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AmorphometricanalysisofDaniellia(FabaceaeCaesalpinioideae)ARTICLEinBOTANICALJOURNALOFTHELINNEANSOCIETYFEBRUARY2009ImpactFactor:2.7DOI:10.1111/j.1095-8339.2008.00894.x

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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

Amultivariate morphometric study of Daniellia, an endemic genus of tropical and subtropical Africa, indicates thatnine 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, weresignificant in species delimitation: petiole indumentum, petiole width, number and position of glands on the lowersurface of the leaflets and presence or absence of glands at the insertion of each pair of leaflets. The rare andscattered material of D. pilosa and D. soyauxii made their classification uncertain, although some qualitativecharacters support their differentiation. 2009 The Linnean Society of London, Botanical Journal of the LinneanSociety, 2009, 159, 268279.

ADDITIONAL KEYWORDS: Africa Leguminosae morphometry savannah taxonomy tropical forest.

INTRODUCTION

Daniellia Benn. (Fabaceae: Caesalpinioideae) is agenus of nine species of medium to large trees foundin tropical and subtropical areas of Africa. The differ-ent species grow from sea level to 1500 m, fromswampy areas to seasonally dry forest (Mackinder,2005). The highest concentration of species and mor-phological variation is found in the Guineo-Congolianregion.Related genera and species are easily distinguished

from Daniellia by its leaf rachis with a minute pair ofglands below the insertion of the petiolules (some-times in each pair of leaflets) visible at low magnifi-cation, by its flowers with four imbricate sepals, fivepetals, 10 stamens, all free or nine shortly connate,and its 1-seeded fruits, with a long funicle and theseed 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 Daniel-lia thurifera Benn., in Sierra Leone. Hutchinson &Dalziel (1928) published the first work that included

keys for the eight accepted species known in WestTropical Africa. Baker (1930), in The Leguminosae ofTropical Africa, presented a key for the then 12accepted species and proposed the subgeneric divisioncurrently accepted with some changes in nomencla-ture; subgenus Daniellia (as subgenus Eudaniellia)includes all known species except D. oliveri(Rolfe) Hutch. & Dalziel, which alone forms subgenusParadaniellia.Lonard (1950), in his tude Botanique des

Copaliers du Congo Belge, tried to establish a classi-fication of the different species within Daniellia, agenus which is easily recognizable from other Caesal-pinioideae, but which has a complicated speciesdelimitation (J. Lonard, pers. comm.). Scattered andpoor materials for some species were some of thebiggest difficulties encountered Keay (1954), prior tothe second edition of Flora of West Tropical AfricaKeay, 1958 checked the species of Daniellia, placedfive previously accepted species into synonymy andestablished guidelines for the currently acceptedspecific subdivision, followed in most respects byAubrville (1968, 1970). According to Bruneau et al.(2001), Daniellia is a monophyletic group sister to theremainder of Detarieae sensu stricto. During ourongoing taxonomic revision of Daniellia, we realized*Corresponding author. E-mail: [email protected]

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

2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159, 268279268

that the taxon described by Lonard (1949) as avariety of D. soyauxii (Harms) Rolfe had sufficientconsistent differences to be recognized as a differentspecies (Estrella, Aedo & Velayos, 2007).The aim of the present work is to carry out a

morphometric study, based mainly on numericalanalysis, as a preliminary step in establishing a com-prehensive taxonomic treatment as part of a mono-graph of the genus.

MATERIAL AND METHODS

This study is based on 90 herbarium specimens (seeSUPPORTING INFORMATION, Appendix), belong-ing to the nine recognized species of Daniellia. Someadditional sheets examined during the study were notanalysed numerically but contributed to validatingour final conclusions (Fig. 1). The following herbariawere 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

were also examined. The specimens were selected inorder to characterize the morphological and geo-graphical range of each species. Some of them werepoorly represented in the herbaria and in such caseswe studied all the available samples. They were con-sidered 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 tax-onomists as representative for recording the morpho-logical range of the species (Rico & Bachman, 2006).Each character was analysed for its mean and medianvalues, range, standard deviation and significance,using the STATISTICA (http://www.statsoft.com)package. To represent the variability of each descrip-tor within species, box plots containing medians andpercentiles were prepared and most informativewidth/length ratios were also plotted. Box plotsshowing the variability of the 14 most discriminant

Figure 1. Generic distribution of Daniellia in Africa.

DANIELLIA: MORPHOMETRIC ANALYSIS 269


characters are shown in Figs. 316. The descriptorswere: leaf total length (LTL); largest leaflet width(LLW); largest leaflet petiolule length (LLPL); apicalleaflet width (ALW); inflorescence lateral branchlength (ILBL); flower pedicel length (FPL); receptaclewidth (RW); sepal length (SL); sepal hairs length(SHL); lateral petal length (LPL); stamen filamentunion length (SFUL); anther length (AL); ovary stipelength (OSL); pod length (PL).These characters were used to perform the multi-

variant analyses using the STATISTICA package (seebelow). The most operative ones should be used to

build the species key in our future monograph inconjunction with some of the 74 qualitative charactersthat we also studied and which could aid in speciesdelimitation (Table 1), but these were not used instatistical analyses.

Daniellia oblonga Oliv. was excluded from the mul-tivariant analysis because of the scarcity of availablesamples. Daniella oliveri has numerous qualitativecharacters which made its inclusion in a multivariantanalysis superfluous (see RESULTS).Principal component analyses (PCA) was carried

out using the matrix of the standardized descriptors.

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.

