![Solanum alternatopinnatum from Paraguay (É. Hassler 9365, lectotype of Solanum juciri Sendtn. forma paraguariense Hassl., BM [incorrectly labelled isolectotype in image]). Reproduced with permission of the Natural History Museum, London.](https://www.researchgate.net/profile/Lynn-Bohs/publication/290196470/figure/fig3/AS:11431281174384674@1689202695610/Solanum-alternatopinnatum-from-Paraguay-E-Hassler-9365-lectotype-of-Solanum-juciri_Q320.jpg)
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A Revision of Solanum Section Aculeigerum (the Solanum wendlandii Group,
Solanaceae)
Author(s): John L. Clark, Michael Nee, Lynn Bohs and Sandra Knapp
Source: Systematic Botany, 40(4):1102-1136.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364415X690148
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Systematic Botany (2015), 40(4): pp. 1102–1136
© Copyright 2015 by the American Society of Plant Taxonomists
DOI 10.1600/036364415X690148
Date of publication December 15, 2015
A Revision of Solanum section Aculeigerum (the Solanum wendlandii group, Solanaceae)
John L. Clark,
1,2
Michael Nee,
3
Lynn Bohs,
4
and Sandra Knapp
5,6
1
Department of Biological Sciences, Box 870345, The University of Alabama, Tuscaloosa, Alabama 35487-0345, U. S. A.
2
Department of Botany, Smithsonian Institution, PO Box 37012, National Museum of Natural History, MRC-166,
Washington, D. C. 20013-7012, U. S. A.
3
The New York Botanical Garden, Bronx, New York 10458-5126, U. S. A.
4
Department of Biology, 257 South 1400 East, University of Utah, Salt Lake City, Utah 84112, U. S. A.
5
Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, U. K.
6
Author for correspondence: s.knapp@nhm.ac.uk
Communicating Editor: Michael J. Moore
Abstract—Solanum section Aculeigerum (the Solanum wendlandii group, Solanaceae) comprises eight species ranging from Mexico and Cen-
tral America to Ecuador and Peru with one species in southeastern Brazil, Paraguay and northern Argentina. One species, Solanum
wendlandii, is commonly cultivated worldwide in tropical and subtropical regions, and one new species from southern Mexico, Solanum
triunfense, is described here. Members of Solanum section Aculeigerum are differentiated from other sections of Solanum by a combination of
plurifoliate sympodial units, branched inflorescences, presence of prickles coupled with absence of stellate trichomes, and a vine-like habit.
Unlike most other prickly species of Solanum, which are placed into Solanum subgenus Leptostemonum, they lack stellate hairs, and phylo-
genetic studies suggest that the section is one of several potential sister groups for the spiny solanums. A key, descriptions, habitat and dis-
tribution data, preliminary conservation assessments, specimens examined and illustrations are provided for all species in Solanum section
Aculeigerum. Lectotypes are designated for the following five names: S. aculeolatum M. Martens & Galeotti, S. mazatenangense Coult. & Donn.
Sm., S. molinarum J. L. Gentry, S. pachyandrum Bitter, and S. sagranum A. Rich., and neotypes are designated for S. glaucescens Zucc. and
S. wendlandii Hook. f.
Keywords—Central America, heterandry, lianas, South America, vines.
Introduction
Solanum L. is one of the ten most species-rich genera of
flowering plants (Frodin 2004) and has approximately 1,400 spe-
cies (Särkinen et al. 2013) occurring on all temperate and
tropical continents. The highest diversity of both groups and
species is in tropical South America, concentrated in circum-
Amazonia (Knapp 2002). Species of Solanum have usually
five-merous flowers with fused sepals and petals, stellate to
pentagonal corollas, stamens with short filaments, and anthers
opening by terminal pores. The genus was one of Linnaeus’s
(1753) larger, with 23 species mostly described from European
or African material. The last time Solanum was monographed
in its entirety was in De Candolle’s Prodromus (Dunal 1852),
which included 901 species (with an additional 19 listed at
the end of the treatment as incompletely or poorly known).
Until the 21st century, the taxonomy of Solanum was largely
limited to rearrangements of infrageneric taxa, species-level
treatments of smaller groups within the genus, and floristic
works. The large size of Solanum and its poorly understood
infrageneric structure has meant that Solanum taxonomy has
proceeded in a piecemeal fashion until relatively recently, and
the genus has acquired a reputation of being intractable. A
project funded by the United States National Science Founda-
tion’s Planetary Biodiversity Inventory (PBI) program begun
in 2004 has sought to accelerate species-level taxonomic work
across the genus as a whole. Ongoing work by participants of
the PBI Solanum project is resulting in a modern monographic
treatment of the entire genus available online in the electronic
resource Solanaceae Source (http://www.solanaceaesource.org).
This treatment is part of that collaborative effort.
Taxonomy of Solanum section Aculeigerum —One of the
principal divisions in Solanum is that between spiny (techni-
cally prickly) and non-spiny species. Early authors distin-
guished all spiny solanums as members of Solanum subgenus
Leptostemonum Bitter (Dunal 1852; Bitter 1911) due to their
tapered anthers, prickles, and pubescence of stellate trichomes.
Phylogenetic studies using DNA sequences confirmed the
monophyly of the spiny solanums with stellate pubescence
(Solanum subgenus Leptostemonum or the Leptostemonum
clade), but the species with prickles and lacking stellate
pubescence did not resolve as members of this strongly
supported group (Bohs 2005; Levin et al. 2006; Weese and
Bohs 2007).
Solanum section Aculeigerum Seithe was broadly circum-
scribed by Seithe (1962) and included all Solanum species
with prickles but lacking stellate trichomes. Child (1983) seg-
regated Solanum section Nemorense A. Child from Seithe’s
Solanum section Aculeigerum andincludedtwoofher
Aculeigerum species (S. barbeyanum Huber and S. hoehnei C. V.
Morton; see Table 1) within it along with S. coriaceum Dunal,
S. nemorense Dunal, and S. reptans Bunbury. Child defined
Solanum section Nemorense as those species with difoliate
sympodial units, scorpoid cyme inflorescences, and deeply
stellate corollas up to 2 cm in diameter; he differentiated
Solanum section Aculeigerum by its plurifoliate sympodial
units, paniculate inflorescences, and rotate to stellate corollas
up to 4 cm in diameter. Child did not consider Solanum sec-
tion Nemorense to be part of Solanum subgenus Leptostemonum,
but was otherwise vague in assigning it to be closely related
or affiliated with any specific section other than his large
Solanum subgenus Solanum; he later included both Solanum
sections Nemorense and Aculeigerum in his Solanum subgenus
Potatoe (G. Don) D’Arcy as “Anomalous prickly taxa”(Child
1990). Whalen (1984) recognized Solanum sections Nemorense
and Aculeigerum as distinct groups, the Solanum nemorense
and Solanum wendlandii species groups respectively. In con-
trast to Child (1983), Whalen (1984) included both these
groups in Solanum subgenus Leptostemonum, but as a “phy-
logenetically isolated line”(Whalen 1984: 210). Nee (1999)
treated these species with prickles but no stellate trichomes
in Solanum subgenus Leptostemonum,withSolanum section
1102
Aculeigerum as “Series 2”of Solanum section Herposolanum
Bitter and S. nemorense Dunal + S. barbeyanum Huber as
“Subsect. 2”of Solanum section Micracantha Dunal. He
suggested the two groups were closely related, and that
the lack of stellate pubescence was the result of reduction of
stellate trichome complexity. See Table 1 for previous infra-
generic classifications of all species treated as related to
the taxa included in our current circumscription of Solanum
section Aculeigerum.
Phylogenetic studies using DNA sequence data have shown
that both Solanum sections Aculeigerum and Nemorense do not
belong to the strongly supported monophyletic Leptostemonum
clade (Solanum subgenus Leptostemonum; Bohs 2005; Weese
and Bohs 2007; Levin et al. 2006; Stern et al. 2011; Särkinen
et al. 2013). Three species of Solanum section Aculeigerum
have been included in molecular phylogenetic analyses
(S. alternatopinnatum,S. bicorne,andS. wendlandii;notethat
S. bicorne is determined as S. refractum in some of these
publications, but all sequence data included in GenBank to
date under S. bicorne and S. refractum are from Iltis et al.
29694,identifiedhereasS. bicorne). In all of these ana-
lyses, the species of Solanum section Aculeigerum form a
strongly supported clade, distinct from Solanum section
Nemorense.WithinSolanum section Aculeigerum Stern et al.
(2011) found S. alternatopinnatum resolved as sister to S. bicorne
(as S. refractum)+S. wendlandii. In the trees resulting from
some phylogenetic analyses S. allophyllum (Miers) Standl. is
sister to Solanum section Aculeigerum, but this relationship
Table 1. Taxonomic position of species of Solanum section Aculeigerum (and other species previously considered related to them). Species treated in
this revision are in bold face type.
Species Seithe 1962
Child 1983, 1990; Child
and Lester 2001 Whalen 1984 Nee 1999 Stern et al. 2011
S. alternatopinnatum
Steud.
S. sect. Aculeigerum
Seithe (as S. juciri
Mart. ex Sendtn.)
S. sect. Aculeigerum
Seithe (as S. juciri
Mart. ex Sendtn.)
Solanum
wendlandii
group
S. sect.
Herposolanum
Bitter (series 2)
S. wendlandii
Group
S. barbeyanum Huber S. sect. Aculeigerum
Seithe (as
S. megistophyllidium
Bitter)
S. sect. Nemorense
A. Child
Solanum
nemorense
group (as
S. viridipes
Dunal)
S. sect.
Micracantha
Dunal
(subsection 2)
–
S. bicorne Dunal –S. sect. Aculeigerum
Seithe (as
S. refractum Hook.)
Solanum
wendlandii
group (as
S. refractum
Hook. & Arn.)
–S. wendlandii
Group (as
S. refractum
Hook. & Arn.)
S. cobanense
J. L. Gentry
–S. sect. Aculeigerum
Seithe (as
S. refractum Hook.)
Solanum
wendlandii
group
––
S. coriaceum Dunal –S. sect. Nemorense
A. Child
–S. sect.
Micracantha
Dunal
(subsection 1,
series 4)
Androceras/
Crinitum
Clade
S. glaucescens Zucc. S. sect. Aculeigerum
Seithe
S. sect. Aculeigerum
Seithe
Solanum
wendlandii
group
S. sect.
Herposolanum
Bitter (series 2)
–
S. hoehnei
C. V. Morton
S. sect. Aculeigerum
Seithe (as S. alatum
Dunal)
S. sect. Nemorense
A.Child(Child1983);
section Herposolanum
Bitter (Child 1990;
Child and Lester 2001)
Solanum
nemorense
group
S. sect.
Herposolanum
Bitter (series 1)
S. nemorense
Group
S. nemorense Dunal –S. sect. Nemorense
A. Child
Solanum
nemorense
group
S. sect.
Micracantha
Dunal
(subsection 2)
S. nemorense
Group
S. pachyandrum
Bitter
–S. sect. Aculeigerum
Seithe
Solanum
wendlandii
group
S. sect.
Herposolanum
Bitter (series 2)
–
S. refractum
Hook. & Arn.
–S. sect. Aculeigerum
Seithe
Solanum
wendlandii
group
S. sect.
Herposolanum
Bitter (series 2)
[as S. aculeolatum
M. Martens &
Galeotti]
–
S. reptans Bunbury S. sect. Herposolanum
Bitter
S. sect. Nemorense
A. Child
Solanum
nemorense
group
S. sect. Herposolanum
Bitter (series 1)
–
S. triunfense
S. Knapp
–– –––
S. wendlandii
Hook. f.
S. sect. Aculeigerum
Seithe
S. sect. Aculeigerum
Seithe
Solanum
wendlandii
group
S. sect.
Herposolanum
Bitter (series 2)
S. wendlandii
Group
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1103
is poorly supported and unexpected on the basis of mor-
phology. Resolution and support is poor in the tree backbone
so that relationships among Solanum sections Aculeigerum and
Nemorense and the Geminata, Brevantherum, Cyphomandra,
and Leptostemonum clades are unclear (see Fig. 1 in Särkinen
et al. 2013), but no analysis places Solanum section Aculeigerum
within the well-supported Leptostemonum clade. Prickles are
only found in Solanum sections Aculeigerum,Nemorense,and
the Leptostemonum clade, while stellate hairs are found in
the Leptostemonum and Brevantherum clades (see Stern et al.
2013). Thus, both prickles and stellate hairs are of ambiguous
origin, depending on the sister relationships of the various
groups involved. Further analyses of Solanum using more
markers will be helpful in resolving this. It is clear, however,
that the species of Solanum section Aculeigerum are not part of
the core spiny solanums and their relationship to the species
of Solanum section Nemorense is not well-resolved.
Materials and Methods
The taxonomic conclusions presented here are a result of field and
herbarium work and are supported by molecular phylogenetic results
(Weese and Bohs 2007; Stern et al. 2011). We examined approximately
1,200 specimens representing more than 700 collections from the follow-
ing herbaria (herbarium acronyms follow Index Herbariorum, found
online at http://sciweb.nybg.org/science2/IndexHerbariorum.asp): AD,
BH, BHCB, BM, BR, C, CAS, CEPEC, CIQ, CM, CORD, CR, CUZ, DS,
DUKE, E, G, GH, INB, JBSD, K, L, LE, M, MCNS, MEDEL, MEXU, MJG,
MO, NY, P, PH, PMA, R, RB, SI, SP, TEX, U, UC, UPR, US, USM, WIS,
XAL, and W. Images of type specimens from the web resource Global
Plants (http://plants.jstor.org) were also used to establish the distribu-
tion of isotypes and to aid in lectotypification choices. Type specimens
with sheet numbers are cited with the herbarium acronym followed by a
dash and the sheet number (i.e. MO–1781232); barcodes are written as a
continuous string (i.e. G00104280). We have cited geographically repre-
sentative specimens for taxa where more than 100 collections are known.
Identities of all numbered collections seen for this study are in the Index
to numbered collections (Exsiccatae; http://dx.doi.org/10.5519/0039503)
and full specimen details are available on the Solanaceae Source website
(www.solanaceaesource.org).
Citation of literature follows BPH-2 (Bridson 2004) with alterations as
implemented in IPNI (International Plant Names Index, http://www
.ipni.org) and Harvard University Index of Botanical Publications (http://
kiki.huh.harvard.edu/databases/publication_index.html). Following
Knapp (2013) we have used the square bracket convention for publica-
tions in which a species is described by one author in a publication edited
or compiled by another, the traditional “in”attributions such as Dunal in
DC. for those taxa described by Dunal in Candolle’sProdromus Systematis
Naturalis Regni Vegetabilis.ThisworkiscitedhereasProdr.[A.P.de
Candolle] and the names are thus attributed only to Dunal. For “ex”
attributions we cite only the publishing author, as suggested in the Code
(McNeill et al. 2012). Standard forms of author names are according to
IPNI (International Plant Names Index, http://www.ipni.org).
We cite page numbers for all previous lectotypifications. Where specific
herbaria have not been cited in protologues we have followed McNeill
(2014) and designated lectotypes rather than assuming holotypes exist.
We have followed the morphological species concept or “morpho-
logical cluster”concept (Mallet 1995) in delimiting species of Solanum
section Aculeigerum. 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. Molecular evidence corroborates the delimitation
of taxa using morphological methods (Stern et al. 2011).
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 collections; in these
cases we retroactively georeferenced the collections using available locality
data. Maps were constructed with the points in the centers of degree
squares in a 1° square grid. Conservation threat status was assessed fol-
lowing the IUCN red list categories and criteria (IUCN 2014) using the
GIS-based method of Moat (2007) as implemented in the online assess-
ment tools in GeoCat (http://geocat.kew.org). The extent of occurrence
(EOO) measures the range of the species, and the area of occupancy
(AOO) represents the number of occupied points within that range based
on the default grid size of 2 km
2
.
Results
Morphology—Habit and Stems—Members of Solanum
section Aculeigerum are all woody or semi-woody vines with
small, broad-based recurved prickles (Figs. 1A, 1B, 1D, 2C).
These prickles are morphologically similar to those of the
S. lanceifolium species group of Whalen (1984) (Micracantha
clade of Stern et al. 2011) and almost certainly help the
plants to clamber over and ascend vegetation. In general the
prickles of Solanum section Aculeigerum are relatively widely
spaced along the stems, averaging 1–5 prickles per cm of
stem length, and are sometimes absent from the uppermost
stems. Prickles in the spiny solanums are interpreted as being
derived from stellate trichomes (Whalen 1984), and so their
occurrence in members of Solanum section Aculeigerum is
either convergent and follows adifferentdevelopmental
pathway, or stellate trichomes have been completely lost in
Solanum section Aculeigerum.
Most species in Solanum section Aculeigerum can begin to
flower when quite small and then they are recorded as shrubs
on herbarium labels, but all species apparently grow to
canopy height with long woody stems.
Leaves—Leaves of members of Solanum section Aculeigerum
vary from simple to pinnatifid, often on the same plant
(e.g. S. wendlandii). The only species with what could be
termed pinnately compound leaves with petiolulate leaflets
is S. alternatopinnatum; others are merely very deeply pin-
natifid with laminar tissue along the midrib. In common
with many members of Solanum subgenus Leptostemonum,
prickles can be borne on the midrib abaxially and occasion-
ally also adaxially. The leaves of most of the species are gla-
brous, or occasionally with a few papillate trichomes when
young; only S. bicorne and S. triunfense have multicellular
trichomes on the leaf lamina. The leaves of members of
Solanum section Aculeigerum often have copious crystal
sand –isolated idioblasts of calcium oxalate that appear as
white dots in dry specimens.
Pubescence—Despite bearing prickles, members of sec-
tion Aculeigerum do not possess the distinctive stellate tri-
chomes of Solanum subgenus Leptostemonum.Trichomes,if
present, are generally simple and uniseriate, usually com-
posed of only a few cells. Solanum triunfense has a few
furcate trichomes mixed with the simple uniseriate pubes-
cence of young stems and inflorescences; these hairs have
a single small branch and are not dendritic as found else-
where in the genus (e.g. S. aureum Dunal, a member of the
Dulcamaroid clade, see Knapp 2013). It has been suggested
(Nee 1999) that the simple trichomes of Solanum section
Aculeigerum are similar to those found in members of Solanum
section Acanthophora Dunal or S. leucopogon Huber (of the
Micracantha clade sensu Stern et al. 2011) and are simply
stellate trichomes that have lost the lateral rays. This loss
occurs in Solanum section Gonatotrichum Bitter (Stern et al.
