A Molecular Phylogeny and Taxonomic Notes in Caamembeca (Polygalaceae)

Author(s): José Floriano Barêa Pastore, J. Richard Abbott, Kurt M. Neubig, W. Mark Whitten, Renata B.
Mascarenhas, Michelle Christine Almeida Mota, and Cássio van den Berg
Source: Systematic Botany, 42(1):54-62.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364417X694935

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Systematic Botany (2017), 42(1): pp. 54–62
© Copyright 2017 by the American Society of Plant Taxonomists
DOI 10.1600/036364417X694935
Date of publication March 1, 2017

A Molecular Phylogeny and Taxonomic Notes in Caamembeca (Polygalaceae)

José Floriano Barêa Pastore,1,7 J. Richard Abbott,2 Kurt M. Neubig,3 W. Mark Whitten,4
Renata B. Mascarenhas,5 Michelle Christine Almeida Mota,6 and Cássio van den Berg5
1
Universidade Federal de Santa Catarina, Campus Curitibanos. Herbário CTBS. Rodovia Ulysses Gaboardi,
Km 3, 89520-000 Curitibanos, Santa Catarina, Brazil.
2
New York Botanical Garden, 2900 Southern Boulevard, Bronx, New York 10458, U. S. A.
3
Department of Plant Biology, Southern Illinois University Carbondale, 1125 Lincoln Drive,
Carbondale, Illinois 62901, U. S. A.
4
Florida Museum of Natural History, 1659 Museum Road, Gainesville, Florida 32611-7800, U. S. A.
5
Universidade Estadual de Feira de Santana, Departamento de Ciências Biológicas, Av. Transnordestina, s.n.,
44036-900 Feira de Santana, Bahia, Brazil.
6
Universidade Federal do Paraná. Centro Politécnico, SCB, 81531-990 Curitiba, Paraná, Brazil.
7
Author for correspondence (jfpastore@hotmail.com)

Communicating Editor: Timothy M. Evans

Abstract—Caamembeca (Polygalaceae) is a genus of 13 species endemic to South America. The genus is morphologically distinctive, e.g.
supported by the putative morphologic synapomorphy of paired glands in a stipular position and on the rachis. However, its monophyly
has not been robustly tested as only four species have been included in previous phylogenetic analyses. We present a phylogenetic analysis
of 11 species based on nrITS, matK, the trnL intron, and trnL-trnF intergenic spacer. Relationships among Caamembeca species are discussed,
and three new combinations are made, Caamembeca amazonensis, Caamembeca autranii, and Caamembeca formosa, providing an
improved understanding of the genus.

Keywords—New combination, Polygala section Ligustrina, South America.

Polygalaceae (Fabales) have a worldwide distribution and
are subdivided into four tribes, Carpolobieae B. Eriksen,
Moutabeae Chodat, Polygaleae Chodat, and Xanthophylleae
Baill., with 27 genera and about 1,200 species. The internal
classification of Polygalaceae has changed greatly during the
last decades, mainly with respect to the boundaries of the
genus Polygala L. The first relevant infrageneric delimitation in
Polygala was presented by Chodat (1893) with 10 sections,
which were almost all elevated to subgenera by Paiva (1998).
Recently, after phylogenetic studies (Eriksen 1993; Persson
2001; Forest et al. 2007; Abbott 2009), several taxa traditionally
considered in a sectional or subgeneric delimitation were
elevated to a generic circumscription. Thus, the genera
Acanthocladus Klotzsch ex Hassk., Asemeia Raf. em. Small.
(formerly Polygala sect. Hebeclada Chodat), Badiera DC.,
Caamembeca J. F. B. Pastore (formerly P. sect. Ligustrina Chodat),
Gymnospora (Chodat) J. F. B. Pastore, Hebecarpa (Chodat) J. R.
Abbott, Heterosamara Kuntze, Phlebotaenia Griseb., Polygaloides
(formerly P. sect. Chamaebuxus (Spach) Chodat), and
Rhinotropis (Chodat) J. R. Abbott were segregated from
Polygala L. (see Paiva 1998; Pastore et al. 2010; Pastore 2012;
Pastore and Abbott 2012; Pastore and de Moraes 2013; Abbott
and Pastore 2015). All non-autonymic subgenera in Polygala,
except subgen. Chodatia Paiva, are now treated as genera.
One of the recently segregated genera, Caamembeca J. F. B.
Pastore (Polygalaceae), is endemic to South America, occur-
ring from Guyana to the State of Santa Catarina in Southern
Brazil. The species grow mostly in savannas of the Amazon
and the Atlantic rainforest of Brazil. Despite the geographic
and morphologic consistency among this clade, the species of
Caamembeca have been placed variously among different taxo-
nomic treatments. In his section Chamaebuxus (Dill. ex Spach)
DC., de Candolle (1824) described Polygala oxyphylla DC. and
P. spectabilis DC., the first species now included in Caamembeca,
together with some unrelated species: Polygala chamaebuxus L.
(=Polygaloides chamaebuxus (L.) O. Schwarz), P. chinensis L.
(from Polygala subgen. Polygala sect. Chloropterae (Chodat)
Paiva), P. paucifolia Willd. (=Polygaloides paucifolia (Willd.) J. R.
Abbott), P. uniflora Michx. (=Polygaloides paucifolia), and
P. venenosa Juss. ex Poir. (from Polygala subgen. Chodatia).
Bennett (1874) included these species now in Caamembeca in
his section ‘A’ together with the species now included in the
genus Gymnospora. Chodat (1893) circumscribed the natural
group now recognized as Caamembeca in his Polygala section
Ligustrina Chodat which excludes Polygala violoides A. St.-Hil. &
Moq. (=Gymnospora violoides A. St.-Hil. & Moq.) and P.
pedicellaris A. St.-Hil. & Moq. (=Gymnospora violoides). More than
a century later, Paiva (1998) elevated the section Ligustrina to a
subgenus, in a general rearrangement of the genus Polygala
according to a morphology-based phylogeny. Despite the con-
tribution made after Chodat´s Ligustrina circumscription (e.g.
Taubert 1896), Chodat was for more than one hundred years
the only reference to the Caamembeca species.
Bernardi (2000) presented a controversial study synonymiz-
ing several names and reducing the number of accepted spe-
cies from 15 to nine in Ligustrina. Bernardi's work is arguably
useful in terms of the graphics and the amount of detailed
discussion, at least on the surface. On the other hand, his
(Bernardi 2000) taxonomy is questionable because of extensive
species-level synonymization, and which has been revised by
later authors such as Rankin-Rodríguez and Greuter (2001),
Marques and Peixoto (2004, 2007), Aguiar et al. (2008), Pastore
(2012), Wendt (2012), Pastore and de Moraes (2013), and
Lüdtke et al. (2013). Bernardi (2000) changed the entire infrag-
eneric classification of neotropical Polygala, as he included all
species of sect. Ligustrina in a new and superfluous sect.
Laureolae Bernardi, which also included Polygala membranacea
(Miq.) Görts and species from sect. Gymnospora.
A few years later, Marques and Peixoto (2007), proposed a
midpoint between Chodat (1893) and Bernardi (2000), recog-
nizing 10 species plus the recently described Polygala
martinellii Marques & E. F. Guim. (Marques and Guimarães
2003). Taxa which had been synonymized by Bernardi (2000),
were revived by Marques and Peixoto (2007) as varieties:

