Pl Syst Evol 270: 183–207 (2008)
DOI 10.1007/s00606-007-0604-4
Printed in The Netherlands
Plant Systematics
and Evolution
Biosystematics of Chamaecrista sect. Absus subsect.
Baseophyllum (Leguminosae-Caesalpinioideae) based
on allozyme and morphometric analyses
A. S. Conceição,1,2 L. P. Queiroz,1 S. M. Lambert,1 A. C. S. Pereira,1 E. L. Borba3
1
Departamento de Ciências Biológicas, Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brazil
Current address: Colegiado de Ciências, Universidade do Estado da Bahia, Campus VIII, Paulo Afonso Bahia,
Brazil
3
Departamento de Botânica, Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil
2
Received 16 February 2007; Accepted 4 September 2007; Published online 4 December 2007
Springer-Verlag 2007
Abstract. We carried out genetic and morphometric
analyses in 33 populations belonging to all nine
described and one putative taxa of Chamaecrista
subsect. Baseophyllum. Genetic variability was low in
all taxa. Morphological and genetic data are consistent
with the status of C. depauperata and C. coriacea.
However, divergence between C. cytisoides var. cytisoides and all the other six conspecific varieties is higher
than the divergence between it and the other two species.
Our data support the recognition of eight species in the
subsection Baseophyllum. Chamaecrista coriacea and
C. depauperata are supported in their original circumscription. However, the varieties ascribed to C. cytisoides are best treated as six different species. We propose
five new combinations in order to raise varieties of C.
cytisoides to specific rank: C. blanchetii, C. brachystachya, C. confertiformis, C. decora, C. unijuga. We also
propose synonimization of one variety, C. cytisoides
var. micrantha, under C. brachystachya.
Keywords: allozymes; Caesalpinioideae; campos
rupestres; Chamaecrista; Fabaceae; flower morphology; genetic diversity; morphometrics
Chamaecrista Moench is one of the largest
genera of the subfamily Caesalpinioideae. It
includes about 330 species widely distributed in
the tropical areas of America, Africa and Asia
(Lewis 2005). The genus is divided in six
sections according to Irwin and Barneby (1982)
classification, diagnosed mainly by combining
characters of the habit, inflorescence and indument. Section Absus is characterized by mostly
shrubby habit with terminal racemes. Most of its
subsections present glandular setose trichomes
and lack extrafloral nectaries. Subsection Baseophyllum differs from all other subsections by the
presence of extrafloral nectaries on the petiole or
leaf rachis, lack of glandular trichomes, leaves
with one to four pairs of leaflets, leaflets 3–9
palmately veined, and inflorescences racemosesubcorymbose (Irwin and Barneby 1978).
Subsection Baseophyllum is mainly distributed in the Espinhaço mountain range in Bahia
and Minas Gerais states, Brazil, usually in
Correspondence: A. S. Conceição, Departamento de Ciências Biológicas, Universidade Estadual de Feira de Santana, Feira de Santana, Bahia,
Brazil; current address: Colegiado de Ciências, Universidade do Estado da Bahia, Campus VIII, Paulo Afonso, Bahia, Brazil
e-mail: adilva_souza@yahoo.com.br
184
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
‘‘campo rupestre’’ vegetation (‘rocky fields’) but
sometimes occurring in ‘‘cerrado’’ (savanna-like
vegetation). It also occurs disjunctly in coastal
‘‘restinga’’ vegetation (low forests or scrubs on
salty and sandy soils) of Eastern Brazil (Pernambuco to Espı́rito Santo states) and at a spot of
‘‘caatinga’’ vegetation (seasonally dry thorny
forest) in Pernambuco state (Irwin and Barneby
1978, Conceição 2006).
Subsection Baseophyllum has a tortuous taxonomic history mainly due to different interpretations of the species limits in the Chamaecrista
cytisoides complex. Bentham (1870) recognized
four independent species and two varieties (as
Cassia): Cassia blanchetii, Cassia brachystachya
(vars. brachystachya and unijuga), Cassia coriacea and Cassia cytisoides. One year later the same
author did not recognize the variety unijuga
(Bentham 1871). Irwin and Barneby (1978)
treated all these taxa as varieties of Cassia
cytisoides, except for Cassia coriacea. Later,
they transferred these taxa to varieties of Chamaecrista cytisoides (Irwin and Barneby 1982).
According to the circumscription of Irwin and
Barneby (1982), the subsection Baseophyllum
includes only two species: Chamaecrista coriacea
(Benth.) H.S.Irwin & Barneby and C. cytisoides
(Collad.) H.S.Irwin & Barneby, the later including seven varieties: C. cytisoides var. blanchetii
(Benth.) H.S.Irwin & Barneby, C. cytisoides var.
brachystachya (Benth.) H.S.Irwin & Barneby, C.
cytisoides var. confertiformis (H.S.Irwin & Barneby) H.S.Irwin & Barneby, C. cytisoides var.
cytisoides, C. cytisoides var. decora (H.S.Irwin &
Barneby) H.S.Irwin & Barneby, C. cytisoides var.
micrantha (H.S.Irwin & Barneby) H.S.Irwin &
Barneby, and C. cytisoides var. unijuga (Benth.)
H.S.Irwin & Barneby.
According to Irwin and Barneby (1982), this
wide circumscription of C. cytisoides is supported by the conservative floral morphology
amongst the putative varieties and the presence of
intermediate morphs between the different taxa,
suggesting a continuous variation pattern in
vegetative traits. Some varieties of the C. cytisoides complex occur in sympatry and sometimes
are syntopic, although they flower in different
periods (according to data presented in Irwin and
Barneby 1978). Conceição (2000) recognized a
‘‘prostrata form’’ of C. cytisoides but did not
ascribe a formal taxonomic rank since it was not
possible to establish its relationships with other
taxa of the complex. Later, Conceição et al.
(2001) described C. depauperata Conc.,
L.P.Queiroz & G.P.Lewis, a species similar to
the ‘‘prostrata’’ morph of C. cytisoides but with
flowers smaller than those found in C. cytisoides
and C. coriacea. A summary of the current
taxonomic situation of the subsection Baseophyllum (sensu Irwin and Barneby 1982) and diagnostic characters of their taxa is presented in
Table 1 and Figs. 1, 2.
The purposes of this work are to explore the
number of taxonomical entities, their taxonomic
status and to provide diagnostic characters for all
taxa currently ascribed to Chamaecrista sect.
Absus subsect. Baseophyllym. In order to achieve
these objectives we assessed the levels of genetic
and morphological variation and sub-structuring
within and between the species and populations
of the taxa in subsect. Baseophyllum using
allozyme markers and morphometric analyses,
so as to expand the knowledge of intra and
interspecific relationships within this group.
We also investigated the correlation between
genetic and morphologic variability. The use of
genetic diagnostic markers, genetic distance/
similarities, and multivariate and statistical analyses of morphometric data can be helpful in
evaluating specific and infraspecific limits,
mainly when the traditional taxonomic procedures are not satisfactorily conclusive (Crawford
1989; van der Bank et al. 2001; Henderson 2006).
A similar approach has been successfully used in
studies of other plant groups native to the
Brazilian ‘‘campos rupestres’’ (Borba et al.
2000, 2001, 2002; Lambert et al. 2006a, b;
Pereira et al. 2007).
Materials and methods
Populations sampled. We collected 600 individuals
from 30 natural populations distributed in the nine taxa
of Chamaecrista subsection Baseophyllum (Table 2,
Figs. 1, 2) at 22 localities in Espı́rito Santo, Minas
Gerais, Bahia, Sergipe and Pernambuco states:
Taxon
(Irwin and
Barneby 1982)
Characters
Leaflet
shape
Leaflet
color
Flower
diameter (cm)
1–2
Suborbicular to
oblanceolate
Orbicular to
suborbicular
Green
1.5–1.8
campo rupestre
N-MG
C. coriacea
Green
0.5–1.0
campo rupestre
C-BA
C. depauperata
Absent or 1
short
Orbicular to
suborbicular
Green
1.5–1.8
campo rupestre,
cerrado, restinga
C. blanchetii
var.
