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203
de Araújo & al. • Ch autems ia calc icola, gen. et sp. nov.TAXON 59 (1) • Februar y 2010: 203–208
INTRODUCTION
Generic boundaries in tribe Gloxinieae were reorganized by
Roalson & al. (2005a) on the ground of phylogenetic analyses
of molecular characters and morphological synapomorphies
(Roalson & al., 2005b). Gloxinieae now include some 20 genera
and approximately 170 species (Boggan & al., 2008; Roalson &
al., 2005a, 2008). The recent studies showed that several genera
of Gloxinieae, particularly Gloxinia L’Hér. as recognized by
Wiehler (1983), were not monophyletic. From the latter genus,
a new tribe Sphaerorrhizeae Roalson & Boggan was segregated
including the new genus Sphaerorrhiza Roalson & Boggan. For
the remaining taxa new generic names were proposed including
Gloxinella (H.E. Moore) Roalson & Boggan, Gloxiniopsis Roal-
son & Boggan and Nomopyle Roalson & Boggan, whereas gen-
era Mandirola Decne. and Seemannia Regel were resurrected.
In the framework of a detailed taxonomic survey of the
Gloxinieae in Brazil, a new taxon was discovered in Minas
Gerais, Brazil. The material was identified as belonging to
Gloxinieae based on the presence of scaly rhizomes, but did
not fit in any of the existing genera. In the present study we
describe the diagnostic features of this plant and explore its
affinity in Gloxinieae using sequences of the rpl16 intron,
rps16 intron, ncpGS, and ITS regions and the published ITS/
trnL-F dataset of Roalson & al. (2005b).
MATERIALS AND METHODS
Taxon sampling and
DNA
sequencing. —
Leaves from live
plants collected in the field or from cultivated specimens held
in the greenhouses of the Conservatoire & Jardin botaniques
de la Ville de Genève (Switzerland) were sampled and immedi-
ately dried in silica gel. Twenty-four accessions representative
of 20 Gloxinieae species and 14 genera (including the newly
described taxon) plus two outgroups were sequenced for the
rpl16 intron, rps16 intron, the trnL-F intron-spacer, a portion
of the plastid-expressed glutamine synthetase gene (ncpGS)
and the ribosomal DNA internal transcribed spacer (ITS).
Two outgroup species, Gesneria humilis L. and Rhytidophyl-
lum vernicosum Urb. & Ekman, were selected from the tribe
Gesnerieae, a sister lineage of Gloxinieae according to previ-
ous phylogenetic analyses (Zimmer & al., 2002; Roalson &
al., 2005b). A list of these 26 accessions and their vouchers is
provided in the Appendix.
DNA was isolated using the CTAB method of Doyle &
Doyle (1987) with the addition of 1% polyphenol polyvinyl-
pyrolidon. The primers used to amplify and sequence the rpl16
intron, trnL-F intron-spacer and the region between exons
7 and 11 of ncpGS are indicated in Perret & al. (2003). The
rps16 intron was amplified and sequenced using the primers
rpsF and rpsR2 (Oxelman & al., 1997). ITS was amplified and
sequenced with primers ITS 5P and ITS 8P (Möller & Cronk,
1997). Both amplifications and sequencing reactions follow
procedures described in Perret & al. (2003). All sequences
were aligned by eye, insertions/deletions (indels) were not
coded as separate characters. Newly acquired sequences have
been deposited in EMBL/GenBank.