54
2017] PASTORE ET AL.: SYSTEMATICS OF CAAMEMBECA
Polygala autranii Chodat as a variety of P. spectabilis (Polygala
spectabilis var. autranii (Chodat) Marques & Peixoto) and
Polygala salicina Chodat as a variety of P. oxyphylla
(P. oxyphylla var. salicina (Chodat) Marques & E. F. Guim.).
Furthermore, Marques and Guimarães (2003) described two
new varieties: Polygala spectabilis var. amazonensis Marques &
E. F. Guim. and P. martinellii var. carnosa Marques & E. F.
Guim. Marques and Peixoto (2007) considered Ligustrina as a
subgenus initially proposed by Chodat (1893). Following
Marques and Peixoto (2007), Pastore (2012) proposed species
delimitations in Caamembeca with lectotypification of all names
and inclusion of Polygala oxyphylla var. salicina as a synonym
of the autonymic variety.
Although Bennett (1874) and Bernardi (2000) associated
Caamembeca and Gymnospora, these genera are not sister
groups. They were considered similar due to sharing a non-
crested keel and triangular or sub-triangular seeds. However,
these genera differ by the following characters of Caamembeca:
seeds with an inflated caruncle, relatively large, and conical
glands in both a stipular position and in the inflorescence at
the base of flowers, whereas Gymnospora has neither a caruncle
on the seeds nor the conical glands. Caamembeca (as section
Ligustrina) was supported as sister to Polygala section Hebeclada
Chodat (now treated as genus Asemeia) by Forest et al. (2007),
with those two genera sister to a larger clade which includes
the genera Heterosamara, Muraltia, Polygala, Polygaloides,
Rhinotropis, and Salomonia. Gymnospora, represented by Polygala
violoides A. St.-Hil. & Moq. (=Gymnospora violoides (A. St.-Hil. &
Moq.) J. F. B. Pastore), was supported as within the clade also
including species now treated under the genera Acanthocladus,
Badiera, Bredemeyera, Hebecarpa, and Phlebotaenia (Fig. 1).
Only three species of Caamembeca have been included in
previous phylogenetic studies (Persson 2001; Forest et al.
2007; Abbott 2009). The purpose of this study is to demon-
strate the monophyly of Caamembeca and the relationships of
its species. We present here a phylogenetic analysis made
with 11 species (+ 1 potentially undescribed one) based on
nrITS, matK, the trnL intron, and trnL-trnF intergenic spacer,
and compare our results with previous treatments (Chodat
1893; Bernardi 2000; Marques and Peixoto 2007). Additionally,
we wish to clarify lingering taxonomic problems in the genus
by providing taxonomic notes with a modified key to the
13 currently recognized species.
Materials and Methods
Taxon Sampling — The Polygalaceae analyses included all genera with
a minimum of two species per genus, except Gymnospora and Heterosamara
sampled here with one species) and the monospecific genera Balgoya
Morat & Meijden, Barnhartia Gleason, Eriandra P. Royen & Steenis, and
Hualania Phil. (Appendix 1). Our Caamembeca analyses included 25 acces-
sions representing 11 known species and one potentially undescribed spe-
cies (Appendix 1). Caamembeca gigantea (Chodat) J. F. B.Pastore, C. martinellii
(E. F. Guim. & Marques) J. F. B.Pastore var. martinellii, and C. warmingiana
(A. W. Benn.) J. F. B.Pastore were not successfully sampled due to low
DNA quality. A member of genus Moutabea Aubl., M. excoriata Mart. ex
Miq. (from Tribe Moutabeae), was selected as the outgroup based on
broader phylogenetic analyses of the plastid loci matK, rbcL, and trnL-F.
DNA Amplification and Sequencing — Total genomic DNA was
extracted from fresh or silica-gel dried leaf material, sodium chloride/
CTAB preserved material (Chase and Hills 1991), and herbarium speci-
mens. An adapted version of the Doyle and Doyle (1987) protocol was
used for genomic DNA extraction or, when available, the DNeasy plant
mini kit (Qiagen) extraction protocol. We amplified the nuclear ribosomal
internal transcribed spacer region (nrITS, including both nrITS1 and
nrITS2 and intervening 5.8S), the plastid 3′trnK-matK (including partial
55
trnK intron and matK coding region) and trnL-F region (including the
trnL intron and the trnL-trnF intergenic spacer). The matK/trnK locus
was amplified in two fragments, using primers matK4La/matK1932R,
and matK1100L/trnK2R (Hu et al. 2000; Wojciechowski et al. 2004).
Some matK were amplified using the barcode primers 3F/1R (Fazekas
et al. 2012). The whole trnL-trnF region was amplified using primers “C”
and “F,” with the use of internal primers “D” and “E” for some problem-
atic samples (Taberlet et al. 1991). The nrITS region was amplified using
the primers 17SE and 26SE of Sun et al. (1994). The plastid loci were
sequenced using the same set of primers used for the amplification,
whereas the nrITS was sequenced using internal primers ITS92 (Desfeux
and Lejeune 1996) and ITS4 (White et al. 1990) with the same PCR pro-
gram. All PCR amplifications were performed in a final volume of 10 μL
containing: 5 μL of TopTap master mix kit (Qiagen, Valencia, California),
2.25 pMol primers each, 5–10 ng of genomic DNA, and ultrapure H2O
(enough to complete the volume to 10 μL). For the nrITS amplification,
2% DMSO (dimethyl sulfoxide) and betaine were added to final concen-
tration of 1 M. The amplicons were amplified using initial denaturation
at 94°C (5 min), 28 (ITS) or 32 (plastid loci) cycles of denaturation at
94°C (1 min), annealing 52°C (ITS) or 54°C (plastid loci) (1 min), elonga-
tion at 72°C (2 min), and a final elongation of 4 min. Amplified products
were purified using precipitation with 11% solution of polyethylene
glycol (PEG) 8000 and ethanol cleaning. Sequencing reactions in both
directions were performed using BigDye Terminator 3.1 (Applied Bio-
systems, Carlsbad, California) chemistry and analyzed on an ABI3130XL
sequencer (Applied Biosystems/Life Technologies Corporation) follow-
ing the manufacturer’s protocol at Universidade Estadual de Feira
de Santana, Bahia, Brazil. Some PCR products were sequenced at the
Interdisciplinary Center for Biotechnology Research at the University of
Florida, Gainesville.
Alignment and Phylogenetic Analyses — The electropherograms were
edited and assembled using Geneious version 6.1.6 (Drummond et al.
2012). After trimming we assembled a matrix in which 7.95% of nucleo-
tides were scored as missing because of trimming the ends in different
positions. Sequences were automatically aligned in MUSCLE with
default settings (Edgar 2004) and then manually adjusted in Geneious.
We performed phylogenetic analyses of separate nrITS and plastid loci
and combined plastid and nuclear DNA datasets. The statistical congru-
ence between nuclear and plastid datasets was assessed by the incongru-
ence length test (ILD) test (Farris et al. 1994) based on the partition
homogeneity test (PHT) implemented in PAUP* 4.0 beta 10 (Swofford
2002) with 1,000 replicates, simple addition of taxa, tree bisection-
reconnection (TBR) branch swapping, and multrees option, saving
10 trees per replicate. The unambiguous indels were coded for all
datasets using the simple indel coding criteria (Simmons and Ochoterena
2000) implemented in the Seqstate software (Müller 2005). The search for
the most parsimonious trees was carried out using a heuristic search
with 1,000 random taxon-addition and tree bisection-reconnection (TBR)
branch swapping, saving 10 trees per replicate. Trees saved in this first
round were used as starting trees in a second search using the same
parameters, but saving a maximum of 10,000 trees. All characters were
considered equally weighted and unordered (Fitch 1971). Nonparametric
bootstrap resampling (BS) implemented in PAUP* 4.0 beta 10 was used
to estimate clade support (Felsenstein 1985) which was assessed through
1,000 bootstrap replicates, each analyzed using the heuristic search
parameters mentioned above and with 10 trees retained per replicate.
Bayesian analyses were carried out using MrBayes v. 3.1.2 (Huelsenbeck
and Ronquist 2001; Ronquist et al. 2012) through the Cyberinfrastructure
for Phylogenetic Research (Cipres Science Gateway; Miller et al. 2010).
The molecular evolution model was selected for each dataset using
Akaike information criterion implemented in jModeltest 2 (Darriba
et al. 2012). The best models were GTR + I + Γ for nrITS, GTR + Γ for
plastid loci (including plastid Polygalaceae general analysis, Fig. 1) and
SYM + Γ for 5.8S (ITS). Heterogeneity of sites was modeled with the
gamma distribution for plastid and 5.8S partitions and gamma plus
proportion of invariant sites for the nrITS partitions. Two separate runs
of a Metropolis-coupled Markov Chain Monte Carlo (MCMC) permuta-
tion of parameters were each initiated with a random tree and eight
simultaneous chains set at default settings (Huelsenbeck and Ronquist
2001). We performed two runs in parallel of four MCMC for 2 ×
106 generations, with trees sampled every 1,000th generation. The first
106 trees were discarded as burn-in. The 50% majority-rule consensus was
calculated in MrBayes for assessing posterior probabilities. All Bayesian
majority-rule consensus trees were visualized and partially edited in
FigTree v. 1.4 (Rambaut 2012). Convergence of runs was assessed using
the program Tracer v. 1.6 (Rambaut et al. 2014). MrBayes v. 3.1.2 was
used to summarize trees sampled from post burn-in generations in a
56 SYSTEMATIC BOTANY [Volume 42