Shrub to
brachystachya
treelet
Absent or 1–2
short
(-3)
Reniform to
widely obovate
Green
1.5–2.0
campo rupestre,
caatinga, restinga
var.
confertiformis
var. cytisoides
Short
2
1.2–2.0
campo rupestre
Short
3–4
1.5–2.0
cerrado
S-MG
C. confertiformis
C. cytisoides
Present
1
Oblong to
Green
oblanceolate
Ovate to
Green
oblanceolate
Obliquely obovate Purple or
Green
Obovate to
Green
oblanceolate
Reniform to
Green
orbicular
Suborbicular
Green
C-BA,
N-MG,
ct-ES
N-MG,
C-PE,
ct-BA
and SE
C-BA
1.5–2.0
campo rupestre
N-MG
C. decora
1.5–1.7
campo rupestre
1.3–1.7
restinga
C. brachystachya
C. unijuga
1.5–2.0
campo rupestre
C-BA,
N-MG
ct-PE, AL
and SE
C-BA
C. depauperata
C. cytisoides
var. blanchetii
var. decora
var. micrantha
var. unijuga
‘‘f. prostrata’’
Petiole
Distribution Proposed
classification
(this work)
Leaflet
pairs
C. coriacea
Habit
Habitat
Prostate
Present
sub-shrub
Prostate
Present
sub-shrub
Shrub to
treelet
Shrub to
treelet
Shrub to
treelet
Shrub to
treelet
Shrub to
treelet
Shrub to
treelet
Prostate
sub-shrub
1
Absent or 2
short
Present
1
Absent or 1
short
C. brachystachya*
C. blanchetii
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Table 1. Diagnostic characters and distribution data of the taxa of Chamaecrista section Absus subsection Baseophyllum according to Irwin and
Barneby (1982) classification with addition of C. depauperata by Conceição et al. (2001) and the unpublished form ‘‘prostrata’’ (Conceição 2000). It is
also presented the proposed classification based on the results of this work. (see discussion in the text). Distribution: C center; N Northern; S Southern; ct
coastal. Acronyms for Brazilian States: AL Alagoas; BA Bahia; ES Espı́rito Santo; MG Minas Gerais; PE Pernambuco; SE Sergipe
* Except populations CBR1, CBR4 and CBR5 (see Table 2 and discussion in the text)
185
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A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
C. coriacea (one population), C. depauperata (two
populations), C. cytisoides var. blanchetii (seven
populations), C. cytisoides var. brachystachya
(five populations), C. cytisoides var. confertiformis
(five populations), C. cytisoides var. cytisoides
(one population), C. cytisoides var. decora (three
populations), C. cytisoides var. micrantha (four
populations), and C. cytisoides var. unijuga (two
populations). We also collected three populations of
a putative taxon for the subsection, called here C.
cytisoides ‘‘f. prostrata’’. The distance among sample
individuals within a population was of approximately
20 meters. The individuals were not maintained in the
same environmental conditions due to the difficulty of
growing them in a greenhouse, as all species are large
long-lived shrubs or small trees. Vouchers for each
population are deposited in the herbarium of the
Universidade Estadual de Feira de Santana (HUEFS).
Electrophoretic procedures. Small sections of
leaf tissue were crushed in 0.5 mL of grinding buffer
(Lambert et al. 2006a). Extracts were absorbed in 1.0
· 0.3 cm Whatman number 3 paper wicks, which were
loaded into 8.5% starch gels (Sigma hydrolyzed
potato starch). For the electrodes and gels four
buffer systems were used: 1) electrode: lithium
hydroxide 0.05 mol L–1 boric acid 0.0935 mol L–1
EDTA 0.0059 mol L–1 pH 8.0; gel electrode solution
diluted 1:10; modified from Ridgway et al. (1970); 2)
electrode: citric acid 0.04M adjusted to pH 6.1 with
N-(3-aminopropyl)-morpholine;
gel:
electrode
solution diluted 1:20; modified from Clayton and
Tretiak (1972); 3) electrode: Tris 0.17 mol/L, citric
acid 0.04 mol/L, pH 8.6; gel: tris 0.02 mol/L, citric
acid 0.0013 mol/L, pH 8.6; 4) electrode: boric acid 0.3
mol/L, NaOH 0.06 mol/L, pH 8.0; gel: Tris 0.01 mol/
L, pH 8.5; modified from Shaw and Prasad (1970).
Standard horizontal eletrophoresis was performed
until the inner marker (bromophenol blue) reached 9
cm from the application site using the following
running conditions: systems 1, 2 and 3 – 25 mA;
system 4 – 15 mA. Nine enzymatic systems gave
enough resolution for reading and were used. System 1
was used for malate dehydrogenase (MDH; EC
1.1.1.37), phosphoglucomutase (PGM; EC 2.7.5.1),
esterase (EST; EC 3.1.1.1), shikimate dehydrogenase
(SKDH; EC 1.1.1.25), and glucose-6-phosphate
dehydrogenase (G6PDH; EC 1.1.1.49); system 2 was
used for isocitrate dehydrogenase (IDH; EC 1.1.1.42);
system 3 was used for leucine aminopeptidase (LAP;
AC 3.4.11.1); and system 4 was used for
phosphoglucose isomerase (PGI; EC 5.3.1.9) and
acid phosphatase (ACPH; EC 3.1.3.2). The staining
procedures were similar to but slightly adjusted from
Brune et al. (1998; ACPH, EST, LAP, SKDH,
G6PDH), Corrias et al. (1991; IDH, PGI) and Soltis
et al. (1983; PGM, MDH). Modifications were mainly
in the amounts of the components used; the exact
recipes can be obtained on request. Enzymatic
systems showing more than one locus were
numbered in ascending order from the locus with
lowest mobility. The alleles were numbered according
their mobility relative to the allele with highest
mobility of a standard individual present in all gels
and designated as 100.
Analyses of allozyme data. Banding patterns
were interpreted by direct observation of the
homozygotes and heterozygotes stained in the gels.
Genetic variability for each population was estimated
by the following parameters: proportion of
polymorphic loci (P; 0.95 criterion), mean number
of alleles per locus (A), observed (Ho) and expected
(He) mean heterozygosity per locus. Deviations from
the expected mean heterozygosity under HardyWeinberg (HW) equilibrium were tested using v2
with a correction for small samples according to
Levene (1949). Partitioning of genetic diversity
among conspecific populations was estimated by F
statistics (FIS, the inbreeding coefficient measures the
reduction in heterozygosity due to nonrandom mating
within a population; FST, measures the differentiation
among populations; Wright 1978).
Matrices of genetic distances (unbiased genetic
distance; Nei 1978) and genetic identities (unbiased
genetic identity; Nei 1978) were calculated for
populations and species. Cluster analysis was performed with the genetic distance matrix of the
populations using UPGMA (unweighted pair-group
method using arithmetical averages) as grouping
algorithm (Sneath and Sokal 1973). We also accomplished analyses with other clustering algorithm
(complete linkage, Ward, weighted pair-group) to test
the consistence of the groups. All analyses were
carried out using the BIOSYS 1.0 software package
(Swofford and Selander 1989), except for the cluster
analysis that was performed using STATISTICA for
Windows 6.0 version (StatSoft 2000).
Morphometric analyses. The individuals sampled
for the allozyme analysis were also used in analyses of
morphological variability. We measured 54 quantitative
morphological characters (except for variable #2, which
is binary), with 38 floral and 16 vegetative characters
(Table 3). All measurements of quantitative characters
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
187
Fig. 1. Distribution map of Chamaecrista sect. Absus subsect. Baseophyllum based on field collection and
herbarium material, and location of the populations studied: A, Buı́que (CBR3); B, Santo Amaro das Brotas
(CU1); C, Pirambu (CU2); D, Itabaiana (CBR5); E, Indiaroba (CB7); F, Esplanada (CBR1); G, Salvador (CBR4);
H, Saúde (CM1); I, Jacobina (CB2); J, Lençóis (CC4); K, Mucugê (CB4, CB6, CC1, CC2, CC3, CC5, CM3, D2);
L, Abaı́ra (CB3, CFP2, CM4, D1); M, Ibicoara (CFP1); N, Barra da Estiva (CFP3); O, Pedra Azul (CB5); P, Grão
Mogol (CD1, CM2); Q, Diamantina (CBR2, CD2, CD3); R, Conceição do Mato Dentro (C1); S, Guarapari
(CB1); T, Sta. Bárbara M. Verde (CCY). See Table 2 for the names of the populations. Acronyms for Brazilian
States: AL Alagoas; BA Bahia; ES Espı́rito Santo; MG Minas Gerais; PE Pernambuco; SE Sergipe
were taken with the aid of a vernier caliper. Based on
morphology and position of the nectaries, we considered
that the leaflets of bifoliolate leaves is homologous to the
second pair of leaflets of tetrafoliolate leaves. We carried
out the following analyses: 1) all populations, using only
vegetative morphological characters; 2) all populations,
188
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Fig. 2. A Chamaecrista depauperata (pop. D2); B C. coriacea (C1); C C. cytisoides var. cytisoides (CCY1); D
C. cytisoides var. blanchetii (CB2); E C. cytisoides var. brachystachya (CBR5, population from coast); F
C. cytisoides var. brachystachya (CBR3, population from interior); G C. cytisoides var. confertiformis (CC1); H
C. cytisoides var. decora (CD1); I C. cytisoides var. micrantha (CM4); J C. cytisoides var. unijuga (CU2); K
flower of C. cytisoides var. blanchetii (CB2); L flower of C. depauperata (D2). See Table 2 for the names of the
populations
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
189
Table 2. Populations of Chamaecrista coriacea, C. depauperata, C. cytisoides var. blanchetii, C. cytisoides
var. brachystachya, C. cytisoides var. confertiformis, C. cytisoides var. cytisoides, C. cytisoides var. decora, C.