Phylogenetic analyses and tests of alternative topolo-
gies. —
For parsimony analyses, all characters were equally
weighted and gaps were treated as missing characters. Heu-
ristic searches were conducted in PAUP* v.4.0b10 (Swofford,
2002), using 1000 random taxon addition sequences with tree-
bisection-reconnection (TBR) branch swapping and keeping
Chautemsia calcicola: A new genus and species of Gloxinieae
(Gesneriaceae) from Minas Gerais, Brazil
Andréa Onofre de Araújo,1 Vinicius Castro Souza1 & Mathieu Perret2
1 Departamento de Ciências Biológicas, Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo,
Av. Pádua Dias, 11, 13418-900, Piracicaba – SP, Brazil
2 Conservatoire & Jardin botaniques de la Ville de Genève, Impératrice 1, C.P. 60, 1292 Chambésy, Genève, Switzerland
Author for correspondence: Andréa Onofre de Araujo, aonofrearaujo@yahoo.com.br
Abstract
A new species of Gesneriaceae discovered in remnants of deciduous forests on limestone outcrops in Minas Gerais,
Brazil, is described and compared with morphologically related taxa. This plant presents the diag nostic features of the tribe
Gloxinieae, but a unique combination of mor phological traits distinguishes this taxon f rom previously described genera. Its
phylogenetic position was inferred based on analyzing DNA sequences variation of five loci: the rpl16 intron, rps16 int ron,
trnL-F intron-spacer, a por tion of the plastid-expressed glutamine synthetase gene (ncpGS ) and the ribosomal DNA internal
transcribed spacer (ITS). Molecular phylogenetic analyses confirm the position of this new species in the Gloxinieae, as a sister
lineage of a clade including the Brazilian genera Mandirola and Goyazia. However, tests using topological constraints do not
reject the alternative relationship that places this taxon with Gloxiniopsis in a monophyletic group. To accommodate this species
in the current generic circumscription of Gloxinieae, the new genus Chautemsia A.O. Araujo & V.C. Souza is created.
Keywords
Brazil; Brazilian Atlantic forest; Chautemsia; Gesneriaceae; Gloxinieae; molecular phylogeny; taxonomy
204
TAXON 59 (1) • Februar y 2010: 203–208de Araújo & al. • Chaute msia cal cicola , gen. et sp. nov.
up to 1000 most parsimonious trees per random addition rep-
licate. To evaluate internal support, heuristic searches were
completed for 1000 bootstrap replicates (BS; Felsenstein,
1985), with TBR swapping. The number of trees saved at each
replicate was limited to 1000.
Bayesian inferences were performed with MrBayes v.3.1
(Ronquist & Huelsenbeck, 2003). The combined dataset was
divided into three partitions: plastidic regions, ncpGS and ITS.
For each partition, the optimal model of DNA substitution
was selected by the Akaike information criterion (AIC) using
Modeltest v.3.7 (Posada & Crandall, 1998). In each case the
optimal model was the General Time Reversible model, with
rate heterogeneity modeled by assuming that some proportion
of sites are invariable and that rate of evolution at other sites is
modeled using a discrete approximation to a gamma distribu-
tion (GTR + I + Γ). All the parameters values were unlinked
across partitions and estimated during the Markov chain
Monte Carlo (MCMC) runs. Two independent analyses were
run from different random trees (Nruns = 2) with temperature
of 0.15. The chains were run for 10,000,000 generations, with
trees sampled every 100th generation. We discarded 25% of
the samples obtained during the f irst 1,000,000 generations.
Posterior probability (PP) and branch lengths were averaged
across remaining sampled trees. To examine potential incon-
gruences between datasets we inspected whether the topology
resulting from the combined analyses of all the regions con-
flicts with the topologies obtained through separate analyses
of the plastidic and nuclear sequences. Only conflicting nodes
with 70% BS or higher were considered.
The relevance of alternative relationships between the new
taxon and other Gloxinieae genera were tested using the “Shi-
modaira-Hasegawa test” (SH test; Shimodaira & Hasegawa,
1999; Goldman & al., 2000) implemented in PAUP* v.4.0b8
(Swofford, 1999). Topologies that force the monophyletic re-
lationships between Chautemsia and the members of the
other Gloxinieae genera were built in MacClade (Maddison &
Maddison, 1992). For each of these topological constraints, a
new heuristic search was performed using our 5-regions matrix
and the maximum parsimony criteria. Then, two subsets of trees
obtained with and without constraint were compared simultane-
ously with the RELL option of the SH test and 1000 bootstrap
replicates as implemented in PAUP* v.4.0b8 (Swofford, 1999).