Fig. 1. Cladogram (A) and phylogram (B) based on the majority-rule consensus tree from the Bayesian analysis of the combined plastid
(matK + trnL-F + rbcL) datasets for the genera of Polygalaceae. Numbers above branches are posterior probabilities (in percentage) and branches in bold
have PP = 100. Numbers below branches are bootstrap support (BS); those marked with an arrowhead have BS < 50. Species name in bold were
included in former delimitation of Polygala (Polygala s.l., sensu Chodat 1893).

majority rule consensus tree that included posterior probabilities (PP) as
branch support estimates.
Results
We produced 64 new sequences (20 of nrITS, 24 of matK,
and 20 of trnL-F) from 27 accessions (25 new accessions in
Caamembeca) of 11 out of 13 species recognized in Caamembeca.
The dataset contains 8 species that are evaluated for the first
time in a molecular phylogenetic analysis.
Phylogeny of Caamembeca — Polygalaceae plastid-based
analyses (Fig. 1) supported all tribes as monophyletic.
Caamembeca is included within tribe Polygaleae, sister to Asemeia
(90% bootstrap/100% posterior probability) in agreement
2017] PASTORE ET AL.: SYSTEMATICS OF CAAMEMBECA 57

Fig. 2. Cladogram (A) and phylogram (B) based on the majority-rule consensus tree from the Bayesian analysis of the combined nuclear (ITS) and
plastid (matK + trnL-F) datasets for Caamembeca, including coded gaps. Numbers above branches are posterior probabilities (in percentage) and
branches in bold have PP = 100. Numbers below branches are bootstrap support (BS) values from the parsimony analysis, branches in bold without
any number have BS = 100% and those marked with an arrowhead have BS < 50. Major clades ordered are discussed in the text.