cytisoides var. micrantha, C. cytisoides ‘‘f. prostrata’’ and C. cytisoides var. unijuga, occurring in northeastern
and southeastern Brazil, used in the study. Vouchers deposited at the herbarium of the Universidade Estadual de
Feira de Santana (HUEFS). Acronyms for Brazilian States: AL Alagoas; BA Bahia; ES Espı́rito Santo; MG
Minas Gerais; PE Pernambuco; SE Sergipe
Population Municipality
C. coriacea
C1
Conceição do Mato Costa Sena-Guritiba
Dentro-MG
Cima
C. depauperata
D1
Abaı́ra-BA
Catolés-Virassaia
D2
Mucugê-BA
Paty-Gerais do
Rio Preto
C. cytisoides var. blanchetii
CB1
Guarapari-ES
P. Estad. Paulo César
Vinha
CB2
Jacobina-BA
Serra do Tombador
CB3
Abaı́ra-BA
Catolés-Bicota
CB4
Mucugê-BA
Projeto Sempre Viva
CB5
Pedra Azul-MG
47 km ao Sul Pedra
Azul
CB6
Mucugê-BA
Gobira
CB7
Indiaroba-SE
2 km da entrada
de Pontal
C. cytisoides var. brachystachya
CBR1
Esplanada-BA
Faz. Morro da Graça,
Bú
CBR2
Diamantina-MG
7 km E de Diamantina
CBR3
Buı́que-PE
Serra de Jerusalém
CBR4
Salvador-BA
APA Stella Mares
CBR5
Itabaiana-SE
Serra de Itabaiana
C. cytisoides var. confertiformis
CC1
Mucugê-BA
Piabinha
CC2
Mucugê-BA
Campo Alegre
CC3
Mucugê-BA
Paty-Gerais do
Rio Preto
CC4
Lençóis-BA
Foz do rio Capivara
CC5
Mucugê-BA
Morro Fervido
C. cytisoides var. cytisoides
CCY1
Santa Bárbara
Três Cruzes-Serra
M. Verde-MG
Negra
C. cytisoides var. decora
CD1
Grão Mogol-MG
Vau-Itacambiruçu
CD2
Diamantina-MG
Beriberi
CD3
Diamantina-MG
Cristais
C. cytisoides var. micrantha
CM1
Saúde-BA
Serra de Saúde
CM2
Grão Mogol-MG
Cancela
CM3
Mucugê-BA
9 km de Igatu
Location
Voucher
18430 3100 S, 43370 2400 W
A.S.Conceição 869
13500 S, 41220 W
12450 49.100 S, 41300 0500 W
A.S.Conceição 775
A.S.Conceição 863
20350 7.800 S, 40250 17.300 W
A.S.Conceição 826
11120 S, 40410 W
13190 23.900 S, 41500 58.700 W
12590 3000 S, 41200 3000 W
16200 20.800 S, 41050 23.300 W
A.S.Conceição
A.S.Conceição
A.S.Conceição
A.S.Conceição
772
664
759
831
13040 3800 S, 41220 3100 W
E.L.Borba 1849
11290 40.400 S, 37260 18.200 W A.S.Conceição 693
12030 3800 S, 37410 3900 W
A.S.Conceição 687
18100 5400 S, 43330 45.500 W
08340 5700 S, 37140 2400 W
12560 S, 38210 W
10450 6.900 S, 42430 38.300 W
A.S.Conceição
A.S.Conceição
A.S.Conceição
A.S.Conceição
540
761
832
819
12590 33.500 S, 41200 27.800 W A.S.Conceição 755
13140 S, 41230 W
A.S.Conceição 834
12460 15.700 S, 41290 29.300 W A.S.Conceição 864
12330 S, 41240 W
A.S.Conceição 833
12590 46.400 S, 41200 17.700 W A.S.Conceição 835
21580 0.300 S, 43490 12.100 W
A.S.Conceição 870
16350 26.900 S, 42540 8.100 W A.S.Conceição 810
18080 53.200 S, 43360 53.100 W A.S.Conceição 828
18100 0.0800 S, 43350 57.300 W A.S.Conceição 827
10560 3700 S, 40240 2900 W
A.S.Conceição 774
16250 7.100 S, 42430 38.300 W A.S.Conceição 818
12560 40.500 S, 41170 20.700 W A.S.Conceição 754
190
Table 2.
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
(Continued)
Population
Municipality
CM4
C. cytisoides
CFP1
CFP2
CFP3
Abaı́ra-BA
‘‘f. prostrata’’
Ibicoara-BA
Abaı́ra-BA
Barra da Estiva-BA
C. cytisoides var. unijuga
CU1
Santo Amaro das
Brotas-SE
CU2
Pirambu-SE
Location
Voucher
Catolés-Bicota
13190 42.300 S, 41510 09.900 W
A.S.Conceição 567
13.2 km de Ibicoara
Catolés-Virassaia
7 km de Barra
da Estiva
13260 37.800 S, 41170 1600 W
13500 S, 41220 W
13400 48.300 S, 41180 40.200 S
A.S.Conceição 757
A.S.Conceição 672
A.S.Conceição 758
Ca. 2 km Rio
Pomonga
8.2 km de Pirambu
10480 2400 S, 36570 5000 W
A.S.Conceição 694
10400 5600 S, 36520 5100 W
A.S.Conceição 820
using only vegetative morphological characters
excluding variables #1 and #2 (both relative to the
number of leaflets, which could force a separation of biand tetrafoliolate groups); 3) using only floral
morphological characters (excluding CD2, CD3, CFP1,
CB1, CC2, CC3, CC4, CC5, which were not found
flowering); 4) only the populations which present two or
three pairs of leaflets, using only vegetative
morphological characters, and including for the first
pair of leaflets the same variables used for the second pair
(variable #3 to 9).
We performed canonical variate analysis (CVA)
and cluster analysis using the software package
STATISTICA 6.0. CVA was performed with population as the categorical variable (individuals were
grouped according to the population to which they
belong). The standardized coefficients for canonical
variables obtained from discriminant analysis were
used to identify the characteristics that most significantly contributed to the resulting patterns observed.
The cluster analysis was carried out using the
Mahalanobis Generalized Distance and UPGMA as
the clustering algorithm (Sneath and Sokal 1973). The
morphological variability of populations was obtained
using the median of the Mahalanobis distances of
individuals to the centroid of the group (D2m)
(Goldman et al. 2004; Lambert et al. 2006a, b). We
used the median of the squared Mahalanobis distances
instead of an average of these distances because of the
non-normal distribution of the data. A multi-response
permutation procedure (MRPP) analysis made using
PCOrd 4.10 program (McCune and Mefford 1999)
was used to calculate the chance-corrected withingroup agreement (AMRPP) among populations of every
species, and AMRPP-values were compared with the
indexes of genetic differentiation among populations
(FST) (Borba et al. 2002; Lambert et al. 2006a, b). The
average Euclidian distance among the individuals of
each population (ED) resulting from de MRPP
analysis was also used as a measure of morphological
variability within populations (Borba et al. 2002;
Lambert et al. 2006a, b). The two indexes of
morphological variability are essentially different, as
D2m is more affected by form and ED is more affected
by size of the characters (Lambert et al. 2006a, b). A
Spearman rank correlation analysis between the
morphological (ED and D2m) and genetic (He)
variability of populations was also carried out with
STATISTICA 6.0.
Results
Variation within populations. Using nine
enzymatic systems 15 loci were obtained with
good resolution and were used in the study
(Table 4). Three loci were monomorphic for all
populations (ACPH, MDH and IDH-1). Five loci
displayed low polymorphism, with only two or
three alleles (PGI-2, EST-1, SKDH, G6PDH and
LAP-2). The remaining loci present four to eight
alleles per locus. PGM-1 was the most
polymorphic, with eight alleles. Some alleles
were exclusive to a single taxon: five to C.
coriacea; one to C. cytisoides var. unijuga; five to
C. cytisoides var. cytisoides; two to C. depauperata.
A few alleles were exclusive to one population.
The percentage of polymorphic loci (P; 0.95
criterion) ranged from 0.0 to 33.3%, the mean
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
number of alleles per locus was 1.1 to 1.4, and
mean heterozygosity (He) ranged from 0.010 to
0.067 (Table 5). Populations CCY1 (C. cytisoides
var. cytisoides), D1 (C. depauperata), CBR1 (C.
cytisoides var. brachystachya), CD3 (C. cytisoides var. decora) and CU1 (C. cytisoides var.
unijuga) displayed the highest genetic variability.
The majority of the populations (72.7%) were in
HW equilibrium in all loci. The other populations
presented significant deviations in only one locus
each. Five loci were not in HW equilibrium in at
least one population: EST-1 (C1), G6PDH
(CCY1), LAP-1 (CM2) and IDH-2 (in all populations of CC) with deficit of heterozygotes, and
LAP-2 (CD3) with excess of heterozygotes. The
low values for FIS reflect the HW equilibrium in
most of the taxa, except for C. cytisoides var.
confertiformis (CC) that presented very high FIS
(Table 6).
In both morphological analyses, the population with the highest variability was CBR1,
which also presented one of the highest genetic
variability (Table 5). The other populations presenting high morphological variability expressed
as D2m were CM3, CBR5 and CU1, while the
lowest variability was found in CC3 (Table 5).
The most variable characters considering all taxa
pooled were leaflet number (variable #1), number
of main veins (#11), length of the leaflet (#3),
width at 1/3 of the leaflet (#5), apex angle of the
leaflet (#9), least distance from the margin to the
main nervure, in the base of the leaflet (#7),
length of the rachis (#12), and length of the
petiole (#10). The least variable character was the
distance of the nectary in relation to the second
pair of leaflets (#16). Spearman rank correlations
between morphological and genetic variability
resulted in a statistically significant positive high
correlation between He and D2m (r = 0.917,
p £ 0.001) and between He and ED (r = 0.901,
p £ 0.001).
