by F.W. Sieber (whom he employed to make
entomological collections in Brazil and whose name he
misspelled as Siber) to Sendtner. The specific epithet
was originally published as hoffmanseggii,andwehave
Fig. 8. Photos of type collection of S. evolvuloides (Giacomin 974, BHCB). A. Habit. B. Flower; note glandular hairs on calyx. C. Flower; note the
elongated lowermost stamen. D. Flower. E. Fruit; note the thin, nearly transparent skin. Scale bars: A = 2 cm. B = 5 mm. C, D, E = 1 cm.
484 SYSTEMATIC BOTANY [Volume 38

Fig. 9. Scan of lectotype of S. hoffmanseggii (Photo credit L. A. Mentz).
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 485

retained the original spelling (as stipulated in Art. 60
of the ICBN; McNeill et al. 2006) rather than correcting
it to hoffmannseggii, as spelling of surnames was quite
fluid in the 19th century (he was also referred to as Graf
von Haffmannsegg).
Additional Specimens Examined—BRAZIL.Para
´:Mun. Belterra, Fordla
ˆndia,
Praia Tabocal, 6 Jan 1948 (fl, fr), Black 48 –2316 (IAN, SP); Rio Tocantins,
nella foresta, Capuera roca presso Itacayuna, 1 Jul 1899 (fr), Buscalioni
3656 (MG, NY); Mun. Altamira, Ilha do Inferno Verde, 28 Nov 1986
(fl, fr), Dias 627 (MG); Mun. Maraba
´,RioTocantins,IlhadaPraia,
26 Jun 1949 (fl, fr), Fro
´es 24667 (IAN); Mun. Itaituba, Sa
˜oLuisdo
Tapajo
´s, margens do Rio Tapajo
´s, 23 Nov 1999 (fl, fr), Lisboa 6782
(MG); Mun. Conceic¸a
˜o do Araguaia, Rio Araguaia, Praia de Santana,
23 Sep 2000 (fl), Lobato 2656 (MG); [Mun. Altamira] Rio Xingu, trecho
compreendido entre o Rio Iriri e a cachoeira da Baleia, calha do Rio
Xingu, 1 Oct, 2007 (fl, fr), Lobato 3282 (MG); Mun. Tucuruı
´,BR-422,
km 45, Breu Branco, margem do Rio Tocantins, 5 Nov 1983 (fl), Ramos
1010 (INPA); Mun. Altamira, Rio Iriri, reserva indı
´gena dos Araras,
12 Jan 1985 (fl, fr), Rosa
´rio 713 (MG); margem esquerda do Rio Tocantins,
montante da Usina Hidrele
´trica de Tucuruı
´,meiahoradebarco,acima
dos canteiros da obra, 7 Dec 1979 (fl, fr), Silva 112 (INPA, MG); Mun.
Altamira, Rio Xingu, Ilha a margem direita subindo orio, em frente
so acampamento de CNEC, cachoeira do Espelho, 6 Oct 1986 (fl, fr),
da Souza et al. 223 (MG). Tocantins: Ilha do Bananal, foz do Rio Javae
´s,
cerra do sujeito a inundaca
˜o perio
´dica, 20 Aug 1978 (fr), da Silva 4868
(MG, MO, NY).
Notes—Solanum hoffmanseggii, although rarely collected,
is morphologically and geographically distinct from the other
species of sect. Gonatotrichum. This species is more robust
than any of the other species except S. lignescens, which
can be easily distinguished by its stellate hairs. Anoma-
lously large plants of S. turneroides may reach nearly simi-
lar sizes; however, S. turneroides has much larger fruits
and heterantherous flowers. The most similar species is
S. olympicum, which occurs in southernmost Brazil, Bolivia,
Paraguay and Argentina. The two species share distinct
geniculate hairs as well as seeds with raised ridges radiat-
ing from the center to the margins. Unlike S. hoffmanseggii,
S. olympicum has much broader, larger leaves and is a
more diminutive plant. Solanum hoffmanseggii is the only
member of sect. Gonatotrichum known from northern Brazil.
The Souza et al. 223 collection is unusual because the leaves
are more ovate than other specimens of S. hoffmanseggii but
other characteristics and its geographic location are consis-
tent with this species. The nearest members of the section
geographically are S. evolvuloides and S. olympicum, which
are found in the drier northeastern state of Bahia and in
deciduous forest of northern Goia
´s respectively.
Solanum hoffmanseggii has been problematic because it is
known from relatively few collections and, until recently,
the only known type was a mixed collection from M. This
sheet contains a sterile leafy twig bearing branched hairs
and a small handwritten label with the number “787” and
a packet of the same material on the lower right hand side
of the sheet; this may belong to a different species of
Solanum. Another packet contains fragments including
leaves, buds, and flowers of a different plant and is labeled
“echtes S. hoffmanseggii Sendtn.” by Bitter. However, the
material in this packet does not seem to match that of the
type of S. hoffmanseggii at BR because the leaves are larger,
ovate, and more densely pubescent. This is likely material
from a collection of S. olympicum, a species found much
further south in Bolivia, Paraguay, and Argentina, that Bitter
thought was S. hoffmanseggii. The material on the M sheet is
in our view different from that of the BR collection and
should be treated with some skepticism. The specimen at
BR was chosen as the lectotype because it is far superior
in quality and is unequivocally original material. Because
of the confusion with the type material of S. hoffmanseggii
prior to the discovery of the excellent BR collection, Stern
and Bohs (2009) erroneously referred to what is now known
as S. olympicum as S. hoffmanseggii. It is now clear, however,
that the widespread species of Bolivia, Paraguay, Argentina
and the southernmost parts of Brazil is S. olympicum, while
this unique species of Para
´, Brazil is S. hoffmanseggii.
5. SOLANUM LIGNESCENS Fernald, Proc. Amer. Acad. Arts 33: 91.
1897.—TYPE: MEXICO. Guerrero: Acapulco, Nov 1894
(fl, fr), E. Palmer 216 (lectotype, here designated,
US-259574!; isolectotypes: F-66593!, GH-GH00077503!,
MO-3378770!, K!, NY-NY00139001!).
Solanum roei Ugent & Iltis, Phytologia 40: 379. 1978.—TYPE:
MEXICO. Chiapas: 6 km NW of Las Rosas, in region
of tropical deciduous vegetation on NE slope of Valley
of Chiapas, ca. 900 m, 8 Aug 1965 (fl, fr), K. Roe et al.
1045 (holotype: WIS).
Herb or subshrub from a lignescent base, 0.5–1 m tall.
Young stems densely pubescent with pale yellow-grey,
porrect-stellate hairs, the stalks nearly absent to 0.5 mm,
uniseriate, 5–6 rays, 0.2–0.5 mm in length, unicellular, mid-
points lacking to 0.2 mm long; older stems glabrescent with
yellow-gray bark. Sympodia 3–plurifoliate. Leaf blades 2 –7
+
1.5–4 cm, ovate, chartaceous to membranaceous, sparsely
and uniformly pubescent adaxially with stalked stellate tri-
chomes 0.5–1 mm, the rays often absent and the trichomes
appearing simple, the base of each trichome somewhat
swollen, densely pubescent abaxially with a mixture of
stalked and sessile stellate trichomes 1– 1.5 mm long, with
2– 5 rays and midpoints small or lacking, the trichomes
yellowish and almost obscuring the leaf undersides; base
truncate to broadly cuneate; apex acute to more or less
rounded; petioles 0.2–1 cm, densely stellate-pubescent like
the leaf undersides. Inflorescences opposite the leaves or
extra-axillary, with (1–) 3–6 (11) flowers, the axes densely
stellate pubescent with a mixture of stalked and sessile
trichomes ca. 0.5 mm, like those of the stems; peduncles
0.3–0.8 cm; rachis nearly absent; pedicels 7 – 12 mm in
flower, 12–20 mm in fruit, expanding at the apex, spaced
1–2 mm apart. Flowers with the calyx 3 – 6 mm long, the
tube 1–1.5 mm, the lobes 2– 4
+
1–2 mm, broadly triangu-
lar, reflexed at anthesis, sparsely stellate-pubescent
abaxially, glabrous adaxially with a few scattered, usually
simple, trichomes along the midvein. Corolla 1.4– 1.8 cm in
diameter, stellate, membranaceous, white, lobed 3
/
4of the
way to the base, the lobes 7– 9.5
+
3–4 mm, planar at anthe-
sis, sparsely stellate-pubescent abaxially, glabrous adaxially,
the trichomes denser at the tips and along the lobe midveins.
Stamens 3.5– 5 mm; filaments to 0.5 mm; anthers 3.5 –5
+
1.5–2 mm, oblong, the base sagittate, the apex emarginate.
Ovary glabrous; style 7–8
+
0.5– 1 mm, exserted beyond sta-
mens; stigma 1–1.5 mm wide. Berries 10–12 mm in diameter,
globose, green when immature, maturing white or purplish
black, semitransparent, drying brown, glabrous, the pericarp
thin, the mesocarp watery and held under pressure until
dehiscing explosively at maturity; calyx lobes not expanding
in fruit. Seeds 15– 20 per fruit, ca. 2
+
1mm,thesurfaces
minutely pitted. Figure 10.