with former phylogenetic studies (see Forest et al. 2007). The
ILD test indicated low incongruence among nuclear and plas-
tid datasets ( p = 0.576, with n = 1,000 replicates). Branch sup-
port across the trees also concurred with the ILD test and we
didn’t find highly supported incongruent nodes when trees
from different partitions were compared. All analyses of indi-
vidual, plastid (tree not shown), nuclear (tree not shown), and
combined datasets (Fig. 2) revealed the same general back-
bone, except for the position of C. grandifolia which appears as
sister group of clade B (Fig. 3) in matK dataset analyses (tree
not shown), or sister to the clade that contains the remaining
Caamembeca species in nrITS analyses (Fig. 2). The indel coding
in all datasets resulted in a general improvement of tree resolu-
tion and branch support. Therefore, the discussion will be
based on the results from the combined analyses of nuclear
and plastid datasets and the associated indels (Fig. 2).
Caamembeca is monophyletic with 100% bootstrap and 100%
PP support in all analyses. Caamembeca grandifolia is sister to
all other Caamembeca species. The species are grouped into A)
C. grandifolia (A. St.-Hil. & Moq.) J. F. B. Pastore and three
major clades (Figs. 1 and 2); B) C. insignis and C. laureola; C)
C. oleifolia and C. oxyphylla; and D) C. amazonensis, C. autranii,
C. formosa, C. martinellii var. carnosa, C. spectabilis, C. ulei, and
one undescribed species.
Discussion
Members of the clade including Caamembeca and Asemeia
(from North, Central, and South America, from the United
States to Paraguay) are endemic to the New World, mostly
from the Neotropical region. Non-molecular synapomorphies
have not been identified for the clade including these genera,
but the conical stipular glands of Caamembeca and the fused
lower sepals of Asemeia may prove to be synapomorphies
once the level of universality has been fully assessed. The
phylogenetic trees supported four basic groups, each cohesive
morphologically (Figs. 2, 3). Caamembeca grandifolia (A) from
the Brazilian Atlantic coastal forest (Mata Atlântica) is the
only species of Caamembeca with acute wings (internal, lateral
sepals). Clade B includes Atlantic Forest species exclusively,
with the wings wide spreading and slightly reflexed in flower.
The larger clade C+D has species in which the wings only
spread less than a 45° angle at anthesis. Clade C includes spe-
cies from both the Atlantic forest and savanna with yellow or
orange internal sepals (wings). This clade is also characterized
by having the leaf blade with the tertiary nerves abaxially
prominent, forming a dense reticulum visible to the naked
eye, and pyriform seeds. Clade D includes species from
savannas, Amazon and Atlantic Forest, and the Andes, all
with more or less quadrangular wings in fruit (Fig. 1).
Three taxa were not sampled: C. gigantea, C. martinellii var.
martinellii, and C. warmingiana. Morphological studies suggest
that C. gigantea is sister to C. spectabilis, C. warminginana is
closely related to C. insignis or C. laureola, and the two distinc-
tive varieties of C. martinellii might be different species
(C. martinellii var. martinellii and C. martinellii var. carnosa).
Given its morphological variation we suggest that the tradi-
tional circumscription of C. spectabilis may represent a meta-
species or a stem group (Pastore in prep.). This analysis, in
conjunction with morphological study of specimens, supports
the idea that there is likely at least one undescribed species of
Caamembeca. This, too, will be dealt with in a separate publica-
tion, as it is clear that the C. spectabilis s. l. complex (including
at least C. formosa and C. gigantea) needs to be studied in
58 SYSTEMATIC BOTANY [Volume 42

Fig. 3. Recovered cladogram of Caamembeca with images overlaid on the main clades. 1. Caamembeca grandifolia. Photo by J. F. B. Pastore.
2. Caamembeca insignis. Photo by M. S. Wängler. 3. Caamembeca laureola. Photo by I. Gonçalves. 4. Caamembeca oleifolia. Photo by D. B. O. S. Cardoso.
5. Caamembeca oxyphylla. Photo by M. C. A. Mota. 6. Caamembeca amazonensis. Photo by L. P. Queiroz. 7. Caamembeca martinellii var. carnosa. Photo by
A. Popovkin. 8. Caamembeca ulei. Photo by H. Moreira. 9. Caamembeca spectabilis. Photo by J. F. B. Pastore.
2017] PASTORE ET AL.: SYSTEMATICS OF CAAMEMBECA 59

greater detail. Given the low support for nodes, which
includes C. formosa, C. spectabilis, and C. ulei, and the great
amount of morphological variation in the complex, it is clear
that C. spectabilis, as traditionally defined, is not monophy-
letic, and even in its narrower sense as treated here it may still
represent a metaspecies or stem species, with various lineages
more closely related to other taxa than to each other. More
sampling is needed.
The species of Caamembeca are distributed in different South
American biomes: Caatinga (included in the Seasonal Tropical
Dry Forest), Central Savannas, Andean, Atlantic Forest, and
Amazon. However, the only clade in which all species share the
same biome, Atlantic Forest, is clade B (C. laureola + C. insignis).
In contrast, clade D includes species of several different biomes,
e.g. with C. spectabilis from the Amazon forest and C. ulei from
the Brazilian Central Savannas as sister groups.
Caamembeca spectabilis s. s. is a suffrutescent herb frequently
collected in the Amazon region, with substantial morphologi-
cal variation in the shape and texture of the leaves and flower
and fruit size. The phylogenetic results here for C. spectabilis
s. s., albeit preliminary, suggest that this morphological varia-
tion reflects the genetic structure of the species. The phylogeny
included only two specimens of C. spectabilis, which are sister
in the phylogeny: Lima 7041 (HUEFS) collected in Carajás (PA)
in the Cangas rock field with iron stone outcrops, and Pignal
3063 (MO) from the Amazon Forest of Belém (PA). The
branch-length distance (0.009) between them in the phylogram
is comparable to the distances between C. oleifolia and
C. oxyphylla (0.006 to 0.011). The Carajás region has plants
with most of the morphological variation found in
C. spectabilis and seems to be key to understanding the species
(Pastore in prep.). The iron-rich Cangas soil seems to prevent
dominance by either Amazon Forest or a grassy vegetation,
diminishing the role of fire as it occurs in typical savanna con-
ditions. These conditions could be the result of a historically
stable climate in Carajás with the retention of morphological
variation of C. spectabilis in a relative small area. Further studies
are needed to explore the possibility of the existence of cryptic
species, especially within the C. spectabilis s. l. complex, and the
role of different vegetation types in the genetic variation.