Genetic structuring. Chamaecrista cytisoides var. blanchetii and C. cytisoides var.
brachystachya displayed very high values of
FST, interpreted as a very high level of genetic
structuring. The other taxa showed moderate to
low average values of FST (Table 6). By
excluding populations CB1 and CB5 of C.
191
cytisoides var. blanchetii and CBR2 of C.
cytisoides var. brachystachya the average values
of FST drop to 0.042 and 0.053, respectively. In
C. cytisoides var. blanchetii this is due to the
allele frequencies of IDH-2, and in C. cytisoides
var. brachystachya to the loci PGM-1 and PGI-1.
Morphological structuring. Chameacrista
cytisoides var. brachystachya also presented the
highest value of morphological structuring
(AMRPP = 0.361), correlated with the highest
values of genetic differentiation (FST). C.
cytisoides var. confertiformis and C. cytisoides
var. micrantha that presented high values of
morphological differentiation had low values of
genetic differentiation (Table 6).
Phenetic relationships – genetic data.
Populations of C. depauperata, C. cytisoides
var. decora, C. cytisoides var. micrantha and C.
cytisoides var. unijuga did not present any intrataxon genetic differentiation. The populations of
C. cytisoides var. blanchetii, C. cytisoides var.
confertiformis, and C. cytisoides ‘‘f. prostrata’’
also presented very high genetic identity among
intra-taxon populations, ranging from 0.93 to
1.00. The lowest intra-taxon genetic identity
(0.87) was found within C. cytisoides var.
brachystachya, between the population CBR2
and the four remaining populations.
The UPGMA dendrogram obtained from the
cluster analysis of Nei’s (1978) unbiased genetic
distances (Fig. 3) reveals the formation of two
main groups. The first contains the population of
C. cytisoides var. cytisoides (CCY) and the two
populations of C. depauperata (D). The high
divergence of C. cytisoides var. cytisoides and C.
depauperata is due to the presence of exclusive
alleles fixed in two enzymatic systems (both in
PGM-2 and LAP-1), besides the same fixed allele
in PGI-1, similar for the two taxa (Table 4). The
second and largest group may be subdivided in
three subgroups: one formed by all of the
populations of C. cytisoides var. decora (CD)
and population (C) of C. coriacea; another
composed of the two populations of C. unijuga
and population CBR2; the third formed by the
populations of the remaining taxa. In this third
subgroup only C. cytisoides var. confertiformis
formed group containing exclusively all of their
192
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Table 3. Characters used in the morphometric analyses of 33 populations of nine described and one
putative taxa of Chamaecrista sect. Absus subsect.
Baseophyllum occurring in northeastern and southeastern Brazil
Characters
Vegetative characters
Leaves (second par of leaflets)
1- Number of leaflets
2- Occurrence of variation in the
number of leaflets
3- Length of leaflet
4- Maximum width of leaflet
5- Width of leaflet at 1/3 of its length
6- Width of leaflet at 2/3
7- Least distance from the margin to the
main vein, in the base of the leaflet
8- Maximum distance from the margin to the
main vein, in the base of the leaflet
9- Apex angle of the leaflet
10- Length of the petiole
11- Number of main veins
12- Length of the rachis
13- Length of the nectary
14- Width of the nectary
15- Number of nectaries
16- Distances of the nectary to the
second par of leaflets
Floral characters
Inflorescence
17- Length
18- Number of flowers per raceme
Sepal
19- Length of the first sepal
20- Width of the first sepal
21- Length of the second sepal
22- Width of the second sepal
23- Length of the third sepal
24- Width of the third sepal
25- Length of the fourth sepal
26- Width of the fourth sepal
27- Length of the fifth sepal
28- Width of the fifth sepal
Petal
29- Length of the first petal
30- Maximum width of the first petal
31- Width of the first petal at 1/3 of its length
32- Width of the first petal at 2/3
33- Maximum distance from the margin to the
main vein, in the base of the first petal
Table 3.
(Continued)
Characters
34- Least distance from the margin to the main
nervure, in the base of the first petal
35- Length of the second petal
36- Maximum width of the second petal
37- Width of the second petal at 1/3
38- Width of the second petal at 2/3
39- Length of the third petal
40- Maximum width of the third petal
41- Width of the third petal at 1/3
42- Width of the third petal at 2/3
43- Length of the fourth petal
44- Maximum width of the fourth petal
45- Width of the fourth petal at 1/3
46- Width of the fourth petal at 2/3
47- Length of the fifth petal
48- Maximum width of the fifth petal
48- Width of the fifth petal at 1/3
50- Width of the fifth petal at 2/3
Anther
51- Length of the anthers
Style
52- Length of the style
Ovary
53- Length of the ovary
54- Number of ovules
intra-taxon populations. Populations of C. cytisoides var. blanchetii, C. cytisoides var. brachystachya, C. cytisoides var. micrantha and
C. cytisoides ‘‘f. prostrata’’ formed mixed groups.
Four populations of C. cytisoides var. brachystachya are nested in the group containing most
of the populations of C. cytisoides var. blanchetii;
the remaining population is very divergent
(CBR2) due to the presence of two exclusive
fixed alleles in two loci (Table 4).
Phenetic relationships – morphological
data. The scatterplots of the scores of
individuals on the first two CVA canonical axes
of the morphological analyses are shown in
Fig. 4. The first and second canonical axes
explained respectively 56% and 16% of the
variation in the analysis including only
vegetative characters. On the first canonical axis
there is a separation of three groups, one with
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
C. cytisoides var. cytisoides, one with the
populations of C. cytisoides var. confertiformis,
C. coriacea and C. cytisoides var. brachystachya
(except for CBR5), and another with C. unijuga,
C. cytisoides var. decora, C. depauperata, C.
cytisoides ‘‘f. prostrata’’, C. cytisoides var.
blanchetii and population CBR5 (Fig. 4A). The
variables that more contributed to that separation
in the first axis were the leaflets number (variable
#1), length of the second pair of leaflets (#3) and
number of main veins (#11). In the second axis C.
coriacea was separated from the remaining taxa
in the second group, and the third group was
divided in two subgroups, one containing C.
unijuga, C. cytisoides var. decora and C.
depauperata and the other with the remaining
taxa. The variables that contributed most to the
separation on this second canonical axis were the
length of the petiole (variable #10) and the least
distance from the margin to the main nervure in
the base of the leaflets (#7). This structure had no
significant changes in the analysis in which the
variables #1 and #2 (both relative to the number
of leaflets) were excluded (data not shown). In
the matrix of classification the percentage of
correct classifications ranged from 38 to 100%.
The incorrect classifications mostly occurred
between populations of the same taxon, except
for eight individuals of CBR1 which were
classified in CU1 and CB7, eight individuals of
CBR5 in CB7 and CB2, and one individual of
CBR3 in CM4.
The first and second canonical axes explained
42% and 16%, respectively, of the variation in
the analysis using only floral characters. In the
first canonical axis the two populations of C.
depauperata are separated from the other taxa
(Fig. 4B). The variables that contributed most to
this separation were maximum width of the
fourth petal (variable #44), length of the anthers
(#51), width at 2/3 of the fourth petal (#46) and
number of ovules (#54). In the second axis
occurred a slight separation of C. cytisoides var.
cytisoides from the remaining taxa.
The same structure observed in the CVA was
found in the UPGMA dendrogram obtained from
the cluster analysis of Mahalanobis distance
among the centroids of the populations. The
193
analysis using vegetative morphological characters resulted in the separation of C. cytisoides var.
cytisoides, the only taxon presenting predominantly three pairs of leaflets, and two other
groups composed by the populations presenting
one or two pair of leaflets (Fig. 5A). Populations
of the same taxon are mostly present only in one
of these two last groups, except for C. cytisoides
var. brachystachya, with three population in one
group and two in the other. In the subgroup
presenting predominantly two pairs of leaflets,
the population CBR4 and C. coriacea displayed
the greatest differentiation, and the remaining
populations were divided into two subgroups, one
including all the populations of C. cytisoides var.
confertiformis and the other composed by all
populations of C. cytisoides var. micrantha plus
the populations CBR2 and CBR3. In the subgroup presenting predominantly one pair of
leaflets, two subgroups were formed: one composed of all populations which do not have a
petiole, intermingled (C. cytisoides var. brachystachya, C. cytisoides ‘‘f. prostrata’’ and C.
cytisoides var. blanchetii), and another of all
populations that possess a petiole (C. unijuga, C.
cytisoides var. decora and C. depauperata; the
populations of each taxon grouping together in
three subgroups).
The morphological cluster analysis carried out
only with the taxa presenting two or three pairs of
leaflets resulted in the same general structure of
the analysis carried out with only the second pair
of leaflets. In the analyses using other clustering
algorithms, the same structuring was found. In the
analyses which variables 1 and 2 were excluded,
the same groups were maintained, except for the
population CBR4 grouping with CCY1.
The UPGMA dendrogram obtained from the
cluster analysis of morphological distance using
only floral characters resulted in the formation of
two main groups (Fig. 5B): a group containing
only the populations of C. depauperata, and
another larger group composed by the populations of all remaining taxa. This larger group
presents high degree of chaining, with the
populations of all taxa intermingled; no subgroups composed primarily by all populations of
the same taxon were formed.
