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A morphometric analysis of Daniellia (Fabaceae

Caesalpinioideae)
ARTICLE in BOTANICAL JOURNAL OF THE LINNEAN SOCIETY FEBRUARY 2009
Impact Factor: 2.7 DOI: 10.1111/j.1095-8339.2008.00894.x

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Natural History Museum, London

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Botanical Journal of the Linnean Society, 2009, 159, 268279. With 19 figures

A morphometric analysis of Daniellia (Fabaceae

Caesalpinioideae)
MANUEL DE LA ESTRELLA*, CARLOS AEDO and MAURICIO VELAYOS
Real Jardn Botnico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain
Received 2 January 2007; accepted for publication 5 June 2008

A multivariate morphometric study of Daniellia, an endemic genus of tropical and subtropical Africa, indicates that
nine species may be recognized: D. alsteeniana, D. klainei, D. oblonga, D. ogea, D. oliveri, D. pilosa, D. pynaertii,
D. soyauxii and D. thurifera. In our study we found that some characters, not previously studied in detail, were
significant in species delimitation: petiole indumentum, petiole width, number and position of glands on the lower
surface of the leaflets and presence or absence of glands at the insertion of each pair of leaflets. The rare and
scattered material of D. pilosa and D. soyauxii made their classification uncertain, although some qualitative
characters support their differentiation. 2009 The Linnean Society of London, Botanical Journal of the Linnean
Society, 2009, 159, 268279.

ADDITIONAL KEYWORDS: Africa Leguminosae morphometry savannah taxonomy tropical forest.

INTRODUCTION
Daniellia Benn. (Fabaceae: Caesalpinioideae) is a
genus of nine species of medium to large trees found
in tropical and subtropical areas of Africa. The different species grow from sea level to 1500 m, from
swampy areas to seasonally dry forest (Mackinder,
2005). The highest concentration of species and morphological variation is found in the Guineo-Congolian
region.
Related genera and species are easily distinguished
from Daniellia by its leaf rachis with a minute pair of
glands below the insertion of the petiolules (sometimes in each pair of leaflets) visible at low magnification, by its flowers with four imbricate sepals, five
petals, 10 stamens, all free or nine shortly connate,
and its 1-seeded fruits, with a long funicle and the
seed dispersed with one valve of the fruit (Cowan &
Polhill, 1981).
Bennett (1854) described Daniellia, named after Dr
W. F. Daniell, collector of the type specimen of Daniellia thurifera Benn., in Sierra Leone. Hutchinson &
Dalziel (1928) published the first work that included

*Corresponding author. E-mail: mestrella@rjb.csic.es

268

keys for the eight accepted species known in West

Tropical Africa. Baker (1930), in The Leguminosae of
Tropical Africa, presented a key for the then 12
accepted species and proposed the subgeneric division
currently accepted with some changes in nomenclature; subgenus Daniellia (as subgenus Eudaniellia)
includes all known species except D. oliveri
(Rolfe) Hutch. & Dalziel, which alone forms subgenus
Paradaniellia.
Lonard (1950), in his tude Botanique des
Copaliers du Congo Belge, tried to establish a classification of the different species within Daniellia, a
genus which is easily recognizable from other Caesalpinioideae, but which has a complicated species
delimitation (J. Lonard, pers. comm.). Scattered and
poor materials for some species were some of the
biggest difficulties encountered Keay (1954), prior to
the second edition of Flora of West Tropical Africa
Keay, 1958 checked the species of Daniellia, placed
five previously accepted species into synonymy and
established guidelines for the currently accepted
specific subdivision, followed in most respects by
Aubrville (1968, 1970). According to Bruneau et al.
(2001), Daniellia is a monophyletic group sister to the
remainder of Detarieae sensu stricto. During our
ongoing taxonomic revision of Daniellia, we realized

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DANIELLIA: MORPHOMETRIC ANALYSIS

that the taxon described by Lonard (1949) as a
variety of D. soyauxii (Harms) Rolfe had sufficient
consistent differences to be recognized as a different
species (Estrella, Aedo & Velayos, 2007).
The aim of the present work is to carry out a
morphometric study, based mainly on numerical
analysis, as a preliminary step in establishing a comprehensive taxonomic treatment as part of a monograph of the genus.