Habitat and Distribution—An herb or subshrub of dry,
deciduous forests and thickets in mountainous regions from
486 SYSTEMATIC BOTANY [Volume 38

Fig. 10. Scan of isolectotype of S. lignescens.
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 487

the Sierra Madre Occidental in the Mexican state of Guerrero
to Honduras and Nicaragua, from 1,000–1,500 m in elevation
(Fig. 7).
Phenology—Flowering collections have been made in
February, March, and June-October. Fruiting collections
have been made in June – October.
Conservation Status—The widespread distribution and
abundant populations of S. lignescens give it an IUCN Red
List Status of Least Concern.
Etymology—The epithet lignescens refers to the lignescent,
woody habit of the species.
Additional Specimens Examined—MEXICO.Chiapas: Mun. Tzimol,
ca. 7 km SE of Comita
´n, ca. 3 km SE of Tzimol, 1,350 m, 7 Jul 1990
(fl, fr), Hampshire et al. 1151 (BM); Mun. La Trinitaria, 4 km E of
La Trinitaria along Mex. 190, 16!080N, 92!020W, 1480 m, 8 Jul 1990 (fl),
Hampshire et al. 1154 (BM); same loc., 8 Jul 1990 (fl, fr), Hampshire et al.
1184 (BM); Las Rosas, Rancho Santa Isabel, 7 km al N de Villa Las
Rosas, 16!2500000 N, 92!2400000 W, 1642 m, 9 Aug 1998 (fl), Martı
´nez
Salas et al. 31192 (NY); Mun. Tzimol, 5 km al SO de Tzimol, 1150 m,
14 Sep 1988 (fl), Reyes Garcia & Uriquijo 794 (MEXU). Guerrero:
Costa Verde, Acapulco, 16 Feb 1941 (fl), Langman 3337 (MEXU). Jalisco:
La Huerta, arroyo Tapeixtes (La Mina), km 55 de la carretera Puerto
Vallarta-Barra de Navidad, a 4 km al SE de la Estacio
´n de Biologı
´a,
19!3000000N, 105!0300000W, 50 m, 25 Jun 1985 (fl), Ayala & Lott 9 (NY);
La Huerta, arroyo Maderas, antiguo camino a Nacastillo, 17.5 km de la
carretera Puerto Vallarta-Barra de Navidad, 22 Aug 1985 (fl, fr),
Ayala 136 (NY); La Huerta, Estacio
´nBiologı
´a “Chamela”, estacio
´nde
Investigacio
´n, Experimentacio
´n y Difusio
´n Chamela, UNAM, 26 Jul 1982
(fl), Magallanes 3650 (NY); same loc., 500 m, 3 Jul 1984 (fl, fr), Magallanes
4230 (NY). Oaxaca: Mun. Pochulta, vicinity of Concordia, San Rafael,
1000 m, 2 Mar 1937 (fl), Makrinius 530 (US).
HONDURAS.Francisco Moraza
´n: Mun. Tegucigalpa, Rı
´o Las Canoas,
ca. 5 km al E de Tegucigalpa, 14!020N, 87!100W, 1020 m, 27 Sep 1996
(fl), Linares 3536 (MEXU); Ta
´mara Valley, between Amarateca and
Ta
´mara, 1,000 m, 10 Oct 1969 (fl, fr), Molina 24551 (BM, NY, S).
NICARAGUA.Estelı
´:Mesas Moropotente, 12.5 km SW of Laguna
Miraflor, along road to Estelı
´,13
!110N, 86!170W, ca. 1310 m, 23 Jun 1982
(fl, fr), Stevens 21623 (BM, NY).
Notes—The leaf upper surfaces of S. lignescens usually
have a mixture of simple and stellate trichomes, and a given
individual may have a mixture of hair types or have only
simple trichomes adaxially. Due to the presence of inter-
mediate hair types between simple and stellate, it is likely
that the simple trichomes represent cases where the rays
have been lost. Stellate trichomes always have a uniseriate,
often unicellular stalk. The presence of these hairs and the
shrubby habit make S. lignescens easily recognizable from
the simple-haired, herbaceous species that comprise the rest
of sect. Gonatotrichum.
The type of S. lignescens was not designated in the
protologue. From the handwriting on the labels of the
duplicates of Palmer 216, it appears that Fernald saw and
used the specimens at GH, MO, NY, and US for the species
description (the handwriting on the material at F appears
to be that of someone else). The material of each of these
collections is excellent; however, most collections just have
buds while the US specimen has open flowers, making this
our choice for the lectotype.
6. SOLANUM MANABIENSE S.Stern, J. Bot. Res. Inst. Texas 3(2):
504. 2009.—TYPE: ECUADOR. Manabı
´: Pacoche Reserve,
road from Manta to San Lorenzo, "2kmWof
El Aromo, 01!04009.500S, 80!5203200W, 350 m, 8 Feb 2009
(fl, fr), S.Stern & E.J. Tepe 377 (holotype: QCNE!;
isotypes: BM!, F-2295591!, QCA!, MO-6262688!, NY!, UT!).
Rhizomatous herb, sometimes slightly woody at the base,
1.5–4 dm tall. Stems sparsely to densely pubescent with two-
celled, unbranched, straight hairs. Sympodial units bifoliate,
geminate. Leaf blades 1 –6
+
0.5–3 cm, elliptic to elliptic-
ovoid, chartaceous to membranaceous, nearly glabrous to
sparsely pubescent adaxially and abaxially with one or
two-celled, unbranched, straight hairs, these lying flat
along the blade, denser along veins; base rounded to
obtuse, often decurrent into petiole; apex acute to obtuse;
petioles 0.5– 1 cm, moderately pubescent with unbranched
straight hairs. Inflorescence with 1– 5 flowers, the axes
sparsely to moderately pubescent with unbranched hairs;
peduncle absent; rachis absent to ca. 1 mm; pedicels 5– 15 mm
in flower, 10– 20 mm in fruit. Flowers with the calyx
3–10 mm long, the tube 1–3 mm, the lobes 2–7
+
0.5–1.5 mm,
linear-lanceolate, moderately to densely pubescent. Corolla
0.4– 1 cm in diameter, rotate with abundant interpetalar
tissue, chartaceous to membranaceous, white, the tube
2–3 mm long, the lobes very short, 1– 2
+
0.5– 1 mm, trian-
gular, acute at apices, sparsely to moderately pubescent
abaxially and on margins with 2 or 3-celled unbranched,
straight hairs, glabrous adaxially. Stamens 2–4 mm long;
filaments up to 1 mm long; anthers 1.5 – 3
+
0.5– 1.5 mm,
oblong, the base cordate, the apex emarginate. Ovary gla-
brous; style 4–6
+
0.5– 1 mm, equal to or exserted beyond
stamens; stigma to 1 mm wide. Berries 5 – 12 mm in diam-
eter, globose, white to yellow when immature, maturing
semitransparent, drying brown, glabrous, the mesocarp
watery and held under pressure until dehiscing explo-
sively at maturity. Seeds 10–35 per fruit, ca. 2.5
+
1.5 mm,
lacking an obvious notch where connected to placenta, the
margin not swollen, the surface with fine raised ridges
radiating from the center to the edges and shallow ridges
running parallel to margin. Figure 11.
Habitat and Distribution—A species known only from
the central coast of Ecuador in Provinces Manabı
´and
Guayas from sea level to 400 m in elevation (Fig. 7).
Phenology—Flowering and fruiting specimens have been
collected in February and July.
Conservation Status—Due to the very restricted geo-
graphic area in which S. manabiense occurs (<5,000 km
2
),
the small number of known populations (<5) and the
few collections (4), it is given an IUCN Red List Status
of Endangered.
Etymology—The epithet manabiense refers to Manabı
´
Province of Ecuador where the type was found.
Additional Specimens Examined—ECUADOR.Guayas: 2 – 4 km E from
Recinto Olon, ca. 10 km N of Manglaralto, 19 Feb 1974 (fl, fr), Gentry
10068 (MO). Manabı
´:Montecristi, Cerro Montecristi, eastern slopes
above town, 18 July 1986 (fl, fr), Plowman & Alcorn 14334 (F, NY);
Cerro Montecristi, E slopes above town, 1!0302700S, 80!3905800 W, 400 m,
7 Feb 2009 (fl, fr), Stern & Tepe 374 (QCNE, QCA, UT).
Notes—Solanum manabiense is unique in sect. Gonatotrichum
due to the combination of its rhizomatous habit; bifoliate,
occasionally geminate sympodia; leaves nearly glabrous on
the abaxial surface; unbranched, straight (non-geniculate)
hairs; small flowers with equal stamens; and seeds with
unexpanded margins and no obvious notch at the attach-
ment point to the placenta. It is apparently restricted to
coastal Ecuador in the Guayas and Manabı
´provinces.