Taxonomic Treatment

Key to CAAMEMBECA Species (Modified from Marques and Peixoto 2007)

1. Leaf blades with the tertiary nerves prominent abaxially, forming a dense reticulum visible to the naked eye; pyriform seeds . . . . . . . . . . . . . . . . 2
2. Coriaceous leaves, frequently deflexed; robust and hirsute inflorescence rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. oleifolia
2. Chartaceous leaves, frequently erect; slender and strigose inflorescence rachis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. oxyphylla
1. Leaf blades with very fine intersecondary veins or immersed on both surfaces, forming a loose reticulum inconspicuous
to the naked eye; campanuliform, elliptic, or subtriangular seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Attenuate or apically acuminate floral buds; campanuliform seeds, apiculate in the center of the base . . . . . . . . . . . . . . . . . . . . . . C. grandifolia
3. Obtuse to rounded floral buds; elliptic or subtriangular seeds, not apiculate in the center of the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Petioles lacking or up to 1 mm long; elliptic seeds, densely sericeous, with hood-shaped and light yellow caruncle. . . . . . . . . . . . . . C. ulei
4. Petioles 2–8 mm long; subtriangular seeds, loosely strigose, with inconspicuous or little evident caruncle,
reddish-brown or yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Floral keels 17–25 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Coriaceous to chartaceous or papery opaque leaf blades; revolute margin; glands on petiole
and rachis 0.3–0.5 mm wide; endemic to Atlantic Forest of Bahia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 (C. martinellii)
7. Dried leaf blades coriaceous to chartaceous and yellowish-brown with obtuse base,
strongly revolute margin; papery flowers and capsules, capsule ca. 15 mm long, oblong. . . . . . . C. martinellii var. martinellii
7. Dried leaf blades papery and dark brown, acute base, lightly revolute margin; fleshy flowers
and capsules, capsule 11–12 mm long, elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. martinellii var. carnosa
6. Membranous leaf blades, opaque or often so thin as to be semi-translucent when dry
(like paper dipped in grease), revolute or plane margin, glands in petiole and on rachis 0.2 mm wide;
widespread, including Caatinga and Savanna of Bahia (for C. autranii and C. spectabilis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Rachis glands 1.2–2 × 0.5–0.7 mm (width at base); yellow caruncle, the appendage little differentiated . . . . . . . . . . C. gigantea
8. Rachis glands 0.3–0.9(–2) × 0.2–0.8 mm (width at base); yellowish-brown caruncle,
white to yellowish well differentiated appendage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Pilose leaf blades with abaxial indumentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. autranii
9. Leaf blades with sparse trichomes on both faces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Membranous flowers; lower outer sepals 2–4 × 2–5 mm and the upper outer
sepal 3.5–6.5 × 8–10 mm, smooth and opaque abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spectabilis
10. Papery flowers; lower outer sepals 5–7 × 5.5–8 mm and the upper outer
sepal 9–12 × 15–16 mm, rugose and shiny abaxially. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. amazonensis
5. Floral keels 6–16 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Internal sepals (wings) up to 1/2 the length of the keel; caruncular appendages inflated, 2/3 length of seed. . . . . C. warmingiana
11. Internal sepals (wings) 2/3 or nearly equal to the length of the keel; flat caruncular appendages,
generally wrinkly, 2/3 or as long as the seed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Leaf blades usually thinly membranous, generally with cuspidate apex; keel 6–10(–12) mm long. . . . . . . . . . . . . . C. laureola
12. Leaf blades usually papery to chartaceous (sometimes membranous in C. formosa), acute, obtuse
or slightly acuminate apex; keel (10–)10.5–15 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Intersecondary veins clearly visible joining with tertiary veins to form a lose reticulum;
leaf margins often plane, sometimes revolute, sparsely (to densely) ciliate;
lateral petiole glands 0.2–0.5(–1) × 0.1–0.2 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. formosa
13. Intersecondary veins usually not visible, not forming a reticulum with tertiary veins,
revolute leaf margins, densely (to sparsely) ciliate; lateral petiole glands 0.6–1 × 0.2 mm. . . . . . . . . . . . . . . C. insignis