194
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Table 4. Allelic frequencies at 15 loci in populations of Chamaecrista coriacea (C), C. cytisoides var. cytisoides
(CCY), C. depauperata (D), C. cytisoides var. decora (CD), C. cytisoides var. unijuga (CU), C. cytisoides
‘‘f. prostrata’’ (CFP), C. cytisoides var. micrantha (CM), C. cytisoides var. blanchetii (CB), C. cytisoides var.
brachystachya (CBR) and C. cytisoides var. confertiformis (CC), occurring in northeastern and southeastern
Brazil. N = sample size. See Table 2 for the name of the populations
Locus C1
ACPH
100
(N)
PGI-1
140
130
115
110
100
90
(N)
PGI-2
100
95
(N)
PGM-1
120
116
107
100
92
83
75
41
(N)
PGM-2
106
100
93
81
(N)
MDH
100
(N)
EST-1
106
100
75
(N)
EST-2
120
105
100
95
90
(N)
EST-3
100
94
85
80
(N)
SKDH
100
93
(N)
G6PD
105
100
95
(N)
IDH-1
100
(N)
IDH-2
100
CCY1 D1 D2 CD1 CD2 CD3 CU1 CU2 CFP1 CFP2 CFP3 CM1 CM2 CM3 CM4
1.00 1.00
19
20
1.00 1.00 1.00
19
20
18
1.00
20
1.00
20
1.00
17
1.00
20
1.00
20
1.00
18
1.00
20
-
1.00
1.00 19
20
1.00
19
-
-
-
1.00
14
-
-
11
1.00
10
1.00
11
1.00
18
1.00 1.00
19
20
1.00 1.00 1.00
19
20
17
1.00
18
1.00
11
1.00
14
1.00
14
1.00
15
-
-
-
-
1.00
18
-
-
-
-
-
0.83 1.00
0.17 18
20
-
1.00
16
0.83
1.00
16
-
1.00
0.17 0.13 18
16
16
-
1.00
8
0.88
-
1.00
-
7
1.00 1.00 19
14
15
12
-
1.00 1.00
14
16
1.00 1.00 1.00
19
20
17
1.00
20
1.00
20
-
19
1.00 16
20
20
20
-
-
-
-
0.82 0.83
0.18 0.18
19
20
0.85 0.88
0.15 0.12
17
16
1.00
12
0.2
0.70
0.10
20
1.00 1.00
15
17
1.00
1.00
1.00
1.00
0.84
0.16
0.07
-
1.00
8
0.05
1.00
17
1.00
6
0.11
1.00
18
1.00
13
1.00
12
1.00
13
1.00
16
1.00
19
1.00
20
1.00
13
1.00
18
1.00
15
1.00
17
-
-
-
-
-
-
1.00
12
1.00
13
-
-
0.03
0.97
14
19
1.00
19
-
-
-
-
-
1.00
1.00
-
1.00
20
1.00
20
1.00
11
20
20
20
8
0.05
13
10
1.00
13
1.00
20
1.00
20
1.00
20
1.00
20
20
20
20
20
-
-
-
0.16
7
1.00
6
0.03
1.00
7
1.00
-
-
-
-
-
0.98
20
0.84
19
1.00
20
1.00
10
1.00
0.05
0.8
0.15
20
0.98 0.16 0.03 16
20
20
20
20
20
20
20
20
20
-
0.96
0.04
14
0.98
0.02
20
0.98
0.02
20
0.95
0.05
19
0.98
0.02
20
0.98
0.02
20
0.96
0.04
13
0.94
0.06
16
0.94
0.06
17
0.94
0.06
18
0.97
0.03
19
0.97 0.98 0.97
0.03 0.02 0.03
18
20
15
-
-
-
-
-
-
-
-
-
-
-
0.96
0.04
14
0.96
0.04
11
0.96
0.04
13
0.94
0.06
16
0.95
0.05
20
0.95
0.05
20
0.93
0.07
20
0.97
0.03
17
0.96
0.04
13
0.97
0.03
19
0.98
0.02
20
1.00 1.00 1.00
15
15
14
1.00
11
1.00
9
1.00
16
1.00
18
1.00
20
1.00
20
1.00
20
1.00
19
1.00
14
1.00
11
1.00
20
1.00
1.00
-
-
-
-
-
-
-
-
-
-
0.94 0.96 0.98
0.06 0.04 0.02
9
14
20
-
1.00
1.00
1.00
0.93
0.05
0.18
0.82
10
0.03
-
0.87
0.03
-
19
0.84
0.81
0.12
-
16
-
0.83
0.12
-
-
-
-
1.00
9
8
-
20
17
0.86
0.07
-
1.00
20
-
8
1.00
-
0.06
-
-
19
0.06
0.94
1.00
20
0.06
0.94
8
1.00
1.00
20
14
19
0.91
0.09
1.00
1.00
1.00
20
0.96
11
1.00 1.00
15
20
-
-
6
-
-
-
1.00
-
1.00
0.81 0.19 18
20
0.80 1.00
0.20 5
19
-
1.00
1.00
8
1.00
20
1.00
-
6
1.00
11
1.00
-
5
1.00
-
8
8
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
195
Table 4. (Continued)
CB1 CB2 CB3 CB4 CB5 CB6 CB7 CBR1 CBR2 CBR3 CBR4 CBR5 CC1 CC2 CC3 CC4 CC5
1.00
20
1.00
20
1.00
18
1.00
20
1.00
20
1.00
15
1.00
20
1.00
20
1.00
20
1.00
20
1.00
16
1.00
20
1.00
20
1.00
11
1.00
17
1.00
12
1.00
19
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1.00
18
1.00
20
1.00
18
1.00
1.00
-
1.00
-
20
20
18
14
20
18
20
13
20
1.00
18
1.00
20
1.00
19
1.00
14
1.00
20
1.00
18
1.00
14
1.00
20
1.00
16
1.00
16
1.00
19
1.00
12
0.96
0.04
13
-
-
-
-
-
-
-
-
-
-
-
-
-
-
20
13
1.00
10
1.00
12
12
-
-
-
-
-
1.00
0.06
0.94
1.00
1.00
1.00
0.04
0.96
1.00
0.14
0.86
1.00
1.00
1.00
1.00
17
20
13
11
16
-
-
-
-
-
-
-
1.00
1.00
1.00
20
17
-
1.00
20
1.00
20
1.00
18
-
-
-
1.00
1.00
-
1.00
1.00
14
1.00
20
-
-
-
1.00
13
-
-
-
-
1.00
20
1.00
13
1.00
20
-
0.97
0.04
15
0.98
0.02
20
0.96
0.04
13
0.97
0.03
18
0.98
0.02
20
-
-
-
-
0.82
0.18
14
0.97
0.03
18
0.96
0.04
13
0.92
0.08
12
1.00
18
1.00
13
1.00
20
1.00
11
-
-
-
1.00
13
1.00
11
-
-
-
1.00
11
1.00
-
9
9
-
0.79
0.21
-
1.00
20
1.00
20
-
-
6
1.00
-
6
0.03
1.00
0.85
0.12
1.00
8
1.00
-
-
1.00
17
0.96
0.04
13
0.96
0.04
14
-
-
-
1.00
14
-
-
-
1.00
-
8
1.00
1.00
11
1.00
-
9
-
1.00
9
-
1.00
1.00
13
1.00
20
1.00
20
1.00
16
1.00
20
20
20
20
20
20
-
-
-
-
-
1.00
16
1.00
20
1.00
20
1.00
15
1.00
16
-
20
-
-
-
-
-
1.00
19
1.00
16
1.00
18
12
13
10
20
1.00
-
1.00
10
1.00
11
1.00
10
-
1.00
5
0.98
0.02
-
1.00
-
6
9
6
9
0.92
0.08
0.89
0.11
0.90
0.10
0.90
0.10
18
1.00
10
-
0.97
0.04
15
0.95
0.05
20
0.85
0.15
17
0.97
0.03
18
0.96
0.04
14
0.95
0.05
20
0.94
0.06
16
0.97
0.04
15
0.97
0.04
15
0.98
0.02
20
0.98
0.02
20
0.96
0.04
11
-
-
-
-
-
-
-
-
-
-
-
-
-
0.92
0.08
13
0.92
0.08
13
0.98
0.02
20
0.95
0.05
19
0.97
0.03
15
0.96
0.04
14
0.98
0.02
20
0.91
0.09
16
1.00
14
1.00
6
1.00
20
1.00
18
1.00
9
1.00
18
1.00
20
1.00
20
-
-
-
-
-
8
1.00
-
8
1.00
-
7
1.00
6
-
1.00
1.00
-
6
5
-
5
1.00
11
1.00
-
10
0.92
0.08
20
7
20
1.00
9
15
-
1.00
1.00
19
-
9
1.00
1.00
14
-
0.17
0.81
0.03
1.00
9
20
12
8
-
1.00
0.96
0.04
7
17
16
1.00
-
-
1.00
-
6
-
1.00
18
1.00
7
1.00
-
-
1.00
0.10
0.90
1.00
8
1.00
12
0.92
0.08
-
1.00
17
8
6
1.00
-
1.00
6
8
1.00
1.00
1.00
-
18
9
1.00
1.00
1.00
13
1.00
-
8
7
1.00
0.917
-
1.00
0.083
12
1.00
10
-
9
13
1.00
5
1.00
7
1.00
11
1.00
12
1.00
6
0.2
0.08
0.64
0.29
0.18
9
196
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Table 4.