Solanum manabiense resembles S. olympicum, but the hairs
of S. olympicum are geniculate whereas those of S. manabiense
are straight. Solanum manabiense also resembles S. turneroides,
but the latter is a more robust herb with heterantherous
flowers that are much larger than those of S. manabiense.
Solanum manabiense is also similar to S. deflexum, but the latter
has leaves pubescent on both surfaces, is not rhizomatous,
488 SYSTEMATIC BOTANY [Volume 38

and has seeds with a swollen margin and pronounced notch
where they connect to the placenta.
7. SOLANUM OLYMPICUM Hassl., Repert. Spec. Nov. Regni
Veg. 9: 116. 1911.—TYPE: PARAGUAY. Alto Paraguay:
Olimpo Berg [Fuerte Olimpo], Dec 1907 (fl, fr) K. Fiebrig
1392 (holotype: G–G00357994!)
Solanum parcistrigosum Bitter, Repert. Spec. Nov. Regni Veg.
12: 75. 1913.—TYPE: PARAGUAY. Estancia Sta. Maria,
18 Feb 1898 (fr), J. D. Anisits 2866 (holotype: S-04– 2965!).
Herb, sometimes slightly woody at base, single- to few-
branched, 1–4 dm tall. Stems sparsely to densely pubes-
cent with two-celled unbranched hairs, these geniculate
between first and second cells and pointing apically, very
rarely with straight hairs. Sympodia 2-foliate, usually
geminate. Leaf blades 1.5–7
+
1–3 cm, elliptic to elliptic-
ovoid, chartaceous to membranaceous, sparsely to moder-
ately pubescent adaxially and abaxially with 1 or 2-celled
unbranched geniculate hairs, these lying flat along blade,
denser along veins; base rounded to obtuse, often decur-
rent into petiole; apex acute; petioles 0.5– 2.5 cm, moder-
ately pubescent with unbranched geniculate hairs. Inflores-
cences with 1–5 flowers, the axes sparsely to moderately
pubescent with unbranched hairs; peduncle absent or nearly
so; rachis absent; pedicels 5–15 mm in flower, 10–20 mm
in fruit. Flowers with the calyx 3 – 10 mm long, the tube
1–3 mm, the lobes 2–7
+
0.5–1.5 mm, linear-lanceolate,
moderately to densely pubescent. Corolla 0.5–1.5 cm in diam-
eter, rotate with abundant interpetalar tissue, chartaceous
to membranaceous, white, the tube 3–6 mm long, the lobes
very short, 1 – 2
+
0.5–1 mm, triangular, acute at apices,
glabrous abaxially and adaxially. Stamens 2 – 4 mm; fila-
ments up to 1 mm long, one filament 0.25 – 0.5 mm longer
then the others; anthers 1.5– 3 mm
+
0.5–1.5 mm, oblong,
the base cordate, the apex emarginate. Ovary glabrous;
style 4– 6
+
0.5– 1 mm, equal to or exserted beyond sta-
mens; stigma to 1 mm wide. Berries 5 – 12 mm in diameter,
globose, white to yellow when immature, maturing semi-
transparent, drying brown, glabrous, the mesocarp watery
and held under pressure until dehiscing explosively at
maturity. Seeds 10– 35 per fruit, ca. 2.5
+
1.5 mm, with
Fig. 11. Photos of type collection of S. manabiense (Stern & Tepe 377, QCNE). A. Habit. B. Flower; note hairs on stem and leaves. C. Fruit, with
skin becoming transparent as turgor pressure increases. D. Roadside habitat in Manabı
´, Ecuador and author with type collection. Scale bars: A = 10 cm.
B, C = 5 mm. (reprinted with permission of the Botanical Research Institute of Texas).
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 489

a small notch where connected to placenta, the margin
not swollen, the surface with fine raised ridges radiating
from center to edges and shallow ridges running parallel
to margin. Figure 12.
Habitat and Distribution—A weedy species of roadsides,
rocky slopes, grassy areas, and forests in central southwest
to eastern Brazil, eastern to central Bolivia, eastern Paraguay,
and northwestern Argentina between 115–2,700 m in eleva-
tion (Fig. 5).
Phenology—Flowering specimens have been collected
January through March and September through November.
Fruiting specimens have been collected January through
May and August through November.
Conservation Status—The widespread distribution and
abundant populations of S. olympicum give it an IUCN Red
List Status of Least Concern.
Etymology—The epithet olympicum refers to the type
locality which was collected in the hills near Fuerte Olimpo.
Additional Specimens Examined—BRAZIL.Bahia: Mun. Urandi, 7 km N
de Urandi, aprox. 14!480S, 42!380W, ca. 600 m, 20 Nov 1992 (fl, fr), Arbo
et al. 5613 (BHCB, CTES, NY, SPF). Goia
´s: Mun. Nova Roma, saı
´da da
cidade em direc¸a
˜o a Iaciara, Fazenda Cachoeira (Proprieta
´rio Sr. Manoel
R. G. de Sousa), 13!4401500S, 46!6201100 W, 710 m, 1 Mar 2000 (fl, fr),
Alvarenga et al. 1295 (IBGE, MO, NY, RB); Mun. Iaciara, Estrada Nova
Roma-Iaciara, Fazenda Sobradinho, 7.6 km de Nova Roma, 13!4705300S,
46!5105100W, 466 m, 4 Nov 2003 (fl, fr), Mello-Silva 2286 (RB, SPF); Mun.
Guarani do Goia
´s, fazenda Forquilha, proprieta
´rio Sr. Vigilato Francisco
dos Santos, 13!4802900S, 46!3201100 W, ca. 450 m, 6 Mar 2001 (fl, fr), Silva
et al. 4860 (NY). Mato Grosso do Sul: Mun. Bonito, Fazenda Vale
Verde, 13 Oct 2003 (fl, fr), Hatschbach et al. 76267 (MBM, NY); Mun.
Miranda, Sede da Fazenda Guaicurus, Pantanal, 13 Jun 1973 (fl, fr),
Silva 194 (SP); Mun. Taquarussu, Rio Baia, margem esquerda, 13 Dec 1992
(fl), Souza 2 (HNUP, RB); Mun. Jateı
´, Rio Ivinhema, Campinho, 11 Dec 1993
(fr), Souza 200 (HNUP, RB).
BOLIVIA.Beni: Prov. Ballivia
´n, Espiritu en la zona de influencia del rio
Yacuma, 200 m, 15 Oct 1980 (fl, fr), Beck 5063 (MO); Prov. Ite
´nez, palmar
de Copernicia alba, 11 Nov 1993 (fl), Moraes et al. 1791 (NY); Prov.
Cercado, a 14 km de San Javier, en la carretera a Trinidad, 14!43014.400S,
64!53051.100W, 200 m, 20 Feb 2000 (fr), Orellana 786 (NY). Santa Cruz:
Prov. N
˜uflo Cha
´vez, Estancia Las Delicias, Hacienda ganadera del
Vicariato Concepcio
´n, 16!290S, 62!030W, 400 m, 14 Nov 2000 (fl, fr),
Beck 25662 (M); vicinity of abandoned old Jardı
´n Bota
´nico along Rio
Piraı
´and roadsides on W side of Santa Cruz, 17!470S, 63!130W,
420 m, 28 Nov 1984 (fr), Nee 30473 (MO, NY); city of Santa Cruz,
17!460S, 63!120W, 400 m, (fl, fr), Nee 30494 (MO, NY); Prov. Andre
´s
Iban
˜ez, along road from Santa Cruz to Samaipata, 1 km SW of
Angostura, 18!090S, 63!310W, 650 m, 13 Jan 1987 (fl, fr), Nee 33478
(NY); Prov. Andre
´s Iban
˜ez, 1 km N of center of Cotoca, 17!450S,
62!590W, 350 m, 15 Jan 1987 (fl, fr), Nee 33545 (NY); Prov. Andre
´s
Iban
˜ez, along Rio Piraı
´, 1 km N of La Guardia, 17!530S, 63!150W, 460 m,
20 Jan 1987 (fl, fr), Nee 33710 (MO, NY, US); Prov. Florida, valley of
Rı
´o Paredones, 0.5 km N of Achiras camping resort, 18!090S, 63!490W,
1350 m, 8 Mar 1998 (fl, fr), Nee 48591 (NY); Prov. Cordillera, S side of
bridge over Rı
´oSeco,18
!40.030S, 63!14.420W, 550 m, 1 May 2001 (fl, fr),
Nee et al. 51724 (BM, MO, NY); Prov. Cordillera, Cabezas, 420 m,
19 Feb 1945 (fl, fr), Pereolo 248 (A); Prov. A. Iban
˜ez, flood plain of the
Rı
´oPiraı
´, ca. 6 km NW of Santa Cruz, vicinity of the sewage settling
ponds, 17!450S, 63!110W, 450 m, 22 Apr 1985 (fr), Solomon 13470 (MO).