Of the 13 currently recognized species of Caamembeca, Caamembeca amazonensis (Marques & E. F. Guim.) J. F. B.
11 were treated in Pastore (2012). Three new combinations Pastore, comb. & stat nov. Caamembeca spectabilis (DC.)
or status changes are addressed here. J. F. B. Pastore var. amazonensis (Marques & E. F. Guim.)
60 SYSTEMATIC BOTANY
J. F. B.Pastore, Kew Bull. 67(3): 441. 2012. Polygala
spectabilis var. amazonensis Marques & E. F. Guim.,
Bradea 9(10): 48. 2003.–TYPE: BRAZIL. Amazonas, Near
Livramento, on Rio Livramento, Humaitá. 12 Oct 1934,
Krukoff 6958 (holotype RB [31569]!, isotypes F!, MO!,
NY!, S!).
Remarks — Marques and Peixoto (2007) mistakenly cited
Kuhlmann as the original collector instead of Krukoff.
Caamembeca amazonensis can be recognized by rugose external
sepals, the upper ones 5–7 × 5.5–8 mm and the lower one
9–12 × 15–16 mm. Phylogenetically, C. amazonensis (endemic
to the Amazon Forest) was recovered as sister to C. martinellii
var. carnosa (endemic to Atlantic Forest in coastal Bahia).
Caamembeca autranii (Chodat) J. F. B. Pastore, comb. nov.
Caamembeca spectabilis (DC.) J. F. B. Pastore var. autranii
(Chodat) J. F. B. Pastore, Kew Bull. 67(3): 441. 2012.
Polygala spectabilis var. autranii (Chodat) Marques &
Peixoto, Rodriguésia 58(1): 131. 2007. Polygala autranii
Chodat, Mém. Soc. Phys. Genève 31: 80. 1893.–TYPE:
BRAZIL. Bahia, Igreja Velha [Egreja Velha], 1841,
Blanchet 3242 (lectotype G!, selected by Pastore 2012;
isolectotypes B† photo F neg. Macbride 12998, BM!, F!,
FI!, HAL!, P[221215]!, P[221214]!, P[221213]!, NY!).
Remarks — Caamembeca autranii was described by Chodat
(1893) based on a specimen from Jacobina, Bahia, in a local-
ity called Igreja Velha. The membranaceous flower and the
abaxially pubescent leaves resemble those of Caamembeca
spectabilis. However, phylogenetically these species are not
closely related to each other; C. autranii is sister to all other
species of clade D, whereas C. spectabilis is sister to C. ulei
and closely related to C. formosa. Caamembeca autranii was
collected in rocky fields with the influence of the Caatinga
biome (which is part of the Seasonally Tropical Dry Forest)
in the states of northwestern Brazil: Alagoas, Bahia, Ceará,
Paraíba, and Pernambuco. On the other hand, C. spectabilis is
from the Amazon Forest and the Cangas vegetation. There-
fore, Caamembeca autranii is here recognized as a legitimate
species, in accordance with Chodat (1893), instead of as a
synonym of C. spectabilis.
Caamembeca formosa (A. W. Benn. ex Britton) J. F. B. Pastore,
comb. nov. Polygala formosa A. W. Benn. ex Britton, Bull.
Torrey Bot. Club 16(1): 19. 1889.–TYPE: BOLIVIA. Mapiri,
May 1886, Rusby 1908 (lectotype, selected by Pastore
2012, NY!, isolectotypes E!, GH n. v., MO!, PH n. v., US!).
Polygala andina A. W. Benn. ex Britton (1889), Bull. Torrey
Bot. Club 16(1): 19. 1889. –TYPE: BOLIVIA. Near La Paz,
April 1885, Rusby 2869 (holotype NY!),
Polygala bangiana Chodat, Bull. Herb. Boissier 4: 234. 1896.
Caamembeca bangiana (Chodat) J. F. B. Pastore, Kew Bull.
67(3): 437. 2012.–TYPE: BOLIVIA. Yungas, 1890, Bang
230 (holotype BM!, isotypes BR!, GH n. v., photo GH!, E
n. v., photo E!, F!, K!, M!, MICH n. v., photo MICH!,
MO!, NY!, US!, WU!).
Remarks — It was described as three separate names:
Polygala andina, P. bangiana, and P. formosa, now associated
with the genus Caamembeca, all of them from the pre-Andean
region in the south of the Bolivian department of La Paz.
Polygala formosa and P. andina were published at the same
time (1889), earlier than P. bangiana (1896). These names
[Volume 42
clearly represent the same species. Marques and Peixoto
(2007) treated the species as P. bangiana Chodat, and Pastore
(2012) transferred it to Caamembeca. We are choosing the
most used name in herbarium collections, P. formosa. In the
phylogenetic analyses, C. formosa is represented by the acces-
sion Grifo 757 (MO) from Guanay, La Paz and has an
unresolved position among other species from clade D. The
other five samples from Bolivia, all from the Department of
Cochabamba, were recovered in a clade (including Brummitt
19367, Teran 996, Teran 2984, Teran 3131, and Teran 3607).
More studies are needed to resolve the relationships within
C. formosa populations and how it fits into the C. spectabilis
s. l. complex.
Caamembeca insignis (Klotzsch ex Chodat) J. F. B. Pastore,
Kew Bull. 67(3): 437. 2012. Polygala insignis Klotzsch ex
Chodat, Mém. Soc. Phys. Genève 31: 81. 1893.–TYPE:
BRAZIL. Minas Gerais, Rio das Pedras, 1818–1820,
Sellow B 2027 and c 1559 (lectotype, designated by
Pastore 2012, G[#300197]!, selected by Marques and
Peixoto (2007); isolectotype (?) B† photo F!, neg. J. F.
Macbride 13029).
Remarks — Bernardi (2000) erroneously lectotypified
Monnina selloi Spreng., which was described explicitly based
on a Sellow specimen (Sprengel 1826). Bernardi (2000) chose
a Caamembeca insignis specimen collected by Beyrich and
proposed the combination Polygala selloi (Spreng.) Bernardi.
Marques and Peixoto (2007) rejected Polygala selloi consider-
ing it a doubtful name, and reestablished Polygala insignis.
Therefore, Polygala insignis was used as the basionym of
Caamembeca insignis by Pastore (2012). The original descrip-
tion of Sprengel (1826) is not sufficient to identify the spe-
cies, or even to assign it correctly to this genus. Furthermore,
the original type collection was not found in visited herbaria
(B, BM, BR, FI, E, G, HAL, K, L, MO, NY, P, W), and might
have been destroyed during the Second World War. The only
relevant information about the identity of M. selloi was pro-
vided by Don (1831) who described M. selloi in more detail,
in addition to the translation from the original Latin diagnosis
to English. Monnina selloi is described by Don (1831), as a pos-
sible synonym of Monnina tristaniana, as “an erect herb, stem
