RESOLVING RELATIONSHIPS WITHIN THE CYPHOMANDRA CLADE (SOLANUM, SOLANACEAE)

Charity Goeckeritz, Christina Kilbane, Katie Sanbonmatsu (Undergraduates), and Elisabeth Morrill (High school student)
Mentor: Lynn Bohs
Department of Biology, University of Utah, 257 South 1400 East, Salt Lake City, Utah 84112, U.S.A

S. roseum INTRODUCTION RESULTS S. melissarum

The Solanaceae is an important flowering plant family that includes well-

The parsimony analysis resulted in 929 trees of 849 steps. There were 685
known economic species such as the tomato (Solanum lycopersicum), potato
characters, of which 183 (26.7%) were parsimony-informative.
(Solanum tuberosum), eggplant (Solanum melongena), and chili pepper
(Capsicum). Solanum, with about 1,500 species, is the largest genus within
All species of the Cyphomandra clade (sensu Bohs, 1994, 2001, 2005) form a
Solanaceae and one of the largest plant genera (Weese & Bohs, 2007). Previous
monophyletic group with the exception of S. graveolens.
molecular phylogenies have identified about 13 well-supported clades within
Solanum (Bohs, 2005). One of these is the Cyphomandra clade, a group consisting
Within the Cyphomandra clade, sections Pachyphylla and Cyphomandropsis
of nearly 50 Neotropical species. The Cyphomandra clade includes the cultivated
do not emerge as monophyletic groups. However, support for relationships is poor
tree tomato (Solanum betaceum), as well as several other species that produce
along the tree backbone and relationships at deep levels within the clade are not
edible fruits. The most consistent character that defines the Cyphomandra clade is
resolved.
the large amount of nuclear DNA in comparison to other Solanum species. S. latiflorum S. betaceum
New accessions of previously sampled species grouped together except for
While all species of the Cyphomandra clade are now part of the genus
the two subspecies of S. endopogon; these, however, were not distantly related to
Solanum, the clade originally obtained its name from the genus Cyphomandra,
each other on the S. splendens clade.
created by Otto Sendtner in 1845. However, several phylogenetic studies revealed
the genus to be nested within Solanum and as a result, all species of Cyphomandra
The analysis revealed the relationships of the species newly added to the
were transferred to Solanum (Bohs, 1995).
phylogeny:
The group is traditionally divided into two sections: Solanum section
Solanum zumbense was sister to S. obliquum. Both are morphologically
Cyphomandropsis and S. section Pachyphylla. Solanum section Pachyphylla
similar species native to the eastern Andean slopes from Colombia to Peru and
represents the members of the old Cyphomandra genus and is characterized by
adjacent areas of Brazil.
enlarged anther connectives that function as osmophores in some species. These
species are visited by male euglossine bees which collect volatile compounds from
Solanum luridifuscescens, S. paralum, and S. sycocarpum form a clade. All
the osmophores, presumably as precursors for their sex pheromones (Gracie,
are species from the Atlantic coastal rain forest from the states of Bahia to So
1993; Sazima et al. 1993). Solanum section Pachyphylla is the only group within the
Paulo. Solanum paralum and S. sycocarpum are morphologically similar and were
Solanaceae that is known to exhibit this male euglossine syndrome. Other Solanum
placed in S. section Pachyphylla. Solanum luridifuscescens was placed in S.
species, including those of S. section Cyphomandropsis, are buzz pollinated by
section Cyphomandropsis and is morphologically divergent from the other two
female bees.
species of the clade.
A previous study using ITS molecular sequences found the Cyphomandra
Solanum graveolens was excluded from Cyphomandra by Bohs (1994) on
clade to be monophyletic (Bohs, 2007). However, S. sections Cyphomandropsis
the basis of morphology. On the phylogeny it occurs outside the Cyphomandra
and Pachyphylla were not monophyletic in the most parsimonious trees. In the
clade in an isolated position among the outgroup taxa. It is morphologically
current study, we have added species and accessions not included in the analysis
enigmatic and does not fit in well with any of the known clades of non-spiny
of Bohs (2007) in order to further resolve species relationships within the
Solanums.
Cyphomandra clade and to provide insight into the monophyly of S. sections
Cyphomandropsis and Pachyphylla.
The relationships of S. hutchisonii, S. ovum-fringillae, S. pelagicum, and S.
jussari are not well resolved or supported on the ITS tree. S. jussari is only
known from its collecting locality in Bahia, Brazil and probably represents a new
species.