2013) in which S. lignescens Fernald is stellate-pubescent
and other members of the section have distinctive genicu-
late trichomes with one short and one long cell. The few-
celled trichomes of S. bicorne are similar to those of Solanum
1104 SYSTEMATIC BOTANY [Volume 40
Fig. 1. Morphology of Solanum section Aculeigerum.A.S. alternatopinnatum inflorescence in bud (Giacomin et al. 1692). B. S. alternatopinnatum imma-
ture fruits; note the armed inflorescence rachis (Giacomin et al. 1692). C. Flower of S. cobanense; note fleshy petals and ellipsoid anthers (Christenhusz
et al. 5584). D. S. cobanense inflorescence on young shoot (Christenhusz et al. 5584), note absence of prickles on the inflorescence rachis. Credits: A–B: L. L.
Giacomin; C–D: K. Watson).
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1105
Fig. 2. Morphology of Solanum section Aculeigerum.A.S. pachyandrum flower; note ellipsoid anthers (Särkinen et al. 4546). B. S. pachyandrum imma-
ture fruit (Särkinen et al. 4546). C. S. pachyandrum inflorescence with buds; note the keeled calyx lobes and prickly pedicels (Särkinen et al. 4546). D. Inflo-
rescence of S. wendlandii pistillate plant (Christenhusz et al. 5372). E. Immature fruit (seeds still very small and not at all well-developed) of S. wendlandii
(Christenhusz et al. 5372). F. S. wendlandii pistillate flower (Christenhusz et al. 5372). Credits: A–C: T. Särkinen; D–F: M. Vorontsova).
1106 SYSTEMATIC BOTANY [Volume 40
section Gonatotrichum, with a long apical cell and a shorter
(pustulate) basal cell (but are not geniculate). Those of
S. triunfense are not of this morphology; they are typical
simple, multicellular uniseriate trichomes like those found
in major groups of non-spiny solanums (e.g. the Geminata
Clade, see Knapp 2008a).
Inflorescences—Members of Solanum section Aculeigerum
usually have branched inflorescences; those of S. wendlandii
can be many times branched and very large (Fig. 2D). Inflo-
rescence branches are highly divaricate, usually spreading
at ca. 90 degree angles so the structure is very open. Flow-
ers are often clustered near the tips of the branches, but
are usually evenly spaced along the inflorescence branches
(0.5–1 cm apart). The rachis can be armed or unarmed; all of
the pinnatifid-leaved species (S. alternatopinnatum,S. pachyandrum,
S. refractum) have densely prickly inflorescences (Fig. 1B, 2C),
while the rest of the taxa (Fig. 1D) only rarely have inflores-
cence prickles (S. wendlandii is variable in this regard).
Flowers—Flowers of members of Solanum section Aculeigerum
are pentamerous and sympetalous, like all of Solanum.Flower
buds are ellipsoid to tapering (turbinate), depending on the
shape of the anthers (see below; Fig. 1A, C). The calyx is usu-
ally campanulate and openly spreading and the buds are usu-
ally exserted from the calyx tube when very young, long
before anthesis. Most species have thickened lobes with scari-
ous sinuses that tear early on in bud growth to leave broadly
deltate regular calyx lobes. Solanum triunfense,however,has
a calyx that completely encloses the buds until much later
in growth and irregularly tears into deep uneven lobes. The
calyx lobes of S. pachyandrum are markedly keeled (Fig. 2C).
All species have copious crystal sand in the more robust
tissue of the calyx lobes.
Members of Solanum section Aculeigerum have relatively
large flowers; in S. wendlandii they can reach 6 cm in diameter
(long-styled flowers, e.g. Christenhusz et al. 5372,seeFig.18).
Corolla shape varies from deeply stellate (S. cobanense;
Fig. 1C, D) to almost rotate (S. wendlandii;Fig.2D,F),and
color varies from deep purple (S. cobanense, Fig. 1C) to
white or greenish white (S. alternatopinnatum,Fig.1A).Spe-
cies that commonly have white or greenish white flowers
(e.g. S. alternatopinatum,S. bicorne,S. refractum) have also been
recorded as having purple or lilac corollas, so it clear that poly-
morphism exists in corolla color as is common in other groups
of solanums (e.g. Knapp 2013). Corolla color in S. wendlandii
changes throughout anthesis (Fig. 2D); when flowers first
open they are deep purple and with age they fade to pale
violet (sometimes described on labels as blue) or white
with a concomitant increase in size (SK, pers. obs.); this
has been observed in other Solanaceae such as Lycianthes
(Dean 2001), in some members of the Dulcamaroid clade
of Solanum (Knapp 2013), and in Solanum section Crinitum
(Whalen) A. Child in the spiny solanums (SK, pers. obs.).
Androecium—Like all species of Solanum,membersof
Solanum section Aculeigerum have poricidal anthers; the pores
are directed distally and in these species usually split with
age along the margins so that the anthers appear to be lon-
gitudinally dehiscent. Solanum cobanense,S. pachyandrum and
S. triunfense have ellipsoid anthers (Figs. 1C, 2A), while the
rest of the species have variously tapering anthers, from
strongly tapering in S. refractum to only slightly tapering in
S. wendlandii (Fig. 2F).
Many species of Solanum have dimorphic stamens (Anderson
et al. 2006; Bohs et al. 2007; Vorontsova et al. 2013) and stamen
dimorphism occurs in almost half of the 13 major clades
of the genus. It is especially diverse in Solanum subgenus
Leptostemonum,withbothanther(e.g.S. rostratum Dunal
from the southwestern USA) and filament (e.g. S. coagulans
Forssk. from Africa) dimorphism present. The dimorphic
stamens of Solanum section Aculeigerum are due to differences
in filament length. Four species of the group (S. alternatopinnatum,
S. bicorne,S. glaucescens,andS. wendlandii) have one anther on
a markedly elongate filament, while S. cobanense,S. pachyandrum,
S. refractum and S. triunfense have filaments of equal size. Occa-
sionally the anther borne on the long filament is slightly larger
than the rest, but all anthers are usually of equal sizes, as are
the remaining four filaments in any given flower.
Anther color in the group varies from cream to yellow or
orange (S. refractum) to pale purplish blue (S. wendlandii).
Label data often describes the anthers of S. wendlandii as
yellowish at the base and blue or purple more distally (see
Fig. 2F).
Gynoecium—The ovary and style of members of Solanum
section Aculeigerum are glabrous, and the stigma is minutely
papillate. Style length varies among flowers in an inflores-
cence, and section Aculeigerum has been interpreted as com-
prising andromonoecious plants (Whalen 1984). Short-styled
flowers have styles that are included in the anther tube,
while those of long-styled flowers are exserted (Fig. 2F). In
S. cobanense, however, the stigma is held at the level of the
tips of the anthers (Fig. 1C), suggesting that the expression
of andromonoecy in this group may be more complex than
previously described. Solanum wendlandii appears to have
all long-styled or all short-styled flowers on any given plant;
it may be dioecious, a breeding system that occurs sev-
eral times in different clades of Solanum (Knapp et al. 1998;
Martine et al. 2009), or sequentially monoecious with younger
plants producing flowers with exclusively male function
(see description of S. wendlandii). This needs further investi-
gation of living plants in the field and greenhouse.
Fruits and Seeds—Fruits of members of Solanum section
Aculeigerum are bilocular berries with fleshy pulp (eaten by
local people in some species; e.g. S. pachyandrum) and thick,
leathery pericarp. The fruit surface appears rugose in dried
specimens, but is smooth, glabrous and matte in fresh mate-
rial (Fig. 2E). Immature fruits are often pale green or greenish
white with darker markings (see Figs. 1B, 2B); mature fruit
colour is quite variable. The pedicel and calyx lobes are thick-
ened and woody in fruit, and the calyx lobes often break
off in herbarium specimens. Fruiting specimens are rarely
collected. Seeds are flattened-reniform, very large and many
(usually > 100) per berry. The surfaces are minutely pitted with
rectangular or pentagonal cells, and at least in S. glaucescens,
the lateral testal cells walls are elongate, giving the seed a
hairy appearance.
Taxonomic Treatment
Solanum section Aculeigerum Seithe, Bot. Jahrb. Syst. 81:
291. 1962.—TYPE SPECIES: S. wendlandii Hook. f.
Solanum section Micracantha Dunal subsection Juciri Bitter
ex Marzell, in Hegi, Flora von Mittel-Europa 5: 2548.
1927.—TYPE SPECIES: S. wendlandii Hook.f.
Perennial woody vines to canopy lianas, usually armed
with recurved prickles on stems, petioles, and midveins of
abaxial leaf surfaces. Stems subwoody to woody, sometimes
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1107
herbaceous, usually glabrous or puberulent with simple hairs,
the prickles to 8 mm long, recurved, stout. Sympodial units
usually plurifoliate, rarely difoliate, not geminate. Leaves
simple and entire or pinnately lobed to compound, usually
petiolate, occasionally decurrent to base. Inflorescence terminal,
often appearing lateral, usually branched, with 5–35 flowers,
the axes with or without prickles. Flowers 5-merous, usually
long- or short-styled, the plants probably andromonoecious
or in one case (S. wendlandii) dioecious. Calyx spreading with
rounded lobes, the lobes usually equal in size but occasion-
ally tearing and markedly unequal in size. Corolla rotate-
pentagonal to stellate, purple, violet (blue), white, or greenish
white. Stamens all equal or dimorphic with four filaments
equal in length and one filament 2–6 times longer than the
rest; anthers variously tapering, usually somewhat attenuate,
ellipsoid in some species. Style cylindrical, glabrous, usually
straight but sometimes slightly curved; stigma truncate or
bilobed. Fruit a globose berry, ca. 2–10(–20 fide Christenhusz
et al. 5372) cm in diameter, the surface glabrous (sometimes
rugose when dry), yellow or light green, less commonly red
or orange, the mesocarp orange-red, green, or white. Seeds
many per berry, strongly flattened-reniform, light brown, the
surface minutely pitted, the testal cells pentagonal to rectan-
gular in outline, the lateral testal cell walls elongate or not.
Solanum section Aculeigerum includes eight species: six
in Mexico and Central America, one in southern Ecuador
and northern Peru, and one in southeastern Brazil, northern
Argentina and Paraguay (see maps of individual species).
Section Aculeigerum is differentiated from other clades of
Solanum by a combination of plurifoliate sympodial units,
branched inflorescences, prickles with an accompanying lack
of stellate trichomes, and a woody vine-like habit.
Section Aculeigerum is best known for the commonly culti-
vated species S. wendlandii (Figs. 2D–F). It is grown in the
Americas and has also been introduced to tropical and sub-
tropical regions of the Old World. It is also widely cultivated
in greenhouses in the northern hemisphere (see description
of S. wendlandii for more details).
Key to the Species of SOLANUM Section ACULEIGERUM
1. Leaves simple, usually shallowly lobed, only deeply lobed on vigorous young shoots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Stem prickles all or mostly paired at base of petioles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. S. glaucescens
2. Stem prickles more or less evenly scattered, not paired at base of petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaves chartaceous, mostly clustered on short shoots, pubescent with stiff simple trichomes with long basal cells;
corolla creamy white (occasionally tinged with violet). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. S. bicorne
3. Leaves coriaceous, evenly distributed along the branches, glabrous or pubescent with simple uniseriate trichomes
with 2–4 cells, the basal cell not markedly longer than the rest; corolla purple, blue or violet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Flowers rotate to pentagonal; filaments unequal; usually some pinnatifid leaves present on plant . . . . . . . . . . . . . . . . . 8. S. wendlandii
4. Flowers stellate; filaments equal; leaves never pinnatifid.
5. Leaves and stems glabrous; leaf bases truncate to acute; inflorescence axes glabrous; buds exserted
from the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. S. cobanense
5. Leaves and stems with scattered simple uniseriate trichomes; leaf bases attenuate; inflorescence axes
densely pubescent with simple 2–4-celled uniseriate trichomes; calyx completely enclosing the buds . . . . . . . . . . 7. S. triunfense
1. Leaves compound or deeply lobed with the lobe sinuses more than half the distance to the midrib, the uppermost
leaves near the inflorescence sometimes only shallowly lobed or entire. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Flowers heterandrous, with one filament markedly longer than the other four.
7. Corolla light blue to purple, rotate to pentagonal; leaves highly variable in shape on the same plant,
often entire near the inflorescence, generally ranging from deeply lobed to pinnately compound. . . . . . . . . . . . . . . . . . . . 8. S. wendlandii
7. Corolla white or cream, stellate, lobed nearly to the base; leaves compound or deeply pinnatifid.
8. Inflorescence axis and pedicels prickly; inflorescence borne on main branches, not borne on short shoots;
leaves compound with the leaflets with distinct petiolules; southern South America . . . . . . . . . . . . . . . . . . . . . 1. S. alternatopinnatum
8. Inflorescence axis unarmed; inflorescence borne or short shoots; leaves deeply pinnatifid, the lobes with
distinct wings of tissue decurrent along the main rachis, distinct petiolules absent; Mexico
and Central America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. S. bicorne
6. Flowers with all filaments equal in length within a flower.
7. Anthers ca. 3 mm wide; flowers white; pedicels armed with stout prickles; southwestern Ecuador
and northwestern Peru. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. S. pachyandrum
7. Anthers 1–1.5 mm wide; flowers greenish white; pedicels unarmed; Pacific slope, southern Mexico, Guatemala
and Honduras. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. S. refractum
1. Solanum alternatopinnatum Steud., Nomencl. Bot.,
ed. 2: 600. 1841. Nom. nov. for Solanum oleraceum Vell.
(1829), non Solanum oleraceum Dunal (1813). Solanum
oleraceum Vell., Fl. Flumin. 89. 1829. —TYPE: BRAZIL.
Rio de Janeiro (no specimens are known to exist; lecto-
type, designated by Knapp et al. 2015, pg. 832: original
parchment plate held in Biblioteca Nacional. Rio de
Janeiro [mss1198651_128]; incorrectly lectotypified by
Matesevach Becerra 2013, pg. 290 on the published
plate in Vellozo, Fl. Flumin. Icon. 2: Table 125. 1831).
Solanum viridipes Dunal, Prodr. [A. P. de Candolle] 13(1):
234. 1852 —TYPE: BRAZIL: Bahia, J. Guillot s. n. (holo-
type: P [P00384877, F neg. 39219]).
Solanum juciri Sendtn. var. inermis Glaz., Bull. Soc. Bot. France
58 (Mem. 3f): 497. 1911. —TYPE: BRAZIL. Rio de Janeiro:
Fazenda de Boa Esperança, Chez M. Bellieni, dans les
cafféiers, 22 Oct 1886, A. F. M. Glaziou 16294 (lecto-
type, designated here: P [P00319401]; isolectotypes: C,
K [K000788574]).
Solanum juciri Sendtn. forma paraguariense Hassl., Repert.
Spec. Nov. Regni Veg. 9: 118. 1911. Solanum viridipes
Dunal var. intermedium Hassl., Repert. Spec. Nov. Regni
Veg. 15: 239. 1918. —TYPE: PARAGUAY. Caaguazú [Depto.
Caaguazú], Amambay [Depto. Amambay] and Paraná
regions, Mar 1905, É. Hassler 9365 (lectotype, designated
by Matesevach Becerra 2013, pg. 290: BM [BM000087604];
1108 SYSTEMATIC BOTANY [Volume 40
isolectotypes: G [G00443009, Morton neg. 8558; G00443010],
GH [no barcode or sheet number], K [K000788653], NY
[NY00172245], P [P000384878, Morton neg. 8362], W
[W-1906–0001280]).
Woody or subwoody vines to 3 + m long or scandent
shrubs, the stems 0.2–0.4 cm in diameter, prickly; inter-
nodes 1–5 cm long, glabrescent; prickles to 2 mm long,
stout, recurved. Sympodial units usually plurifoliate, rarely
difoliate, not geminate. Leaves pinnately compound, the
blades in outline (5–)15–20 × (4–)6–12 cm, with 5–7pairsof
opposite to subopposite petiolulate leaflets, chartaceous,
glabrous adaxially and abaxially, with calcium oxalate crys-
tals commonly present, often armed on rachis with recurved
prickles; major lateral veins 5–7oneachside,corresponding
to the leaflets; petioles (1–)3–5 cm, glabrous, often armed
with recurved prickles; leaflets 3–10 × 1–3cm,oblongto
elliptic, widest at the base; base cuneate and oblique; margin
entire; apex acuminate; petiolules 0.3–1 cm. Inflorescences
(10–)20–30 cm long and wide, terminal or appearing lateral,
several times branched, with 20–35 flowers, the axes glabrous,
densely prickly; peduncle (2–)5–15 cm; rachis 4–10 cm; pedi-
cels 10–15 mm, slender at anthesis, articulated at the base;
fruiting pedicels to ca. 20 × 3 mm and becoming subwoody.
Flower buds ellipsoid to somewhat pointed, the corolla soon
exserted from the calyx tube. Calyx campanulate, spreading,
the tube 2–4 mm, the lobes ca. 3 × 2 mm, rounded, apiculate
at tips, glabrous, reflexed at anthesis. Corolla 2.5–4cmin
diameter, spreading at anthesis, uniformly white or greenish
white, sometimes tinged with violet (Nee 54960, plant culti-
vated in Bolivia), membranous to slightly fleshy, stellate,
lobed ca. 3/4 way to the base, the lobes 1–2×0.3–0.5 cm,
narrowly triangular, acute to acuminate at apices, glabrous
abaxially and adaxially, the tips densely papillate. Stamens
5–10 mm long; filaments unequal, with four 1–1.5 mm long,
the fifth 3–4 mm long, glabrous; anthers nearly equal, 5–7×
1.5–2 mm, the one on the longer filament slightly larger,
tapering at the tips, connivent, yellow or orange, poricidal
at the tips, the pores lengthening to slits with age. Ovary
glabrous; style in long-styled flowers ca. 10 mm long,
straight, glabrous, in short-styled flowers ca. 4 mm long;
stigma truncate. Fruit a globose berry, 3–4cmindiameter,
green with light green streaks when immature, yellow with
white streaks when ripe, the pericarp glabrous, rugose when
dry. Seeds > 20 per berry, ca. 3 × 3 mm, strongly flattened-
reniform, light brown, the surface foveolate. Figs. 1A–B, 3.
Distribution and Habitat—Solanum alternatopinnatum (Fig. 4)
occurs in Argentina (Province of Misiones), Brazil (States of
Bahia, Distrito Federal, Espírito Santo, Mato Grosso, Minas
Gerais, Paraná, Rio de Janeiro, São Paulo) and Paraguay
(Provinces of Alto Paraná, Canindeyú, San Pedro) in the
wet forests of “mata atlântica”or “Selva Paranäense”at
elevations from 100–950 m.
Phenology—Solanum alternatopinnatum is most commonly
collected in flower from November to April, and fruiting col-
lections are most common from June to August.
Common Names—Brazil: “juciri”(Martius s. n.); “jiquiri”
(Warming s. n.). Paraguay: “yuá-pará”(Woolston 826).
Preliminary Conservation Status (IUCN 2014)—Least
Concern (LC); EOO 2,998,232 km
2
(LC); AOO 240 km
2
(EN);
75 localities. Although the AOO suggests that S. alternatopinnatum
is of some conservation concern, the large number of localities
and wide habitat preferences of this species lead us to give
it the assessment of LC.