We used the GTR model with proportion of invariant sites (I)
and gamma shape (G) parameters (six substitution types: A/C =
0.8511, A/G = 2.2387, A/T = 0.8533, C/G = 0.5712, C/T = 3.5860,
G/T = 1.0000, I = 0.5094, G = 0.6706, freqA = 0.3041, freqC
= 0.2038, freqG = 0.1995, freqT = 0.2926). The model and all
parameters were selected based on the results of analyses using
Modeltest v.3.7 and AIC (Posada & Crandall, 1998).
To test the inf luence of taxon sampling on the phylogenetic
position of Chautemsia, newly generated sequences for this
taxon were added to the ITS/trnL-F dataset of Roalson & al.
(2005b). New sequences for Gloxiniopsis racemosa (Benth.)
Roalson & Boggan, Goyazia petraea (S.M. Phillips) Wiehler
were also added to this dataset. Accessions for which either
ITS or trnL-F sequences were not available were excluded
from the dataset (i.e., Bellonia spinosa Swartz, Gloxinia
planalta Wiehler and Solenophora tuerckheimiana J.D. Sm.)
The resulting matrix includes an extensive sampling in the
tribe Gloxinieae with 59 species representing all the 19 genera
recognized by Roalson & al. (2005a). Phylogenetic analyses
of this combined ITS/trnL-F dataset were performed using
MrBayes v.3.1 with each region treated as a single data parti-
tion and following the procedures defined above.
RESULTS
Phylogenetic analyses. —
The combined matrix of all
DNA regions comprised a total of 4124 characters with 294
variable sites and 96 parsimony-informative characters. Of
these, 2273 (294 variable, 96 parsimony-informative) were
derived from plastidic sequences, 701 (185 variable, 68 par-
simony-informative) from ncpGS, and 650 (244 variable, 141
parsimony-informative) from ITS.
The
Bayesian 50% majority rule consensus tree of the
total combined dataset
is shown in Fig. 1. This topology is
identical to the one obtained using parsimony analyses. Values
of posterior probability and maximum parsimony bootstrap are
indicated at each node. The comparison of this tree with re-
sults derived from separate analyses of the nuclear and plastid
datasets indicated two supported topological incongruences
(BS > 70%). Both are in the plastid-based topology: one is re-
lated with the placement of Mandirola cf. rupestris (Gardner)
Roalson & Boggan within the Mandirola/Goyazia clade and
the other concerns the monophyly of the two Gloxinia species
with the Gloxinia/Seemannia clade. Because these topological
incongruences do not involve the newly described genus, only
results obtained from the complete dataset will be discussed.
Chautemsia appears sister of a clade including the Brazilian
genera Goyazia Taub. and Mandirola (Fig. 1). This relationship
has a posterior probability of 99% but a bootstrap support of
only 63%. This lineage is in turn sister with the Seemannia/
Gloxinia clade (PP = 100%, BS = 63%). The other relation-
ships within Gloxinieae agree with the topology presented in
Roalson & al. (2005b: f ig. 6) and Roalson & al. (2008: fig.
2). Unfortunately, the low supports for the basal relationships
within the Gloxinieae observed by Roalson & al. (2005b) were
little improved in our analyses despite the larger number of mo-
lecular characters used. The SH-tests of alternative topologies
forcing Chautemsia to be monophyletic with other Gloxinieae
genera, indicate significant differences in the likelihood scores
for all genera (P < 0.05) except Gloxiniopsis (P ≥ 0.06). In
further Bayesian analysis of the extended ITS/trnL-F matrix
of Roalson & al. (2005b), Chautemsia appears in an unresolved
position, as a single branch at the base of a large polytomy that
groups all the major lineages of South American Gloxinieae
(result not shown; see Roalson & al., 2005b).