MATERIAL AND METHODS

This study is based on 90 herbarium specimens (see
SUPPORTING INFORMATION, Appendix), belonging to the nine recognized species of Daniellia. Some
additional sheets examined during the study were not
analysed numerically but contributed to validating
our final conclusions (Fig. 1). The following herbaria
were consulted: A, AAU, B, BM, BR, BRLU, C, COI,
E, G, H, HBG, K, L, LISC, M, MA, MO, NY, P, U,
UPS, US, WAG and Z. Digital photographs from FI

269

were also examined. The specimens were selected in

order to characterize the morphological and geographical range of each species. Some of them were
poorly represented in the herbaria and in such cases
we studied all the available samples. They were considered as operational taxonomic units (OTUs).
Seventy quantitative characters were recorded and
measured using a Mitutotyo CD-15CD digital calliper
(see Fig. 2 for some leaf and flower measurements).
When possible, at least 10 specimens were measured,
a number which has been considered by other taxonomists as representative for recording the morphological range of the species (Rico & Bachman, 2006).
Each character was analysed for its mean and median
values, range, standard deviation and significance,
using the STATISTICA (http://www.statsoft.com)
package. To represent the variability of each descriptor within species, box plots containing medians and
percentiles were prepared and most informative
width/length ratios were also plotted. Box plots
showing the variability of the 14 most discriminant

Figure 1. Generic distribution of Daniellia in Africa.

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270

M. DE LA ESTRELLA ET AL.

Figure 2. Scheme of characters meassured in Daniellia and floral diagrams. A, leaf total length. B, largest leaflet width.
C, largest leaflet petiolule length. D, apical leaflet width. E, inflorescence lateral branch length. F, flower pedicel length.
G, receptacle width. H, sepal length. I, lateral petal length. J, stamen filament union. K, anther length. L, pod length.
M, floral diagram of D. thurifera. N, floral diagram of D. oliveri.

characters are shown in Figs. 316. The descriptors

were: leaf total length (LTL); largest leaflet width
(LLW); largest leaflet petiolule length (LLPL); apical
leaflet width (ALW); inflorescence lateral branch
length (ILBL); flower pedicel length (FPL); receptacle
width (RW); sepal length (SL); sepal hairs length
(SHL); lateral petal length (LPL); stamen filament
union length (SFUL); anther length (AL); ovary stipe
length (OSL); pod length (PL).
These characters were used to perform the multivariant analyses using the STATISTICA package (see
below). The most operative ones should be used to

build the species key in our future monograph in

conjunction with some of the 74 qualitative characters
that we also studied and which could aid in species
delimitation (Table 1), but these were not used in
statistical analyses.
Daniellia oblonga Oliv. was excluded from the multivariant analysis because of the scarcity of available
samples. Daniella oliveri has numerous qualitative
characters which made its inclusion in a multivariant
analysis superfluous (see RESULTS).
Principal component analyses (PCA) was carried
out using the matrix of the standardized descriptors.

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DANIELLIA: MORPHOMETRIC ANALYSIS

271

Figures 316. Box plots representing the variability of the most discriminant quantitative characters in Daniellia.

Several combinations were analysed and those plots

which showed groups of OTUs in accordance with
qualitative data were selected. The correlation matrix
was obtained from the initial matrix, eigenvectors

were extracted and the OTUs plotted (Fig. 17).

Although these analyses are not designed for clustering, a general tendency between main groups could be
traced.

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M. DE LA ESTRELLA ET AL.

Figures 316. Continued.

The relationships between the different groups

were investigated by way of discriminant analysis
(DA; Sneath & Sokal, 1973), which requires the a
priori assignment of OTUs to groups and allows the

determination of whether the recognized groups are

statistically definable entities or whether there is too
much variation within groups to permit classification.
For DA, the raw matrix was obtained, the results

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DANIELLIA: MORPHOMETRIC ANALYSIS

273

Figures 316. Continued.

were assorted in to discrete groups and calculations

were carried out.