Tarija: between entre Rios and Cerere
´,2700m,Mar1952(fl,fr),
Cardenas 4929 (US).
PARAGUAY.Alto Paraguay: Chaco, linea 3 (Oeste), km 50 [19!420S,
61!180W], 44 km al este de la pista de aviacio
´n de Cabrera (19!420S,
61!190W), 9 Nov 1992 (fl), Ramella et al. LR2924 (G); Amambay: ruta
3 y rı
´o Aquidaba
´n, 23 Dec 1980 (fr), Schinini & Bordas 25042 (G).
Caaguazu
´:cerca y al Norte de Yhu
´, 21 Feb 1982 (fl, fr), Casas FC6384
(G, MO); cerca y al Sur de Yhu
´, 24, Sep 1980 (fl), Casas FC3910 (NY);
unos 5 km al Norte de Yhu
´, en una zona inundable en mayor o menor
grado, 320 m, 12 Dec 1982 (fl, fr), Casas & Schinini 7460 (MO, NY);
15 km al N de Caaguazu
´,cauuiuoaYhu
´, 8 Feb 1966 (fl, fr), Krapovickas
et al. 12568 (US); 10–15 km N of Caaguazu
´,19Feb1994(fl,fr),
Pedersen 16075 (G); Arroyo Cambay, 22!250S, 55!550W, 10 Nov 1990
(fl, fr), Zardini & Vela
´zquez 23807 (MO); Arroyo Yuquyry-Arroyo
Taruma, 4 km N of Arroyo Yuquy ry, 25!130S, 55!550W, 12 Jan 1991
(sterile), Zardini & Vela
´zquez 25858 (MO). Canendiyu
´:Mbaracayu
´
Natural Reserve, administered by Fundacio
´nMoise
´sBertoni,around
N
˜andurokai, 23!5903900S, 55!2804400 W, 27 May 1999 (fl), Zardini &
Chaparro 50809 (NY). Concepcio
´n: Estancia n
˜uApua,110kmN
of Concepcio
´n, a 1500 m W. de la Adm., 19 Mar 1991 (fl), Eliceche
44 (MO); Paso Horqueta, Rı
´oAquidaba
´n, 41 km N de Concepcio
´n,
140 m, 17 Dec 1983 (fl, fr), Vanni et al. 377 (G, NY); Estancia Bello
Horizonte, Arroyo Tagatiya
´-Guazu
´,22
!4504000S, 57!2601500W, 13 Oct 1994
(fr), Zardini & Guerrero 41286 (NY); Paso Horqueta, Rio Aquidaban,
20!070S, 57!200W, 18 Nov 1993 (fr), Zardini & Tilleria 37468 (G, MO,
NY). Cordillera: Salto Pirareta
´,25
!300S, 56!550W, 18 Oct 1994 (fl, fr),
Krapovickas et al. 45675 (G); 4 km SE of Emboscada on road to Nueva
Colombia, 25!090S, 57!140W, 18 Nov 1991, (fl, fr), Zardini & T. Tillerı
´a
28888 (MO); Tobatı
´“Ybytu
´Silla” mesa, southern area, 25!120S, 57!070W,
297 m, 3 Mar 1991 (fr), Zardini & Vela
´zquez 26963 (MO). Paraguari:
National Park Ybycu’ı
´,Northeasternarea,26
!010S, 56!460W, 12 Mar 1992
(fr), Zardini & Guerrero 31042 (G, MO, NY); Tucangua, Cordillera de
Altos, 25!310S, 57!090W, 9 Dec 1943, (fr), Rojas 10731 (MO). Without
Dept.: N. Paraguay, zwischen Rio Apa und Rio Aquidaban, 1908/1909
(fl, fr), Fiebrig. 4408 =4827 (BM, G, GH); Paraguay, 1885–1895 (fl),
Hassler 1010 (G); in arenosis pr. Hacurubi, Dec 1885–1895 (fl), Hassler
1585 (G); Sapucay, Dec 1885–1895 (fl), Hassler 1638 (G); in silva pr.
Cordillera de Altos, Jan 1885–1895 (fl), Hassler 1737 (G); in arenosis
pr. Estero Troxler, Jan 1885– 1895 (fl), Hassler 1793 (G); Paraguaria
Centralis, in campo “Intacurabi”, Jan 1900 (fl), Hassler 3801 (BM, G,
GH, NY); Yerbales” montium “Sierra de Maracayu
´”, in regione fluminis
Tapiraguay, Aug (fl, fr), Hassler 4293 (G); Paraguay, (fl, fr), Hassler 4396
(UC); inter ad “Yerbales” montium “Sierra de Maracayu
´”, in regione
vicine “Igatimı
´”, Oct 1898–1899 (fl, fr), Hassler 4841 (G); Cerros de
Tobaty, (fl), Hassler 8069 (G, GH, MO, NY, S, UC, US); in regione lacus
Ypacaray, Jan 1913 (fl, fr), Hassler 12141a (G); Paraguaria Centralis, in
regione lacus Ypacaray, Apr 1913 (fl), Hassler 12441 (BM, G, GH, K,
MO, NY, S, UC, US).
ARGENTINA.Jujuy: Dept. Valle Grande, ca. 10 km N from San Francisco
on road to Valle Grande, 23!35.7130S, 64!58.3960W, 1230 m, 17 Apr 2000
(fr), Nee & Bohs 50808 (NY); Dept. Valle Grande, along gravel and
sandbars of Rı
´o Valle Grande, directly below (W of) San Francisco,
23!37.50S, 64!57.50W, 915 m, 20 Apr 2000 (fr), Nee & Bohs 50825 (NY).
Salta: Hill San Bernardo just outside city of Salta, shady woods among
rocks, 5 Mar 1983 (fr), Bohs 2109 (GH); Dept. Ora
´n, a orillas del
rı
´o Colorado, 30 Jan 1945 (fl), Krapovickas 1565 (US). Tucuma
´n: Dept.
Capital, El Cadillal, Jardı
´n de las Americas, 27 Mar 1975 (fr), Krapovickas
et al. 27853 (MO, WIS).
Notes—Solanum olympicum is the one of the two mem-
bers of sect. Gonatotrichum, along with S. hoffmanseggii, with
characteristic geniculate hairs. These hairs always bend
between the first and second cells and point upwards. This
trait is absent in some specimens from the northern extent
of the species’ range in Bahia and Goia
´s, Brazil (including
Arbo et al. 5613, Alvarenga et al. 1295, Mello-Silva 2286, and
Silva et al. 4860). These specimens are disjunct in their
distribution and have a mixture of hairs that are held per-
pendicular to the stem (similar in orientation but shorter
than those of S. deflexum; see Fig. 1) and hairs that are
adpressed and lie flat along the stem. However, other mor-
phological characteristics of these specimens correspond
to the concept of S. olympicum, and molecular phylogenies
place one of these Bahian specimens as sister to the other
populations of S. olympicum. One specimen from Mato
Grosso do Sul, Hatschbach et al. 76267, has a shrubby habit
with a mixture of geniculate and glandular hairs. This might
represent a new species or simply a shrubby example of
S. olympicum; however, since this form is only represented
by one specimen we have included the collection within
S. olympicum.
Solanum olympicum is similar to S. hoffmanseggii, but these
species can be distinguished by the narrow, lanceolate
leaves and shrubby habit of S. hoffmanseggii compared with
the elliptic to elliptic-ovoid leaves and herbaceous habit
of S. olympicum. The flowers of S. olympicum are weakly
heterantherous with one filament slightly elongated (Fig. 12),
490 SYSTEMATIC BOTANY [Volume 38

Fig. 12. Solanum olympicum (Bohs 3194, UT). A. Habit. B. Inflorescence with flower buds; note adpressed, geniculate hairs on stem. C. Mature fruit
prior to dehiscence; note the skin becoming transparent as turgor pressure increases. D. Fruit after explosive dehiscence. Scale bars = 1 cm.
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 491

which is typically only visible in living plants and differen-
tiates it from the larger, strongly heterantherous flowers of
S. turneroides and S. evolvuloides.Solanum olympicum also
resembles S. adscendens,butthelatterspecieshascordate
leaves, a more branched, spreading growth form, and is
restricted to Rio Grande do Sul in Brazil and neighboring
parts of Argentina. The Bahian collections of S. olympicum
lack glandular hairs on the stems and calyx, unlike the
Bahian species S. evolvuloides, which has abundant glan-
dular pubescence.
The protologue describing S. olympicum placed it in sect.
Morella Dumort. (now sect. Solanum) and until this revision
the name S. olympicum has not been associated with sect.
Gonatotrichum. All specimens of S. olympicum cited in this
revision have been recently annotated as “S. parcistrigosum,”
which was the accepted name until the earlier S. olympicum
was discovered. The protologue describing S. olympicum clearly
indicates that the specimen in the Hassler Herbarium in
Geneva is the holotype. Bitter’s description of S. parcistrigosum
cited only one specimen at the Stockholm herbarium, making
this the holotype.