S. betaceum S. sciadostylis

S. graveolens S. splendens

Fig. 1. 50% majority rule consensus tree from parsimony

analysis of 106 taxa resulting in 929 trees of 849 steps. Numbers S. jussari S. betaceum S. amotapense
S. latiflorum S. glaucophyllum above the branches are bootstrap values from 100 pseudoreplicates.
Members of S. section Cyphomandropsis according to Bohs (2001)
are italicized; species of S. section Pachyphylla are in plain font. New
MATERIALS AND METHODS taxa or accessions not included in the phylogeny of Bohs (2007) are FUTURE DIRECTIONS
marked with asterisks.
Sampling.
All accessions of Cyphomandra clade species sampled in Bohs (2007) were included as well as a subset of
We are continuing to add taxa and sequence data to the phylogeny. In
outgroup taxa from that paper. We added 13 new accessions of species previously sampled and nine species not particular we are sequencing the waxy nuclear gene and the chloroplast trnT-F
included in the previous phylogeny. Most of these are rare species from Brazil or Peru that have been newly collected region to construct trees to compare to the ITS phylogeny.
in the field, including one newly described species, S. zumbense Bohs, and a potentially new Brazilian species here
called S. Jussari after its collecting locality.
REFERENCES CITED
Lab and analytical methods.
DNA was extracted from silica-dried leaves or from herbarium specimens and the ITS region was amplified Bohs, L. 1994. Cyphomandra (Solanaceae). Flora Neotropica. 63: 1-175.

via PCR using the primers described in Bohs (2007). Samples were cleaned and sequenced on an ABI automated
sequencer at the University of Utah. Sequences were edited and contigs assembled using Sequencher (Gene Codes
Corp.) and manual alignments were performed using Se-Al (Rambaut, 1996). Phylogenetic trees were produced
using PAUP* 4.0b10 (Swofford, 2002). Parsimony analyses were performed using a heuristic PAUP* search using the
TBR and MulTrees options with branch swapping limited to 1,000,000 rearrangements per replicate. A parsimony
bootstrap analysis was conducted using 100 pseudoreplicates with random addition replicates = 50, TBR, MulTrees,
and rearrangements limited to 1,000,000 per replicate.
Bohs, L. 1995. Transfer of Cyphomandra (Solanaceae) and its species to Solanum. Taxon 44: 583-587.
Bohs, L. 2001. A revision of Solanum section Cyphomandropsis (Solanaceae). Syst. Bot. Monogr. 61: 185.
S. stuckertii S. unilobum S. maternum Bohs, L. 2005. Major clades in Solanum based on ndhF sequence data. Pp. 2749 in: Keating, R.C., Hollowell, V.C. & Croat, T.B. (eds.), A Festschrift for William G.
DArcy: The Legacy of a Taxonomist. Missouri Botanical Garden Press, St. Louis, Missouri [Monogr. Syst. Bot. Missouri Bot. Gard. 104].
Bohs, L. 2007. Phylogeny of the Cyphomandra clade of the genus Solanum (Solanaceae) based on ITS sequence data. Taxon 56: 1012-1026.
Bohs, L. & Olmstead, R.G. 2001. A reassessment of Normania and Triguera (Solanaceae). Pl. Syst. Evol. 228: 3348.