Solanum alternatopinnatum differs from other species of
Solanum section Aculeigerum in its compound leaves with
clearly petiolulate leaflets, heterandrous flowers, and nearly
herbaceous stems. Solanum pachyandrum and S. refractum have
deeply pinnatifid leaves but do not have clearly petiolulate
leaflets and S. refractum occasionally has leaves that are very
shallowly lobed. We have only seen a single specimen of
S. alternatopinnatum (Glaziou 16294,oneofthetypecollec-
tions of S. juciri var. inermis) with simple leaves. The stems
of S. alternatopinnatum are herbaceous or rarely subwoody
in contrast to the subwoody stems of S. refractum.These
species are also completely allopatric, with S. refractum
restricted to southern Mexico, S. pachyandrum found in the
Huancabamba depression of northern Peru and southern
Ecuador and S. alternatopinnatum ranging from Brazil and
Paraguay to northern Argentina.
Leaves of S. alternatopinnatum are eaten as a vegetable in
Brazil (as jiciri or juciri; Mentz and Oliveira 2004); appar-
ently the prickles soften with cooking.
This species was known for a long time as S. juciri Sendtn.,
a name coined as a replacement for S. oleraceum Vell. (Vellozo
1829), a later homonym of S. oleraceum Dunal (Dunal 1813;
asynonymofS. americanum Mill. of the Morelloid clade).
Solanum juciri is an illegitimate name (McNeill et al. 2012) as
S. alternatopinnatum was cited in synonymy, with the com-
ment “nomen ineptissimum plantae ab auctore non visae
pinni tam alternantibus quam oppositis instructe datum”(this
is a silly name given to this plant by the author who did not
see alternate but opposite leaflets); Sendtner (1846) recognized
the existence of the name but considered it inappropriate.
In describing S. viridipes, Dunal (1852) cited two elements:
an herbarium specimen Guillot s.n. from Paris, and a living
plant “v.v. in hort. Monsp.”cultivated in Montpellier. Since
he only cited a single herbarium specimen, this is the holo-
type (McNeill 2014). He also further specified and annotated
material as “S. laxiflorum Dunal mss. in herb. Schad. nec
Sendtn.”The holotype specimen in P (P00384877) has a number
of annotations on it, including “S. laxiflorum Dun. 1842 nec
Sendtn. 1848.”We suspect “in herb. Schad.”is a misprint for
“in herb. Sched.”and refers to an annotation on a herbarium
specimen and not to a particular herbarium.
Names coined by Glaziou (1911) in his list of plants from
Central Brazil are sometimes indicated as not validly pub-
lished. Here we are treating the varietal name S. juciri var.
inermis as validly published as it is accompanied by a (very
short) description that distinguishes it from the species.
Glaziou (1911) cited only his collection number 16294 in
the protologue, but a compound locality (“Fazenda de Boa
Esperança, près Cantagallo et Serra da Estrella RIO-JAN”).
In Paris there are two sheets of this number that are clearly
collected in two places —one in Serra da Estrella (P00319400,
Feb 1880, Glaziou 16294a) and the other from Fazenda de Boa
Esperança (P00319401, 22 Oct 1886, Glaziou 16294). The ‘a’has
been added to the number in Glaziou’shandwriting.Glaziou
16294a has typical pinnately compound leaves, while the
other sheet has unusual simple leaves (see above). We select
P00319401 as the lectotype of this varietal name as there is
a clear duplicate at Kew as evidenced by the simple leaves
on the K sheet.
Hassler described S. juciri forma paraguariense using his
collection Hassler 9365 and seven years later used the same
collection (together with Fiebrig 6157) to describe S. viridipes
var. intermedium as a new name for his forma paraguariense
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1109
Fig. 3. Solanum alternatopinnatum from Paraguay (É. Hassler 9365, lectotype of Solanum juciri Sendtn. forma paraguariense Hassl., BM [incorrectly
labelled isolectotype in image]). Reproduced with permission of the Natural History Museum, London.
1110 SYSTEMATIC BOTANY [Volume 40
(Hassler 1911, 1918). He described var. intermedium in terms
of his forma paraguariense (“Caule foliis ut in Fedde Rep. l. c.
pro forma paraguariensi nob. indicatus”[stem and leaves from
Fedde Rep. description of the new forma paraguariense]:
Hassler 1918: 239). We consider that he was explicitly coin-
inganewnameusingthesamematerialtoreplaceforma
paraguariense, and therefore treat these two names as homo-
typic as did Matesevach Becerra (2013). It is unfortunate
that Matesevach Becerra (2013) did not use material at
G annotated by Hassler, but her lectotypification is correctly
made and stands.
Representative Specimens Examined —BRAZIL. Bahia: Mun. Jussari,
Palmira, entrada ca. 7.5 km de Jussari, Faz. Teimoso, RPPN Serra do
Teimoso, 21 Apr 1999 (fl), Amorim et al. 2915 (CEPEC, MO, NY); Juçarí,
27 May 1966 (fl), Belém & Pinheiro 2329 (US); Mun. Almadina, Faz. Beija-
Flora, ramal com entrado no km 3 da Rod. Almadina/Floresta Azul,
9 Nov 1978 (fl), dos Santos et al. 3382 (F); Rodovia BR-330, Barragem do
Funil (Rio de Contas), 33 km SE de Ipiau, 6 Mar 1978 (fl), Mori et al. 9549
(MO, NY, WIS). Distrito Federal: Côrrego Landim, ca. 20 km N of
Brasília, 950 m, 16 Dec 1965 (fl), Irwin et al. 11343 (AAU, K, MO, NY,
WIS); Côrrego Landim, ca. 25 km N of Brasília, 950 m, 16 Mar 1966 (fl),
Irwin et al. 13998 (AAU, DS, K, MO, NY, WIS). Espírito Santo: Mpio.
Itaguassú, Jatiboca, 28 May 1946 (fl), Brade et al. 18467 (K, RB);
Aldeiamento dos Indios, Rio Pancas, 5 Jul 1942 (fl), Bueno & Emygdio
208 (R); Fazenda Salvador, Rio Doce, 17 Jul 1942 (fl), Bueno & Emygdio
209 (R); Reserva Florestal de Linhares, 31 Oct 1995, Folli 2668 (RB).
Mato Grosso: Mun. Chapada dos Guimarães, rodovia Chapada dos
Guimarães-Campo Verde, km 10, 11 Aug 1997 (fl), Hatschbach et al. 66672
(BHCB); Vale do Caxipó, 28 May 1983 (fl), Guilherme 8 (R). Minas Gerais:
Parque Estadual do Rio Doce, Marliéria, 15 Jun 1995 (fl), Lombardi &
Temponi 784 (F); Mun. Belo Horizonte, Serra do Taquaril, 31 Apr 1933
(fl), Mello Barreto 7826 (F); Fazenda de Aguada, 19 Sep 1930 (fl), Mexia
5066a (US); Paraopeba, 31 Mar 1957 (fl), Pereira & Pabst 2731 (RB, US,
WIS); Mun. Juiz de Fora, Morro do Imperador, 8 May 2003 (fl, fr), Pifano
et al. 511 (BHCB); Belo Horizonte, Sep 1988 (fl), Requens s.n. (BHCB);
Caratinga, Aug 1991 (fl), Stehmann 19535 (RB); Mun. Caratinga, Fazenda
Macadonea/Cenibra-Ipaba, 22 Aug 1992 (fl), Stehmann & Soares s.n.
(BHCB). Paraná: Foz do Iguaçu, Parque Nacional, 20 Feb 1963 (fl),
Hatschbach 9749 (US);Mun.CerroAzul,Cab.Rib.doTigre,24April
1987 (fl) Hatschbach & Cervi 51229 (MO, US); Porto de Cima, 24 Jun
1914 (fl), Jönsson 585a (GH, NY); Jaguariahyva, 25 Oct 1910 (fl), Dusén
16559 (GH, NY). Rio de Janeiro: Tres Fomosos, Alampaio 3181A (US);
Fig. 4. Distribution of Solanum alternatopinnatum.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1111
Itaipuaçú, Pico Alto Moirão 14 Apr 1982 (fr), Andreata 413 (RB);
Mangaratiba, Reserva Ecológica do Rio das Pedras, trilha do Cambucá,
17 Aug 1996 (fr), Bovini et al. 1011 (RB); Guaratiba, May 1929, Brade
10516 (US); Paraíba do Sul, Mar 1954 (fl), Duarte & Ahumada 3691 (RB);
Barra da Tijuca, 15 Oct 1961 (fl), Duarte 9880 (RB); Corcovado, Rio de
Janeiro, 20 Apr 1922 (fl), Ducke & Kuhlmann 396 (RB); Rio de Janeiro,
1867 (fl), Glaziou 1333 (BR); mata do Horto Florestal, Rio de Janeiro,
14 May 1929 (fl), Kuhlmann 1117 (RB); Vista Chineza, 20 May 1958 (fl),
Liene et al. 3793 (RB); Morro do Pão de Açúcar, 17 Oct 1977 (fl), H. C.
de Lima 117 (RB); Serra dos Órgãos, 22 Oct (fl), Riedel s.n. (NY); Rio
Janeiro, May 1832 (fr), Riedel s.n. (NY); Niterói, Côrrego dos Colibris,
30 Jun 1998 (fl,fr), Santos et al. 168 (RB); Tejuco, Schott 5447 (F, W); Serra
do Mendanha, 450 m, 27 Nov 1969 (fl), Sucre et al. 6395 (RB); Rio
Janeiro, 18 Jan 1864 (fl), Warming s.n. (C). Santa Catarina: Tapora,
Tibeirão, 19 Jun 1968 (fl), Klein et al. 7961 (US). São Paulo: Serra do Mar
above Ubatuba, 100–200m,21Aug1976(fl),Davis et al. 59838 (E);
Barreiro, Expedicão do Rio Feio, Edwall 15378 (US); Mun. Iepe, Fazenda
C.A.P.I. 4 km N of Rio Paranapanema & 6 km E of Porto Alvorada,
22°44′S, 57°70′W, 9 Feb 1965 (fl), Eiten et al. 5956 (US); Ubatuba,
Parque Estadual da Serra do Mar, Núcleo Picinguaba, trilha saindo da
Base Cambucá para a sede da fazenda (Base de Pesquisa), 3 m, 23°19′39″S,
44°56′21″W 26 Jan 2012, Giacomin & Almeida 1692 (BHCB); Campinas,
8 Jan 1905 (fl), Heiner 377 (MO); Serra de Caraguatatuba, 15 May 1938 (fl),
Kuhlmann & Gehrt s.n. (F, NY, SP); Mun. Aguas de Prata, Rod. Gobernador
Adhemar P. Barros, ca. 1 km do centro Aguas de Prata, 21 Mar 1994 (fl),
Martins et al. 31435 (BHCB); Guaratinguetá, Martius s.n. (M); Ilha Seca,
1940 (fl), Santos s.n. (R); Mun. Campinas, Fazenda Santa Elisa, Feb 1996
(fl), Vasconcellos-Neto s.n. (BHCB); Reserva Municipal da Santa Genebra,
near Campinas, 750 m, 1 Feb–21 Mar 1987 (fl), Vogel 59 (MJG).
PARAGUAY: in regione fluminis Alto Paraná, 1909–1910 (fl), Fiebrig
6157 (BM, E, GH, K, L, US). Alto Paraná: Reserva Limoy, 10 Nov 1981
(fl), Binacional 896 (MO); Centro Forestal Alto Paraná, 12 km W del
Puerto Presidente Stroessner, 25°28’s, 54°42′W, 16 Apr 1986 (fl), Brunner
1816 (PY). Canindeyú: Jejui-mí, 23 km NE de Ygatimí, 24 Apr 1996 (fl),
Jiménez et al. 1212 (BM, MO, PY). Caazapá: National Park Caaguazú,
26°05′52″S, 55°26′35″W, 2 4 N o v 1 997 (fl ) , Zardini & Benitez 47379 (NY).
San Pedro: Alto Paraguay, Primavera, Rozados, 8 Jun 1957 (fl), Woolston
826 (C, K, NY, US); Yaguareté forest (Sustainable Forest Systems Site),
23°46′16″S, 55°59′37″W, 20 Jun 1995(fr), Zardini & Balbuena 42892 (NY).
ARGENTINA. Misiones: Dtto. Cainguás, Capiovysiño, 6 Apr 1949 (fl),
Schwarz 7674 (CORD); ruta 7, 5 km W de acceso a Aristóbulo del Valle,
30 Jul 1987 (fr), Vanni et al. 886 (GH).
2. Solanum bicorne Dunal, Prodr. [A. P. de Candolle] 13:
232. 1852.—TYPE: MEXICO, “Pavón s. n.”[a collec-
tion made by M. Sessé and J. Mociño and distributed
from Pavon’s herbarium, see Knapp 2008b] (holotype: G
[G000343458, F neg. 34109]).
Solanum bicorne Dunal var. angustifolium Dunal, Prodr. [A. P.
de Candolle] 13(1): 233. 1852.—TYPE: MEXICO, “Pavón
s. n.”[a collection made by M. Sessé and J. Mociño
and distributed from Pavon’s herbarium, see Knapp
2008b] (holotype: G [G000343452, F neg. 34110]; iso-
type: F [F-846743]).
Solanum dichotomum Sessé & Moc., Pl. Nov. Hisp. 35. 1888.—
TYPE: MEXICO. Guerrero: Mazatlán, M. Sessé & J. Mociño
5348 (lectotype, designated by Knapp 2008b, pg. 11: MA
[MA-604601, F neg. 48342]; isolectotypes: F [F-845066,
fragment], MA [MA-604599, F neg. 48341; MA-604600]).
Solanum guamuchilense Cast.-Campos, Acta Bot. Mex. 27: 34,
Fig. 1. 1994.—TYPE: MEXICO. Nayarit: Mpio. Bahía de
Banderas, El Guamúchil, between Bucerías and Sayulita,
ejido of Punta Mita, 300 m, 10 Oct 1990, G. Castillo
Campos 6103 (holotype: XAL [XAL0106656]; isotype:
XAL [XAL0106655]).
Woody vine to 15 + m long, the stems 0.25–0.9 cm in
diameter; internodes 5–10 cm long on main stems, 0.5–1cm
long on short shoots, glabrous or puberulent with simple
hairs to 0.5 mm long with enlarged pustulate bases; prickles
2–5 mm long, stout, recurved, occasionally very sparse on
long internodes. Sympodial units plurifoliate, not geminate,
the leaves clustered on short shoots to 5 cm long. Leaves
usually simple (one specimen seen with deeply pinnatifid
juvenile leaves on vigorous shoots), the blades 10–28 × 5–10 cm,
oblanceolate to lance-elliptic, chartaceous, puberulent adaxially
and abaxially with simple 2–4-celled hairs 1–1.5 mm long with
an elongate basal cell, often armed abaxially on midrib with
recurved prickles; major lateral veins 3–5oneachside;base
long-attenuate; margin usually entire, occasionally sinuate with
2–3 pairs of shallow lobes ca. 3 × 2.5 cm (leaves on very young
shoots possibly more deeply lobed); apex acute to acuminate;
petioles broadly winged to base and nearly sessile. Inflores-
cences ca. 30 × 20 cm, terminal or lateral on young growth
of short shoots, many times branched, with 20–55 flowers,
the axes glabrous or occasionally minutely puberulent with
simple 1–2-celled trichomes to 0.5 mm long, not prickly, or
sometimes with a few prickles at the base (Soto N. 10188);
peduncle 6–15 cm; rachis (5–)15–20 cm; pedicels 10–15 mm
and slender at anthesis, elongating in fruit to 2–2.5 × ca. 1 cm
and becoming subwoody, terminally clustered, articulated
at the base. Flower buds narrowly ellipsoid to pointed, the
corolla early exserted from the calyx tube. Calyx campanu-
late, spreading, the tube 2–4 mm, the lobes ca. 1.5 × 2 mm,
rounded, apiculate at tips, glabrous. Corolla 2.5–4cmin
diameter, spreading at anthesis, creamy white, membranous,
deeply stellate, lobed nearly to the base, the lobes 1–2×
0.3–0.5 cm, narrowly deltate to triangular, acute to acumi-
nate, glabrous abaxially and adaxially, densely short white
pubescent at the hooked cucullate tips. Stamens 5–10 mm
long; filaments unequal, with four ca. 1 mm long, and one
ca. 5–6 mm long (at approximately the same height as the
tips of the other four anthers), glabrous; anthers nearly
equal, 4–6 × 1.5–2 mm, the one on the longer filament slightly
larger, usually somewhat tapering, connivent, yellow or
orange, poricidal at the tips, the pores lengthening to slits
with age. Ovary glabrous; style in long-styled flowers ca.
10 mm long, approximately equal in height to the long
stamen, straight, glabrous, in short-styled flowers ca. 5 mm
long; stigma capitate. Fruit a globose berry, ca. 5.5 cm in
diameter, globose, red or orange, the pericarp glabrous,
rugose when dry. Seeds > 50 per berry, ca. 5 × 5.5 mm,
strongly flattened-reniform, brown, the surfaces minutely
pitted. Fig. 5.
Distribution and Habitat (Fig. 6) —Solanum bicorne is endemic
to Mexico (States of Colima, Durango, Guanajuato, Guerrero,
Jalisco, Michoacán, Morelos, Nayarit, Sinaloa, and Zacatecas),
in deciduous forests, grasslands, and humid montane forests
at elevations from sea level to 2,000 m.
Phenology—Solanum bicorne appears to flower and fruit
throughout the year, but more fruiting speciens have been
collected from November to April (e.g. Gentry 5248).
Uses—Mexico: “The fruits are red when ripe in March
and April and eaten by birds”(Gentry 5248).
Common Names—Mexico. Guerrero: “chilacayote”(Halbinger
s.n.); “tomate de chichalaca”(Halbinger s.n.); Sinaloa: “toronjo”
(Montes & Salazar 591 and Ortega 4021).
Preliminary Conservation Status (IUCN 2014)—Least
Concern (LC); EOO 545,195 km
2
(LC); AOO 404 km
2
(EN);
126 localities. Solanum bicorne is a commonly collected and
apparently relatively common species where it occurs.
This species is easily differentiated from others in section
Aculeigerum by the presence of clustered leaves on short shoots
1112 SYSTEMATIC BOTANY [Volume 40
Fig. 5. Representative specimen of Solanum bicorne from Mexico (C. Pringle 6396, BM). Reproduced with permission of the Natural History
Museum, London.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1113
(Fig. 5). Dunal, in describing Solanum bicorne, mistook the
clustered stem leaves as being the rosette of a possibly
acaulescent herb (Dunal 1852: 232) when in fact the plant
is a high-climbing, robust, woody vine. This species is simi-
lar to S. glaucescens, especially with respect to the white,
stellate flowers with dimorphic stamens, but differs by its
leaves that are narrowed to the base with definite wings on
the petiole and short shoots with densely crowded leaves.