(Continued)
Locus C1
CCY1 D1 D2 CD1 CD2 CD3 CU1 CU2 CFP1 CFP2 CFP3 CM1 CM2 CM3 CM4
93
90
(N)
LAP-1
100
93
90
89
(N)
LAP-2
117
100
94
(N)
17
15
15
-
-
14
10
1.00
17
19
1.00
1.00 12
9
-
-
-
-
-
1.00 -
17
1.00
19
1.00
1.00
17
1.00
19
1.00
20
0.17
0.83
12
11
9
1.00
-
1.00
-
8
7
0.17
0.83
0.25
0.75
20
12
-
-
1.00
16
1.00
18
0.07
0.93
20
1.00
14
1.00
10
1.00
12
-
0.19
0.81
0.19
0.81
-
-
16
Discussion
Genetic variability. The genetic variation found
within populations for all taxa of the subsection
Baseophyllum are low when compared with
values found by Hamrick and Godt (1992) for
plants of regional and endemic geographical
distribution, respectively (P = 36.4, A = 1.55,
and
P = 26.3,
A = 1.39,
He = 0.118
He = 0.063). However, the variability levels
found in this study were similar to the values
found for other genera of Leguminosae (e.g.
Baatout et al. 1991, Liston 1992, Chamberlain
1998, Casiva et al. 2002, Sotuyo et al. 2004). The
low variability presented within populations for
all taxa of the subsection Baseophyllum may be
associated with the pollinator behavior and
mating systems. These plants are selfcompatible and pollinated by social bees
presenting a behavior which favors endogamy
(Costa 2007, Silva-Pereira 2007).
The populations of C. cytisoides var. micrantha and C. depauperata which had the highest
diversity (CM3 and D1, respectively) are located
in preserved areas, where human activity is
restricted. Conversely, population CC2 of C.
cytisoides var. confertiformis occurring in a site
under anthropogenic action displayed the lowest
genetic diversity of the taxon, in spite of its
moderate morphological variability. The populations of C. cytisoides var. blanchetii with slightly
higher genetic diversity are located in disjunct
areas, CB1 in the restinga of Espı́rito Santo state,
and CB5 in the savanna of Minas Gerais state,
whilst the other populations occur in areas of
16
1.00
12
-
-
-
-
1.00
20
1.00
20
1.00
19
1.00
14
1.00
9
0.05
0.95
20
1.00
1.00
10
1.00
19
0.95
1.00
10
1.00
16
-
-
-
-
8
1.00
20
1.00
10
1.00
16
-
0.05
20
20
1.00
15
1.00
16
‘‘campos rupestres’’ in Bahia state. These two
disjunct populations presented an exclusive fixed
allelle in the locus IDH-2, indicating a restricted
gene flow among these three major areas of
occurrence of the taxon.
Chamaecrista cytisoides var. confertiformis
was the only taxon presenting high values of FIS,
which could be the result of endogamy or substructuring of the populations. This taxon displays a singular spatial distribution for subsection
Baseophyllum, almost always occurring in valleys along the margins of rivers, forming dense
linear populations. In addition to this spatial
distribution, self-compatibility and the behavior
of its pollinator favors endogamy and nonrandom mating (Costa 2007), probably leading
to genetic sub-structuring of the populations.
Morphologic variability. Studies of intraand inter-populational multivariate morphological variability are apparently absent in
Chamaecrista and scarce in Leguminosae
(Riggins et al. 1977, Casiva et al. 2002, Leht
and Jaaska 2002, Boonkerd et al. 2005). The
number of leaflets is not the only characteristics
that forced the separation of the three groups in
the multivariate analyses. The length of the
rachis, apex angle of the leaflet, length of the
leaflets, and least distance from the margin to
the main vein in the base of the leaflet sustained
the structure of three groups, as the same groups
were maintained in the analysis excluding the
variables 1 and 2. It is important notice since that
these characters were not evidenced by Irwin and
Barneby (1978) or by Bentham (1870, 1871) in
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
197
Table 4. (Continued)
CB1 CB2 CB3 CB4 CB5 CB6 CB7 CBR1 CBR2 CBR3 CBR4 CBR5 CC1 CC2 CC3 CC4 CC5
18
1.00
6
1.00
12
-
-
-
1.00
13
1.00
17
1.00
12
1.00
13
1.00
16
1.00
10
-
-
-
-
-
0.97
0.03
15
1.00
14
12
1.00
9
1.00
-
1.00
-
7
7
1.00
6
-
-
-
-
1.00
20
1.00
18
1.00
9
1.00
18
1.00
20
1.00
9
1.00
20
1.00
11
1.00
11
1.00
16
-
1.00
9
-
-
-
-
-
1.00
20
-
1.00
12
1.00
16
the distinction of the taxa in the subsection
Baseophyllum.
Taxonomic
implications. Chamaecrista
coriacea was recognized (as Cassia) by
Bentham (1840) as a peculiar species,
characterized by one or two pairs of palmately
veined leaflets, and its status has never been
objected (Irwin and Barneby 1978, 1982). Both
allozyme and morphological data supported its
distinction at specific rank. Chamaecrista
coriacea differs from the other taxa of the
subsection Baseophyllum in at least one
allozyme locus, with a genetic identity of 0.78
to C. cytisoides var. cytisoides. In the discriminant
analysis C. coriacea differed from C. cytisoides
var. cytisoides mainly is its leaves predominantly
with two pairs of leaflets, smaller rachis and
smaller leaflets.
The recently described C. depauperata was
the only taxon with clearly distinctive flowers,
as evidenced in the morphological analyses of
floral characters, mainly due to the width of
fourth petal, length of the anthers and fewer
ovules. In the analyses using vegetative morphological data, C. depauperata also differed of
the other taxa by presenting petiolate leaves with
two pairs of smaller leaflets. Furthermore, this
species is very distinctive genetically in at least
three allozyme loci (two of them being diagnostic and the other similar to C. cytisoides var.
cytisoides). Chamaecrista depauperata and
C. cytisoides var. cytisoides are quite different
in both habit and habitat (prostrate sub-shrubs
from ‘‘campo rupestre’’ vegetation vs. shrubs to
1.00
16
0.05
0.95
20
1.00
1.00
-
5
6
-
1.00
10
1.00
16
-
-
-
-
-
0.8
10
0.92
12
0.36
14
0.71
14
0.82
11
1.00
16
1.00
13
1.00
11
1.00
11
-
-
-
-
-
-
1.00
16
1.00
13
1.00
10
1.00
11
1.00
9
1.00
15
treelets from ‘‘cerrado’’), and in number and
form of the leaflets (Table 1). Besides, C.
depauperata occurs exclusively in the state of
Bahia in the southern portion of the Chapada
Diamantina, while C. cytisoides var. cytisoides is
endemic to southern portion of Minas Gerais
state (Fig. 1).
The high similarity in floral traits has probably lead Irwin and Barneby (1978) to treat some
of the former species of the subsection Baseophyllum as varieties of C. cytisoides. This picture
was possibly reinforced by the occurrence of
hybridization in some taxa, such as between C.
cytisoides var. confertiformis and C. cytisoides
var. blanchetii and between C. cytisoides var.
confertiformis and C. depauperata (Conceição
et al. unpubl.). These hybrids are very rare in
nature due to several isolation barriers between
the sympatric species, such as different flowering
period in C. cytisoides var. confertiformis and
C. cytisoides var. blanchetii (Costa 2007), but
may create confusion among taxonomists
when they examine only herbarium collections.
In spite of being reproductively viable, apparently these hybrids are not stable and do not
compromise taxon circumscription in this group
(Costa 2007).