Morphological traits. —
The new taxon displays the
following floral characters: frondose florescence, corolla
infundibular-cylindrical, erect in the calyx, floral anthocy-
anin absent (yellow only in the throat), corolla tube internally
glabrous except at the throat with glandular trichomes, nectary
divided into five finger-like glands. The fruit structure found
205
de Araújo & al. • Ch autems ia calc icola, gen. et sp. nov.TAXON 59 (1) • Februar y 2010: 203–208
Fig. . Bayesian 50% majority
rule consensus tree resulting
from the combined analysis of
the
rpl16 intron, rps16 intron,
trnL-F region, ncpGS and ITS
sequenced for 24 accessions in
Gloxinieae and two outgroups
in Gesnerieae
. Numbers above
the branches are PP values;
those below are bootstrap
values when ≥50%. Asterisks
indicate conflicting nodes with
BS > 70% between this tree and
the plastid-derived tree.
Table .
Morphological comparison of Chautemsia with other genera in Gloxinieae.
Character Chautemsia Gloxinia Gloxiniopsis Mandirola Seemannia Goyazia Diastema
Florescence Frondose Bracteose,
rarely frondo-
bracteose
Bracteose Frondose Fondose,
rarely frondo-
bracteose
Frondose Bracteose
Corolla color White with
yellow throat
White, purple,
pinkish or
partly maroon
White with
purple stripes
in throat
White and
partly purple,
violet or
pinkish
Red, orange
or dark pink
White or white
with violet
stripes, pink-
ish, purple
White with
purple mark-
ings on lobes
or reddish
Corolla shape Infundibular-
cylindrical
Campanulate,
campanulate-
infundibular
or cylindrical-
bilabiate
Campanulate Campanulate-
infundibular
or hypocrateri-
form
Cylindrical-
ventricose or
infundibular-
ventricose
Infundibular-
cylindrical
Infundibular-
cylindrical
Nectary 5 finger-like
glands
Absent or
annular with
5 lobes
Absent or
vestigial
5 lobed glands,
or annular
with 5 lobes or
unlobed
Annular
unlobed
5 lobed glands
or annular
unlobed
5 finger-like
glands
Capsule
consistence
Fleshy Dry Fleshy Dry Dry Dry Fleshy
Capsule
dehiscence
Dorsal;
splitting the
hypanthium
to the base
Apical Dorsal;
splitting the
hypanthium
to the base
Apical Apical Apical Dorsal;
splitting the
hypanthium
to the base
206
TAXON 59 (1) • Februar y 2010: 203–208de Araújo & al. • Chaute msia cal cicola , gen. et sp. nov.
in this species matches the definition of horizontal follicular
fruits (Weber, 2004). This kind of fruit is f leshy at maturity,
horizontally held with the hypanthium splitting dorsally and
completely to the base, exposing the seeds (Fig. 2). None of
these characters is exclusive of this taxon, but this set of char-
acters is not found in any other genus of Gloxinieae (Table 1).
DISCUSSION
According to the classification of Wiehler (1976) and his
identification key to tribes of Gesnerioideae (Wiehler, 1983),
Chautemsia fits in the tribe Gloxinieae. Indeed, the new taxon
displays all the diagnostic traits of Gloxinieae such as rhi-
zomes with f leshy leaf scales, opposite leaves and inferior
ovary. Moreover, our phylogenetic result clearly confirmed
that Chautemsia is nested within Gloxinieae (Fig. 1).
Combined analyses of five DNA regions support Chautem-
sia as the sister lineage of a clade comprising the genera Goya-
zia and Mandirola (Fig. 1). This grouping is geographically
consistent since species in this clade are endemic to central or
southeastern Brazil, outside of the range of other Gloxinieae
species. Chautemsia can be easily distinguished from other
Brazilian taxa based on features such as its white corolla,
horizontal follicular fruit and nectary composed of five long
finger-like and separate glands (Table 1; Fig. 2). The phylo-
genetic affinities of Chautemsia with Brazilian taxa should
be considered with caution. Indeed, the SH-tests indicated
that our data do not allow to reject the alternative relationship
that places Chautemsia and the monotypic genus Gloxiniop-
sis in a same clade (P = 0.06). Analysis of the phylogenetic
relationship of Chautemsia based on the taxonomically more
exhaustive ITS/trnL-F dataset of Roalson & al. (2005b) did
not provide further information about the generic affinities
of Chautemsia. These appeared as unresolved at the base of a
large polytomy including the main lineages of South American
Gloxinieae (Roalson & al., 2005b, 2008). Therefore, a more
confident placement of Chautemsia in Gloxinieae will need
additional data to help improve the supports of the basal rela-
tionships among the main Gloxinieae lineages.