A pair of prickles at the base of the petiole is present in
S. glaucescens butabsentinS. bicorne.
A single specimen identified here as S. bicorne from Sinaloa
(Sanders et al. 4576, MO-3279500) has quite deeply lobed
leaves that resemble those of S. refractum, but these seem to
be from a very young shoot and have fewer lobes than typical
specimens of S. refractum. The non-prickly inflorescence
rachis arising from a short shoot and distribution suggest
this specimen is S. bicorne with juvenile foliage. Other Solanum
species have pinnatifid juvenile foliage (e.g. members of the
Dulcamaroid clade such as S. dulcamaroides Poir., see Knapp
2013) that is rarely collected; this needs further investiga-
tion in the case of S. bicorne and other members of this
group of woody vines.
In his description of S. bicorne Dunal (1852) cited a speci-
men from “Nova Hispania (Pavon in hb. Boiss. sub nom.
S. bicorne)”; he used the name on the specimen that almost
certainly came from the original collection made in Mexico
by Martin Sessé and José Mociño. A single specimen from
the Boissier herbarium (now part of the general collections
at G) is therefore the holotype for this name. The complex
typification of names for this species based on the Mexican
and Central American collections of Martin Sessé and José
Mociño is discussed in detail in Knapp (2008b).
Representative specimens examined —MEXICO. Colima: 14 mi WNW
of Santiago, road to Cihuatlán, Jalisco, 5 m, 25 Jul 1957 (fl), McVaugh
15741 (MEXU); Mpio. Comala, Rancho El Jabalí, 22 km NNW of Colima,
19°27′N, 103°41′W, 1,325 m, 10 Jul 1991 (fl), Vázquez V. & Phillips 822
(K, MEXU). Durango: Tamazula [westernmost point, almost in Sinaloa],
1,000 ft, 16 Dec 1939 (fr), Gentry 5248 (DS, F, GH, MO, NY). Guanajuato:
Mpio. Acámbaro, 3 km NW de Irámuco, 1,950 m, 15 Aug 1993 (fl),
Fig. 6. Distribution of Solanum bicorne.
1114 SYSTEMATIC BOTANY [Volume 40
Rzedowski 52029 (MEXU). Guerrero: 36 km by road N of coast road
(Hwy 200) on road to Ciudad Altamirano, 600 m, 26 Sep 1983 (fl),
Anderson 12827 (CAS, MEXU, MO); Mpio. José Azueta, 1.8 km SO del
caserío La Vainilla, 17°42′N, 101°31′W, 350 m, 26 Oct 1989 (fl), Gallardo
Hernández et al. 162 (MEXU); Mochitlán al E de Chilpancingo, Aug 1962
(fl,fr), Halbinger s.n. (MEXU); Cruz de Ocote, adelante de Filo de Caballo,
1,800 m, Apr 1975 (fr), Halbinger s.n. (MEXU). Jalisco: 5–10 km SW of
Talpa de Allende on road toward Tomatlán, 23 Sep 1983 (fl, fr), Anderson
12773 (NY); La Manzanilla, 4.5 km E de la carretera Barra de Navidad–
Puerto Vallarta, 17 Jan 1986 (fr), Ayala & Bullock 531 (MEXU); 16 km N
de Garita, 8 Jul 1967 (fl), Cedillo Trigos 31 (MEXU); below SE edge of Cerro
Toxín in bottom of Arroyo Pitahayas, 1 km NE of Toxín, barely N of
Puerto de Toxín–La Loma crossroads, along El Sauz–San Pedro Toxín
road, 19°30′N, 104°00′W, 1,180 m, 13 Jan 1990 (fr), Cochrane et al. 12266
(WIS-2 sheets); Reserva Biósfera Sierra de Manantlán, E end of the Sierra
de Manantlán in the pass between Cerro Grande and Cerro en Medio,
midway between Puerto de Toxín and Piedras Cuatas, 19°33′N, 103°59′W,
1,260–1,290 m, 14 Jan 1990 (fr), Cochrane et al. 12279 (WIS); S of Sayola,
304–914 m, Sep 1961, Faberge s.n. (TEX); Mpio. Jocotepec, 5 km S de
Huejotitlán, entre Huejotitlán y San Marcos, 1,500 m, 15 Oct 1989 (fr),
Flores M. et al. 1718 (MEXU); Mpio. Zapotitlán, 82 km de Cd. Guzmán,
carr. a San Antonio (pasando por Quesería), a 8 km de San Antonio por
brecha a Zapotitlán, 12 Jun 1988, Gaona P. 418 (NY); 18–20 miles SW of
Autlán, 23 Jul 1951, Gentry 10953 (LL); west of Magdalena, 25 May
1849 (fl, fr), Gregg 880 (MO); Cerro Viejo, above Zapotitlán de Hidalgo,
about 25 mi S of Guadalajara, 1,800 m, 17 Jun 1956 (fl,fr), Gregory &
Eiten 229 (MO);Mpio.TalpadeAllende,15kmSdeTalpadeAllende,
1,300–1,400 m, 4 Oct 1982 (fr), Hernández M. et al. 9058 (MEXU, NY);
between Nacastillo and Chamela, 13 Jan 1979 (fr), Iltis & Nee 1567 (WIS);
1 km S of N end of Tecopatlán valley, 4.5 km SSE of El Chante on road
to Manantlán, 19°41′N, 104°11′W, 1,200 m, 11 Jan 1980 (fr), Iltis & Schatz
2548 (WIS-2 sheets); Reserva Biósfera Sierra de Manantlán, path to La
Lima from El Rodeo, 19°33′N, 104°03′W, 1,350 m, 15 Oct 1982 (fr), Iltis
et al. 28923 (WIS-2 sheets); Mpio. Venustiano Carranza, 4 km N de
V. Carranza, camino a Tapalpa, puente sobre el Río Jiquilpan, 30 Jun 1981
(fl), Lott et al. 428 (MEXU); Mpio. La Huerta, Rancho Cuixmala, Cumbres
1, E of Puerto Vallarta–Barra de Navidad hwy, 19°31′N, 104°56′W, 14 Jan
1991 (fr), Lott et al. 3261 (MEXU); Mpio. La Huerta, Estación Biología
“Chamela”,4Sep1982(fl),Magallanes 3674 (MEXU, NY); 11 mi N of
bridge of Río Cihuatlán on road from Santiago, Colima, to Durazno,
Jalisco, 500–550 m, 1 Aug 1957 (fl,fr), McVaugh 15968 (MEXU, US); hills
2–3milesEofLaManzanilla,settlementontheSEshoreofBahía
Tenacatita, 150–200 m, 11 Nov 1960 (fr), McVaugh 20967 (NY-2
sheets, US); Mpio. Zapopan, “La Piedrera”,10kmNWdeTesistán,
1,700 m, 9 Oct 1980 (fl), Nieves Hernández 53 (F); Barranca, near
Guadalajara, Jun 1886 (fl), Palmer 108 (BM, GH-2 sheets, NY-2
sheets, P, U); Tequila, 6 Jul 1893 (fl), Pringle 7918A (E,F,MO,NY;
as 2918A in F, US); Mpio. Cuautitlán, al NE de Ayotitlán, 950 m,
10 Aug 1986 (fl), RamírezD.&RodríguezC.430(WIS); Mpio. Tala, a lo
largo del Arroyo Caliente y Los Letreros, Bosque Escuela, La Primavera,
1,450 m, 13 Aug 1988 (fl), Rodríguez C. & Reynoso D. 1432 (MEXU, NY);
Mpio. Zapotitlán, La Joya area of Rancho El Jabali, 22 km NNW of the
city of Colima, 19°27′30″N, 103°40' W, 24 Sep 1988, Sanders et al. 8236
(CAS, MO, NY, TEX); Mpio. Autlán, 14–16 km NE de Casimiro Castillo,
2kmNEdeAhuacapan,19°14′N, 104°19′W, 5 Aug 1988 (fl), Santana
M. & Lorente 3732 (MEXU, WIS); 15 km al SW de Autlán, Barranca
del Tecolote, 5–8 km al NE de Casimiro Castillo, 19°37′N, 104°23′W,
800–900 m, 22 Jul 1987 (fl), Vázquez 4510 (WIS); Mpio. Tamazula, 16 km
al SE de Tamazula, brecha a Uña de Gato, 1,500 m, 14 Dec 1989, Vill a
C. & Chávez L. 452 (NY). México: 3 km NW de Los Pinzanes, Tejupilco,
29 Jan 1973 (fr), González-Medrano 5498 (MEXU); Mpio. Temascaltepec,
Salitre Cañitas, 12 Jul 1933, Hinton 4312 (BM,K,UC,US);Mpio.
Temascaltepec, Luvianos, 14 Jul 1933 (fl), Hinton 4320 (K, GH, MO).
Michoacán: Santa Ana Maya, Sep 1955 (fl), Castillo 2735 (MEXU);
lower north-facing slopes of Cerro Santa María, 8–10 km SW of Jiquilpán
and 5 km NE of Quitupán, 2,000 m, 5–7 Aug 1959 (fr), Feddema 102
(TEX); Villa Victoria, District Coalcomán, 700 m, 11 Jul 1939, Hinton
13906 (GH, LL, US, W); “El Salto”, E of Morelia on road to Ciudad
Hidalgo, 2,250 m, ca. 19°14'N [sic], 100°50'W, 11 Aug 1960 (fl), Iltis
et al. 839 (WIS); Mpio. Hidalgo, 2 km S de Mil Cumbres, camino a
San Antonio Villalongín, 2,500 m, May 1985 (fl), Soto N. et al. 8494
(MEXU, US); Mpio. Aguililla, 11 km NO de Aguililla, 1180 m, 11 Jul
1985 (fl), Soto N. et al. 9250 (MEXU, TEX); Mpio. Lázaro Cárdenas,
10 km N de Playa Azul, carretera a Nueva Italia, 250 m, 22 Aug 1985
(fl), Soto N. & Román G. 10188 (MEXU). Morelos: Cuernavaca, 5,000 ft,
24 Jul 1896 (fl), Pringle 6396 (BM, CM, E, F, GH, K, MO, NY, P-3 sheets,
PH, UC, US-2 sheets, W); near Cuernavaca, 10 Sep 1903, Rose 6947 (US).
Nayarit: San Blas, 2 Oct 1925 (fr), Ferris 5341 (DS, GH, US); Mpio.
Nayar, 10 km NE de La Guerra, camino La Guerra–Huejuquilla, 22°15′N,
104°15′W, 1,300 m, 10 Nov 1990 (fr), Flores F. & Ramírez R. 2388 (MEXU);
Mpio. Ahuacatlán, 8 km W de Jala, camino a la estación de Microondas,
Volcán El Ceboruco, 21°05′N, 104°30′W, 1,700 m, 18 Mar 1991 (fr), Flores
F. & Ramírez R. 2596 (MEXU); Mpio. Xalisco, 1 km SW de Palapitas,
21°25′N, 105°03′W, 790 m, 16 Sep 1994 (fr), Flores-Franco et al. 3830
(MEXU); Tres Marías Islands, María Madre, 24 Jul 1932 (fl,fr), Howell
10440 (CAS); Tepic, 14 Feb 1927 (fr), Jones 22925 (F,UC);TresMarías
Islands, María Madre, 17 May 1925 (fr), Mason 1732 (CAS, K); road
from Tepic to Calixcillo, 1,000 m, 13 Sep 1926 (fl,fr), Mexia 588 (CAS,
F, GH, MO, UC, US); Barranca, SE of Trapichillo, 650–700m,19Aug
1935 (fl), Pennell 19826 (PH); Acaponeta, 11 Apr 1910 (fl), Rose et al.
14372 (NY, US); Mpio. Compostela, 20–25 km S de Tepic, carr. a
Compostela, 21°20′N, 104°51′W, 31 Ju l 1 990 (fl ), Téllez V. 12766 (K,
MEXU, MO, NY). Sinaloa: Mpio. Concordia, 3 km above El Coco
on Mex. Hwy. 40, 1,020 m, 6 Aug 1980 (fl), Breedlove & Almeda 45058
(CAS, MEXU, MO); Labradas, 22 Sep 1925 (fl,fr), Ferris & Mexia 5246
(DS-2 sheets, GH); Mpio. Mazatlán, Rancho de Los Cannobio, Juantillos,
comunidad San Marcos, Sindicatura de la Noria, 10 Apr 1994 (fr), Guízar
Nolazco 3150 (MEXU); Mpio. Escuinapa, La Campana, 100 m, 30 Aug
1990 (fr), Hernández A. et al. 930 (MEXU); Rancho de Campanillas, San
Ignacio, 13 Sep 1918, Montes & Salazar 591 (US); San Agustia, Ortega
4021 (US); Culiacán, 27 Aug–15 Sep 1891 (fl,fr), Palmer 1534 (F, GH, NY,
UC); Rosario, 22 Jul 1897, Rose 1819 (GH, US); vicinity of Villa Union,
2 Apr 1910, Rose et al. 13909 (US); Mpio. Concordia, along Hwy. 40,
0.6 mi. above Cantil and 21 mi. NE of Concordia, ca. 600 m, 31 Dec
1983, Sanders et al. 4576 (MO); Mpio. Culiacán, Sanalona a 30 km E de
Culiacán, 200 m, 13 Nov 1989 (fr), Vega A. & Hernández 3586 (MEXU);
rd. between El Salto and Concordia, 10.5 mi by road N of Chupaderos,
23°30′N, 105°50′W, 3500 ft, 1 Jul 1969 (fl), Webster & Breckon 15545
(GH, MEXU, MO, TEX). Zacatecas: Mpio. Moyahua, 3 km W de Las
Palmas, 3 Sep 1992 (fl), Enríquez E. 201 (MEXU).
3. Solanum cobanense J. L. Gentry, Phytologia 26(4): 276.
1973.—TYPE: Nom. nov. for Cyphomandra aculeata Donn.
Sm. (1914), non Solanum aculeatum (Jacq.) O. E. Schulz.
Cyphomandra aculeata Donn. Sm., Bot. Gaz. 57: 423. 1914.—
TYPE: GUATEMALA. Alta Verapaz: Cobán, Apr 1882,
F. Lehmann 1334 (holotype: US [US-1324994, barcode
US00036715]; isotypes: BM [BM000514914], G [G00442620]).
Robust woody vine or canopy liana to 18 m long, the
stems 0.3–0.8 cm in diameter, prickly, sometimes densely so;
internodes 3–10 cm long, completely glabrous; prickles to
3 mm long, stout, recurved. Sympodial units plurifoliate, not
geminate. Leaves simple, the blades (6–)12–17 × 3–5(–9) cm,
ca. 3 times as long as wide, oblong or elliptic to narrowly
elliptic, widest at the middle, coriaceous, glabrous on both
surfaces, armed abaxially on midrib with recurved prickles;
major lateral veins 6–9 on each side; base truncate to acute;
margin entire; apex acuminate to acute; petioles (1–)3–7 cm,
glabrous, armed with recurved prickles. Inflorescences 10–15 cm
long and wide, terminal, few-branched, the axes glabrous,
not prickly; peduncle 3–5cm;rachis(3–)5–10 cm; pedicels
(10–)25–50 mm and slender at anthesis, elongating in fruit
to ca. 50 × 6 mm and becoming woody, articulated at the
base. Flower buds ellipsoid, the corolla strongly exserted from
the calyx when very young. Calyx campanulate, spreading,
the tube 5–7 mm, the lobes ca. 2 × 4 mm, rounded, the tips
apiculate, minutely papillate on the thickened margins. Corolla
4–6 cm in diameter, spreading at anthesis, thick and fleshy,
blue to violet, deeply stellate, lobed ca. 3/4 of the way to
the base, the lobes 1.2–2×0.3–0.6 cm, deltate to triangular,
glabrous adaxially, mostly glabrous abaxially, the tips deeply
cucullate, minutely papillate at the tips and margins. Stamens
equal, to 12 mm long; filaments ca. 2 mm long, glabrous;
anthers ca. 10 × 1 mm, ellipsoid, yellow, poricidal at the tips,
the pores lengthening to slits with age. Ovary glabrous;
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1115
Fig. 7. Representative specimen of Solanum cobanense from Guatemala (Förther et al. 10950, BM). Reproduced with permission of the Natural History
Museum, London.
1116 SYSTEMATIC BOTANY [Volume 40
style in long-styled (?) flowers ca. 6 mm long; stigma clavate
and somewhat bilobed. Fruit a globose or apically pointed
(“turnip-shaped”fide Steyermark 42873) berry ca. 3.5 × 4 cm,
green, glabrous with rugose surface when dry, the flesh white.
Seeds < 30 per berry, ca. 4 × 5 mm, strongly flattened, light
brown, the surface foveolate. Figs. 1C–D, 7.
Distribution and Habitat—Solanum cobanense (Fig. 8)
occurs in the cloud forests of Honduras (Depts. of Lempira,
Comayagua), Nicaragua (Dept. of Estelí), and Guatemala
(Depts. of Alta Verapaz, Huehuetenango) at elevations from
900–2,550 m.
Phenology—Solanum cobanense is most commonly collected
in flower from March to May, and fruiting specimens have
been collected from October to January.
Preliminary Conservation Status (IUCN 2014)—Vulnera-
ble (VU); EOO 6,668 km
2
(LC); AOO 28 km
2
(EN); 8 locali-
ties. Solanum cobanense would merit a status of endangered
based on the AOO, but given that it is a canopy liana, it is
likely that it is overlooked by many general collectors.
Solanum cobanense differs from all species of Solanum sec-
tion Aculeigerum except S. truinfense by its woody canopy
liana habit, coriaceous leaves, and large, fleshy flowers
(4–6 cm in diameter) with stamens and anthers of equal size
(Fig. 1C). Some specimens of S. wendlandii have simple leaves
and might be confused with S. cobanense,buttherotate
flowers of S. wendlandii with unequal anthers are distinctive.
Solanum cobanense differs from S. triunfense,withwhich
it is nearly sympatric, in its regular calyx lobes that do not
completely enclose the bud at late stages, the completely
glabrous stems and new growth, and the acute, rather
than attenuate, leaf bases. The flowers of S. cobanense
are more deeply stellate than those of S. triunfense,but
this character can be difficult to assess in herbarium
sheets where flowers are collected at different stages
of development.
Specimens examined —GUATEMALA. Alta Verapaz: Sierra de Chamá,
Montaña Yalijux, Finca Chelem-há, Aufsteig vom Tal des Riachuelo Yalijux
[15°23′09″N-90°03′45″W] zum nördlichen Bergkamm [15°23′32″N-
90°04′12″W], ca. 15 km Luftline NE von Tucurú, 27 Mar 2001 (fl), Förther
Fig. 8. Distribution of Solanum cobanense.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1117
et al. 10950 (BM); Pansamala, May 1887 (fl), von Türckheim 746 (US).