base with branched pili; leaves oblong or obovate-lanceolate,
tapering to both ends, but much more so at base, with ciliate
revolute margins, young leaves pilose on nerves beneath;
native of Brazil (. . .) shrub 2 to 6 feet.” The only Brazilian spe-
cies of Monnina matching the description by Don (1831), after
considering Sellow’s original itinerary and the plant size
between 60–220 cm, would be Monnina tristaniana. The
remaining features described by Don for M. selloi Spreng. are
also found in M. tristaniana A. St.-Hil. & Moq. However, the
circumstance of Don (1831) having ‘improved’ the original
description of Sprengel (1826) is not clear to us now. Never-
theless, we support Marques and Peixoto (2007) and Pastore
(2012) treating Monnina selloi as a doubtful name.
Undescribed Species—Caamembeca sp. nov. (Pastore 2533)
is a common species in the rocky fields of Rio de Contas and
other municipalities in the region of Chapada Diamantina
(Bahia State, Brazil), with several specimens, e.g. Harley
55065 (HUEFS), Harley 55768 (HUEFS), and Sano 14859
(HUEFS). It has been identified as Polygala insignis by
Marques in herbarium specimens. Phylogenetically, it is
quite distant from C. insignis, being sister to the clade of
C. martinelli var. carnosa (from Atlantic Forest) and
2017] PASTORE ET AL.: SYSTEMATICS OF CAAMEMBECA
C. amazonensis (from the Amazon Forest). Morphologically, it
does resemble C. insignis (from Atlantic Forest) by the ciliate
leaf margins. However, its phylogenetic position indicates
that it most likely represents an undescribed species. Its sta-
tus needs to be addressed within the context of a detailed
analysis of C. spectabilis s. l.
Acknowledgments. The first author would like to thank Alex
Popovkin, Domingos Cardoso, Henrique Moreira, Luciano Paganucci,
and Marilia Wängler for the photos used to compose Figure 1. We are also
grateful to Paulo Schwirkowski for kindly sending us a sample of
Caamembeca insignis in silica. We thank Alina Freire-Fierro and an
anonymous reviewer for their careful reading and relevant suggestions
on final manuscript. Finally, we thank the herbarium HUEFS for
permission to extract material from their Caamembeca specimens, and
PRONEX-Núcleo de Excelência em Sistemática e Variabilidade de Plantas
e Fungos – PNX0014/2009).
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Appendix 1. Voucher information for this study, presented in the
following order: Taxon; Voucher specimen: collector and number
(herbarium acronym), locality; GenBank accessions: ITS; rbcL; matK;
trnL-F. Long dash (—) = sequence not available,* = sequence newly
generated for this study.
Acanthocladus albicans A.W.Benn.; Fonseca 2973 (CEN), Brazil, Goiás;
*KU682359; —; *KU682399; *KU682396. Acanthocladus brasiliensis
(A.St.-Hil. & Moq.) Klotzsch ex Hassk.; Brunner et al. 945 (MO), Brazil; —;
AM234209; —; AF366973. Asemeia floribunda (Benth.) J.F.B.Pastore &
62 SYSTEMATIC BOTANY
J.R.Abbott; Abbott 19659 (FLAS), Mexico; —; —; —; AF366985. Asemeia
monninoides (Kunth) J.F.B.Pastore & J.R.Abbott; Harley 55602 (HUEFS),
Brazil, Bahia; *KU682361; —; *KU682405; *KU682395. Asemeia violacea
(Aubl.) J.F.B.Pastore & J.R.Abbott; D'Arcy & D'Arcy 6036 (MO), Pastore 242
(CEN), Brazil; —; EU644686; EU604035; *KU682423. Atroxima afzeliana
Stapf; Jongkind 4281 (WAG), Africa; —; AM234175; EU604049; AM234273,
AM234310. Atroxima liberica Stapf; Adams 24834 (MO), Reitsma et al. 1115
(NY), Africa; —; AM234174; —; AF366941. Badiera fuertesii Urb.; Abbott
20901 (FLAS), Thompson 10465 (NY), Dominican Republic; —; AM234200; —;
AF366979. Badiera penaea (L.) DC.; Thompson 9763 (FLAS), Dominican
Republic; —; AM234201; KJ012710; AF366982. Balgoya pacifica Morat &
Meijden; MacPherson 3394 (US), New Caledonia; —; —; —; AF366942.
Barnhartia floribunda Gleason; Boom et al. 8594 (K), Brazil; —; AM234168; —;
AM234271, AM234308. Bredemeyera floribunda Willd.; Belo 742 (K),
Irwin et al. 27995 (NY), Brazil; —; EU644699; EU596520; AF366945.
Bredemeyera martiana A.W.Benn.; Harley 55577 (HUEFS), Brazil, Bahia;
*KU682360; —; *KU682400; *KU682386. Bredemeyera microphylla
(Griseb.) Hieron.; Maas 8181 (U), Argentina; —; AM234173; —; AF366948.
Caamembeca amazonensis (Marques & E.F.Guim.) J.F.B.Pastore; Queiroz
13920 (HUEFS), Brazil, Amazonas; *KU682368; —; *KU682401; —.
Caamembeca autranii (Chodat) J.F.B.Pastore; Brummitt 19367 (MO),
Bolivia, Cochabamba; *KU682376; —; —; —. Caamembeca autranii
(Chodat) J.F.B.Pastore; Schwartsburd 2349 (HUEFS), Brazil, Bahia; —; —;
*KU682410; —. Caamembeca autranii (Chodat) J.F.B.Pastore; Pereira 1752
(HUEFS), Brazil, Bahia; —; —; *KU682419; —. Caamembeca autranii
(Chodat) J.F.B.Pastore; Agra 4845 (MO), Brazil, Paraiba; *KU682366; —;
*KU682416; *KU682398. Caamembeca formosa (A.W.Benn.) J.F.B.Pastore;
Teran 3131 (MO), Bolivia, La Paz; *KU682373; —; —; —. Caamembeca
formosa (A.W.Benn.) J.F.B.Pastore; Teran 996 (MO), Bolivia, La Paz;
*KU682377; —; —; —. Caamembeca formosa (A.W.Benn.) J.F.B.Pastore;
Grifo 757 (MO), Bolivia; *KU682371; —; —; *KU682390. Caamembeca
formosa (A.W.Benn.) J.F.B.Pastore; Teran 3607 (MO), Bolivia, Cochabamba;
*KU682374; —; —; *KU682391. Caamembeca formosa (A.W.Benn.) J.F.B.
Pastore; Teran 2984 (MO), Bolivia, La Paz; *KU682375; —; *KU682417;
*KU682392. Caamembeca grandifolia (A.St.-Hil. & Moq.) J.F.B.Pastore;
Pastore 2110 (HUEFS), Brazil, Bahia; *KU682363; —; *KU682406;
*KU682387. Caamembeca insignis (Chodat) J.F.B.Pastore; Schwirkowski s.n.
(MBM), Brazil, Santa Catarina; —; —; *KU682418; —. Caamembeca
laureola (A.St.-Hil. & Moq.) J.F.B.Pastore; Castro 1017 (HUEFS), Brazil,
Rio de Janeiro; *KU682364; —; *KU682407; —. Caamembeca laureola
(A.St.-Hil. & Moq.) J.F.B.Pastore; Borges 627 (HUEFS), Brazil, Bahia; —; —;
*KU682409; —. Caamembeca laureola (A.St.-Hil. & Moq.) J.F.B.Pastore;
Meireles 354 (UEC), Brazil, São Paulo; —; —; *KU682420; —. Caamembeca