Morphologically, Chautemsia most closely resembles
Diastema Benth., which also includes weak herbs with an
Fig. .
Chautemsia calcicola.
A,
habit;
B,
calyx and finger-
like nectar y glands (ovary and
corolla removed);
C,
longitu-
dinal section of inner corolla,
showing stamens and stami-
node;
D,
longitudinal sect ion
of outer corolla;
E,
lateral and
superior views of stigma;
F,
mature dehiscent fruit (seeds
removed). Scales: A = 1 cm;
B = 0.5 mm; C, D = 5 mm;
E = 1 mm; F = 2 mm. (Illustra-
tor Samira Rolim).
207
de Araújo & al. • Ch autems ia calc icola, gen. et sp. nov.TAXON 59 (1) • Februar y 2010: 203–208
infundibular-cylindrical corolla, a nectary of five long, finger-
like, separate glands, and horizontal follicular fruit, dehiscing
dorsally and splitting the hypanthium to the base (Weber, 2004).
Chautemsia can be distinguished from Diastema by the follow-
ing characters: frondose f lorescence with solitary flowers (vs.
bracteose), corolla white with unspotted lobes, and stigma sto-
matomorphic (vs. bilobed; Table 1). Furthermore, Chautemsia
occurs only in southeastern Brazil whereas Diastema is found
from Bolivia to Mexico and rarely occurs in Brazil (only in
Acre near the border of Peru). Our result confirms the finding of
Roalson & al. (2005b) that D. vexans H.E. Moore is segregated
from the rest of the genus (Fig. 1). However, neither of these two
lineages was found related with Chautemsia indicating that their
morphological similarities could be due to convergences. Beside
Diastema, other taxa share with Chautemsia a fleshy fruit with a
splitting hypanthium, e.g., Gloxiniopsis, Kohleria Regel, and the
monophyletic group comprising Gloxinella, Monopyle Moritz
ex Benth., Nomopyle and Phinaea Benth. (Kvist & Skog, 1992;
Roalson & al., 2008). None of these taxa have a nectary similar
to that of the new genus, however.
In their revision of the genus Kohleria, Kvist & Skog (1992)
stated: “As is often the case in genera of Neotropical Gesneria-
ceae, no single feature distinguishes the genus Kohleria from
related and similar genera in the tribe Gloxinieae, but the ge-
nus is set apart by a suite of characters.” In a similar way, the
unique combination of characters that define this new taxon
and its lack of close relatives in the tribe, leads us to recognize
this taxon as a new genus. With this addition, tribe Gloxinieae
now comprises 21 genera.
TAXONOMICAL TREATMENT
Chautemsia calc icola A.O. Araujo & V.C. Souza, gen. &
sp. nov. – Type: Brazil. Minas Gerais, Mun. Iguatama,
Fazenda Faroeste, próximo ao município de Arcos,
20°15′45.1″ S, 45°40′10.3″ W, 690 m alt., 13 Jan. 2005, A.O.
Araujo, V.R. Scalon, A. Chautems, G. Barriera & R. Tsuji
500 (holotype: ESA!; isotypes: G!, SPF!, SPFR!). – Fig. 2.
Affinis Diastemati Benth. nectario quinque glandulis di-
git i formibus et capsula carnosa dorsaliter omnino secedenta,
sed floribus axillaribus solitariis, lobis corollae plene candidis
et stigmate stomatoformi differt.
Perennial herbs, 4.0–8.5 cm high, with scaly rhizomes.
Stems terete, erect, unbranched, usually reduced to 3–4 nodes.