Huehuetenango: Nentón, Nuevo San José Frontera, along trail to Laguna
Yolnojal (=Laguna Brava), 16°03′N, 91°33′W, 900–1,100 m, 6 Mar 2009 (fl),
Christenhusz et al. 5584 (BM, NY). Zacapa: Sierra de Las Minas, between
Cerro de Monos and upper slopes of Monte Virgen, 6,562–8530 ft., 17 Jan
1932 (fr), Steyermark 42873 (F, NY).
HONDURAS. Comayagua: Campo Naranja, trail from Gracias
“El Mahon,”10 km SW of Gracias, 13°32′N, 88°39′W,2,550m,13May
1992 (fl), D’Arcy 17923 (MO). Lempira: Quebrada Naranjo, W of Campo
Don Tomás, 10 km SW of Gracias, Parque Nacional de Celaque, 14°33′N,
88°40′W, 1,950 m, 29 Jan 1992 (fr), Hawkins et al. 140 (MO); Río Naranjo,
Parque Nacional de Celaque, 14°33′N, 88°40′W, 2,100 m, 24 May 1991
(fl), House & Andino 968 (MO); W of Campo Don Tomás, 10 km SW
of Gracias, Parque Nacional de Celaque, 14°32′N, 88°40′W, 2,20 0 m,
11 Feb 1993 (fr), Mejía 252 (MO).
NICARAGUA. Estelí: Salto de Estancuela, Río Estancuela, ca. 6 km S
of Estelí, 13°01’N, 86°20′W, 920–1,020 m, 1 Oct 1979 (fr), Stevens et al.
14381 (MO).
4. Solanum glaucescens Zucc., Abh. Math.-Phys. Cl. Königl.
Bayer. Akad. Wiss. 2: 325. 1837.—TYPE: Grown in the
Munich Botanical Garden by Zuccarini from seeds
collected in Mexico by Karwinsky [Wilhelm Friedrich
Karwinsky von Karwin], 1835, Anonymous s.n. (neotype,
designated here: M [M0124264, Morton neg. 8691];
isoneotype: BR [BR0000008366221]).
Solanum hamatile Brandegee, Univ. Calif. Publ. Bot. 6: 192.
1915.—TYPE: MEXICO. Oaxaca: Río San Gerónimo,
Jul 1914, C. Purpus 7164 [“6164”in the protologue]
(holotype: UC [UC175042]; isotypes: BM [BM000514922],
F [F-424575], MO [MO-555119], NY [NY00138991], US
[US-467485]).
Solanum oaxacanum Dunal, Prodr. [A. P. de Candolle] 13(1):
204. 1852.—TYPE: MEXICO. Oaxaca: Tehuántepec, Aug,
G. Andrieux 189 (holotype: G-DC [G00154874, F neg.
6832, IDC microfiche 800–61.2078:I.6]; isotypes: K
[K000063703, K000195650], W [W0003308]).
Solanum sagranum A. Rich., Hist. Fis. Cuba 11: 124. 1850, as
“sagraeanum”.Solanum lanceifolium Jacq. var. sagranum
(A. Rich.) M. Gómez, Anal. Hist. Nat. Madrid 23: 268.
1894, as “sagraeanum”—TYPE: CUBA. Sin. loc., R. De la
Sagra s.n. (lectotype: designated here: P [P00370903];
isolectotypes: F, K [K000449492], P[P00371124, P00371122,
P00371123, P00445150], W).
Woody vine or scandent shrub 2–5 m long, the stems
0.25–0.6 cm in diameter, usually very sparsely prickly; inter-
nodes 1–5 cm long, glabrous, with paired prickles at base of
each petiole, these 2–8 mm long, stout, recurved. Sympodial
units plurifoliate, not geminate. Leaves deciduous in the dry
season, simple, the blades 3.5–9 × 1.5–5.5 cm, 1.6–2.3 times
as long as wide, narrowly oblong to ovate, chartaceous,
glabrous adaxially and abaxially, often armed abaxially on
midrib with recurved prickles; major lateral veins 3–6on
each side; base cuneate to decurrent; margin entire; apex
acuminate; petioles 1–4 cm, glabrous, often prickly. Inflores-
cences 7–10 × 15–20 cm, terminal or lateral, unbranched or
few-branched, with 5–12 flowers, the axes glabrous, not
prickly; peduncle 1–2cm;rachis2–7cm;pedicels5–10 mm
and slender at anthesis, articulated at the base; fruiting
pedicels elongating to 3–4 cm and becoming woody. Flower
buds ellipsoid, the corolla exserted from the calyx tube
long before anthesis. Calyx campanulate, spreading, the
tube 2–4 mm, nearly entire with five shallowly subcaudate,
broadly deltate, acuminate lobes, glabrous except for the
puberulent tips. Corolla 2–4 cm in diameter, spreading at
anthesis, pale yellow to yellow-green, membranous, deeply
stellate, lobed nearly to the base, the lobes ca. 1.5 × 0.5 cm,
lanceolate to broadly deltate, acute at apices, glabrous
abaxially, glabrous adaxially except puberulent at tips. Sta-
mens 10–15 mm long; filaments unequal, with four 1.5–2mm
long, the fifth 3.5–4.5 mm long, glabrous; anthers nearly
equal, 6–9 × 1.5–2 mm, the one on the longer filament slightly
larger, slightly tapering, connivent, white to yellowish-orange,
poricidal at the tips, the pores lengthening to slits with age.
Ovary glabrous; style in long-styled flowers ca. 15 mm long,
straight, glabrous in short-styled flowers ca. 2 mm long;
stigma truncate. Fruit a globose berry, ca. 3.5 cm in diameter,
mottled dark green and light green when immature, turning
orange when mature with orange-red pulp, the pericarp gla-
brous, rugose when dry. Seeds > 50 per berry, 4–5×3–5mm,
flattened-reniform to somewhat ovoid, light brown, the sur-
faces minutely pitted, the testal cells pentagonal in shape,
the lateral cell walls elongate and the seed appearing hairy
when outer testal walls break down. Fig. 9.
Distribution and Habitat (Fig. 10)—Endemic to southern
Mexico (States of Chiapas, Guerrero, Oaxaca, and Puebla) in
deciduous and semi-deciduous forests; 10–1,400 m.
Phenology—Solanum glaucescens appears to flower and
fruit at the same time; flowering specimens are most com-
monly collected from April to July and fruiting specimens
from July to January.
Common names—Mexico. Puebla: “cuatomate”(Huerta Z. &
Huerta Z. 1); Oaxaca: “zarza”(Elorsa 301).
Preliminary Conservation Status (IUCN 2014)—Least
Concern (LC); EOO 221,062 km
2
(LC); AOO 324 km
2
(EN);
95 localities. Solanum glaucescens is known from a large range,
and is relatively commonly collected within it.
Solanum glaucescens differs from other members of section
Aculeigerum by the presence of paired recurved prickles
flanking the petiole base (Fig. 9). These appear stipular, but
stipules are absent in the Solanaceae, and these are prickles
like those found on the rest of the plant. The similar situation
occurs in the unrelated spiny solanum species S. microphyllum
(Lam.) Dunal of the West Indies (Knapp 2009). Solanum
glaucescens differs from S. bicorne, with which it is partly
sympatric, in its widely spaced leaves (not clustered as in
S. bicorne), smaller yellowish green rather than white flowers,
and in its glabrous leaf undersides. Solanum glaucescens is usu-
ally a more delicate plant than S. bicorne. Trichomes, if pres-
ent, on all parts of S. glaucescens are small and unicellular, not
the uniseriate trichomes with elongate basal cells of S. bicorne.
Zuccarini (1837) mentions no herbarium specimens in
the protologue of S. glaucescens, and states only “Crescit in
imperio mexicano, unde semina misit clar. de Karwinski.
Floret nobis caldariis Augusto, Septembri (v.v.)”,although
the herbarium of the Munich garden is mentioned in the
title of his article. The indication “(v.v.)”indicates he saw a
living plant, from which the extant herbarium specimens
were presumably made. We therefore designate the sheet
in the Munich herbarium (M0124264) taken from plants cul-
tivated there in 1835 as the neotype of S. glaucescens; the
sheet in BR labelled as coming from “h. Mon.”(garden
[hort.] or herbarium [herb.] of Munich) with no date is a
probably an isoneotype.
The epithet sagranum honoring Ramón De la Sagra was
originally spelled sagraeanum, a correctable orthographic error
following Art. 60 Rec. 60C.1of the Code (McNeill et al. 2012).
1118 SYSTEMATIC BOTANY [Volume 40
Fig. 9. Representative specimen of Solanum glaucescens from Mexico (J. Rivera H. et al. 1928, NY). Reproduced with permission of the New York
Botanical Garden, Bronx, NY.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1119
The illustration of Solanum sagranum in the plates (De la Sagra
1855: pl. 62) accompanying the botany volumes of De la
Sagra’sHistoria fisica, politica y natural de la isla de Cuba pub-
lished five years after the protologue (Richard 1850) is clearly
S. glaucescens. The paired recurved prickles at the base of the
petioles are distinctly shown, but the heterandrous flowers
clearly described in the protologue (“Uno de sus estambres es
constantamente mayor que los otros cuatro.”[One of the sta-
mens is always longer than the other four]; Richard 1850: 124)
are not; specimens at P and K collected by De la Sagra (see
below) have heterandrous flowers and paired prickles. No
other collections of S. glaucescens from Cuba have been seen,
and its occurrence on Cuba has not been reconfirmed. The
type of Solanum sagranum may represent a cultivated collec-
tion and/or a mis-labeling of the collecting locality. Several
duplicates of a collection attributed to Ramón De la Sagra
exist in the Paris herbarium. We have selected that labeled
as “Herbarium Richard”and “Solanum punctulatum nob.
Sagræanum nob.”(P00370903) as the lectotype as it was
clearly part of A. Richard’s herbarium and is indicated as
a new species on the sheet ([Solanum punctulatum new
Sagræanum new]; Richard clearly changed his mind about
the name for this taxon and crossed his original epithet,
punctulatum, out). Other sheets attributed to De la Sagra
(see synonymy) appear to be duplicates and are here recog-
nized as isolectotypes.
Representative Specimens Examined—MEXICO.Chiapas:Mpio.
Jiquipilas, 8 miles E of Cintalapa, along hwy. 190, 2,200 ft, 12 Jun 1965
(fl), Breedlove 10304 (F, US), 10317 (F,TEX,US);Mpio.Chiapade
Corzo, at the Río Grijalva, 10 km W of Chiapa de Corzo, Mex. Hwy.
190, 500 m, 22 Jul 1972 (fl), Breedlove 26219 (CAS, MEXU, MO); Mpio.
Villa Corzo, 56 km S of Mex. Hwy. 190 near junction to Jérico on road
to Nueva Concordia, 700 m, 10 Sep 1974 (fl), Breedlove 37464 (MEXU,
MO); Mpio. Cintalapa, NW of Cintalapa along road to Colonia
Francisco I. Madero, 560 m, 30 Dec 1980 (fr), Breedlove 48993 (MEXU);
Mpio. Acala, along Río Grijalva, 20 km N of Acala, 550 m, 30 Jul 1981
(fl), Breedlove 51893 (CAS, DUKE, MEXU, MO, TEX); Mpio. Jiquipila,
Cerro San Lorenzo o La Chumpa, 580 m, 8 Jul 1998 (fl), Farrera S. 1004
Fig. 10. Distribution of Solanum glaucescens.
1120 SYSTEMATIC BOTANY [Volume 40
(MEXU; Mpio. Venustiano Carranza, along the road from Acala to
Venustiano Carranza 15 km SE of Acala, 3 May 1966, Laughlin 860
(US); San Nicolás, cerca de Cupía, SE de Tuxtla Gutiérrez, 500 m,
19 Feb 1950 (fl,fr), Miranda 6061 (MEXU-3 sheets); Mpio. Chiapa de
Corzo, carretera a La Concordia, cerca del Rancho Chuquiyaca, 23 Oct
1991 (fl), Palacios E. 2004 (MEXU). Guerrero: 3 km NW of Zumpango,
1,000 m, 25 May 1973 (fl), Hansen et al. 1503 (MEXU, US, WIS); 2.5 km
W de Puerto Márquez, 10 Aug 1977 (fl,fr), López F. 467 (MEXU, TEX);
Mpio.Chiapilla,VicenteGuerrero,6kmNWofFloresMagón,16°25'
N, 92°43' W, 475 m, 21 Dec 1985 (fr), Nee 32311 (CORD, MO, NY,
P, XAL). Oaxaca: Dto. Tlacolula, Mpio. San Pedro Totolapan, 5 km
NW de Las Margaritas, 17 Jul 1987 (fl), Aguilar S. 249 (MEXU, NY);
Ixtepec, 11 Jan 1945 (fl), Alexander 326 (NY); Puerto Ángel, 25 Dec
1974 (fr), Boege 3317 (MEXU); Mpio. San Miguel Soyaltepec, Presa
Miguel Alemán, Temascal, 350 m, 20 May 1987 (fl), Calzada 9250 (MEXU);
Mpio. Tehuantepec, recorrido a Cerro El Arenal, NO de Buenos Aires,
16°19′N, 95°32′W,950m,28Jun1991(fl),Campos V. 3700 (MEXU);
Mpio. Huatulco, camino a Barra de Boca Vieja, bajos de Coyula, 15°04′N,
96°18′W, 5 m, 31 Oct 1992 (fl), Castillo Campos et al. 9208 (MEXU); Mpio.
Huatulco, NE de playa de Cacaluta, 15°45′N, 96°10′W, 20 m, 3 Nov 1992
(fr), Castillo Campos et al. 9352 (MEXU); Distr. Pochutla, 1 km S de la
desviación a la Bahía de Santa Cruz Huatulco, 27 Jun 1982 (fl), Cedillo
Trigos & Torres 1558 (MEXU, MO); Distrito Tuquila, Chacahua, 5 m,
17 Dec 1921 (fr), Conzatti 4486 (MEXU, US); Mex. 190, 8 mi E of Gramal,
17 Sep 1978 (fl), D’Arcy 11993 (MEXU, MO); Mpio. Santiago Astat, Dto.
Tehuantepec, Zimatán, 4.5 km S del puente, playa El Mojón, 15°48′44″N,
95°59′18″W, 21 Jun 1998 (fl), Elorsa C. 301 (NY); Mpio. San Miguel del
Puerto, Dto. Pochutla, Petatengo, 3.5 km S, Rancho Zimatán, 15°56′23″N,
96°01′33″W, 30 Jun 1998 (fl), Elorsa C. 319 (NY); Dto. Tehuantepec, Mpio.
Santiago Astata, Campamento Tortuguero, 4 km SE de Barra de la Cruz,
15°50′70″N, 95°56′55″W, 10 Oct 2000 (fr), Elorsa C. 3699 (NY); Dto.
Tehuantepec, Mpio. San Pedro Huamelula, Carretera vieja a Mazcalaco,
3 km NE de Playa Grande, 15°51′24″N, 95°53′39″W, 10 Aug 2000(fr),
Elorsa C. 3374 (NY); Dto. Juchitán, Mpio. Asunción Ixtaltepec, 1 km N de
Nizanda, 16°40′N, 95°02′W, 200 m, 2 Jun 1996 (fl), Gallardo H. & Reyes
D. 1149 (MEXU); Mpio. Asunción Ixtaltepec, 2.2 km SO de Nizanda,
16°38′N, 95°00′W, 200 m, 9 Jul 1995 (fl), Gallardo H. & Pérez-García 1464
(MEXU); Dto. Juchitán, Mpio. Asunción Ixtaltepec, bosque riparia hacia el
“Agua Tibia”, 500 m al N de Nizanda, 16°40′N, 95°00′W, 115 m , 1 A u g
1995 (fl,fr), Gallardo H. & Pérez-García 1562 (MEXU); Dto. Juchitán, Mpio.
Asunción Ixtaltepec, camino hacia el cerro de la “Piedra Azul”, cercano a
Nizanda, 16°39′N, 95°00′W, 180 m, 22 Jul 1996 (fl,fr), Gallardo H. & Pérez-
García 1876 (MEXU, MO); Agua Blanco, 21 mi S of Mitla road junction
along hwy, 16°45′N, 96°22′W, 3,400 ft, 21 Oct 1967 (fl), Gentry 21361C
(A, MEXU); Mpio. San Pedro Pochutla, 15 km E de San Pedro Pochutla,
5 Oct 1980 (fl), González O. 595 (MEXU); 0.5 km E of Tehuantepec, 50 m or
less, 30 Jun 1958 (fl), King 279 (MEXU, TEX, WIS); La Ventosa, 7 km E
of Salina Cruz, 50 m or less, 3 Jul 1959 (fl), King 1277 (NY, TEX, UC, US);
3–4 km W of Ixtaltepec, ca. 50 m, 10 Jul 1959 (fl), King 1519 (NY, TEX, UC,
US); Mpio. Huatulco, Dto. Pochutla, Bahía de Tangolunda, 11 Oct 1984
(fr), López G. 156 (NY); Tehuantepec, 15 Oct 1953 (fl), MacDougall s.n.
(MEXU); Mpio. Tehuantepec, Dto. Tehuantepec, El Limón–El Milagrito,
El Limón está a 17 km al O de Tehuantepec entrando por Hierba Santa,
8 Jul 1985 (fl), Martínez R. 23 (MEXU); Dto. Tehuantepec, Mpio. Salina
Cruz, Carnero, entrando por Rincón Bamba, carr. a Pochutla, O de Salina
Cruz, 23 Jun 1986 (fl), Martínez R. 584 (MEXU); Dto. Yautepec, Mpio.
Santa María Ecatepec, 15 km SO de La Reforma camino a Santa María
Ecatepec, 16°20′N, 95°50′W, 15 Jul 1988 (fl), Martínez R. 1470 (MEXU,
MO); Distr. Tehuantepec, Mpio. San Pedro Huamelula, 2 km N de Sta.
María Huamelula, 15°59′N, 95°42′W, 15 Aug 1988 (fl), Martínez R. 1714
(MEXU, MO); Mpio. Asunción Ixtaltepec, Dto. Juchitán, alrededores del
vía del tren Transístmico, Nizanda, 16°40′N, 95°01′W,110m,15Sep1995
(fr), Meave del Castillo & Pérez-García 1925 (MEXU); Dto. Juchitán, Mpio.
Asunción Ixtaltepec, camino hacia el EVX 1, cercano a Nizanda, 16°39′N,
95°00′W, 180 m, 16 Sep 1995 (fr), Meave del Castillo & Pérez-García 1945
(MEXU); Dist. Yautepec, Mpio. Nejapa, vic. Camarón, 13 km NW of San
Juan la Jarcia, along hwy. Mex. 190, 16°34' N, 96°02' W, 700 m, 19 Dec
1985 (fr), Nee & Andres 32280 (G, K, MO, NY, TEX, XAL); Dto. Pochutla,
Mpo. San Miguel del Puerto, Zimatán, 3 km LR N del puente, por el
Chorro, 15°52′31″N, 96°00′17″W, 27 Aug 1999, Rivera H. et al. 1818 (NY);
70 km (by road) SE of Pinotepa Nacional on road to Puerto Escondido,
16°10'N, 97°40'W, 150 m, 23 Jul 1965 (fl), Roe et al. 523 (WIS); Dto.