RESULTS
Daniellia oliveri is the most widespread species in
lowland savannah and it is easily distinguished from
other species as it has one big petal (lateral, Fig. 2N)
and the other four reduced [rarely two big petals
(lateral ones) and three reduced]; filaments glabrous
(rarely few hairs on basal extreme), free between
them; whereas the other species (subgenus Daniellia)
have two big petals, one medium sized and two
reduced (Fig. 2M); filaments pubescent to villous at
least 1/32/3 of their length, with nine united into a
tube and one free (Table 1). This combination of qualitative characters was used by Baker (1930) to establish the subgeneric division of the genus. Daniellia
oblonga is probably the most poorly understood
species, known only from a few incomplete specimens
from Cameroon, Equatorial Guinea and Gabon. The
description and species delimitation will be improved
when more samples become available. Daniellia
oblonga is similar to D. ogea (Harms) Rolfe ex
Holland, but the former has a glabrous ovary and
stipe (only two or three hairs have been found at the
insertion of the stipe with the ovary), whereas in D.
ogea the ovary is densely villous to pubescent. The
same indumentum differences are seen in the sepals:
those from D. oblonga are glabrous, but have a ciliate
margin and a tuft of hairs at the top; in contrast, D.
ogea has densely pubescent sepals (Table 1). One last
difference is the presence of a ring of hairs around the
glands at the insertion of each pairs of leaflets in
D. ogea, whereas in D. oblonga they are completely
glabrous.

All other species (subgenus Daniellia except D.

oblonga) were included in the PCA and the scatter
plot of two principal components axes is presented in
Fig. 17, where 75.73% of variance was accounted for
by the first two eigenvectors. The third eigenvector
only accounted for 5.79% of variance, so it was not
represented. The descriptors used for this analysis
were LTL, LLW, LLPL, RW, LPL, AL and OSL. The
highest loading on the first principal component corresponded to characters RW, LPL and OSL. This
component is related to the flower characteristics.
The variables with the highest loadings on the second
principal component are LTL, LLW, AL and LLPL.
These variables are more related to leaf structure.
Although we did not find any discrete group (Fig. 17),
we observed two main tendencies: first, specimens
with bigger flowers and leaves on the left side of the
scatter plot, which correspond with D. alsteeniana
Duvign. and D. klainei Pierre ex A.Chev. are bordered
by some specimens of D. pynaertii De Wild. and D.
thurifera; second, specimens with smaller flowers and
leaves, D. soyauxii (Harms) Rolfe, D. pilosa (Lonard)
Estrella and D. ogea are also associated with some
specimens of D. pynaertii and D. thurifera.
Two DAs were carried out to study the separation
between these groups using the descriptors listed
above in MATERIAL AND METHODS. In DA1 we
include the four species found in the first group in our
PCA, D. alsteeniana, D. klainei, D. pynaertii and D.
thurifera. The plot of root 1 against root 2 shows a
significant separation between the species OTUs.
Daniellia pynaertii is placed near D. thurifera and D.
klainei, whereas D. alsteeniana is placed closest to
the latter (Fig. 18). The characters contributing most
to this separation were LLPL, LL, PL, SFUL and
ALW (Table 2).

2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159, 268279

Stamen
filament
indumentum
Ovary
indumentum

Leaflet
indumentum
(midrib on
lower
surface)
Leaflet glands
(on lower
surface)
Inflorescence
indumentum
Inflorescence
lateral
branches
Pedicel
indumentum
Flower bud
width (mm)
Sepal
indumentum

Pairs of
leaflets
Number of
main lateral
veins
Petiole
indumentum
Rachis glands

Glabrous

Pubescent
glabrous
(6.7)7.4
8.7(-9.5)
Pubescent
margins
and apex
Pubescent at
least at 1/3
of its length
Glabrous to
pubescent
in margins
(6.9)7.4
8.8(-9.6)
Glabrous
except
margins
Pubescent at
least at 2/3
of its length
Glabrous