The values of genetic identities found among
populations of the C. cytisoides complex were
similar to the values usually found between
populations of distinct species (Thorpe 1982,
Crawford 1989, van der Bank et al. 2001). The
average genetic identities between the population
where the holotype of C. cytisoides var. cytiso-
198
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Table 5. Genetic variability at 15 allozymic loci and morphological variability (D2m and ED) based on the
morphometric analyses of 54 characters in 33 populations of nine described and one putative taxa of
Chamaecrista sect. Absus subsect. Baseophyllum, occurring in northeastern and southeastern Brazil. N = mean
sample size per locus; genetic parameters - A = mean number of alleles per locus, P = percentage of
polymorphic loci, Ho = observed and He = expected mean heterozygosity per locus (Nei, 1978; unbiased
estimate); morphological parameters - D2m = median of the Mahalanobis Generalized Distance of the
individuals to the centroid of the population, ED mean of the Euclidean distance between the individuals of the
population. Standard deviations in parentheses. A locus was considered polymorphic if the frequency of the most
common allele did not exceed 0.95. See Table 2 for the name of the populations
Population
N
P
Ho
He
(0.1)
33.3
0.092 (0.041)
0.010 (0.039)
8.98
-
(0.1)
(0.1)
26.7
13.3
0.075 (0.034)
0.040 (0.020)
0.067 (0.029)
0.038 (0.018)
5.88
6.40
12.26
10.62
(0.1)
(0.1)
(0.1)
(0.1)
(0.1)
(0.1)
(0.2)
13.3
0.0
6.7
13.3
13.3
6.7
13.3
0.036
0.011
0.026
0.033
0.042
0.015
0.030
(0.025)
(0.006)
(0.012)
(0.021)
(0.030)
(0.011)
(0.021)
0.032
0.011
0.025
0.031
0.037
0.014
0.028
(0.021)
(0.006)
(0.012)
(0.019)
(0.025)
(0.011)
(0.019)
9.34
12.19
5.91
15.88
5.94
6.73
17.47
18.11
19.37
14.37
21.37
15.55
13.52
24.28
(0.2)
(0.1)
(0.1)
(0.1)
(0.1)
26.7
0.0
0.0
6.7
20.0
0.066
0.011
0.010
0.010
0.033
(0.033)
(0.006)
(0.006)
(0.007)
(0.016)
0.058
0.011
0.010
0.010
0.032
(0.028)
(0.006)
(0.006)
(0.007)
(0.015)
38.13
7.17
17.76
14.34
20.70
32.47
18.02
16.24
19.55
21.44
(0.1)
(0.1)
(0.1)
(0.1)
(0.1)
13.3
13.3
13.3
13.3
20.0
0.021
0.020
0.017
0.022
0.032
(0.012)
(0.016)
(0.014)
(0.014)
(0.015)
0.043
0.029
0.048
0.049
0.052
(0.024)
(0.017)
(0.033)
(0.030)
(0.024)
5.33
4.25
3.35
5.75
8.47
12.81
16.38
13.88
12.46
16.01
(0.2)
20.0
0.053 (0.029)
0.066 (0.038)
7.12
-
(0.2)
(0.2)
(0.2)
13.3
13.3
13.3
0.049 (0.028)
0.056 (0.031)
0.068 (0.048)
0.045 (0.025)
0.050 (0.027)
0.058 (0.033)
13.61
11.93
11.47
20.71
18.62
16.85
(0.1)
(0.1)
(0.2)
(0.1)
6.7
13.3
13.3
6.7
0.012
0.013
0.031
0.014
0.012
0.019
0.034
0.013
(0.009)
(0.010)
(0.023)
(0.008)
10.89
4.55
20.90
5.57
17.05
9.10
19.64
14.10
(0.1)
(0.1)
20.0
13.3
0.034 (0.014)
0.037 (0.024)
0.034 (0.014)
0.033 (0.021)
5.38
8.52
15.38
19.90
A
C. coriacea
C1
15.6 (1.1)
1.3
C. depauperata
D1
17.6 (0.7)
1.3
D2
17.0 (0.7)
1.3
C. cytisoides var. blanchetii
CB1
16.7 (1.1)
1.2
CB2
16.3 (1.0)
1.2
CB3
14.4 (1.1)
1.3
CB4
12.3 (1.0)
1.2
CB5
13.6 (1.4)
1.2
CB6
11.0 (1.3)
1.1
CB7
16.1 (1.5)
1.3
C. cytisoides var. brachystachya
CBR1
15.9 (1.2)
1.3
CBR2
13.5 (1.3)
1.2
CBR3
14.5 (1.6)
1.1
CBR4
14.7 (1.5)
1.1
CBR5
14.8 (1.1)
1.3
C. cytisoides var. confertiformis
CC1
11.1 (1.3)
1.3
CC2
10.6 (1.1)
1.2
CC3
13.1 (1.4)
1.2
CC4
12.3 (1.2)
1.3
CC5
11.6 (1.2)
1.3
C. cytisoides var. cytisoides
CCY1
18.6 (0.4)
1.3
C. cytisoides var. decora
CD1
13.3 (1.4)
1.3
CD2
12.3 (1.7)
1.3
CD3
10.8 (1.6)
1.3
C. cytisoides var. micrantha
CM1
14.2 (1.6)
1.1
CM2
12.3 (1.7)
1.2
CM3
13.4 (1.4)
1.3
CM4
15.3 (1.4)
1.2
C. cytisoides ‘‘f. prostrata’’
CFP1
13.3 (1.8)
1.3
CFP2
15.3 (1.8)
1.3
(0.0009)
(0.0009)
(0.021)
(0.008)
D2m
ED
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Table 5.
Population
199
(Continued)
N
CFP3
13.9 (1.8)
C. cytisoides var. unijuga
CU1
14.7 (1.3)
CU2
13.7 (1.5)
A
P
Ho
He
D2m
ED
1.3 (0.1)
13.3
0.040 (0.023)
0.036 (0.020)
8.54
18.02
1.4 (0.2)
1.3 (0.2)
20.0
26.7
0.059 (0.029)
0.050 (0.027)
0.054 (0.026)
0.046 (0.023)
20.47
15.94
23.48
23.22
Table 6. F statistics (Wright 1978) at 15 loci and AMRPP from morphometric analysis in populations of seven
taxa and one morphospecies of Chamaecrista sect. Absus subsect. Baseophyllum occurring in northeastern and
southeastern Brazil
Taxon
No. of Pops
FIS
FST
AMRPP
C.
C.
C.
C.
C.
C.
C.
C.
2
7
5
5
3
4
3
2
–0.141
–0.122
–0.105
0.476
–0.177
0.084
–0.111
–0.141
0.016
0.539
0.648
0.109
0.005
0.049
0.035
0.007
0.128
0.182
0.361
0.335
0.055
0.321
0.070
0.080
depauperata
cytisoides var. blanchetii
cytisoides var. brachystachya
cytisoides var. confertiformis
cytisoides var. decora
cytisoides var. micrantha
cytisoides ‘‘f. prostrata’’
cytisoides var. unijuga
ides was collected and all other taxa in this
complex are lower than the values found between
conspecific populations, ranging from 0.64 to
0.78. Conversely, the genetic identity of that
population with both C. depauperata and C.
coriacea was higher. Chamaecrista cytisoides
var. cytisoides was also the most divergent
population in the morphological analyses based
on vegetative characters, mainly due to its leaves
with three pairs of leaflets. Both low values of
genetic identity and high morphological differences of the type taxon for the complex with the
other conspecific varieties reinforce the hypothesis of the existence of distinct species in the
complex. These results point out that only the
population of C. cytisoides var. cytisoides sampled, among the 33 populations studied, should
remain in the species.
The populations of C. cytisoides var. confertiformis, C. cytisoides var. decora and C. cytisoides var. unijuga formed three very distinctive
groups in both genetic and morphological analyses. These three taxa present different geographic distribution, C. cytisoides var.
confertiformis being endemic to interior Bahia,
C. cytisoides var. decora endemic to inland
Minas Gerais, and C. cytisoides var. unijuga
endemic to coastal areas of Sergipe and Alagoas
states. Therefore, we propose they should be
treated as distinct species, which may be easily
recognized by morphological differences in their
leaves: shortly petiolate leaves with two pairs of
oblong to oblong-lanceolate leaflets in C. confertiformis, petiolate leaves with one pair of
obliquely obovate leaflets in C. decora, and
petiolate leaves with one pair of orbicular to
reniform leaflets in C. unijuga.
The populations of C. cystisoides var. blanchetii, C. cytisoides var. brachystachya, C. cystisoides var. micrantha and C. cytisoides ‘‘f.
prostrata’’ presented a reticulate pattern of grouping in both genetic and morphological analyses,
but with different topology of phenetic relationships in each analysis. Almost all populations
presented very high genetic identities, except for
three disjunct populations of C. cystisoides var.
blanchetii (CB1 and CB5) and C. cytisoides var.
brachystachya (CBR2). The differences pointed
out by Conceição (2000) in the populations
named as C. cytisoides ‘‘f. prostrata’’ are not
200
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
Fig. 3. Dendrogram showing the phenetic relationships among 33 populations of nine described and one
putative taxa of Chamaecrista sect. Absus subsect. Baseophyllum occurring in northeastern and southeastern
Brazil. Constructed using the matrix of genetic distances (Nei, 1978; unbiased estimate) based on 15 allozymic
loci with UPGMA as clustering algorithm. See Table 2 for the names of the populations
consistent with our analyses, and we suggest that
these populations should not be treated as a
distinct taxon.
Populations of C. cytisoides ‘‘f. prostrata’’ are
morphologically very similar to C. cytisoides var.
blanchetii, and the only genetic distinction was
the absence of the loci EST-1 and EST-3 in C.
cytisoides ‘‘f. prostrata’’. A similar picture was
found in Acianthera (Orchidaceae) species of
campo rupestre vegetation by Borba et al. (2001).
Populations CBR2 (from which the holotype was
collected) and CBR3 of C. cytisoides var.
brachystachya are very similar morphologically
to C. cytisoides var. micrantha, in spite of the
genetic differences of the former population.