laureola (A.St.-Hil. & Moq.) J.F.B.Pastore; Harley 55833 (HUEFS), Brazil,
Santa Catarina; *KU682365; —; *KU682408; *KU682397. Caamembeca
martinellii (Marques & E.F.Guim.) J.F.B.Pastore var. carnosa (Marques &
E.F.Guim.) J.F.B.Pastore; Carvalho 2796 (HUEFS), Brazil, Bahia; *KU682378; —;
*KU682413; *KU682384. Caamembeca oleifolia (A.St.-Hil. & Moq.)
J.F.B.Pastore; Queiroz 13290 (HUEFS), Brazil, Minas Gerais; *KU682379; —;
*KU682403; *KU682393. Caamembeca oxyphylla (DC.) J.F.B.Pastore ;
Pastore 5101 (CTBS), Brazil, Minas Gerais; *KU682380; —; *KU682414; —.
Caamembeca oxyphylla (DC.) J.F.B.Pastore; Forzza 3668 (HUEFS), Brazil,
Minas Gerais; —; —; *KU682415; —. Caamembeca oxyphylla (DC.) J.F.B.
Pastore; Harley 20751 (K), Brazil, Bahia; GQ888978; AM234202; EU604054;
AM234343.1, AM234285.1. Caamembeca oxyphylla (DC.) J.F.B.Pastore;
Hatschbach 49863 (MO), Brazil, Espírito Santo; GQ889010; —; —;
GQ889189. Caamembeca oxyphylla (DC.) J.F.B.Pastore; Messias 2248
(HUEFS), Brazil, Minas Gerais; *KU682381; —; *KU682404; *KU682394.
Caamembeca sp. nov.; Pastore 2533 (HUEFS), Brazil, Bahia; *KU682372; —;
*KU682412; *KU682383. Caamembeca spectabilis (DC.) J.F.B. Pastore;
Lima 7041 (HUEFS), Brazil, Pará; *KU682370; —; *KU682402; *KU682382.
Caamembeca spectabilis (DC.) J.F.B.Pastore; Pignal 3036 (MO), Brazil,
Pará; *KU682369; —; —; *KU682389. Caamembeca ulei (Taub.) J.F.B.
Pastore; Pastore 2343 (HUEFS), Brazil, Goiás; *KU682367; —; *KU682411;
*KU682388. Carpolobia alba G.Don.; Cable 747 (K), Africa; —; AM234176;
[Volume 42
EU604053; AF366949. Carpolobia goetzei Gürke; Goyder et al. 3722 (K),
Africa; —; AM234177; —; GQ889065, AM234311. Cercis canadensis
L. (outgroup); Bruneau 802 (MT), Gervais 1397-91 (MT), Canada; —;
U74188; JN881410; FJ801162. Comesperma calymega Labill.; Donner 10317
(MO), Australia; —; —; —; AF366951, GQ889067. Comesperma
esulifolium Gand.; Telford 12350 (CANB), Australia; —; AM234179;
EU596516; AM234312, AM234307. Diclidanthera bolivarensis Pittier;
Rosales & Valles 123 (US), Venezuela; —; —; —; AF366954. Diclidanthera
penduliflora Mart.; Beck 16906 (NY), Bolivia; —; —; —; AF366954.
Epirixanthes pallida T.Wendt; CM 009 (L), Papua New Guinea; —; —;
KR002175; —. Epirixanthes papuana J.J.Sm.; CM 011 (L), Asia; —; —;
KR002176; —. Eriandra fragrans P.Royen & Steenis; Pullen 7234 (K), Asia;
—; AM234170; EU604051; AM234309, AM234272. Gymnospora violoides
(A.St.-Hil. & Moq.) J.F.B.Pastore; Ratter et al. 6425 (K), Brazil; —;
AM234205; —; AM234286, AM234323. Hebecarpa macradenia (A.Gray)
J.R.Abbott; Abbott 19546 (FLAS), United States; —; —; —; AF366980.
Hebecarpa obscura (Benth.) J.R.Abbott; Abbott 14583 (FLAS), Holmgren
8022 (NY), United States; —; AM234207; —; AF366981. Heterosamara
tatarinowii (Regel) Paiva; Sino-Amer. Exped. 1688 (MO), China; —;
AM234208; —; GQ889208, AF366995. Hualania colletioides Phil.;
Guaglionone et al. 1587 (NY), South America; —; AM234171; —; AF366947.
Monnina exalata A.W.Benn.; Pastore 742 (CEN), Brazil, Distrito Federal;
*KU682362; —; *KU682422; —. Monnina insignis A.W.Benn.; Harley 19055
(NY), Brazil, Bahia; —; —; *KU682425; AF366959. Monnina stenophylla
A.St.-Hil. & Moq.; Pastore 1089 (CEN), Brazil, Goiás; —; —; *KU682426;
*KU682424. Moutabea aculeata (Ruiz & Pav.) Poepp. & Endl.; Smith 1522
(US), South America; —; —; —; GQ889089, GQ888789. Moutabea
excoriata Mart. ex Miq.; Pastore 1446 (CEN), Brazil, Distrito Federal;
*KU682358; —; —; *KU682385. Moutabea guianensis Aubl.; Baraloto 8041
(LSID), French Guiana; —; JQ625841; JQ626362; AF366966. Muraltia
spinosa (L.) F.Forest & J.C.Manning ; Chase 281 (K), Africa; —; AJ829700;
KR002178; AF366970. Muraltia thunbergii Eckl. & Zeyh.; Forest 250 (K),
Africa; AJ812637; GQ248650; AM889730; AJ842822, AJ829611. Polygala
arillata Buch.-Ham. ex D.Don; Bartholomew et al. s.n. (US), China; —;
AM234210; —; AF366975. Polygala comosa Schkuhr; Wallnöfer 13839
(FLAS), Wieringa 3462 (WAG), Europe; GQ888941; —; EU362027,
AM234344; AF365036. Polygala cyparissias A.St.-Hil. & Moq.; Carvalho
et al. 1238 (MO), Pastore 2384 (HUEFS), Brazil; —; —; *KU682427;
GQ889127. Polygala myrtifolia L.; Forest 162 (K), South Africa; —;
AJ829699; EU604043; AJ842835, AJ829624. Polygala paniculata L.; Abbott
19614 (FLAS), Guatemala; —; EU644684; EU596518; GQ889174,
GQ888848. Polygala rhinostigma Chodat; Forest & Nanni 295 (NBG),
Africa; —; AM234223; —; AM234335, GQ889185. Polygala scoparia
Kunth; Abbott 19548 (FLAS), Mexico; —; —; —; GQ888857, GQ889192.
Polygala vulgaris L. ; Fay 316 (K), Wallnöfer 13828 (FLAS), Europe; —;
JN894300; EU604046; GQ889221, GQ888870. Polygaloides chamaebuxus
(L.) O.Schwarz; Jacobs-Brouwer s.n. (U), Zpl:02814, Wallnöfer 13860 (FLAS),
Europe; —; AM234197; FR865062; GQ889118, GQ888809. Polygaloides
paucifolia (Willd.) J.R.Abbott; Abbott 25292 (FLAS), United States; —;
HQ590215; HQ593391; GQ889175. Quillaja saponaria Molina (outgroup);
Morgan 2146 (WS), Chile; —; U06822; AY386843; AF367008. Rhinotropis
californica (Nutt.) J.R.Abbott; Abbott 13010 (FLAS), Wojciechowski & Steele
887 (ASU), United States; —; —; AY386842; GQ888807, GQ889116.
Rhinotropis subspinosa (S.Watson) J.R.Abbott; Abbott 14667 (FLAS),
United States; —; —; —; AF366978. Salomonia cantoniensis Lour.; Zhanuo
91-450 (MO), Wilde & Wilde-Duyfjes 21409 (L), Asia; —; —; KR002180;
AF366996. Salomonia ciliata (L.) DC.; K'tung 78 6390 (not informed), Huq
10219 (US), Asia; —; —; KR002181; AF366997. Securidaca diversifolia (L.)
S.F.Blake; Hill 22078 (NY), Guadamuz et al. 322 (not informed), Costa Rica,
Dominica; —; JQ593516; JQ588837; AF366998. Securidaca longipedunculata
Fresen.; OM1965, OM3358 (not informed), Africa; —; JX572980; JX517755;
GQ889228, GQ888874. Xanthophyllum hainanense Hu; tree tag 1916081,
SCBGP276_2 (not informed), Asia; —; HQ415112; KP093726; —.
Xanthophyllum octandrum (F.Muell.) Domin; Forster 9554 (NY), Costion
1818 (not informed), Australia; —; AM234229; JN564163; AF367004.