Glabrousslightly
pubescent
79

Pubescent
glabrescent
512

Glabrous

One gland in
the midrib

Glabrous

With glands

With glands
basally
Glabrous

One gland in
the midrib

With glands

Glabrous

Glabrous

Ciliate
margin and
apex
Pubescent at
least at 2/3
of its length
Glabrous

Glabrous
glabrescent
7.39.2

Slightly
pubescent
7

One gland in
the lamina

Pubescent at
least at 2/3
of its length
Densely
villous

Velvety
pubescent
(4.5)5.2
6.2(-7.5)
Pubescent

512

One or two
glands in
the lamina
Tomentose

Glabrous

816

Glabrous
pubescent
With glands
basally
Tomentose
villous

1016

1626

69

D. ogea

918

78

D. oblonga

47

D. klainei

59

D. alsteeniana

Table 1. Qualitative and quantitative characters in Daniellia

Glabrous

Pubescent at
least at 2/3
of its length
Densely
villous

Velvety
pubescent
Glabrous,
ciliate
margin
Glabrous

611

Velvety
pubescent
45.5

Very long
velvety
912

One gland in
the lamina

Without
glands
Midrib
pubescent

Pubescent

916

810

D. pilosa

Glabrous

Glabrescent
tomentose
616

Two glands in
the lamina

Pubescent

Glabrous
pubescent
With glands

917

611

D. oliveri

Pubescent
margins
and apex
Pubescent at
least at 2/3
of its length
Pubescent
glabrescent

(4)5.48(-9)

Glabrescent

Slightly
pubescent
411

One gland in
the lamina

Midrib
pubescent

Pubescent
glabrous
With glands

1018

510

D. pynaertii

Ciliate
margin and
apex
Pubescent at
least at 2/3
of its length
Few hairs
along
sutures

Velvety
pubescent
3.24.5

Very long
velvety
56

One gland in
the lamina

Without
glands
Glabrous

Pubescent

816

79

D. soyauxii

Glabrous, but
ciliate
margin
Pubescent at
least at 2/3
of its length
Glabrous

(6.2)89.4

Glabrous

610

Glabrous

Two glands in
the lamina

With glands
basally
Glabrous

Glabrous

1025

69

D. thurifera

274
M. DE LA ESTRELLA ET AL.

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DANIELLIA: MORPHOMETRIC ANALYSIS

275

Figure 17. Plot of first two axes of the principal component analyses (PCA). 1, Daniellia alsteeniana; 2, D. klainei;
3, D. ogea; 4, D. pilosa; 5, D. pynaertii; 6, D. soyauxii; 7, D. thurifera.

Figures 18, 19. Plots of the discriminant analyses (DAs) in Daniellia.

In DA2, D. ogea, D. pilosa, D. soyauxii, D. thurifera

and D. pynaertii were included (Fig. 19). The first four
appear clearly separated from each other; D. pynaertii
is placed among D. ogea, D. pilosa and D. thurifera.
The most discriminant characters were LLW, LLPL,
LL, SL and PL (Table 2).
The OTUs used in DA1 were correctly classified in
all cases, except one of D. alsteeniana that was misclassified as D. klainei and one of D. thurifera that
was misclassified as D. pynaertii. In DA2, some predicted classifications were erroneous. One OTU of

D. ogea was misclassified as D. pynaertii, one OTU

of D. pilosa as D. pynaertii, two OTUs of D. pynaertii
as D. ogea and two OTUs of D. thurifera as
D. pynaertii (Table 3).

DISCUSSION
Hutchinson & Dalziel (1928) published the first work
that included keys for the eight accepted species
found in West Tropical Africa; they used the number
of lateral nerves as a significant character in species

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276

M. DE LA ESTRELLA ET AL.

Table 2. Standardized coefficients obtained in discriminant analyses (DAs) for canonical variables
DA1

Leaf length
Largest leaflet width
Largest leaflet petiolule length
Apical leaflet width
Inflorescence lateral branch lenght
Pedicel length
Receptacle width
Sepal length
Sepal hair length
Lateral petal length
Stamen filament union length
Anther length
Ovary stipe length
Pod length
Eigen values
Total cumulative proportion

DA2

Root 1

Root 2

Root 1

Root 2

-0.64914
-0.24551
1.11395
-0.04235
0.49774
0.36537
0.54468
-0.03427
0.16673
-0.16856
-0.11682
0.37161
0.25591
0.63612
6.34754
0.76864

0.52263
-0.30511
0.00474
0.51413
-0.10209
-0.05699
0.06549
-0.49918
-0.27217
0.05097
-0.71725
-0.27287
0.46709
0.05870
1.05295
0.89615