8. SOLANUM TURNEROIDES Chodat, Bull. Herb. Boissier se
´r. 2,
2: 814. 1902.—TYPE: PARAGUAY. In campis arenosis
pr. fl. Capibary, 5 Sep 1898–1899 (fl), E. Hassler 4396
(lectotype, here designated, G-G00076272!; isolectotypes,
BM-BM000074102!, NY-NY00172221!, UC-944864!).
Solanum gonatotrichum Bitter, Repert. Spec. Nov. Regni Veg.
11: 230. 1912.—TYPE: BOLIVIA. Tarija: Chiquiaca
´, in
silva, 1000 m, 8 Mar 1904 (fl), K. Fiebrig 2732 (lectotype,
here designated, M-M0090347!, photo of lectotype
[Morton neg. 8692]: F!, GH!, UC!; isolectotypes: B
(destroyed), CORD!, SI!, W-W1922– 0001732!, photo
of B isolectotype [F neg. 2713]: GH!).
Solanum geniculatistrigosum Bitter, Repert. Spec. Nov. Regni
Veg. 11: 232. 1912.—TYPE: PARAGUAY. Yaguaron, en
los campos, 1 Dec 1879 (fl), R. Balansa 3132 (holotype: B
(destroyed); photos of holotype [F neg 2751]: F!, G!, GH!,
WIS!; lectotype, here designated, BM-BM000087589!;
isolectotypes: LE!, P-P00384651!).
Solanum flavistrigosum Bitter, Repert. Spec. Nov. Regni Veg.
12: 74. 1913.—TYPE: PARAGUAY. Campo cerrado,
Estancia Sta. Maria, 2 Jan 1897 (flower bud), J. D. Anisits
2018 (holotype: S-05–10!).
Herb, sometimes slightly woody, few- to many-branched,
2– 7 dm tall. Stems sparsely to densely pubescent with straight
one-celled or geniculate two-celled hairs, the straight hairs
common on older growth, the geniculate hairs dense on
new growth. Sympodia 2-foliate, usually geminate. Leaf blades
1–7
+
0.75– 3 cm, elliptic to elliptic-ovoid, chartaceous to
membranaceous, sparsely to moderately pubescent adaxially
and abaxially with 1- or 2-celled unbranched hairs, these
lying flat along blade, denser along veins; base rounded to
obtuse, often decurrent into petiole; apex acute; petioles
0.5–2.5 cm, moderately pubescent with unbranched hairs.
Inflorescences with 1–5 flowers, the axes sparsely to mod-
erately pubescent with unbranched hairs; peduncle absent
or nearly so; rachis absent; pedicels 5– 15 mm in flower,
12– 20 mm in fruit. Flowers with the calyx 3–6 mm long, the
tube 1–3 mm, the lobes 2– 4
+
0.5–1.5 mm, linear-lanceolate,
moderately to densely pubescent. Corolla 1– 2.5 cm in diame-
ter, rotate with abundant interpetalar tissue, chartaceous to
membranaceous, white to purple, the tube 4–10 mm, the
lobes 2– 4
+
1– 2 mm, triangular, acute at apices, glabrous
abaxially and adaxially. Stamens 3–6 mm; upper, shorter
filaments 1– 2 mm, the lowermost, longer filament 3–7 mm,
glabrous or pubescent; anthers 4– 6
+
1– 2 mm, oblong, the
base cordate, the apex emarginate. Ovary glabrous; style
7–10
+
0.5–1 mm, longer than the smaller stamens, closely
appressed to larger stamen, curved near the apex; stigma to
1 mm wide. Berries 10–20 mm in diameter, globose, white to
yellow when immature, maturing semitransparent, drying
brown, glabrous, the mesocarp watery and held under pres-
sure until dehiscing explosively at maturity. Seeds 10–35 per
fruit, ca. 2.5
+
1.5 mm, the entire seed twisted, not flattened,
with a small notch where connected to placenta, the surface
with raised cell walls forming netlike projections. Figure 13.
Habitat and Distribution—A weedy species of roadsides,
grassy pastures, gallery forest, alluvial flats, forest edges,
and open shrubby vegetation that occurs from central Bolivia
to eastern Paraguay and the Brazilian state of Mato Grosso
do Sul and south into northwestern Argentina between
300–1,950 m in elevation (Fig. 5).
Phenology—Flowering specimens have been collected in
all months except August with a peak from October through
February. Fruiting specimens have been collected in January
through May and September through December.
Conservation Status—The widespread distribution and
abundant populations of S. turneroides give it an IUCN Red
List Status of Least Concern.
Etymology—The epithet turneroides presumably refers to
a resemblance to the genus Turnera (Turneraceae), possibly
due to the five-parted flowers that usually open in the
morning and last for only a few hours. In the description,
Chodat mistakenly likens the habit of S. turneroides to that of
S. caripense Dunal, a viny member of Solanum sect. Basarthrum.
Additional Specimens Examined—BRAZIL.Mato Grosso do Sul: Mun.
Corumba
´, Morro Bocaina, 18 Oct 1991 (fl), Damasceno 178 (COR, UEC);
Bairro Aeroporto, morro defronte ao aeroporto, rua Alan Kardec,
19!010S, 57!390W, 220 m, 25 Jan 2001 (fl) Gomes 39 (SPF); Mun. Bela
Vista, 10 km W, 17 Mar 1985 (fl), Hatschbach & Zelma 49158 (MBM,
NY); Mun. Aquidauana, Piraputanga, 4 Jun 1994 (fl), Hatschbach et al.
60696 (MBM, NY); Mun. Porto Murtinho, rodovia Bonito-Campo do
I
´ndios, Fazenda A
´gua Doce, 10 Nov 2002 (fl, fr), Hatschbach 74004
(MBM); Mun. Bonito, Fazenda Nossa Senhora do Perpe
´tuo Socorro,
12 Oct 2003 (fl, fr), G. Hatschbach et al. 76260 (CTES, MBM, NY); Mun.
Porto Murtinho, rodovia Jadim-Porto Murtinho, BR-267, pro
´ximo do
Rio Perdido, 250 m, 15 Mar 2004 (fl), Hatschbach 77410 (MBM); Mun.
Corumba
´,Jul1911(fl),Hoehne 3749 (R, US); Estrada para a Cha
´cara
Sa
˜o Marcos, Bairro entre Cristo Redentor e Cravo Vermelho, 19!02016.800S,
57!37041.700W, 29 Nov 2000 (fl, fr) Moraes 554 (UEC); Corumba
´,Fazenda
of Dr. Romeu, 19!010S, 57!390W, 20 Nov 1987 (fl), Ratter et al. 6043 (MO);
15 km from Corumba
´, 29 Jan 1991 (fl) Ratter 6513 (CPAP, MBM); Mun.
Corumba
´, Morro de Azeite, 10 Apr 1992 (fr), Resente 667 (BHCB, CGMS);
Corumba
´, 18 Dec 1902 (fl, fr), Robert 721 (BM); Mun. Miranda, Salobra,
Dec 1941 (fl) Santos s.n. (R 79588); Mun. Bela Vista, Rio Guaviral,
12 Nov 2006 (fl, fr), Silva et al. 5266 (MBM).
BOLIVIA.Beni: Prov. Cercado, campus of the Universidad Tecnica
del Beni, 2.5 km N of center of Trinidad, 14!480S, 64!530W, 200 m, 13 Dec
1988 (fl, fr), Nee 37160 (NY). Chuquisaca: Oropeza, ca. 5 km below
Chuquichuqui in Rı
´o Chaco Valley, 1800 m, 19 Jan 1997 (fl, fr), Wood
11679 (K, LPB). Cochabamba: Rı
´o Caine, 1,180 m, Jan 1949 (fl, fr),
Ca
´rdenas 4098 (US). Santa Cruz: Prov. Nuflo de Chavez, Concepcio
´n,
500 m, 18 Feb 1995 (fr), Abbot 16233 (WU); Santa Cruz, 1200 ft, 23 Sep
1964 (fl, fr), Badcock 416 (K); Prov. A. Iban
˜ez, ca. 15 km hacia el N de
Santa Cruz, por el Nuevo aeropuerto Viru Viru, 420 m, 19 Mar 1981
(fl, fr), Beck 6666 (M); Prov. Cordillera, Alto Parapetı
´, 800 m, 16 Jan 1980
(fl), de Michel 44 (NY); Prov. Caballero, Estancia Lanza-Lanza, 2 km
del Rı
´o Comarapa sobre ladera con 30!de inclinacio
´n exposicio
´nW,
18!0200400S, 64!3500000 W, 1600 m, 22 Jan 1995 (fl, fr), Gutie
´rrez et al. 1521
(NY); Prov. Andre
´s Iba
´n
˜ez, 1 km N from Pedro Lorenzo, 20 km along
492 SYSTEMATIC BOTANY [Volume 38

the Camiri highway, 9 May 2000 (fr), Kuroiwa &. Maeda 1596 (NY);
Alto Parapetı
´, 850 m, 8 Jan 1982 (fl), Michel 98 (LPB); Prov. Andre
´s
Iba
´n
˜ez, along Rı
´o Pantano [=Rı
´o Chore-Chore], 7 km SE of Palmar del
Oratorio and 18 km SE of center of Santa Cruz, 17!560S, 63!060W,
380 m, 9 Dec 1988 (fl, fr), Nee 37085 (LPB, TEX, NY, WIS); Prov.