These two populations occur in the interior of
Brazil, as also C. cytisoides var. micrantha, near
the central (Pernambuco state, CBR3) and northern (center of Minas Gerais state, CBR2)
extremes of distribution of the group (C. cytiso-
c
Fig. 4. Representation of the scores on the two first canonical axes of the CVA using 16 vegetative (A) and 38
floral (B) characters in 33 populations of nine described and one putative taxa of Chamaecrista sect. Absus
subsect. Baseophyllum occurring in northeastern and southeastern Brazil. See Table 2 for the names of the
populations
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
201
202
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
c
Fig. 5. Dendrogram showing the phenetic relationships among 33 populations of nine described and one
putative taxa of Chamaecrista sect. Absus subsect. Baseophyllum occurring in northeastern and southeastern
Brazil. Constructed using the matrix of Mahalanobis generalized distance based on 16 vegetative (A) and 38
floral (B) characters. See Table 2 for the names of the populations
ides var. micrantha occupies an intermediate
distribution, occurring in Bahia and northern
Minas Gerais states). The other three populations
of C. cytisoides var. brachystachya are very
similar morphologically to C. cytisoides var.
blanchetii, differing by their occurrence near the
coast (versus interior of Bahia in C. cytisoides
var. blanchetii) and by the presence of two pairs
of leaflets. These three populations have been
identified as C. cytisoides var. brachystachya due
to the presence of two pair of leaflets and
probably by second pair being similar in morphology to that found in individuals of the
population CBR2. However, the first pair of
leaflets is very similar to the single pair found in
C. cytisoides var. blanchetii.
Based on our present data, we suggest a broad
circumscription of two polymorphic species in
this group: (1) C. brachystachya, including the
populations CBR2 and CBR3 (C. cytisoides var.
brachystachya s.str. Crom.) and C. cystisoides
var. micrantha populations, and (2) C. blanchetii,
including the populations of C. cytisoides ‘‘f.
prostrata’’ and populations CBR1, CBR4 and
CBR5 formerly identified as C. cytisoides var.
brachystachya. This decision is based mainly on
morphology, but it is also supported by genetic
data and by the occurrence of these two taxa in
sympatry with no interbreeding (Costa 2007). In
the multivariate morphological analyses the
major characters which separated the populations
of these two species were number of leaflets,
width of leaflet at 1/3 of its length, angle of the
apex of the leaflet, least distance from the margin
to the main vein in the base of the leaflet, and
length of the petiole.
Taxonomic conclusions. The results of
multivariate morphological and allozyme data
in populations of all taxa of Chamaecrista sect.
Absus subsect. Baseophyllum showed that several
distinct species exist in the subsection. Genetics
and morphology confirmed the circumscription
and specific rank for C. coriacea and C.
depauperata, corroborating the delimitation
proposed by Bentham (1870, 1871) and
Conceição et al. (2001), respectively.
However, the present analyses did not support
the delimitation proposed by Irwin and Barneby
(1978, 1982) for the Chamaecrista cytisoides
complex. Our data showed that the type taxon of
the complex, C. cytisoides var. cytisoides, is
genetic and morphologically very distant from all
the other recognized varieties, and thus the
complex may be re-arranged in six species based
on C. cytisoides var. blanchetii, C. cytisoides var.
brachystachya, C. cytisoides var. confertiformis,
C. cytisoides var. cytisoides, C. cytisoides var.
decora and C. cytisoides var. unijuga. On the
other hand, C. cytisoides var. micrantha is
proposed as synonym of C. cytisoides var.
brachystachya.
Chamaecrista blanchetii is a variable species,
with sessile leaves or bearing a minute petiole,
with one to two (rarely three) pairs of coriaceous
leaflets, the proximal pair reniform to sub-orbicular, base reniform to obtuse, frequently embracing the stem, and the distal pair orbicular with
base obtuse. Chamaecrista brachystachya is also
a variable species, with petiolate leaves and two
pairs of coriaceous to semi-succulent leaflets, the
proximal pair orbicular to obovate with cordate to
cuneate base, the distal pair oblong to suborbicular with cuneate base. The genetic differences
presented by the population CBR2, as generally
observed in allozyme differentiation, are probably a result of genetic drift or directional
selection. The presence of two fixed exclusive
alleles in this population can be interpreted as a
result of its geographic isolation; gene flow
between CBR2 and the remaining C. brachystachya populations is possibly very low or nonexistent, and genetic drift could be responsible
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
203
204
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
for this differentiation. This process can quickly
lead to speciation in small, geographically isolated populations (Levin 2000).
Below we present a synopsis of the taxa
belonging to Chamaecrista sect. Absus subsect
Baseophyllum as recognized in this work. The
taxonomic treatment of the group, including
descriptions, illustrations and identification
keys, is being published elsewhere.
1. Chamacrista blanchetii (Benth.) Conc., L.P.
Queiroz & G.P. Lewis, comb. nov.
Basionym: Cassia blanchetii Benth., J. Bot.
(Hooker) 2: 78 (1840). Type: ‘‘Serra de
Jacobina’’, s.d., Blanchet 2649 (holotype: K!
[photo: HEFS!]; isotypes [numbered 2549]
B, BM!, C, F, G!, K!, LE, MG, MO, NY!, P,
SP!, [photo: HUEFS!]).
: Cassia cytisoides var. blanchetii (Benth.)
H.S.Irwin & Barneby, Mem. New York Bot.
Gard. 30: 13 (1978).
: Chamaecrista cytisoides var. blanchetii
(Benth.) H.S.Irwin & Barneby, Mem. New
York Bot. Gard. 35: 647 (1982).
2. Chamaecrista brachystachya (Benth.) Conc.,
L.P. Queiroz & G.P. Lewis, comb. nov.
Basionym: Cassia brachystachya Benth., J.
Bot. (Hooker) 2: 78 (1840). Type: ‘‘Tejuco,
Herb, Acad. Petrop.’’, s.d., Riedel 566 (lectotype: K-Benth.!; isolectotype: LE!).
: Cassia cytisoides var. brachystachya
(Benth.) H.S. Irwin & Barneby, Mem. New
York Bot. Gard. 30: 13 (1978).
: Chamaecrista cytisoides var. brachystachya (Benth.) H.S. Irwin & Barneby, H.S.Irwin & Barneby, Mem. New York Bot. Gard.
35: 647 (1982).
= Cassia cytisoides var. micrantha H.S.
Irwin & Barneby, Mem. New York Bot.
Gard. 30: 12 (1978). Type: Brazil, Bahia,
Andarahy, Chapada Diamantina, ChiqueChique (Xique-Xique), April 1938, Bondar
2610 (holotype: SP!). : Chamaecrista cytisoides var. micrantha (H.S.Irwin & Barneby)
H.S.Irwin & Barneby, Mem. New York Bot.
Gard. 35: 647 (1982).
3. Chamaecrista confertiformis (H.S. Irwin &
Barneby) Conc., L.P.Queiroz & G.P. Lewis,
comb. et stat. nov.
Basionym: Cassia cytisoides var. confertiformis H.S.Irwin & Barneby, Mem. New
York Bot. Gard. 30: 12 (1978). Type: Brazil,
Bahia, ± km S. of Mucugê, along rio Cumbuca, R.M. Harley 15936 (holotype: NY!;
isotypes: K!, P!).
: Chamaecrista cytisoides var. confertiformis (H.S. Irwin & Barneby) H.S. Irwin &
Barneby, Mem. New York Bot. Gard. 35:
647 (1982).
4. Chamaecrista coriacea (Benth.) H.S. Irwin &
Barneby, Mem. New York Bot. Gard. 35(2):
647 (1982). Type: ‘‘Inter Serra del frio et
Cachoeira, Herb., Acad. Petrop.’’, 01 Jan.
1825, Riedel 563 (holotype: LE [photoK!];
isotypes: A, K! [photos: HUEFS!, IPA!]).
5. Chamaecrista cytisoides (Collad.) H.S Irwin
& Barneby, Mem. New York Bot. Gard. 35:
647 (1982). Type: ‘‘Hab. in Brasilia’’ (holotype: G-DC!; probable isotype: P!)
6. Chamaecrista decora (H.S.Irwin & Barneby)
Conc., L.P. Queiroz & G.P. Lewis, comb. et
stat. nov.
Basionym: Cassia cytisoides var. decora
H.S. Irwin & Barneby, Mem. New York
Bot. Gard. 30: 14 (1978). Brazil, Minas
Gerais, Serra do Espinhaço, ±18 km E of
Diamantina, 19 March 1970, Irwin et al.
27894 (holotype: UB; isotypes: C, F, GH,
K!, NY!, RB!, UC, US).
: Chamaecrista cytisoides var. decora
(H.S.Irwin & Barneby) H.S.Irwin & Barneby, New York Bot. Gard. 35: 647 (1982).
7. Chamaecrista depauperata Conc., L.P. Queiroz & G.P. Lewis, Sitientibus Sér. Ci. Biol.
A. S. Conceição et al.: Biosystematics of Chamaecrista subsect. Baseophyllum (Leguminosae)
1(2): 114 (2001). Type: Brazil, Bahia, Abaı́ra,
distrito de Catolés, Campo do Virassaia, 30
Dec. 1993, W. Ganev 2722 (holotype: HUEFS!; isotypes: K!, SPF!).
8. Chamaecrista unijuga (Benth.) Conc.,
L.P.Queiroz & G.P.Lewis, comb. et stat. nov.
Basionym: Cassia brachystachya var. unijuga Benth., Fl. Bras. (Martius) 15(2): 139.
1870. Type: ‘‘in maritimis prope Maceio
prov. Alagoas’’, April 1848, Gardner 1282
(lectotype: K!; isolectotype: BM!).
: Cassia cytisoides var. unijuga (Benth.)
H.S. Irwin & Barneby, Mem. New York Bot.
Gard. 30: 14 (1978).
: Chamaecrista cytisoides var. unijuga
(Benth.) H.S. Irwin & Barneby, New York
Bot. Gard. 35: 647 (1982).
We thank the Fundação de Amparo à Pesquisa do
Estado da Bahia (FAPESB) for financial support
(#245/04). ASC received a fellowship from Universidade do Estado da Bahia (PAC-UNEB). ELB and
LPQ are supported by a grant (PQ2) from Conselho
Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq).
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