-0.79948
1.08569
0.90353
-0.62804
-0.00955
0.19058
0.28071
0.61141
-0.00421
-0.52099
0.43291
0.15073
-0.10173
-0.34432
3.54284
0.49164

-0.44039
0.35456
-0.89617
0.56807
0.35357
-0.74487
-0.35115
0.78235
-0.17748
0.20015
0.00132
0.32697
0.32651
0.56595
2.25548
0.80463

Table 3. Correct classifications and values of P obtained in discriminant analyses (DAs) of Daniellia

D.
D.
D.
D.
D.
D.
D.
D.
D.

alsteeniana
klainei
pynaertii
thurifera
ogea
pilosa
pynaertii
soyauxii
thurifera

DA

No. of
OTUs

Correct predicted
classifications (%)

Incorrect predicted
classifications

1
1
1
1
2
2
2
2
2

12
14
11
14
15
5
11
4
14

91.7
100
100
92.9
93.3
80
81.8
100
85.7

1 OTU = D.

1 OTU = D.
1 OTU = D.
1 OTU = D.
2 OTU = D.

2 OTU = D.

0.23529
0.27451
0.21569
0.27451
0.30612
0.10204
0.22449
0.08163
0.28571

klainei

pynaertii
pynaertii
pynaertii
ogea
pynaertii

OTU, operational taxonomic unit.

delimitation, which we found in our study to be of

little value (Table 1). They also used the number of
pairs of leaflets, but as shown in Table 1 these values
are normally overlapping between species. The ovary
indumentum and shape of leaflets were also used.
We found that these characters are quite variable
within a species; for example, in D. thurifera the
leaflets are reported to have different sizes and
shapes on depending of the tree age (Lonard, 1950).
Baker (1930) used the same characters for species
delimitation.
Lonard (1950) used almost the same characters in
species delimitation as previous authors, but he also
introduced two new characters that have been found
relevant to our study: pedicel length (Fig. 8) and the

length of petiolules in some species (although he did

not state which leaflet was measured).
Keay (1958) established the guidelines for the currently accepted species limits. These were followed
largely by Aubrville (1968, 1970), who recognized
nine species and one variety. Both authors used some
characters that we consider too variable between
species, such as number of leaflets, leaflet shape,
ovary indumentum and flower colour. However, they
also used some characters that appear to be significant, e.g. pedicel length (Fig. 8), size and indumentum of sepals (Figs 10, 11) and indumentum on the
lower surface of the midrib of the leaflets (Table 1).
In our study we found that some characters, not
previously studied in detail, were significant in

2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159, 268279

DANIELLIA: MORPHOMETRIC ANALYSIS

species delimitation: petiole indumentum, petiole
width, number and position of glands on the lower
surface of the leaflet and presence or absence of
glands at the insertion of each pair of leaflets.
Daniellia alsteeniana is the most southern species
of the genus. Our analyses show that D. alsteeniana
and D. klainei overlap to some extent. The former is
easily distinguished from the majority of Daniellia
species by its comparatively long lateral branches in
the inflorescence (Fig. 7) and the presence of one
gland on the lower surface on the midrib 13 cm from
the petiolule. These two characters are shared with
D. klainei. Most specimens of D. alsteeniana have
some pubescence on the petiolules, rachis, leaflets
and inflorescence, which allows differentiation from
D. klainei, a completely glabrous species (Table 1).
Additionally, D. alsteeniana has larger sepals and
petals (Figs 10, 12); the leaflets are usually larger in
D. klainei and in this species we frequently found
petiolules thickened by glands. Daniellia alsteeniana
filaments are slightly pubescent for 1/3 of their length
in contrast with D. klainei, the filaments of which are
densely pubescent for 2/3 of their length.
In our morphometric study, we found that D.
pynaertii is the species with most overlap with the
other Daniellia species included in the analyses
(Table 3). This is probably a result of the great variability in leaflet texture and size found in D.
pynaertii, which nevertheless is usually readily distinguished from other Daniellia species by means of
its pubescent midribs (Table 1). Daniellia pynaertii
shares this feature with D. pilosa; the former is
distinguished by the presence of a pair of glands at
the insertion of each pair of leaflets and its usually
simple, paniculate inflorescence, whereas, in D.
pilosa, rachis glands have not been found and the
inflorescence is usually a compound panicle.
Material of D. soyauxii in herbaria is limited. This
species is poorly known and its classification will
remain a problem until more collections are obtained.
In our previous work (Estrella et al., 2007) several
characters allowed us to present D. pilosa as a distinct species closely related to D. soyauxii. Daniella
pilosa has a pubescent midrib on the lower surface (in
some specimens a few leaflets were glabrous) and the
sepals are velvety pubescent, whereas in D. soyauxii
they have only a ciliate margin and a tuft of hairs on
the top. Other features concern flower size or number
of lateral branches in the inflorescence (912 in D.
pilosa vs. 56 in D. soyauxii) which also distinguish
these two species (Table 1).
Daniellia thurifera is one of the species with greatest variability in leaflet shape, size and texture.
Lonard (1950: 100) said that, according to field
observations by Melville, this species had small leaflets in older trees. Daniella thurifera is the only