Internodes 0.2–3.2 cm long. Leaves opposite, anisophyllous,
sometimes strongly unequal in a pair, petiole 0.3–3.2 cm long,
blades 0.5–5.5 × 0.4–3.3 cm, elliptic, the basal ones frequently
broadly elliptic or circular, apex acute or obtuse, base sym-
metrical, cuneate or attenuate, margin serrate, sometimes
irregularly, adaxial and abaxial surface pubescent, discolor
green, 4–5 pairs of secondary veins. Florescence frondose,
epedunculate, flowers solitary, 1 or 2 flowers in the upper node,
sometimes another flower in the lower node, subtending leaves
1.3–3.1 × 0.3–1.7 cm. Flowers without bracteoles, with pedicel
erect-ascending, 0.9–1.5 cm long, glabrescent, zygomorphic,
floral tube conic, ca. 2 mm; calyx aestivation open, sepals free,
lobes equal, 0.1–0.3 × 0.1–0.2 cm, green, inside glabrous near
base and sparsely pubescent toward apex, outside pubescent to
densely pubescent, elliptic, sometimes obovate, with margin
entire; corolla infundibular-cylindrical, aestivation imbricate,
tube 0.9–1.5 cm long, inside glabrous, sparsely pubescent only
in the throat, with glandular, uniseriate, capitate, short or long
trichomes, outside sericeous, near base glabrous; tube white
to white-yellowish; throat not constricted, yellow, 0.3–0.5 cm
diameter; lobes white, broadly depressed-ovate with entire or
slightly sinuate margins, glabrous inside, outside glabrous to
sparsely sericeous, 0.3–0.5 cm long; stamens 4, included in the
corolla, 0.9–1.3 cm long, filaments glabrous, anthers all coher-
ent opening by longitudinal slits, staminode ca. 0.8 cm long.;
nectary of 5 long, finger-like, separate glands; ovary inferior;
style 1.2–1.4 cm long, sparsely pubescent, glabrous toward
apex; stigma stomatomorphic. Capsule 0.8–1.2 × 0.2–0.6 cm,
pubescent, ellipsoid, apex erect, fleshy, dehiscing dorsally and
splitting the hypanthium to the base, capsule without prominent
costae. Seeds without enlarged funicle.
Etymology. – The name of this genus (Chautemsia) honors
Dr. Alain Chautems, curator at the Conservatoire & Jardin
Botaniques de la Ville de Genève, who investigated Brazilian
Gesneriaceae for over 20 years. He deserves this homage in
recognition to his important contribution to the knowledge
of this family and to his deep involvement in training new
researchers.
Distribution and ecology. – Despite extensive investiga-
tions of the limestone flora in central and southeastern Brazil
made by the first collector of this taxon (Melo, 2008), the spe-
cies has been so far only found in the localities Arcos, Pains
and Iguatama in the region of Formiga, in the western part
of Minas Gerais. The species is restricted to deciduous forest
remnants on limestone outcrops between 600 and 800 m. We-
ber (2004) stated that horizontal follicular fruits are probably
associated with a rain-splash dispersal mechanism: when seeds
are exposed, they are washed away by water drops. In view of
the occurence of the new species on dripping rocks, this kind
of dispersal strategy seems quite appropriate.
Conservation status. – Endangered, based on the occur-
rence in an area smaller than 5000 km² and known by fewer
than five localities (IUCN, 2001).
Phenology. – The peak f lowering period was observed in
January and fruiting specimens were found in February and
March, matching the rainy season. During the dry season from
May to September, plants are dormant and reduced to scaly
rhizomes hidden in rock crevices.
Paratypes. – Brazil. Minas Gerais, Mun. Arcos, Fazenda
Faroeste, margem direita do Rio São Miguel, 1 Jan. 2003,
P.H.A.Melo & J.A. Lombardi 408 (BHCB, G); Mun. Pains, Fa-
zenda Amargoso, MG 439 Km 16, 20°15′05.7″ S, 45°39′46.7″ W,
6 Mar. 2003, P.H.A. Melo 446 (BHCB, G); Mun. Pains, fronteira
entre os municípios de Pains e Formiga, região da nascente do
Rio São Miguel, 20°27′22″ S, 45°39′15″ W, 700–800 m, 25 Jan.