Andre
´s Iba
´n
˜ez, 13 km SE of Palmar del Oratorio and 6 km SE of
Rio Chore-Chore [= Rı
´o Pantano] 17!580S, 63!040W, 375m, 22 Jan 1989
(fl, fr), Nee 37668 (G, MO, NY); Prov. Andre
´s Iba
´n
˜ez, NW side
of “Valle Sanchez,” 4 km W of Aeropuerto Internacional Viru-Viru,
15 km N of Santa Cruz, 17!380S, 63!100W, 375 m, 26 Jan 1989 (fl, fr),
Nee 37741 (NY); Prov. Caballero, W side of Rı
´o Comarapa, 0.5 km W of
center of Comarapa, 17!540S, 64!320W, 1825 m, 12 Dec 1992 (fl), Nee
43093 (NY); Prov. Andre
´s Iba
´n
˜ez, along highway from Santa Cruz to
Abapo
´, 3 km S of crossing of railroad and 2 km S of bridge over
Quebrada Peji, 17!580S, 63!110W 450 m, 25 Apr 1998 (fl, fr), Nee 49117
(CORD, G, NY); Prov. Vallegrande, 4 km SW of El Trigal on road
to San Juan del Chaco, S side of Rı
´o Ariruma, 19 km (by air) NNW of
Vallegrande, 18!290S, 64!070W, 1950 m, 31 Jan 1987 (fl), Nee & Coimbra
33932 (NY); Prov. Vallegrande, 10 km (by air) NNW of Vallegrande,
18!230S, 64!080W, 1850 m, 1 Feb 1987 (fl, fr), Nee & Coimbra 33944
(G, NY, US); Prov. Florida, 7 km (by air), 10.2 km (by road) NNW of
Mataral on road to San Juan del Potrero, 18!0204500S, 64!1402500 W,
1475 m, 30 Jan 1994 (fl, fr), Nee & Vargas 44789 (NY); Prov. Andre
´s
Iba
´n
˜ez, along highway from Santa Cruz to Abapo
´, 3 km S of crossing
of railroad and 2 km S of bridge over Quebrada Peji, 17!580S, 63!110W,
450 m, 27 Feb 1998 (fr, fl), Nee et al. 48485 (NY); Prov. Andre
´s Iba
´n
˜ez,
along highway from Santa Cruz to Abapo
´, 3 km S of crossing of
railroad and 2 km S of bridge over Quebrada Peji, 17!580S, 63!110W,
450 m, 1 May 2001 (fl), Nee et al. 51716 (NY); Prov. Florida, 3 km
(by air), 7 km (by road) NE of Mairana on road to Parque Nacional
Fig. 13. Solanum turneroides (Bohs 2715, UT). A. Stem with nearly mature fruits. B. Flower bud; note densely pubescent calyx and long calyx lobes.
C. Flower on first day of opening; note that lowermost filament has not expanded. D. Flower after first day; note that the filament of the lowermost
anther is now greatly elongated. Scale bars: A, C, D = 1 cm. B = 5 mm.
2013] STERN ET AL.: SOLANUM SECTION GONATOTRICHUM 493

Amboro
´camp “Yunga de Mairana,” 18!060S, 63!560W, 1850 m, 28 Mar
2002 (fl, fr), Nee et al. 52003 (NY); Prov. Cordillera, Cabezas, 420 m,
19 Feb 1945 (fl), Pareolo 253 (A, NY); Prov. Cordillera, Cabezas, 420 m,
18 Mar 1945 (fl, fr), Pareolo 444 (NY); Serranias de Chiquitos, camino
a Tucavaca 2 Feb 2005 (bs), Solis Neffa 1790 (BHCB, CTES); Prov. Sara,
“Arenales del Gwenda” [sandy areas in savanna of Rı
´o Guenda
´, now
forming border of Prov. Ichilo and Prov. Andre
´s Iba
´n
˜ez, M. Nee 2001]
450 m, 8 May 1921 (fl, fr), Steinbach 5625 (GH, NY); Prov. Sara,
Arenales, Rı
´o Perdix, 400 m, 23 Oct 1924 (fl), Steinbach 6627 (G, GH,
K); Prov. Sara, Rı
´o Perdix, 400 m, 29 Dec 1925 (fl), Steinbach 7367 (F, G,
GH, MO); Prov. Chiquitos, Cerro Mutu
´n, 7 km al NE de la pista de
aterrizaje del campamento minero (25 km al S de Puerto Sua
´rez),
18!11.30S, 57!52.70W, 750 m, 17– 20 Oct 1994 (fl), Vargas et al. 3328
(CTES, NY, USZ); ca. 4 km E of San Isidro (Palizada) on road to
Mataral, 18!20S, 64!250W, 1400 m, 3 Jan 2000 (fl), Wood 15770 (LPB).
Tarija: Prov. Gran Chaco, 5– 5.7 km W of the center of Villa Montes
on road to Entre Rı
´os and Tarija (not the new segment under construc-
tion now), 0– 0.7 km toward Villa Montes from the highway bridge
over the Rı
´o Pilcomayo, 21!1502800S, 63!3004000 W, 400 m, 9 Feb 2006
(fr), Nee & Linneo 54034 (G); Gran Chaco, San Francisco de Inti, camino
de Yacuiba a Villa Montes, a 22 km N de la rotonda de Yacuiba,
21!4805600S, 63!3500500 W, 586 m, 16 Jan 2004 (fl), Neffa et al. 989 (CTES,
G); Prov. O’Connor, hills north of Entre Rios, 1400 m, 6 Feb 1937
(fl, fr), West 8254 (GH, MO, UC).
PARAGUAY.Alto Paraguay: Chaco, linea 3 (Oeste), km 50 [19!420S,
61!180W], 44 km al E de la pista de aviacio
´n de Cabrera (19!420S,
61!190W), 9 Nov 1992 (fl), Ramella et al. LR2924 (G). Amambay: Ruta 3 y
Rı
´o Aquidaba
´n, 23 Dec 1980 (fr), Schinini & Bordas 25042 (G). Caaguazu
´:
cerca y al N de Yhu
´, 21 Feb 1982 (fl, fr), Ferna
´ndez Casas FC6384 (G, MO);
cerca y al S de Yhu
´, 24, Sep 1980 (fl), Ferna
´ndez Casas FC3910 (NY); unos
5 km al N de Yhu
´, en una zona inundable en mayor o menor grado,
320 m, 12 Dec 1982 (fl, fr), Ferna
´ndez Casas & Schinini 7460 (MO, NY);
15 km al N de Caaguazu
´, camino a Yhu
´, 8 Feb 1966 (fl, fr), Krapovickas
et al. 12568 (US); 10– 15 km N of Caaguazu
´, 19 Feb 1994 (fl, fr), Pedersen
16075 (G); Arroyo Cambay, 22!250S, 55!550W, 10 Nov 1990 (fl, fr),
Zardini & Vela
´zquez 23807 (MO); Arroyo Yuquyry-Arroyo Taruma, 4 km
N of Arroyo Yuquyry, 25!130S, 55!550W, 12 Jan 1991 (sterile), Zardini &
Vela
´zquez 25858 (MO). Canendiyu
´:Nandurocai, Reserva Natural de
Bosque Mbaracayu
´, 13 km S de Ipe
´-hu
´, 3 Dec 1997 (fl, fr), Schinini 33262
(CTES); Mbaracayu
´Natural Reserve, administered by Fundacio
´n
Moise
´s Bertoni, around N
˜andurokai, 23!5903900S, 55!2804400 W, 27 May
1999 (fl), Zardini & Chaparro 50809 (NY). Concepcio
´n: Estancia N
˜u
Apua, 110 km N of Concepcio
´n, a 1500 m W de la Adm., 19 Mar 1991
(fl), Eliceche 44 (MO); Paso Horqueta, Rı
´o Aquidaba
´n, 41 km N de
Concepcio
´n, 140 m, 17 Dec 1983 (fl, fr), Vanni et al. 377 (G, NY); Estancia
Bello Horizonte, Arroyo Tagatiya
´-Guazu
´, 22!4504000S, 57!2601500W, 13 Oct
1994 (fr), Zardini & Guerrero 41286 (NY); Paso Horqueta, Rı
´o Aquidaban,
20!070S, 57!200W, 18 Nov 1993 (fr), Zardini & Tilleria 37468 (G, MO, NY).