Tehuantepec, Mpio. San Pedro Huamelula, Mascalco, N de la laguna,
2.5 km S de la carretera costera, 15°52′20″N, 95°52′22″W, 24 Jul 1998 (fr),
Salas M. et al. 1896 (NY); Dto. Tehuantepec, Mpio. Santiago Astata, Barra
de la Cruz, 4 km NE LR, ladera arriba del puente del Curuelo, 15°52' N,
95°57′15″W, 15 Sep 1999 (fr), Salas M. et al. 2321 (NY); 1.2 km S de la
desviación a Bahía de Santa Cruz, la desviación a 15.2 km SW de
Copalita, 150 m, 30 Jul 1984 (fl), Torres C. & Martínez 5763 (MEXU); Distr.
Tehuantepec, Arroyo de las Minas, W de El Limón, entrando por la
desv. a Buenos Aires, 17 Jun 1985 (fl), Torres C. & García M. 6849 (MEXU);
Distr. Yautepec, 5.5 km SW de la entrada a Santa María Ecatepec, SW
de la Reforma, 19 Jun 1985 (fl), Torres C. & M. Laubin 6888 (MEXU, MO,
NY); Distr. Pochutla, Mpio. Santa Cruz Huatulco, 6 km hacia el E de
Bahía de Santa Cruz, hacia Chahue, 30 m, 15 Jun 1986 (fl), Torres C. & O.
Téllez 8405 (MEXU). Puebla: Mpio. Izúcar de Matamoros, Paraje La Poza
de Malpaso, Los Amates, 1,240 m, 18 Jun 1998 (fl), Guízar Nolasco & Herrera
4023 (MEXU);Mpio.Tocomatlán,Tezoquipán,1,000m,Aug1988(fl,fr),
Huerta Z. & Huerta Z. 1 (MEXU); 12 miles E of jnct. of hwy. 190 and 160 out
of Izúcar Matamoros, 3400 ft, 2 Aug 1975 (imm fr), Torke et al. 364 (MO).
5. Solanum pachyandrum Bitter, Repert. Spec. Nov. Regni
Veg. 13: 88. 1914.—TYPE: ECUADOR. Manabí: El Recreo
[“El Keereo”], 200 m, 6 Mar 1897, H. von Eggers 15730
(lectotype, designated here: US [US-1324689, barcode
US00027720]; isolectotypes: B, †destroyed [F neg. 2672],
F [F-143295], K [K000788525], L [L-897313–55], LE, M
[M0124267], P [P00367219]).
Woody vine to 5 + m long, the stems 0.4–0.7 cm in diame-
ter, densely prickly; internodes (2–)4–9 cm long, glabrous;
prickles 0.5–1.2 mm long, stout, recurved. Sympodial units
plurifoliate, not geminate. Leaves pinnatifid to deeply pinnati-
fid with 4–6 pairs of lobes, the blades in outline 12–19(–27) ×
5–10(–17) cm, oblong, chartaceous, glabrous on both surfaces,
the midrib with recurved prickles abaxially; major lateral veins
4–6 on each side, corresponding to the lobes; petioles 2–3cm,
glabrous, often armed with recurved prickles; leaflets 2–4×
ca. 1 cm, widest at the middle or in the basal third, basally
decurrent on the midrib and with a wing of tissue to 3 mm
wide connecting all leaflets to some degree, not petiolulate;
base rounded to acute; margins deeply pinnatifid, entire, the
terminal leaflet confluent with lateral leaflets; apex acuminate
to rounded. Inflorescences 12–20 cm long and wide, lateral
or terminal, several times branched, with 5–20 flowers, the
axes glabrous or minutely papillate, densely prickly; peduncle
5–8cm;rachis4–8cm;pedicels4–5 mm, prickly, slender at
anthesis, elongating in fruit to ca. 2.5 × 0.2 cm and becom-
ing subwoody, articulated at the base. Flower buds ellip-
soid to oblong, the corolla soon exserted from the calyx
tube. Calyx campanulate, spreading, the tube 1–2 mm, the
lobes ca. 3 × 2 mm, deltate, glabrous, the central part a pro-
truding green knob, this more prominent in fresh plants.
Corolla to 3.5 cm in diameter, spreading at anthesis, white,
fleshy, stellate, lobed 1/2–2/3 of the way to the base, the
lobes 1–1.2 × 0.5–0.6 cm, narrowly triangular, acute to acu-
minate at apices, glabrous on both surfaces, minutely papil-
late at the tips. Stamens equal in size, 6–8 mm long; filaments
ca. 2 mm long, glabrous; anthers 5–6×2.5–3mm,ellipsoid,
connivent, yellow, poricidal at the tips, the pores lengthening
to slits with age. Ovary glabrous; style in long-styled flowers
ca. 10 mm long, straight, glabrous, in short-styled flowers ca.
5 mm long; stigma truncate. Fruit a globose berry, 3–5cmin
diameter, purple when immature and yellow when mature
(Rodriguez et al. 1607), the pericarp glabrous. Seeds ca. 100 per
berry, ca. 2 × 4 mm, strongly flattened, light brown, the sur-
faces minutely pitted, the testal cells rectangular. Figs. 2A–C, 11.
Distribution and Habitat (Fig. 12) —Solanum pachyandrum
occurs on the western slopes of the Andes and in the
Huancabamba Depression of Ecuador (Prov. Manabí) and
Peru (Depts. of Cajamarca, La Libertad, and Tumbes) in moist
forests on slopes; 200–1,000 m. The Ecuadorian type collection
is somewhat disjunct from the rest of the species distribution.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1121
Fig. 11. Isolectotype specimen of Solanum pachyandrum from Ecuador (H. Eggers 15730, K). © Trustees of the Royal Botanic Gardens, Kew.
Reproduced with permission of the Royal Botanic Gardens, Kew.
1122 SYSTEMATIC BOTANY [Volume 40
Phenology—Solanum pachyandrum appears to flower and
fruit at the same time; flowering and fruting specimens have
been collected from April to September.
Common Names—Peru. Cajamarca: “bombona”(Rodriguez
et al. 1607).
Preliminary Conservation Status (IUCN 2014) —Near
Threatened (NT); EOO 30,173 km
2
(LC); AOO 24 km
2
(EN);
7 localities. Although the EOO would not place S. pachyandrum
into one of the threatened categories (CR, EN or VU) the single
point that expands the range is derived from the type collec-
tion. Solanum pachyandrum has not been collected again in
coastal Ecuador since. If that point is removed from the analy-
sis the threat status is EN. Searches for S. pachyandrum in
southern Ecuador are a priority.
Solanum pachyandrum differs from all other species in sec-
tion Aculeigerum by the presence of stout armed prickles on
its pedicels and non-tapering (ellipsoid) anthers. Addition-
ally, S. pachyandrum is restricted to the western lowlands of
Ecuador and northwestern Peru. This species is geographi-
cally isolated from all other members of the section. The
large juicy fruits are eaten by local people and are consid-
ered a great treat by children (J. Campos, pers. comm.).
No herbarium was cited in the protologue of S. pachyandrum.
We have therefore chosen the best preserved of the duplicates
of Eggers 15730 as the lectotype (US00027720). It is often
assumed that when Bitter cited no herbarium he was citing a
specimen in Berlin; no internal evidence to that effect is to be
found in the publication, so we are including a reference to the
B sheet photographed by MacBride among the isolectotypes.
Specimens Examined —PERU. Cajamarca: Prov. San Ignacio, Distrito
Huarango, San Martín, 5°16’s, 78°43′W, 900 m, 18 May 1996 (fl,fr), Campos
et al. 2806 (MO, MOL, NY, USM); Prov. San Ignacio, San Martín del
Chinchipe, 5°19′16”S, 78°41′05″W, 1,000 m, 15 Sep 1999 (fr), Flores et al.
160 (BM, NY, USM-2 sheets); Prov. Jaén, Huayhuaya–Peña Blanca
(Tamborapa–Tabaconas), 890 m, 1 Aug 1994 (fr), Leiva G. et al. 1231
(HAO†[destroyed], NY); Prov. San Ignacio, distrito Namballe, Namballe,
arriba del margen derecho del Río Namballe, 4°58’s, 79°10′W, 760–800 m,
6 Jul 1997 (fl,fr), Rodríguez et al. 1607 (HAO†[destroyed], HUT, MO, NY,
Fig. 12. Distribution of Solanum pachyandrum.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1123
USM); Prov. Cutervo, on road from La Capilla to Jaen, in Laguna, km
1575, 1,126 m, 9°09’57.77”S, 78°51′14″W, 14 Apr 2013 (fl, fr), Särkinen et al.
4546 (BM, E, USM). Tumbes: Prov. Zarumilla, entre puesto de vigilancia
El Caucho–Campoverde, 700 m, 9 Jul 1992 (fr), Díaz et al. 4739 (HUT,
MO, USM).
6. Solanum refractum Hook. & Arn., Bot. Beechey Voy.
304. 1838.—TYPE: MEXICO. Nayarit: Tepic, F. Beechey
s.n. (holotype: K [K000195651, MICH neg. 751]).
Solanum aculeolatum M. Martens & Galeotti, Bull. Acad. Roy.
Sci. Bruxelles 12: 143. 1845.—TYPE: MEXICO. Oaxaca:
“Près la Mer Pacifique”, 1840, H. G. Galeotti 1170 (lecto-
type, designated here: BR [BR0000005538768, BH neg.
2394]; isolectotype: P [P00445058]).
Solanum molinarum J. L. Gentry, Phytologia 26: 276. 1973.—
TYPE: HONDURAS. Comayagua: edge of Humuya
River bank, vicinity El Edén, Comayagua Valley, 500 m,
28 Sep 1969, A. Molina R. & A. R. Molina 24532 (lecto-
type,designated here: F [F-1724479]; isolectotypes: EAP
[EAP70442], F [F-1964597, F-1724480, F-1964595,], NY
[NY00139010)].
Woody or subwoody vine to 15 m long, the stems 0.2–0.3 cm
in diameter, prickly; internodes (2–)4–10 cm long, glabrous
or puberulent with simple hairs ca. 0.2 mm long; prickles
0.5–1.5 mm long, stout, recurved. Sympodial units plurifoliate,
not geminate. Leaves pinnatifid to deeply pinnatifid, the
blades in outline 6–12(–16) × 3–10(–16) cm usually ovate
(rarely elliptic), widest at the base, chartaceous, glabrous
adaxially and abaxially, or rarely both surfaces puberulent
with minute simple papillae, often armed abaxially on midrib
with recurved prickles; major lateral veins 3–5oneachside,
corresponding to lobes; petioles 1–2.5(–5) cm, glabrous, often
armed with recurved prickles; lobes (1.5–)3–6×(1–)1.5–3cm,
widest at the middle of in the basal 1/3, confluent with the
midrib and with a wing of tissue to 1 cm wide connecting all
leaflets; base acute; margin deeply pinnatifid with 3–4pairsof
lobes, the terminal and adjacent lateral lobes broadly conflu-
ent; apex acuminate to rounded; petiolules absent, the lobes
confluent with the midrib. Inflorescences 10–15 cm long and
wide, terminal, several times branched, with 20–40 flowers,
the axes glabrous or puberulent with minute papillae, moder-
ately to densely prickly, occasionally with very few prickles
only at the base (type specimen); peduncle 3–5(–10) cm; rachis
3–10 cm; pedicels 3–5 mm and slender at anthesis, articulated
at the base; fruiting pedicels elongating to ca. 2.5 × 0.4 cm
and becoming woody. Flower buds sharply pointed (turbi-
nate), the corolla strongly exserted from the calyx tube before
anthesis. Calyx campanulate, spreading, the tube 1–3mm,the
lobes ca. 1 × 1.5 mm, broadly rounded and the calyx almost
truncate, glabrous, the lobe tips somewhat rounded and swol-
len. Corolla to 2.5 cm in diameter, spreading at anthesis, green
or yellowish green, sometimes tinged with purple (Hinton
10967 “flowers purple and yellow”), membranous, stellate,
lobed to the base, the lobes 0.9–1.5 × 0.2–0.3 cm, narrowly tri-
angular to lanceolate, acute to acuminate at apices, glabrous
abaxially and adaxially. Stamens equal in size, 7–9mmlong;
filaments ca. 0.5 mm long, glabrous; anthers 6–8 × 0.75–1mm,
strongly tapering, connivent, yellow, poricidal at the tips, the
pores only somewhat lengthening to slits with age. Ovary gla-
brous; style in long-styled flowers 12–13 mm long, straight,
glabrous, in short-styled flowers ca. 6 mm long; stigma trun-
cate. Fruit a globose berry, ca. 3–5 cm in diameter, light green,
the pericarp glabrous, rugose surface dry. Seeds 20–50 per
berry, ca. 3 × 5 mm, strongly flattened-reniform, light brown,
the surfaces minutely pitted. Fig. 13.
Distribution and Habitat (Fig. 14)—Solanum refractum
occurs in Mexico (States of Chiapas, Guerrero, Oaxaca),
Guatemala (Depts. of Chiquimula, Zacapa, Jalapa), and
Honduras (Depts. of Comayagua, La Paz) in deciduous and
semi-deciduous forests at elevations from sea level to 1,100 m.
Phenology—Solanum refractum appears to flower and fruit
at the same time throughout the year, with a small peak in
flowering specimens from June to August (but this is possi-
bly a collecting artefact) and in fruiting specimens from
September to November.
Preliminary Conservation Status (IUCN 2014)—Least Con-
cern (LC); EOO 133,321 km
2
(LC); AOO 112 km
2
(EN);
31 localities. Solanum refractum is widespread in the habi-
tats where it occurs.
Solanum refractum is similar to S. alternatopinnatum of south-
eastern Brazil, Paraguay, and northern Argentina but the ter-
minal leaflets in the latter are usually petiolulate and not
broadly confluent with the uppermost pair of lateral leaflets.
The leaves of S. refractum are occasionally puberulent with
unbranched papillae on both surfaces in contrast to the con-
sistently glabrous leaves of S. alternatopinnatum.Theinflo-
rescence structure, flowers, and fruits, are very similar in
S. alternatopinnatum and S. refractum.Solanum pachyandrum of
Ecuador and Peru shares consistently pinnatifid or deeply
pinnatifid leaves with S. refractum, but differs from it in its
white flowers with ellipsoid anthers; the yellow-green flowers
of S. refractum aremoredelicatethanthoseofS. pachyandrum
and have strongly tapered rather than ellipsoid anthers.
Solanum alternatopinnatum,S.pachyandrum and S. refractum
all share a prickly inflorescence rachis, a character that
can be used to differentiate these species from cultivated
S. wendlandii with pinnately compound leaves.
Confusion over the application of the names S. refractum
and S. bicorne has resulted in many specimens of S. bicorne
being annotated in herbaria as S. refractum (in these cases the
name S. aculeolatum was used for the plants here recognized
as S. refractum). Considerable care must be exercised in
using pre-2005 annotations on specimens not re-annotated
for this monograph.
The type locality for Solanum refractum is given by Hooker
and Arnott (1838: 275) on the page at the beginning of their
Mexico section in the account of the Beechey voyage as:
“When no habitat is mentioned, the specimens are under-
stood to have been collected at Tepic.”
Martens and Galeotti (1845) mentioned no specific her-
barium in the protologue of S. aculeolatum. We have selected
the specimen at BR (BR0000005538768) as the lectotype
because it has more copious inflorescence and leaf material
than the duplicate of Galeotti 1170 at P (P00445058); both are
relatively scrappy specimens.
We have selected F-1724479 from among the many duplicates
of Molina R. & Molina 24532 as the lectotype of S. molinarum
because it is annotated as “holotype”in J. L. Gentry’shand,
and the species name is also written by Gentry on the sheet.
The other duplicate labelled “holotype”(F-1964595) has a
typed species name on the label.
Representative Specimens Examined—MEXICO. Chiapas: Cintalapa
de Figueroa, km 22 de la carretera Tapanatepec–Tuxtla Gutiérrez,
16°27'N, 94°05'W, 700 m, 4 Jun 1995 (fl), Panero et al. 5750 (NY). Guerrero:
Atoyac, Distr. Galeano, 40 m, 25 Nov 1937 (fl), Hinton et al. 10967 (GH, K,
LL, MEXU, NY, U, US); Mpio. Petatlán, 22 km NW de Petatlán, camino a
El Camalote, 180 m, 23 Oct 1983 (fr), Martínez S. et al. 5116 (MEXU, NY);
1124 SYSTEMATIC BOTANY [Volume 40
Fig. 13. Holotype specimen of Solanum refractum from Mexico (Beechey s.n., K). © Trustees of the Royal Botanic Gardens, Kew. Reproduced with
permission of the Royal Botanic Gardens, Kew.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1125
15 km N de La Unión, camino a Coahuayutla, 150 m, 24 Oct 1983 (fl), Soto
N. et al. 6026 (MEXU); Mpio. La Unión, 2 km N de La Unión, 150 m,
22 Oct 1985 (fr), Soto N. et al. 11284 (MEXU). Oaxaca: Mpio. Santa María
Huatulco, Dto. Pochutla, 5 km al E de Santa Cruz Huatulco, 15°46′N,
96°07′W, 15 Aug 19 8 8 (fl) , Campos V. & Cortes 2288 (NY); Mpio. Sto
Domingo Tehuantepec, Dto. Tehuantepec, Cerro El Arenal, 17°19′N,
95°30′W, 3 Sep 1991 (fr), Campos V. 3954 (NY); Mpio. Huatulco, NE de
la playa de Cacaluta, 15°45′N, 96°10′W, 20 m, 3 Nov 1992 (fr), Castillo
Campos et al. 9347 (MEXU); Mpio. Santiago Astata, Dto. Tehuantepec,
BarradelaCruz,2.5kmSLR,veredahacialaMareñaalEdelabocana
del Río Zimatán, 15°49′10″N, 95°58′24″W, 17 Sep 1998 (fl), Elorsa C. 737
(NY); Mpio. Chivela, Dto. Juchitán, Nizanda entrando por La Pedrera
rumbo a Chivela, 15 km NE de La Ventosa, 16°37′N, 95°00′W, 18 Jul 1988
(fl), Martínez R. 1540 (MEXU, NY); Mpio. Sta. María Ecatepec, Dto.