2005, P.H.A. Melo & N.L. Assunção 1257 (BHCB, G); Mun.
Iguatama, Fazenda Faroeste, margem esquerda do Rio São
Miguel, 20°15′45″ S, 45°40′10″ W, 700–800 m, 25 Mar. 2005,
P.H.A. Melo & J.F. Martins 1315 (BHCB, G).
208
TAXON 59 (1) • Februar y 2010: 203–208de Araújo & al. • Chaute msia cal cicola , gen. et sp. nov.
Boggan, J.K., Skog, L .E. & Roalson, E.H. 2008. A review of the
Neotropical genera Amalophyllon, Niphaea, and Phinaea (Gesneri-
aceae-Gloxinieae). Selbyana 29: 157–176.
Doyle, J.J. & Doyle, J.L . 1987. A rapid DNA isolation procedu re for
small quantities of fresh leaf tissue. Phytochem. Bull. 19: 11–15.
Felsenstein, J. 1985. Confidence limits on phylogenies: An approach
using the bootstrap. Evolution 39: 783–791.
Goldman, N., Anderson, J.P. & Rodrigo, A.G.. 2000. Likelihood-
based tests of topologies in phylogenetics. Syst. Biol. 49: 652–670.
IUCN. 2001. IUCN Red List categories and criteria, version 3.1. IUCN
Species Survival Commission. Gland, Switzerland & Cambridge,
U.K.: IUCN
Kvist, L.P. & Skog, L.E. 1992. Revision of Kohleria (Gesneriaceae).
Smithsonian Contr. Bot. 79: 1–83.
Maddison, W.P. & Maddison, D.R.. 1992. MacClade, version 3.01.
Sunderland, Massachusetts: Sinauer.
Melo, P.H.A. 2008. Flora vascular relacionada aos afloramentos de
rocha carbonática no interior do Brasil. Master thesis, Universi-
dade Federal de Lavras, Minas Gerais, Brazil.
Möller, M. & Cronk, Q.C.B. 1997. Origin and relationships of Saint-
paulia (Gesneriaceae) based on ribosomal DNA internal tran-
scribed spacer (ITS) sequences. Amer. J. Bot. 84: 956–965.
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ACKNOWLEDGEMENTS
We are g rateful to Pablo Hendrigo Melo for com mun icating the
discovery of this plant, sending pictu res as well as dried and pick-
led mater ial and for guiding us to one of its localities. John Boggan
kindly provided material of Seemannia purpurascens g rown at the
Smith sonian Institution. We also thank Er ic H. Roalson for provid-
ing his alignment of DNA sequences and Fadil Avdija for his help in
the laboratory work. T he first author thanks Fundação de Amparo a
Pesquisa do Estado de São Paulo, International Association for Plant
Taxonomy and the Elvin McDonald Research Endowment Fund of the
Gesneriad Society, Inc. for suppor ti ng this research.
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Appendix.