Cordillera: Salto Pirareta
´,25
!300S, 56!550W, 18 Oct 1994 (fl, fr),
Krapovickas et al. 45675 (G); 4 km SE of Emboscada on road to Nueva
Colombia, 25!090S, 57!140W, 18 Nov 1991, (fl, fr), Zardini & Tillerı
´a28888
(MO); Tobatı
´“Ybytu
´Silla” mesa, southern area, 25!120S, 57!070W,
297 m, 3 Mar 1991 (fr), Zardini & Vela
´zquez 26963 (MO). Paraguari:
Pirareta
´,14Nov1969(fl,fr),Pedersen s.n. (CTES 315405); National
Park Ybycu’ı
´, Northeastern area, 26!010S, 56!460W, 12 Mar 1992 (fr),
Zardini & Guerrero 31042 (G, MO, NY); Tucangua, Cordillera de Altos,
25!310S, 57!090W, 9 Dec 1943, (fr), Rojas 10731 (MO). Without Dept.:
N Paraguay, zwischen Rio Apa und Rio Aquidaban, 1908/1909 (fl, fr),
Fiebrig 4408 =4827 (BM, G, GH); Paraguay, 1885–1895 (fl), Hassler 1010
(G); in arenosis pr. Hacurubi, Dec 1885–1895 (fl), Hassler 1585 (G);
Sapucay, Dec 1885–1895 (fl), Hassler 1638 (G); in silva pr. Cordillera
de Altos, Jan 1885–1895 (fl), Hassler 1737 (G); in arenosis pr. Estero
Troxler, Jan 1885– 1895 (fl), Hassler 1793 (G); Paraguaria Centralis,
in campo “Intacurabi,” Jan 1900 (fl), Hassler 3801 (BM, G, GH,
NY); “Yerbales” montium “Sierra de Maracayu
´,” in regione fluminis
Tapiraguay, Aug (fl, fr), Hassler 4293 (G); Paraguay, (fl, fr), Hassler 4396
(UC); Inter ad “Yerbales” montium “Sierra de Maracayu
´”, in regione
vicine “Igatimı
´,” Oct 1898–1899 (fl, fr), Hassler 4841 (G); Cerros de Tobaty,
(fl), Hassler 8069 (G, GH, MO, NY, S, UC, US); in regione lacus Ypacaray,
Jan 1913 (fl, fr), Hassler 12141a (G); Paraguaria Centralis, in regione
lacus Ypacaray, Apr 1913 (fl), Hassler 12441 (BM, G, GH, K, MO, NY,
S, UC, US).
ARGENTINA.Jujuy: Dept. Capital, Alto La Vin
˜a, ruta 56, 4 km al NE
de Jujuy, 1250– 1300 m, 29 Dec 1989 (fr), Novara 9287 (G). Salta: Dept.
Capital, Chachapoyas, Sierra de Ve
´lez, cerros al E de la Univers.
Cato
´lica, 1200 m, 30 Jan 1987 (fl), Novara 5884 (G); same locality and
date, Novara 5919 (G).
Notes—Strongly heterantherous flowers are only found in
two members of the section, S. turneroides and S. evolvuloides.
Solanum turneroides can be distinguished from S. evolvuloides
by its lack of glandular hairs, its larger flowers, and its
more widespread South American distribution. As noted
by Nee (1989), the flowers of S. turneroides are not open
in the heat of the day, as is also the case for S. evolvuloides
(L. Giacomin, pers. obs.). When grown in the greenhouse
at UT the flowers were open and very fragrant during the
night and early morning. As the day gets warmer the flowers
take on a wilted appearance. The pollinators of these flowers
are unclear. Fragrance is unusual in the genus and is com-
monly only found in S. sect. Pachyphylla (Bohs 1994). The
floral development is also of interest, with the stamens of
equal length on the first day the flower is open and then
the filament of the lowermost anther doubling in length the
following day (Figs. 13C, D).
Although herbarium sheets rarely display the feature,
S. turneroides seems to spread rhizomatously. The hairs of
S. turneroides are also unique in the section. While they super-
ficially appear to be geniculate, like those of S. olympicum
and S. hoffmanseggii, closer inspection shows that the hairs are
simply bent downward on the stem and lack the 90!elbow
bend that characterizes the hairs of the latter two species.
Using scanning electron microscopy, the base of these hairs
appears to have a ring of small lateral cells, hinting that
perhaps these hairs are reduced stellate hairs. If so, it may
suggest that the simple hairs found in this and other spe-
cies of sect. Gonatotrichum may represent evolutionary reduc-
tions from stellate hairs.
The protologue of S. turneroides cites many Hassler speci-
mens as syntypes, all from Paraguay, but gives no herbarium
locations. Hassler 4396 has been chosen from among the
many syntypes cited because of the quality of the collection
and its wide distribution in herbaria. The sheet from the
Hassler herbarium at G is chosen as the lectotype.
Bitter cited four syntypes in his protologue for S. gonatotrichum,
including Fiebrig 2732 and three un-numbered specimens
collected in Salta, Argentina by Hieronymus & Lorentz,all
from Berlin and destroyed. A duplicate specimen of the
only numbered syntype, Fiebrig 2732,isatMunichandwe
have designated this as the lectotype because of the quality of
the material and widespread photographs of the specimen.
Bitter’s protologue of S. geniculatistrigosum cited only a
single specimen from B, the holotype, which was destroyed
in 1942. Photos of this sheet remain at F, G, GH and WIS;
a duplicate at BM has been designated the lectotype and
isolectotypes have been seen at LE and P.
Bitter also cited only one specimen of S. flavistrigosum at
S, making it the holotype. The only unusual aspect of this
is that Bitter describes the flowers in detail, but the specimen
only has buds.
Acknowledgments We thank the following herbaria for hospitality
during visits and/or for loans of specimens used in this study: A, BH,
BHCB, BM, BR, CEPEC, CESJ, CORD, CPAP, CTES, ESA, FUEL, G, GH,
HAS, HB, HNUP, HUEFS, IAC, IAN, IBGE, ICN, INB, INPA, JPB, K,
LPB, M, MBM, MBML, MEXU, MO, NY, P, PACA, PEL, QCA, QCNE,
R, RB, SI, SP, SPF, SPSF, TEX, UC, UEC, UPCB, US, USZ, UT, VIC,
WIS, W, WU. We also thank Lilian Mentz, Livia Echternacht, and Eric
Tepe for assistance in the field, and E. Tepe for spotting S. manabiense;
Eric Tepe and Terri Weese for laboratory assistance; Lilian Mentz for the
photo of S. hoffmanseggii; Ann Kelsey at UT and Alexandre Salino for
help managing herbarium loans; and Juliana Ordones, Miriam Pimentel,
and Ine
ˆs Ribeiro for greenhouse assistance at the Jardim Bota
ˆnico da
494 SYSTEMATIC BOTANY [Volume 38

Fundac¸a
˜o Zoo-Bota
ˆnica de Belo Horizonte. We also thank Tom Ranker
and two anonymous reviewers for improving the manuscript. This
work was supported by NSF through the PBI: Solanum grant, DEB-
0316614, to LB and SK and by FAPEMIG (APQ-01600-08) and CNPq
(305589/2009-1) to JS.
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Appendix 1. Herbarium vouchers and seed provenances for breeding
studies.
S. adscendens, BRAZIL: Est. Rio Grande do Sul, Stehmann 6003
(BHCB); S. deflexum, COSTA RICA: Prov. Guanacaste, Bohs 2715 (UT);
S. evolvuloides, BRAZIL: Est. Bahia, Giacomin 974 (BHCB); S. manabiense,
ECUADOR: Prov. Manabı
´,Stern & Tepe 374 (UT) & 379 (UT); S. olympicum,
BOLIVIA: Dept. Santa Cruz, Bohs 2738 (UT); S. turneroides,BOLIVIA:
Dept. Santa Cruz, Nee et al. 51716 (UT);
Numerical List of Species—1. S. adscendens Sendtn.; 2. S. deflexum
Greenm.; 3. S. evolvuloides Giacomin & Stehmann; 4. S. hoffmanseggii
Sendtn.; 5. S. lignescens Fernald; 6. S. manabiense S.Stern; 7. S. olympicum
Hassl. 8. S. turneroides Chodat
Index to Numbered Collections—The numbers in parenthesis refer to
the corresponding species in the text and in the Numerical List of Spe-
cies presented above. Abbot, J. R. 16233 (8), Alvarenga, D. et al. 1295 (7),
Anisits, J. D. 2018 (8), 2866 (7), Arbo, M. M. et al. 5613 (7), Ayala, M. G. 136
(5), Ayala, M. G. & Lott, E. J. 9 (5), Badcock, W. J. 416 (8), Balansa, B. 3132
(8), Barboza, G. et al. 1494 (1), Beck, S. G. 5063 (7), 6666 (8), 25662 (7), Black,
G. A. 48-2316 (4), Bohs, L. 2109 (7), 3194 (7), Brack, P. 1714 (1), Brandegee,
T. S. 412 (2), Breedlove, D. 52762 (2), Bueno, O. L. 344 (1), Buscalioni, L.
3656 (4), Bye Jr., R. 171 (2), Cabrera, A. et al. 29445 (1), Caldero
´n, S. 1169
(2), Carballo, R. A. 371 (2), Carballo, R. A. & Carrillo, M. 433 (2), Cardenas,
M. 4098 (8), 4929 (7), Carneiro, A. 443 (1), Carranza, E. 3413 (2), Carter,
A. 4991 (2), 5621 (2), Carter, A. & Moran, R. 5376 (2), Carvalho, A. M. de
1591 (3), Chavarrı
´a, U. 1503 (2), 1766 (2), Chaves, D. 2 (2), Co
´bar & Garcı
´a
385 (2), 520 (2), Cowan, C. C. et al. 5455 (2), da Souza, M. C. et al. 223 (4),
D’Arcy, W. G. 11896 (2), Damasceno, G. 178 (8), Davidse, G. et al. 30657 (2),
35391 (2), 35103 (2), Dias, A. T. G. 627 (4), Dı
´az, A. L. 63 (2), Eliceche, A.