Yautepec, Cerro San Lorenzo, 15 km O de La Reforma, 16°20′N, 95°50′W,
3 Oct 1988 (fl), Martínez Calderón 1900 (MEXU); Mpio. Santiago Astata,
Dto. Tehuantepec, Puente Zimatán, 26 Aug 1999 (fl, fr), Martínez S. 32499
(NY); Mpio. San Pedro Huamelula, Dto. Tehuantepec, Tapanalá, 1.5 km
NNE, 15°58′23″N, 95°43′33″W, 28 Jul 1999 (fl), Rivera H. et al. 1642 (NY).
GUATEMALA. Chiquimula: between Ramírez and Cumbre de
Chiquimula, road between Chiquimula and Zacapa, 400–600 m, 15 Oct
1940 (fr), Standley 74557 (F, US). Jalapa: Jalapa, 4,450 ft, 7 Jan 1908 (fr),
Kellerman 7052 (F). Zacapa: road between Agua Blanca and Cumbre de
Chiquimula, 350–500 m, 15 Oct 1940 (ster), Standley 74426 (F).
HONDURAS. Comayagua: Comayagua, Comayagua valley, Río Humuya,
El Edén, 28 Sep 1969, Molina R. 24532 (NY); NW of Comayagua, 2,100 ft,
1 Aug 1962 (fl), Webster et al. 12723 (WIS). La Paz: 4 km O de la ciudad
de La Paz, 700 m, 2 Sep 1979 (fr), Soto Valerio 81A (MEXU, MO).
Lempira: Gualan, along Honduras Road, 20 Jun 1909 (fl), Deam 6375
(GH, MO, US).
7. SOLANUM TRIUNFENSE S. Knapp, sp. nov.—TYPE: MEXICO.
Chiapas: Mcpio. Jaltenango/Mapastapec, Reserva El
Triunfo (Campamento/HQ 15°39′N, 92°48′W), path to
Palo Gordo, 1,900 m, May 1989 (fl), M. Heath & A. Long
MA129 (holotype: MEXU [MEXU-877597]; isotype: BM
[BM001118119]).
Diagnosis—Like S. cobanense J. L. Gentry but differing in
densely puberulent new growth and buds, long-attenuate
leaf bases, calyces completely enclosing the buds until just
before anthesis and tearing into irregular lobes at flowering.
Fig. 14. Distribution of Solanum refractum.
1126 SYSTEMATIC BOTANY [Volume 40
Woody vine or scandent shrub to 3 + m long, the stems
0.2–0.7 cm in diameter; internodes (2–)4–10 cm long, puberu-
lent with 0.2 mm long simple uniseriate few-celled tri-
chomes, these occasionally furcate; prickles 0.5–1.5 mm long,
stout, recurved, sometimes absent on young shoots near
the inflorescence. Sympodial units plurifoliate, not geminate.
Leaves simple, the blades 6–15 × 2.5–6.5 cm, elliptic, widest
at the middle, chartaceous, both surfaces minutely puberu-
lent with simple or occasionally furcate uniseriate 2–5-celled
trichomes ca. 0.5 mm long, often armed abaxially on midrib
with recurved prickles; major lateral veins 6–9 on each side;
base acute; margin entire; apex acute; petioles 0.5–3 cm,
minutely puberulent with simple uniseriate trichomes like
those of the leaf surfaces, sometimes armed with recurved
prickles. Inflorescences 5–10 cm long and wide, terminal,
few-branched, with 10–15 flowers, the axes puberulent with
simple or furcate uniseriate 2–7-celled trichomes, not prickly;
peduncle 1–3 cm; rachis 1–3 cm; pedicels 17–35 mm, articu-
lated at the base, evenly spaced 2–5 mm apart; fruiting pedi-
cels not known. Flower buds ellipsoid, the corolla completely
enclosed in the calyx until shortly before anthesis. Calyx
campanulate, spreading, tearing into irregular lobes, the
tube 1–3 mm, the lobes 5–8×4–6 mm, pubescent with simple
or occasionally furcate uniseriate 2–4-celled trichomes; fruiting
calyx not known. Corolla to 4.5 cm in diameter, spreading
at anthesis, purple or violet, membranous or slightly fleshy,
stellate, lobed 1/2 way to the base, the lobes 1–1.5 × 0.6–1cm,
broadly deltate, glabrous on both surfaces, the tips cucul-
late and minutely papillate abaxially. Stamens equal in size,
8–11 mm long; filaments 1–2 mm long, glabrous; anthers
7–10 × 1–1.5 mm, ellipsoid, connivent, yellow, poricidal
at the tips, the pores lengthening to slits with age. Ovary
glabrous; style in long-styled flowers ca. 13 mm long, straight,
glabrous, in short-styled flowers ca. 3 mm long; stigma capi-
tate. Fruit and seeds not known. Fig. 15.
Distribution and Habitat (Fig. 16)—Solanum triunfense
is endemic to the Sierra Madre Occidental in the State of
Chiapas, Mexico, in evergreen humid forests at elevations
from 1,800–1,900 m. Most collections are from the El Triunfo
Biosphere Reserve.
Phenology—Solanum triunfense has been collected in flower
from February to June; fruiting specimens are not known.
Common Names—Mexico. Chiapas: “quistan”(Heath &
Long MA129).
Preliminary Conservation Status (IUCN 2014)—Endan-
gered (EN) B1,2a,b iii; EOO 493 km
2
(EN); AOO 16 km
2
(EN); 4 locations. Solanum triunfense is assessed as endan-
gered based on the low extent of occurrence and area of
occupancy and the threats to the habitat in the region
where it occurs. It does occur in a protected area (Reserva
El Triunfo, a UNESCO Biosphere Reserve) so is afforded
some protection there.
Solanum triunfense is very similar to S. cobanense, and has
also been identified in herbaria as S. wendlandii. It differs
from S. cobanense in its buds that are completely enclosed in
the calyx until just before anthesis (Fig. 15), the pubescence
of simple uniseriate and sometimes furcate trichomes on the
new growth, and in the attenuate leaf bases. The flowers of
S. triunfense are stellate, as opposed to the rotate to pentago-
nal flowers of S. wendlandii. Plants of S. wendlandii collected
at very early anthesis could be mistaken for S. triunfense, but
lack the pubescence on the new growth that is characteristic
of this new species.
Specimens Examined —MEXICO. Chiapas: SW side of Cerro Mozotal,
11 km NW of the junction of the road to Montozintla along the road to
El Porvenir and Siltepec, 25 Jun 1972 (fl), Breedlove 25708 (DS, MEXU,
MO); Mpio. La Concordia, El Triunfo Reserve, trail WSW from Palo
Gordo towards Finca Catarina, 15°40′N, 92°51′W, 1850 m, 25 Feb 1990
(fl), Hampshire et al. 694 (BM); El Triunfo, S edge, 29 Apr–1 May 1960 (fl),
MacDougall s.n. (MEXU); Mpio. Escuintla, Santa Rosa, Ojo de Agua, 2
Jun 1948 (fl), Matuda 17838 (DS, F).
8. Solanum wendlandii Hook. f., Bot. Mag. 113: pl. 6914.
1887.—TYPE: Cultivated at Kew in the Water Lily House,
28 Jul 1886, Anonymous s.n. (neotype, designated here: K
[K000195647]; isoneotype: K [K000195646]).
Solanum mazatenangense Coult. & Donn. Sm., Bot. Gaz. 37:
421. 1904.—TYPE: GUATEMALA. Retalhuleu: San Felipe,
2050 ft, Apr 1892, J. Donnell Smith 2669 (lectotype, des-
ignated here: US [US-1324650, barcode US00840548];
isolectotypes: F [F-267128], GH [GH0077513], K
[K000195648, K000195649], NY [NY00139006], US [US-
1324649, US-258628]).
Solanum tlacotalpense Sessé & Moc., Fl. Mexic., ed. 2. 1894.—
TYPE: MEXICO.Veracruz: “ad fluviorum Tuxtlentium”
[on the banks of Río Tuxtla, presumably near San
Andrés Tuxtla], M. Sessé & J. Mociño s.n. (lectotype, des-
ignated by Knapp 2008b, pg. 20: MA [MA-604681, F neg.
48334]; isolectotypes: MA [MA-604680, F neg. 483333,
MA-604682, F neg. 48335]).
Solanum unguis-cati Standley, Publ. Field Columbian Mus.,
Bot. Ser. 4: 320. 1929.—TYPE. HONDURAS. Atlántida:
Lancetilla Valley, near Tela, 50 m [“20–600”on label],
9 Mar 1928 [“Dec. 6, 1927–Mar. 20, 1928”on label],
P. Standley 56726 (holotype: F [F-581029]).
Woody (sometimes herbaceous) vine or canopy liana to
20 + m long, the stems 0.25–0.8 cm in diameter, prickly, the
prickles often lacking on herbarium collections of flower-
ing material; internodes 1–10 cm long, glabrous; prickles to
1 mm long, stout, recurved, sometimes absent on terminal
shoots near the inflorescence. Sympodial units plurifoliate,
not geminate. Leaves highly variable in shape on the same
plant, ranging from simple (usually near the inflorescences)
to pinnately compound, the blades 10–20 × 8–15 cm, usually
ovate (rarely elliptic), widest at the base, chartaceous, gla-
brous adaxially and abaxially, or occasionally the abaxial sur-
face puberulent with unbranched hairs, often armed abaxially
on midrib with curved prickles; major lateral veins 5–8on
each side; base rounded to acute; margin entire or pinnatifid
or pinnate with 3–7 lobes or leaflets, the terminal and adjacent
lateral leaflets broadly confluent; apex acuminate to rounded;
petioles 1.5–7 cm, glabrous, often armed with curved prickles.
Inflorescences 10–20 cm long and wide, terminal, many times
branched (rarely unbranched), with 20–30 flowers, with either
staminate or pistillate flowers on a single individual, gla-
brous, not prickly, or occasionally with a few prickles at
the very base; peduncle 5–10 cm; rachis 5–20 cm; pedicels
15–20 mm and slender at anthesis, articulated at the base;
fruiting pedicels elongating to 3–5 × 0.3 cm and becoming
woody. Flower buds narrowly ellipsoid, the corolla long
exserted from the calyx tube, the calyx tube in bud unlobed
with deep papery sinuses regularly tearing before anthesis.
Calyx campanulate, spreading, the tube 2–9 mm, the lobes
ca. 3 × 2 mm, rounded or quadrate with a distinct apiculate
tip, glabrous to puberulent on the tips. Corolla 2.5–6(–9) cm
in diameter, spreading at anthesis, pale lavender to deep
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1127
Fig. 15. Isotype specimen of Solanum triunfense from Mexico (M. Heath & A. Long MA129, BM). Reproduced with permission of the Natural History
Museum, London.
1128 SYSTEMATIC BOTANY [Volume 40
purple, membranous, rotate-pentagonal, lobed ca. 1/4–1/3
of the way to the base, the lobes 0.7–1.2 × 1.2–1.7 mm,
broadly deltate, acuminate at the tips, glabrous abaxially
and adaxially. Stamens 5–10 mm long; filaments unequal,
with four 1–1.5 mm long, the fifth 2.5–4 mm long, glabrous;
anthers nearly equal, 4–10 × 1.5–2 mm, the one on the
longer filament slightly larger, somewhat to strongly taper-
ing, connivent, cream or pale purplish blue, sometimes
yellowish cream at base and pale purple distally, poricidal
at the tips, the pores lengthening to slits with age. Ovary
glabrous; style in long-styled flowers ca. 12 mm long, straight,
glabrous, in short-styled flowers 2–4 mm long; stigma trun-
cate. Fruit a globose berry, 4.5–10 cm (dry, to 20 cm in diame-
ter when fresh fide Christenhusz et al. 5372) in diameter,
globose, green with green to white pulp, the pericarp gla-
brous, rugose when dry. Seeds 50–100 per berry, 4–5×
6–8 mm, strongly flattened-reniform, light reddish brown, the
surfaces minutely pitted, the testal cells pentagonal. Chromo-
some number: n = 12 (Fedorov 1974). Figs. 2D–F, 17 –19.
Distribution and Habitat—Solanum wendlandii is proba-
bly native from southern Mexico to Panama in wet and
semi-deciduous forests at elevations from 400–2,200 m. It is a
commonly cultivated and sometimes naturalized vine in tropi-
cal and subtropical areas worldwide. Most cultivated plants
apparently never bear female-fertile (long-styled) flowers and
are possibly clones of plants with only staminate flowers
(see below).
Phenology—Solanum wendlandii flowers throughout the
year; fruiting specimens have been collected in its putative
native range in January and February.
Common Names—Mexico. Chiapas: “kishtan”(Matuda
17481); Tabasco: “cola de gato”(Rovirosa 885); Guatemala.
Retalhuleu: “kishtan”(Cosminsky 112 and 128).
Preliminary Conservation Status (IUCN 2014)—Least
Concern (LC). We have not evaluated S. wendlandii using
GeoCat due to the large number of cultivated and/or natu-
ralized specimens whose status is not clear from collect-
ing information. We do consider it of some conservation
Fig. 16. Distribution of Solanum triunfense.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1129
Fig. 17. Staminate-flowered specimen of Solanum wendlandii from Panama (R. Hampshire & C. Whitefoord 307, BM). Reproduced with permission of
the Natural History Museum, London.
1130 SYSTEMATIC BOTANY [Volume 40
Fig. 18. Pistillate-flowered specimen of Solanum wendlandii from Guatemala (M.J.M. Christenhusz et al. 5372, BM). Reproduced with permission of
the Natural History Museum, London.
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1131
Fig. 19. Lithograph of Solanum wendlandii illustrating the plant from which the neotype specimen was probably taken (Curtis’s Botanical Magazine
113: pl. 6914. 1887). Reproduced with permission of the Library of the Natural History Museum, London.
1132 SYSTEMATIC BOTANY [Volume 40
concern, however, in its presumed area of native distribu-
tion, given the few fruiting specimens (and thus contributors
to future native populations) we have seen.
Solanum wendlandii is quite variable in appearance. Most
herbarium specimens, collected from flowering twigs, have
only simple, entire leaves while much of the rest of the vine
has pinnatifid to pinnate leaves. Simple-leaved collections
of S. wendlandii can be confused with S. cobanense, but the
leaves of the latter are more coriaceous and the flowers are
stellate and very fleshy rather than pentagonal in outline
with thin petal tissue like those of S. wendlandii.Non-
flowering pinnately-leaved specimens of S. wendlandii can
be very similar to S. refractum, but lack the densely prickly
inflorescence of that species.
Flower size in S. wendlandii is extremely variable, but this
is in part due to flower growth throughout anthesis. Flowers
of this species last approximately three days (Shelly et al.
2000), and on the first day they are both darker purple and
smaller than on subsequent flowering days (see Figs. 2D, 19).
Plants with small-flowered inflorescences (young ones with
flowers only one day old) can be confused with S. triunfense
and S. bicorne, but both of these species have stamens of
equal size.
The breeding system of this species has not been closely
investigated. Nearly, if not all, cultivated material seems to
bear only staminate flowers with vestigial styles and fruit from
cultivated collections is almost never seen (although immature
fruits are described in the protologue; Hooker 1887). Fruiting
material is seldom collected from the wild, perhaps because
the plants only produce their large fruit upon reaching the
canopy. The species is possibly dioecious, this may in part
account for the paucity of fruiting collections.
Solanum wendlandii is apparently native from southern
Mexico to western Panama in wet forests of the Atlantic
Coast, but the exact range is difficult to discern since it is
seldom collected in the wild, is widely planted for the
showy panicles of large flowers, and label data usually do not
specify whether the plants were cultivated or truly wild. Col-
lections outside this range seem to be either escapes from cul-
tivation or merely cultivated plants, although this is not
always stated clearly on herbarium labels. We have not cited
these numerous collections here, but full details can be found
on the Solanaceae Source website (www.solanaceaesource.org).
We have seen cultivated or perhaps naturalized material from
the following countries outside its native range: Argentina,
Australia,Bermuda,Brazil,Cameroon,China,Colombia,Cuba,
Dominican Republic, Ecuador, Egypt, Haiti, India, Jamaica,
Kenya, Malawi, Paraguay, Peru, Philippines, Puerto Rico,
Singapore, Sri Lanka, Tanzania, United Kingdom, United
States of America, Venezuela, and Zimbabwe.
Solanum wendlandii is reportedly used in cooking in
Guatemala (“Leaves and flowers cooked”fide Cosminsky
112 and 128) and the plants are reported in the Flora of
Guatemala to be edible (Gentry and Standley 1974). In at
least one collection (Nee & Taylor 28770) from Veracruz,
Mexico,however,itwasnotedthatthefruitsarenoteaten.
In this location, where the plants seem to be native, the
green fruits were pecked open by birds, which remove the
seeds and green pulp, leaving only an empty shell (Nee &
Taylor 29770).
No herbarium specimens were cited in the protologue
of S. wendlandii (Hooker 1887). The neotype collection was
made by an unknown collector from a cultivated plant
grown in England at the Royal Botanic Gardens, Kew (see
Fig. 19). The species was described from living plants by
Hooker (1887) as “Living plants of this beautiful Solanum
were sent to the Royal Gardens [Kew] in 1882 by Dr.
Wendland, Director of the Royal Gardens at Herrenhausen,
Hanover, with the information that it is a native of the cold
regions of Costa Rica, where it climbs trees. This habit it has
retained at Kew, where it ascends to the rafters of the Water
Lily House, and flowers profusely.”
Two specimens at Kew are labeled “Type specimen of Bot.
Mag. t. 6914!”and dated 28 July 1886 in an unknown hand-
writing (not Joseph Dalton Hooker’s); one (K000695647) has
stem, leaves and one opened out flower, while the other
(K000695646) has only stem and leaves. We select the more
complete K000195647 as the neotype for S. wendlandii.Another
specimen collected from the Water Lily House in 1892
(K000195646) may have come from the same plant, but is not
type material. Edmonds (2012) incorrectly cited a Kew holo-
type for S. wendlandii, and did not effectively typify the name.
Six Guatemalan collections were cited in the protologue of
S. mazatenangense:Shannon 618,Shannon 624,Heyde & Lux
4735,Heyde & Lux 3442,Donnell Smith 2263 and Donnell
Smith 2669. We have chosen the US duplicate (US-1324650)
of Donnell Smith 2669 as the lectotype of this species, as it is
the best preserved and most widely distributed of the collec-
tions cited. It is likely that Donnell Smith used the US sheet
in his description of the material annotated by Coulter as
this species.
Representative Specimens Examined —(putative native range only;
cultivated and naturalized specimens outside this range can be viewed
in Solanaceae Source, http://www.solanaceaesource.org) —MEXICO.