277

species in which the adaxial petal (medium sized,

1216 mm long) is the same size as the lateral ones
(big petals, 1216 mm long), which, united with the
characters presented in Table 1, allows the differentiation of this species.
Daniellia ogea is widely dispersed in West and West
Central Africa. This is the most problematic species
with respect to its delimitation from D. thurifera.
Daniellia ogea also shows great variability in leaflet
shape, size and texture (as a result of the difference
between young and mature leaflets). Daniellia ogea is
distinguished from the remaining Daniellia species
by its usually glabrous leaflets, a ring of hairs around
the glands at the insertion of each pair of leaflets and
flowers with densely pubescent sepals and a densely
villous ovary.
In view of these data, we conclude that nine species
should be recognized in the genus Daniellia. We also
reflect the need for future explorations to improve
the species description and delimitation within this
African genus.

ACKNOWLEDGEMENTS
The authors wish to thank the staff of the cited
herbaria for their support in our visit and/or loan of
selected material, and also J. L. Castillo and A.
Martn for their technical support. We are indebted to
J. J. Wieringa for his help and B. Mackinder for her
advice and critical review of the manuscript. This
work was financed by the Flora of Equatorial Guinea
project (CGL 2006-01223). M. de la Estrella was
funded by a Universidad Complutense de Madrid
pre-doctoral grant and visited BR under FPVI
European-funded Integrated Infrastructure Initiative
grant SYNTHESYS, BE-TAF 2142 project.

REFERENCES
Aubrville A. 1968. Flore du Gabon 15: Lgumineuses
Caesalpinoides. Paris: Musum National dHistoire
Naturelle.
Aubrville A. 1970. Flore du Cameroun 9: Lgumineuses
Caesalpinoides. Paris: Musum National dHistoire
Naturelle.
Baker EG. 1930. The Leguminosae of Tropical Africa. Ostend:
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Bennett JJ. 1854. Description of the Bungo, or Frankincense
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APPENDIX
LIST
Species

OF SPECIMENS MEASURED AND USED FOR THE MORPHOLOGICAL AND NUMERICAL STUDIES

Collections studied

Daniellia alsteeniana Duvign.