Taxa sampled for the st udy. Samples ta ken from cultivated material with indicated accession numbers were collected in CJB (Conservatoire
et Jardin botaniques de la Ville de Genève greenhouse), GRF (Gesneriad Resea rch Foundation, Sarasota, U.S.A.), MP (Mauro Peixoto private collection,
Brazil), or USBRG (Botany Resea rch Greenhouses of the Smithsonian Institution). Voucher number s are given when present. N.A., no data
Species, accession numbers and/or voucher specimen (herbar ium), GenBank accessions (ITS, ncpGS, trnL-F, rps16, rpl16)
Achimenes admirabilis Wiehler, CJB AC-2510, A. Chautems & M. Perret 01-033 (G), GQ344504, AJ459689, AJ439827, N.A., AJ487784; Chautemsia calcicola
A.O. Araujo & V.C. Souza, A.O. Araujo & al. 500 (ESA), GQ344506, GQ383515, GQ383543, GQ383581, GQ383631; Diastema sp., MP 0872, GQ344510,
GQ383517, GQ383546, GQ383585, GQ383634; D. vexans H.E. Moore, GRF 840 306, GQ344511, GQ383518, GQ383547, GQ383586, GQ383635; Gesneria
humilis L., CJB s.n., A. Chautems & M. Perret 97-020 (G), GQ344513, AJ459683, AJ439821, GQ383588, AJ487778; Gloxinella lindeniana (Regel) Roalson &
Boggan, A.O. Araujo 604 (G), GQ344515, GQ383521, GQ383550, GQ383590, GQ383638; Gloxinia erinoides (DC.) Roalson & Boggan, CJB s.n., A. Chautems
& M. Perret 01-034 (G), GQ344519, AJ459690, AJ439828, GQ383593, AJ487785; G. perennis (L.) Fritsch, A.O. Araujo & al. 536 (ESA), GQ344516, GQ383522,
GQ383551, GQ383591, GQ383639; Gloxiniopsis racemosa (Benth.) Roalson & Boggan, A.O. Araujo 548 (ESA), GQ344514, GQ383520, GQ383549, GQ383589,
GQ383637; Goyazia petraea (S.M. Phillips) Wiehler, A.O. Araujo & al. 457 (ESA), GQ344517, GQ383523, GQ383552, N.A., GQ383640; G. rupicola Taub.,
A.O. Araujo & al. 545 (ESA), GQ344518, GQ383524, GQ383553, GQ383592, GQ383641; Kohleria spicata (Kunth) Oerst., CJB s.n., A. Chautems & M. Perret
97-018 (G), GQ344520, AJ459682, AJ439820, GQ383594, AJ487777; Mandirola ichthyostoma (Gardner) Seem. ex Hanst., A.O. Araujo 514 (ESA), GQ344527,
GQ383527, GQ383559, GQ383601, G Q383648; A.O. Araujo 518 (ESA), GQ344528, GQ383528, GQ383560, GQ383602, GQ383649; A.O. Araujo & al. 531
(ESA), GQ344522, G Q383526, GQ383555, GQ383596, GQ383643; A.O. Araujo 544 (ESA), GQ344521, GQ383525, G Q383554, GQ383595, GQ383642; M.
cf. rupestris (Gardner) Roalson & Boggan, A.O. Araujo 530 (ESA), GQ344529, GQ383529, GQ383561, GQ383603, GQ383650; Nomopyle dodsonii (Wiehler)
Roalson & Boggan, GRF 9769, GQ344535, GQ383530, GQ383565, GQ383609, GQ383654; Phinaea albolineata (Hook.) Benth. ex Hemsl., A.O. Araujo & al.
538 (ESA), GQ344538, GQ383531, GQ383566, GQ383612, GQ383655; Rhytidophyllum ver nicosum Urb. & Ek man, A.O. Araujo & al. 600 (G), GQ344539,
GQ383532, GQ383567, GQ383613, GQ383656; Seemannia gymnostoma (Griseb.) Toursark., A.O. Araujo & al. 549 (ESA), GQ344542, GQ383534, GQ383569,
GQ383616, GQ383658; S. nematanthodes (Kuntze) Schum., A.O. Araujo & al. 550 (ESA), GQ344541, GQ383533, GQ383568, GQ383615, GQ383657; S. aff.
purpurascen s Rusby, CJB AC-1481, A. Chautems & M. Perret 97-019 (G), GQ344540, AJ459684, AJ439822, GQ383614, AJ487779; S. purpurascens Rusby,
USBRG 02-197, Ibisch 960111 (US), GQ344544, GQ383535, GQ383571, N.A., GQ383659; S. sylvatica (Kunth) Hanst., CJB s.n., A.O. Araujo & al. 603 (G),
GQ344546, GQ383537, GQ383573, GQ383618, GQ383661; Smithiantha laui Wiehler, GRF s.n., GQ344552, AJ459685, AJ439823, GQ383623, AJ487780.