Chiapas: Mpio. Unión Juárez, Rancho La Libertad, km. 22 carr. á Unión
Juárez, 25 Mar 1992 (fl), Aquino V. 110AA (MEXU); Mpio. Tuxtla Gutiérrez,
El Sumidero, 22 km N of Tuxtla Gutiérrez, 4,500 ft, 9 Apr 1965 (fl),
Breedlove 9613 (F, US); Mpio. Ocozocoautla, Curva de la Gringa, 2 km
adelante de Ocozocoautla, carretera Ocozocoautla–Tuxtla Gutiérrez, 600 m,
10 May 1983 (fl), Calzada et al. 10125 (MEXU); Mpio. Tapachula, 7.5 km
de Tapachula, carretera Costera, 14°55′N, 92°29′W, 160 m, 17 Jul 1986 (fl),
Cigarroa MA2 (MEXU); Mpio. Unión Juárez, volcán Tacaná, camino de
Talquián a la cima del volcán, 2,200–2,400 m, 6 Feb 1987 (fr), Martínez S.
et al. 19677 (MEXU); Trapichito, Comitán, 1,350 m, 2 Jun 1945 (fl), Matuda
15671 (F, MEXU, MO); Mpio. Tenejapa, río de “Cruz Pilal”, 900 m, 1 May
1982 (fl), Shilom Ton 4230 (MEXU, NY); Mpio. Tapachula, Cantón El
Tesoro, 250 m, 17 Apr 1985 (fl), Ventura & López 1527 (MEXU); Mpio.
Tuxtla Chico, Ejido Francisco Villa, 4 Apr 1986 (fl), Ventura & López
3421 (MEXU). Oaxaca: Dtto. Miahuatlán, Mpio. San Jerónimo Coatlán,
Espuelas de San Antonio, 13.5 km SW de San Jerónimo Coatlán, 16°12′N,
96°57′W,1,950m,17May1988(fl),Campos V. 1824 (MEXU); Distr.
Ixtlán, 4 km N de Vista Hermosa entre Metates y Vista Hermosa, 1,340 m,
17 Apr 1982 (fl), Cedilla T. et al. 1241 (MEXU); carretera Oaxaca–Puerto
Escondido, km. 183, 16 Apr 1963 (fl), MacDougall s.n. (MEXU); carretera
Oaxaca a Tuxtepec, km. 100, 21 Apr 1963 (fl), MacDougall 512 (MEXU);
Mpio. Comaltepec, Vista Hermosa, km. 131 carr. Tuxtepec–Oaxaca, 1,700 m,
24 May 1966 (fl), Martínez C. 838 (MEXU); Mpio. San Jerónimo Coatlán,
Dto. Miahuatlán, Espuelas de San Antonio, 13.5 km SW de San Jerónimo
Coatlán, brecha a Piedra Larga, 2,050 m, 27 Jun 1990 (fl), Tor i z & C a m p o s
936 (MEXU); Cerro Espino, Finca Montecristo entrando por Chacalapa
5 km N de Pochutla carr. a Oaxaca, 31 May 1984 (fl), Torres C. et al. 5239
(MEXU); Mpio. Santiago Lachiguiri, Dto. Tehuantepec, E de crucero
Guadalupe, 12.5 km NE de Santiago Lachiguiri, 16°43′N, 95°30′W,
1,400–1,700 m, 9 May 1991 (fl), Torres C. 13967 (MEXU). Puebla: Bosque
Ajenibre, May 1952 (fl), Matuda 1 (MEXU).QuintanaRoo:Mpio.F.
Carillo Puerto, F. Carillo Puerto, 8 Jun 1989 (fl), cult., Cabrera 16494 (CIQ,
MEXU). San Luis Potosí: Mpio. Aquismón, Tampaxal, 1 Aug 1979 (fl),
Alcorn CO1 (MEXU). Tabasco: Mpio. Tacotalpa, 3.5 km adelante de la
panga de Oxolotan camino hacia Oxolotan, 22 May 1983 (fl), Cowan
et al. 3994 (MEXU, NY); Cerro de Coconá, 19 Jul 1890, Rovirosa 885
(K). Veracruz: Mpio. San Andrés Tuxtla, Cerro Mastagaga, ca. 13 km
NE de San Andrés Tuxtla, 18°35'N, 95°09'W, 1,200 m, 1 Mar 1972 (fl),
Beaman 5818 (F, XAL); Mpio. Hidalgotitlán, brecha Hnos. Cedillo–La
Escuadra, 17°16'N, 94°37'W, 152 m, 6 May 1974 (fl), Brigada Vázquez 583
2015] CLARK ET AL.: SOLANUM SECTION ACULEIGERUM 1133
(MEXU, MO, XAL); Mpio. Hidalgotitlán, km. 1–6 camino Cedillo–La
Laguna, 140 m, 17°17′N, 94°30′W, 12 Jun 1974 (fl), Dorantes 3193 (F),
Basuras, km. 18 Catemaco–Las Palmas, González Leija & Garza 3126
(MEXU); Mpio. San Andrés Tuxtla, Est. Biol, 95°04–09′W, 150 m,
8 Aug 1968 (fl), Martínez Calderón 1740 (F, GH, INIF, MEXU, NY, XAL);
Mpio. Naolinco, 3 km. W of San Antonio Paso del Toro, 19°35'N,
96°52'W, 850 m, 14 Jan 1984 (fr), Nee & Taylor 28770 (CORD, F, G,
MO, NY, XAL); Mpio. Hidalgotitlán, Campamento La Laguna, 100 m,
17°17'N, 94°30'W, 1 Mar 1984 (ster), Nee 29728 (F, NY, XAL); Mpio.
Xico, San Marcos de León, 19°26'N, 96°57'W, Ortega O. 2311 (XAL);
1kmSofMontePío,18°38′N, 95°05′W, 10 Jul 1978 (fl), Poole et al.
1433 (MEXU); Mpio. San Andrés Tuxtla, Cd. Madero, carretera a San
Martín, 24 Apr 1983 (fl), Ramamoorthy 3552 (MEXU); Mpio. Uxpanapa,
Esfuerzo Nuevo, 17°10′N, 94°21′W, 230 m , 2 M ay 1996 (f l ), Rivera H. &
Escobedo 66 (MEXU);Mpio.SantiagoTuxtla,LomaQuemada,350m,
11 May 1977 (fl), Ventura A. 14022 (F,MEXU),Mpio.Naolinco,Almolonga,
700 m, 15 Dec 1981 (fr), Ventura A. 19238 (MEXU, XAL), Mpio. Tlapacoyan,
El Embarcadero, 150 m, 9 Jun 1983 (fl), Ventura A. 20274 (MEXU, XAL);
Cerro de Tuxpango Ixtaczotitlán, 10 km SE of Orizaba, 29 May 1944 (fl),
Vera S . 2968 (MEXU). Yucatán: 6 km antes de Oxholom Samahil, 20°40'N,
89°W, 17 Mar 1987 (fl), Góngora 377 (CIQ, MEXU).
GUATEMALA. Alta Verapaz: near Tactic, above bridge across Río
Frío, 1400–1500 m, 30 Mar 1941 (ster), Standley 90308 (F). Guatemala:
Arrazola, 1600 m, May 1892 (fl), Heyde & Lux 3442 (F, US); Guatemala
City, 1,530 m, 3 May 1970 (fl), Harmon 2280 (MO); Calderas, 5 Jul 1941
(fl), Johnston 1896 (F). Escuintla: Escuintla, 1,100 ft, Apr 1890 (fl), Donnell
Smith 2263 (BM, US). Huehuetenango: Barillas, along the road between
Barillas and Malpais, 1,554 m, 28 Feb 2009 (fl, fr), Christenhusz et al. 5372
(BM, NY); between Xoxlac and Nucapuxlac, Sierra de los Cuchumatanes,
1,650–2,500 m, 17 Jul 1942 (ster), Steyermark 48926 (F). Jutiapa: Cuesta
de la Conora, between San José Acatempa and Río de los Esclavos,
900–1,200 m, 21 Dec 1938 (ster), Standley 60614 (F). Petén: Chichi, 9 Apr
1933 (fl), Lundell 2751 (F); La Libertad, 23 Apr 1933 (fl), Lundell 2984 (F,
GH). Quetzaltenango: Fuentes Georginas, W slope of Volcán de Zunil,
2,850 m, 4 Mar 1939 (ster), Standley 67476 (F); Los Positos, SW of San
Martín Chile Verde, 1,500 m, 8 Mar 1939 (ster), Standley 67932 (F);
above Mujulia, between San Martín Chile Verde and Colomba, 1,800 m,
1 Feb 1941 (ster), Standley 85541 (F); Finca Pirineos, between Santa
María de Jesus and Calahuache, 1,300 m, 20 Jan 1940 (fl), Steyermark
34499 (F). Quiché: Chichicastenango, 8 Apr 1949 (fl), Beer s.n. (CM).
Retalhuleu: Finca San Luis, Dec 1974 (fl), Cosminsky 112 (F); Finca San
Luis, San Sebastián, 1,000–1,500 ft, Jan 1976 (ster), Cosminsky 128 (F);
Las Delicias, S of Retalhuleu, 200 m, 22 Feb 1941 (fl), Standley 88039 (F).
Sacatepéquez: 5 km E of Antigua, 1700 m, 24 May 1970, Harmon 2358
(MO); Embaulada, 5,500 ft, Nov 1891 (fl), Heyde & Lux 4735 (F-2 sheets,
US); Las Lajas, 1,200 m, 28 Nov 1938 (ster), Standley 58087 (F). San
Marcos: between Canjula and La Unión Juárez, SE portion of Volcán
Tac a na, 2,0 0 0–3,000 m, 22 Feb 1940 (fl), Steyermark 36504 (F, US); between
Tajumulco and Tecutla, 9 miles S and W of Tajumulco, NW slopes of
Volcán Tajumulco, 1,800–2,500 m, 27 Feb 1940 (ster), Steyermark 36751 (F);
Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo,
1,800–2,400 m, 10–18 Dec 1963 (fl), Williams et al. 25977 (F); Tajumulco
Volcano, 8–10 km W of San Marcos, 2,300 m, 31 Dec 1964–1 Jan 1965
(fl), Williams 26935 (F). Santa Rosa: Cuilapa, 895 m, 20–27 Nov 1940
(fr), Standley 78130 (F). Sololá: Santa Bárbara, 410 m, Aug 1891, Shannon
618 (US). Suchitepéquez: Las Animas, 200 m, Sep 1891, Shannon 624
(US); SW slopes of Volcán Zunil, Finca Montecristo, SE of Santa María
de Jesús, 1,200–1,300 m, 31 Jan 1940 (ster), Steyermark 35231 (F). Zacapa:
Zacapa, 225 m, 23 Apr 1939 (ster), Standley 71996 (F); Quebrada
Alejandria, summit of Sierra de las Minas, Finca Alejandria, 2,500 m,
13 Oct 1939 (ster), Steyermark 29864 (F).
HONDURAS. Atlántida: Lancetilla Valley, near Tela, 20–600 m, 6 Dec
1927–20 Mar 1928 (ster), Standley 53328 (F, GH, US). Comayagua:
Comayagua Valley, 22 Sep 1973 (fl), Hazlett 853 (MO); Río Selguapa,
El Taladro, 650 m, 27 Jun 1964 (fl), Molina R. 14320 (F, NY); Valle
Comayagua entre Las Mercedes y Villa de Flores, 650 m, 28 Jun 1964 (fl),
Molina R. 14367 (F, NY); Comayagua Valley between Comayagua and
Villa San Antonio, 570 m, 1 May 1947 (fl), Williams & Molina R. 12610
(F). Morazán: drainage of the Río Yeguare, entre San Antonio del Mico y
La Estancia, camino Río Cobre, 1,600 m, 29 Sep 1948 (fl), Molina R. 1152
(F); Suyapa, 1,100–1,200 m, 1 Aug 1950 (ster), Standley 22266 (F). Yoro:
entre Yoro y Morazán, 700 m, 11 May 1956 (fl), Molina R. 6924 (F);
Coyoles, 30 Jun 1938, Yuncker et al. 8160 (F, NY).
EL SALVADOR. Ahuachapán: P. N. El Imposible, camino al cerro
Dávila, 13°49′N, 89°56′W, 26 Sep 1991 (fr), Morales et al. 1284 (MEXU).
La Libertad: Mpio. Antiguo Cuscatlán, Laderas de la Laguna, 13°40′N,
89°15′W, 830 m, 18 May 1989 (fl), Cruz 195 (MEXU). San Salvador: San
Salvador, 650–850 m, 30 Mar–24 Apr 1922 (fl), Standley 22402 (GH, NY).
Santa Ana: Cerro Las Mesas, 10 km NO de la ciudad de Santa Ana,
14°02′N, 89°36′W, 850 m, cult., Linares & Martínez 2623 (MEXU).
NICARAGUA. Estelí: road to Laguna de Miraflores, 8 Jun 1975 (fl),
Atwood 320 (NY). Jinotega: Sierra W of Jinotega, along trail to Cerro de la
Cruz, 1,050–1,350 m, 27 Jun 1947 (fl), Standley 10277 (F); La Montanita
and Las Mesitas, W of Jinotega, 1,100–1,400 m, 29 Jun 1947 (fl), Standley
10416 (F). Madriz: Cerro Quisuca, 13°31′N, 86°31′W, 1,100–1,250 m,
Stevens & Montiel 17336 (MEXU).Managua:RepartoBelmonte,NOde
la ciudad de Managua, 49 m, 10 Aug 1979 (fl), Araquistain 6 (MEXU); Casa
Colorada near El Crucero, summit of Sierra de Managua, 800–900 m,
14–25 May 1947 (fl), Standley 8664 (F). Matagalpa: Finca La Castilla,
23 km NE de Matagalpa, carretera al Tuma, 6 May 1982 (fl), cult.,
Sandino 2810 (MEXU).
COSTA RICA. Alajuela: R.B. Monteverde, Río Peñas Blancas, 10°19′N,
84°43′W, 700 m, 27 Mar 1988 (fl), Haber & Bello 8350 (CR); Cantón de
San Ramón, Cordillera de Tilarán, Monteverde, San Gerardo Biological
Station, Sendero Congo, 1 km W of station, 10°22′0″N, 84°48′0″W,
1,200–1,300 m, 14 Mar 1995 (fl), Pennys 249 (CR, INB, MO); Arenal
Volcano, S slope of Cerro Chato, trail to summit, 25 Feb 1989 (fl),
Russell et al. 988 (CR, US). Guanacaste: El Dos de Tilaran, 54 km N,
Cerro La Chirripa, Atlantic slope rain forest, 10°25′N, 84°50′W, 1,000 m,
12 Apr 1986 (fl), Haber et al. 4422 (MO); Río Chiquito de Tilarán, 10 km N
Monteverde, 12 Nov 1986 (fl), Haber & Bello 6366 (CR). Heredia: Cantón
de Santo Domingo, town of Santo Domingo, 9°59”N, 84°06′W, 1,150 m,
2Sept1990(fl),Solomon 19142 (CR, INB, MO). Limon: near Rangalle,
Telire River watershed, 400–900 m, 25–29 Mar 1983 (ster), Hazlett 5211
(MO); Telire, Talamanca, 24–28 Mar 1983 (ster), Ocampo 4011 (CR).
Puntarenas: Zarcero de Álfaro Ruiz, 1,550 m, 21 Dec 1924 (fl), Brenes 4157
(F); Cantón de Coto Brus, P. I. La Amistad, Cordillera de Talamanca,
Estación Pittier, 9°01’30′N, 82°57′40″W, 1,680 m, 28 Jan 1995 (fl), Donzo
et al. 3 (CR, INB, MO); Zarcero, 5300 ft, 7 Sep 1937 (fl), Smith 319 (F). San
José: Sabanilla de Montes de Oca, Residencias Palas Atena, S side of Rte.
202, ca. 0.5 km W of Cristo, 9°57′N, 84°02′W, 1,310 m, 16 Jan 1987 (fl),
Grayum 7967 (CR, MO); San Ignacio de Acosta, Carretera 4, km 27–28,
cult.,, 1100 m, 17 Jan 1984 (fl), Khan et al. 292 (BM, CR); Cerro Pico Blanco,
4 km S of Escazu, 2,000 m, 12 Dec 1976 (fr), Lent 3987 (CR, F, MO, NY);
Cantón de Aserrí, Z. P. Cerros de Escazú, Cuenca del Río Poás, Mesón, en
la cuenca media del Río Lajas, 4 km aguas arriba, 9°52′15”N, 84°05′50″W,
1,500 m, 30 Jan 1994 (fl), Morales et al. 2312 (INB, MO).
PANAMA. Bocas del Toro: Caribbean slopes of Cerro Fabrega at foot
of ‘Falso Fabrega’in Palo Seco Reserve, second N-most tributary (on
map) of Río Culebre, Pavón Camp, 9°09’51”N, 82°39′41″W, 1,300 m,
20 Mar 2005 (fl), Monro & Cafferty 4802 (BM, INB, MEXU, MO, PMA).
Chiriquí: Palo Santo, 3 miles N of Volcán, 14 Feb 1971 (fl), Croat 13547
(F, MO); between Los Planes de Hornito and Fortuna Lake, 8°40′N,
82°14′W, 1,200 m, 4 Mar 1985 (fl), Hampshire & Whitefoord 307 (BM, MEXU).
Names not Validly Published
Solanum dichotomum Pav. ex Dunal, Prodr. [A. P. de Candolle]
13(1): 233. 1852. Nom. nud. pro syn. S. bicorne Dunal var.
angustifolium Dunal.
Solanum juciri Mart. ex Sendtn., Fl. Bras. [Martius] 10: 110.
1846. Nom. illegit. superfl. with S. alternatopinnatum
Steud. (1841), in synonymy.
Solanum laxiflorum Dunal, Prodr. [A. P. de Candolle] 13(1):
234. 1852. Nom. nud. pro syn. S. viridipes Dunal.
Solanum shannoni Coult., nom. nud. in sched. (Heyde & Lux
4735). This collection is a syntype for S. mazatenangense
Coult. & Donn. Sm. and the epithet also appeared as
a name only in the original description of S. unguis-cati
(Standley 1929), but with no description it does not
meet the requirements for valid publication.
Acknowledgments. We thank the herbaria cited in the text for
making their material (as specimens and digital images) available;
Nicholas Fumeaux and Lorenzo Ramella (G) for help finding and citing
specimens; Frances Crawford (K), Leandro Giacomin (UFOP), Tiina Särkinen
(E), Kim Watson (NY), and Maria Vorontsova (K) for photographs of
1134 SYSTEMATIC BOTANY [Volume 40
specimens and live plants; Patrick Meyer-Higgins and Pawel Ficinski
(BM) for assistance in geo-referencing; Mindy Syfert (BM) for producing
the maps; and Paloma Rojas (BM) for databasing much of the material
cited here. This project was funded by the National Science Foundation,
Planetary Biodiversity Inventory (PBI) Solanum: A worldwide treatment
(DEB-0316614) to LB, SK and MN; SK travel to Madrid was funded
by the SYNTHESYS Project http://www.synthesys.info/ financed by
European Community Research Infrastructure Action under the FP6
“Structuring the European Research Area”Programme. The Department
of Botany at the Smithsonian Institution’s National Museum of Natural
History kindly hosted JLC during his postdoctoral work as part of the
PBI Solanum project (2005–2006).
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