44 (8), Elorsa, C. M. 700 (2), 3117 (2), Ferna
´ndez Casas, J. FC3910 (8),
FC6384 (8), Ferna
´ndez Casas, J. & Molero, J. F. C. 6384 (7), Ferna
´ndez Casas,
J. &, Schinini, A. A. 7460 (8), Fiebrig, K. 2732 (8), 4408=4827 (8), Fish, J. 73
(2), Flores M. A. 2025 (2), Fro
´es, R. L. 24667 (4), Garcı
´a et al. 715 (2), Gentry,
A. H. 10068 (6), 14430 (2), Giacomin, L. L. 974 (3), Gilman, M. F. 80 (2),
Gomes, C. G. 39 (8), Grings, M. 340 (1), Gutierrez, E. et al. 1521 (8), Hammel,
B. & Grayum, M. 19933 (2), Hampshire, R. J. et al. 1151 (5), 1154 (5), 1184
(5), Hansen, B. F. & Nee, M. 7447 (2), Happ, G. B. 101 (2), Harmon, W. E. &
Dwyer, J. D. 3085 (2), 3399 (2), Harmon, W. E. & Fuentes, J. A. 6016 (2),
Harrison, G. J. 4777 (2), 8147 (2), Hassler, E. 1010 (8), 1585 (8), 1638 (8),
1737 (8), 1793 (8), 3801 (8), 4293 (8), 4396 (8), 4841 (8), 8069 (8), 12141a
(8), 12441 (8), Hatschbach, G. 74004 (8), 77140 (8), Hatschbach, G. & Zelma
49158 (8), Hatschbach, G. et al. 60696 (8), 76260 (8), 76267 (7), Heithaus,
E. R. 274 (2), Hinton, G. B. et al. 4219 (2), 4470 (2), 6481 (2), 9143 (2),
10557 (2), 12057 (2), Hoehne, F. C. 3749 (8), Holm, R. & Iltis, H. H. 295
(2), Howell, J. T. 10253 (2), Jardim, J. 1843 (3), Jarenkow, J. A. 720 (1),
Jarenkow, J. A. & Bueno, O. L. 1171 (1), Jesus, J. A. 367 (3), Jime
´nez, A. 721
(2), Kearney, T. H. & Peebles, R. H. 10387 (2), Keller, H. A. 3746 (1), King,
G. 446 (2), 555 (2), Kral, R. L. 69080 (2), Krapovickas, A. 1565 (7),
Krapovickas, A. et al. 12568 (8), 25819 (1), 27853 (7), 45675 (8), Kuroiwa,
N. & Maeda, N. 1596 (8), Langman, I. K. 2145 (2), 3337 (5), LaSalle, J.
et al. 810629-2 (2), Leavenworth, W. C. 473 (2), Leavenworth, W. C. &
Hoogstraal, H. 1279 (2), 1344 (2), Lehmann, F. C. 1671 (2), Lehto, E. 24716
(2), Leite, J. E. 658 (1), 1864 (1), Liesner, R. et al. 2798 (2), Linares, J. L.
3536 (5), Lisboa, R. 6782 (4), Lobato, L. C. B. 2656 (4), 3282 (4), Magallanes,
A. S. 3650 (5), 4230 (5), Makrinius, E. 530 (5), Martı
´nez, C. R. 1461 (2),
Martı
´nez, C. R. & A
´guilar 36935 (2), Martı
´nez Salas, E. M. et al. 31192
(5), Maxon, W. R. et al. 7138a (2), 7191 (2), McVaugh, R. 15658 (2),
15796 (2), Mexia, Y. 673 (2), Michel, R. 44 (8), 98 (8), Molina, A. R.
2535 (2), 24551 (5), 27185 (2), Montalvo, A. M. 6380 (2), Moraes, M. 554
(8), Moraes, M. et al. 1791 (7), Moreno, P. P. 2157 (2), 9222 (2), 9280
(2), 9294 (2), 16703 (2), Nee, M. 30473 (7), 30494 (7), 33478 (7), 33545 (7),
33710 (7), 37085 (8), 37160 (8), 37668 (8), 37741 (8), 43093 (8), 48591 (7),
49117 (8), Nee, M. & Bohs, L. 50808 (7), 50825 (7), Nee, M. & Coimbra, S.
33932 (8), 33944 (8), Nee, M. & Linneo, I. I. 54034 (8), Nee, M. & Vargas,
I. 44789 (8), Nee, M. et al. 48485 (8), 51716 (8), 51724 (7), 52003 (8),
Neffa, V. et al 989 (8), Nelson, E. W. 2876a (2), Neill, D. A. 2174 (2), 2460
(2), Novara, B. 5884 (8), 5919 (8), 9287 (8), Opler, P. 860 (2), Orcutt, C. R.
4210 (2), 4389 (2) 5286 (2), Orellana, R. 786 (7), Ortiz, J. J. 1029 (2), Palmer,
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E. 216 (5), Pedersen, T. M. 16075 (8), Peebles, R. H. et al. 5625 (2), Pennell,
F. W. 19496 (2), Pereolo 248 (7), 253 (8), 444 (8), Pfeifer, H. W. 1245 (2),
Pittier, G. E. 3646 (2), Plowman, T. & Alcorn, P. 14334 (6), Pringle, C. G.
6400 (2), 6729 (2), 7508 (2), Purpus, C. A. 3563 (2), 6100 (2), 7509 (2), 7860
(2), 8016 (2), 8498 (2), 10771 (2), 12055 (2), 13000 (2), Ramella, L. et al.
LR2924 (8), Ramı
´rez, N. & Ve
´liz, M. 948 (2), Ramos, B. 1010 (4), Ratter,
J. A. 6513 (8), Ratter, J. A. et al. 6043 (8), Resente, U. M. 667 (8), Reyes
Garcia, A. & Uriquijo 794 (5), Robert, A. 721 (8), Rodrı
´guez, J. V. 940 (2),
1013 (2), 2126 (2), 3800 (2), Rodrı
´guez, J. V. et al. 3909 (2), Rojas, T. 10731
(8), Rosales, J. M. 837 (2), Rosa
´rio, C. S. 713 (4), Rose, J. N. 1852 (2), Rose,
J. N. & Hay, R. 6288 (2), Rose, J. N. et al. 8587 (2), Rueda, R. 7420 (2),
Rusby, H. H. 85 (2), Salinas, A. et al. 7242 (2), Sanders, A. C. et al. 8033
(2), Sandino, J. C. 3134 (2), Schery, R. W. 107 (2), Schinini, A. A. 33262 (8),
Schinini, A. A. & Bordas. E. 25042 (8), Sehnem, A. 1546 (1), Silva, M. F. F.
112 (4), Silva, N. T. et al. 4860 (7), 4868 (7), 5266 (8), Silveira, N. 1699 (1),
8734 (1), 9634 (1), Smith, A. 403 (2), Smith, J. & Rojas, J. 348 (2), 561 (2),
Sobral, M. & Almeida, S. C. 7911 (1), Solis Neffa, V. 1790 (8), Solomon, J. C.
13470 (7), Soto Nu
´n
˜ez, J. C. 952 (2), Souza, M. C. 2 (7), 200 (7), Spellman,
D. L. et al. 95 (2), Standley, P. C. 1654 (2), 1811 (2), 3833 (2), 11148
(2), 13024 (2), 16093 (2), 19647 (2), 21306 (2), 22063 (2), 22210 (2), 22258
(2), 22678 (2), 73719 (2), Standley, P. C. & Chaco
´n, J. P. 5407 (2), Standley,
P. C. & Molina, A. R. 13233 (2), 14278 (2), Standley, P. C. & Padilla, E. V.
3808 (2), Stehmann, J. R. 473 (1), Stehmann, J. R. et al. 6001 (1), 6002 (1),
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S. & Tepe, E. J. 374 (6), 377 (6), Stevens, W. D. 2610 (2), 4708 (2), 9363
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496 SYSTEMATIC BOTANY [Volume 38