ANGOLA. Lunda Norte: Mungo (Carumbo), confluencia do Luxico com o Lule, 14.ix.1927, Carrisso & Mendoza 537
(COI); entre Maludi e Chiafua, 11.viii.1965, Mendes dos Santos 1605 (LISC). Lunda Sul: Saurimo, Gossweiler 14088
(BM). Moxico: Moxico, R. Cassai, between R. Cassai and Vila Luso, v.1937, Exell & Mendoza 1478 (COI); Luena
(Vila Luzo), Cassai, v.1937, Gossweiler 11282 (LISC). DEMOCRATIC REPUBLIC OF CONGO. Katanga: Mwene-Djungu,
Lamanga, iv.1958, Franois 48 (BR). Kasai-Oriental: Tshibombo, territoire Bakwanga, 16.xi.1956, Liben 1930 (BR),
Bakwanda (au N de Katabaie, Territoire Mwene-Ditu), 12.v.1957, Liben 2938 (BR); Mwene-Ditu, savane de
Kabwele, 26.v.1951, Simon 30B (BR). Kinshasa: Kahemba-Kwanbo, Mikondo, 30 iv 1955, Devred 1849 (BR).
Sud-Kivu: Masisi, Kahemba, ix.1949, Dubois 1491 (BR)
GABON. Doudou Mountains, ca. 35 km SW of Doussala, 27.viii.1985, Reitsma 1414 (MA).
Daniellia klainei Pierre ex A.Chev.
ANGOLA. Cabinda: Maiombe, Buco-Zau, 14.x.1916, Gossweiler 6746 (BM); Buco Zau, entre Chion e Chiaca, 22.ix.1958,
Monteiro, Santos & Murta 337 (LISC). DEMOCRATIC REPUBLIC OF CONGO. Bas-Congo: Luki, Mayumbe, 1959,
Hombert 568 (BR); Lopoldville, Boma, Luki, 23.x.1948, Madoux 91 (BR); Luki, Mayumbe, 1959, Wagemans 2440
(BR). GABON. 10.xii.1900, Klaine 1925 (BR). Estuaire, environs Libreville, x.1900, Klaine 1440 (G); Mbilagon
(Bilagone), 27.viii.1938, Thomson 12 (K). Moyen-Ogoou: environs dAdouma, sur lOrimbo, affluent de lOgoou,
29.vii.1912, Fleury & Chevalier 26540 (P). Ngouni: Mission de St. Martin, 7.ix.1938, Walker s.n. (P). Ogoou-Lolo:
Makande surroundings, about 65 km SSW of Boou. Makande, 26.ii.1999, Breteler, Caball, Issembe, Moussavou &
15223 (WAG). Ogoou-Maritime: Nguessi, 24.viii.1918, Le Testu 2283 (BM). Tchibanga, 14.ix.1914, Le Testu 1784
(BM).
Daniellia oblonga Oliv.
CAMEROON. Yaound (Jande), i.1914, Mildbraed 8031 (K); Likomba pflanzung, 1535 km NE von Victoria, xii.1928,
Mildbraed 10759 (A). GABON. 530 km NNW of Ndjol, 23.iv.1992, Breteler, Jongkind & Wieringa 11046 (WAG).
EQUATORIAL GUINEA: Bioko (Fernando Po), Barter 2074 (K).
Daniellia ogea (Harms) Rolfe ex Holland
CAMEROON. Ndiki, xi.1938, Jacques-Flix 2536 (WAG). GHANA. Gold Coast, 27.iii.1909, Imperial Institute s.n. (K).
IVORY COAST. Agnby, Agboville, Abreville 2775 (K). LIBERIA. Nimba, New Bapa, 19.ix.1964, Adames 561 (UPS);
Gbanka (Banga), 25.x.1926, Linder 1233 (K); Zle (Tiatown), 10 miles E of Tapeta, 8.x.1961, Voorhoeve 517 (WAG),
Bong, Loma National Forest near Basiweng, 15.xii.1961, Voorhoeve 721, 735 (WAG). NIGERIA. Without locality,
Kenedy 328 (BM). Central Province: Agogidigbo, 11.xii.1907, Unwin 179 (K). Lagos: Ibadan Forest Reserve,
25.xi.1900, Punch 115 (K). Mamu Reserve, 19.ii.1906, Foster 156 (K); Ijebu Lagos, 1895, Millen 191 (K). Benin,
Okomu Forest Reserve, 15.i.1948, Brenan, Jones & Richards 8817 (K). SIERRA LEONE. Gola forest, 13.iii.1909,
Unwin & Smythe 45 (K).
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279

APPENDIX Continued
Species

Collections studied

Daniellia oliveri (Rolfe) Hutch. & Dalziel

BURKINA FASO. Between Ramba & Laye on Ouaga-Ouhigouya road, 18.iii.1999, van Slageren & Lessina MSSL851
(K). CAMEROON. about 5 km S of Ngaoundr, 5.xii.1964, Wilde & Wilde-Duyfjes 4618 (WAG). CENTRAL AFRICAN
REPUBLIC. Haute-Kotto, Yalinga, 26.i.1923, Le Testu 4510 (BM); Bamingui-Bangoran, 29.i.1980, Spinage 310 (K).
GHANA. Mole National Park, 30.xi.1995, Schmidt, Amponsah & Welsing 1870 (UPS). GUINEA BISSAU. Bissau, Brene,
24.i.1945, Espirito Santo 1675 (LISC). NIGERIA. Jamtari, on Jamtari-Kamari old motor road, 22.xii.1954, Latilo &
Daramola 28931 (K). SENEGAL. Tambacounda, Parc National du Niokolo Koba, Gu de Damantan-Dabala, km 13,
22.i.1993, B, Madsen, Sambou, Goudiaby, Traor, Sa, 1246 (AAU); Fatick, Sine Saloum National Park, fort
Fathala, 26.i.1994, Madsen, Goudiaby & Traor 3052 (AAU). SENEGAMBIA. Without locality, Heudelot 364 (K).
WITHOUT LOCALITY. Mann 978 (K).
Daniellia pilosa (J.Lonard) Estrella
GABON. Nyanga: Mayumba (Mayoumba), 24.v.1915, Le Testu 2062 (LISC). Ogoou-Lolo: rgion de la fort des
Abeilles, campement rivire Makand (2 km en amont de son embouchure dans lOffou), 8.iii.1999, Hall 4610
(WAG); regin de Lastoursville, Nzambi, 29.viii.1930, Le Testu 8292 (BR); 28 km NE of Lastoursville, CEB
explotation, 23.viii.1992, Wieringa & van de Poll 1462 (WAG); fort des Abeilles, 13 km SE of the confluence of
Gongu-Offou river, 2.viii.1993, Wilks & Dibata 2703 (MA).
Daniellia pynaertii De Wild.
DEMOCRATIC REPUBLIC OF CONGO. quator: quateur, Forestier Central, Eala, 13.iv.1934, Corbisier Baland 1032 (BR);
Eala, 1931, Corbisier Baland 1176 (A); Eala, 1921, Goossens 1639 (K); Eala, 1936, Lemans 220 (BR); Eala, route de
Coq, 3.xii.1946, Lonard 1090 (BR); Forestier Central, Dundusana, xii 1913, Mortehan 931 (BR); Eala, route de
Coq, 1947, Poucet 1090b (BR); Eala, 20.xi.1906, Pynaert 679 (BR). Kakenge, 4.ix.1958, Dechamps 56 (BR),
Bena-Lungu, 20.xi.1958, Dechamps 94 (BR). NIGERIA. Southern Province: Lagos Colony, v.1883, Moloney s.n. (K).
Daniellia soyauxii (Harms) Rolfe
GABON. Estuaire: Munda, Sibange Farm, 2.vi.1880, Soyaux 90 (K). Moyen-Ogoou: 26 km ENE of Lambarn, 6 km
ENE of Bellevue, 2.iv.1994, Wieringa & Haegens 2614 (WAG). Ogoou-Maritime: Rabi-Kounga, road to Divangui,
23.ix.1992, Wieringa & Epoma 1627 (WAG); Rabi, 11 km on road to Divangui, 29.ix.1994, Wieringa & Nzabi 2800
(WAG).
Daniellia thurifera Benn.
GUINEA. route Longuery, 1.xii.1905, Caille 14827 (P). GUINEA BISSAU. Tombali, Cati, 24.vi.1945, Espirito Santo 2099
(LISC). IVORY COAST. Lagunes, Fort dI.D.E.R.T. Along the Lagune Ebri, near O.R.S.T.O.M., W of Abidjan,
3.viii.1963, Wilde 625 (WAG). LIBERIA. from vicinity of Firestone Plantation, along Dukwai River, Monrovia,
27.x.1928, Cooper 95 (K); Dukwai river, Monrovia, 5.iv.1929, Cooper 350 (A); Bong, Bong Range, 32 km N of
Kakata, 7.i.1962, Voorhoeve 753 (WAG). SIERRA LEONE. Without locality, 12.x.1795, Afzelius s.n. (UPS); 1854,
Daniell s.n. (BM); the evergreen rain forest of Sierra Leone, Lane Poole s.n. (K). Eastern: Kenema, Nongowa
chiefdom, 20.xii.1965, Samai 261 (K). Western Area: road to Kent from main road aroun the Peninsula, 6.xii.1963,
Morton 197 (K).

2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159, 268279