270 M. DE LA ESTRELLA ET AL.


Several combinations were analysed and those plotswhich showed groups of OTUs in accordance withqualitative data were selected. The correlation matrixwas obtained from the initial matrix, eigenvectors

were extracted and the OTUs plotted (Fig. 17).Although these analyses are not designed for cluster-ing, a general tendency between main groups could betraced.

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



The relationships between the different groupswere investigated by way of discriminant analysis(DA; Sneath & Sokal, 1973), which requires the apriori assignment of OTUs to groups and allows the

determination of whether the recognized groups arestatistically definable entities or whether there is toomuch variation within groups to permit classification.For DA, the raw matrix was obtained, the results

Figures 316. Continued.



were assorted in to discrete groups and calculationswere carried out.

RESULTS

Daniellia oliveri is the most widespread species inlowland savannah and it is easily distinguished fromother 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 betweenthem; whereas the other species (subgenus Daniellia)have two big petals, one medium sized and tworeduced (Fig. 2M); filaments pubescent to villous atleast 1/32/3 of their length, with nine united into atube and one free (Table 1). This combination of quali-tative characters was used by Baker (1930) to estab-lish the subgeneric division of the genus. Danielliaoblonga is probably the most poorly understoodspecies, known only from a few incomplete specimensfrom Cameroon, Equatorial Guinea and Gabon. Thedescription and species delimitation will be improvedwhen more samples become available. Danielliaoblonga is similar to D. ogea (Harms) Rolfe exHolland, but the former has a glabrous ovary andstipe (only two or three hairs have been found at theinsertion of the stipe with the ovary), whereas in D.ogea the ovary is densely villous to pubescent. Thesame indumentum differences are seen in the sepals:those from D. oblonga are glabrous, but have a ciliatemargin and a tuft of hairs at the top; in contrast, D.ogea has densely pubescent sepals (Table 1). One lastdifference is the presence of a ring of hairs around theglands at the insertion of each pairs of leaflets inD. ogea, whereas in D. oblonga they are completelyglabrous.

All other species (subgenus Daniellia except D.oblonga) were included in the PCA and the scatterplot of two principal components axes is presented inFig. 17, where 75.73% of variance was accounted forby the first two eigenvectors. The third eigenvectoronly accounted for 5.79% of variance, so it was notrepresented. The descriptors used for this analysiswere LTL, LLW, LLPL, RW, LPL, AL and OSL. Thehighest loading on the first principal component cor-responded to characters RW, LPL and OSL. Thiscomponent is related to the flower characteristics.The variables with the highest loadings on the secondprincipal 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, specimenswith bigger flowers and leaves on the left side of thescatter plot, which correspond with D. alsteenianaDuvign. and D. klainei Pierre ex A.Chev. are borderedby some specimens of D. pynaertii De Wild. and D.thurifera; second, specimens with smaller flowers andleaves, D. soyauxii (Harms) Rolfe, D. pilosa (Lonard)Estrella and D. ogea are also associated with somespecimens of D. pynaertii and D. thurifera.Two DAs were carried out to study the separation

between these groups using the descriptors listedabove in MATERIAL AND METHODS. In DA1 weinclude the four species found in the first group in ourPCA, D. alsteeniana, D. klainei, D. pynaertii and D.thurifera. The plot of root 1 against root 2 shows asignificant separation between the species OTUs.Daniellia pynaertii is placed near D. thurifera and D.klainei, whereas D. alsteeniana is placed closest tothe latter (Fig. 18). The characters contributing mostto this separation were LLPL, LL, PL, SFUL andALW (Table 2).

Figures 316. Continued.



Tab

le1.

Qualitativean

dqu

antitative

characters

inD

anie

llia

D.

alst

een

ian

aD

.kl

ain

eiD

.ob

lon

gaD

.og

eaD

.ol

iver

iD

.pi

losa

D.

pyn

aert

iiD

.so

yau

xii

D.

thu

rife

ra

Pairs

ofleafl

ets

59

47

78

69

611

810

510

79

69

Numbe

rof

mainlateral

veins

918

162

610

16

816

917

916

101

88

1610

25

Petiole

indu

men

tum

Glabrou

spu

bescen

tGlabrou

sGlabrou

sGlabrou

sGlabrou

spu

bescen

tPube

scen

tPube

scen

tglab

rous

Pube

scen

tGlabrou

s

Rachis

glan

dsWithglan

dsba

sally

Withglan

dsba

sally

Withglan

dsWithglan

dsWithglan

dsWithou

tglan

dsWithglan

dsWithou

tglan

dsWithglan

dsba

sally

Lea

flet

indu

men

tum

(midribon

lower

surface)

Tomen

tose

villou

sGlabrou

sGlabrou

sGlabrou

sPube

scen

tMidrib

pube

scen

tMidrib

pube

scen

tGlabrou

sGlabrou

s

Lea

flet

glan

ds(onlower

surface)

Oneglan

din

themidrib

Oneglan

din

themidrib

Oneglan

din

thelamina

Oneor

two

glan

dsin

thelamina

Twoglan

dsin

thelamina

Oneglan

din

thelamina

Oneglan

din

thelamina

Oneglan

din

thelamina

Twoglan

dsin

thelamina

Inflorescence

indu

men

tum

Pube

scen

tglab

rescen

tGlabrou

sslightly

pube

scen

tSligh

tly

pube

scen

tTo

men

tose

Glabrescent

tomen

tose

Verylong

velvety

Sligh

tly

pube

scen

tVerylong

velvety

Glabrou

s

Inflorescence

lateral

bran

ches

512

79

75

126

169

124

115

66

10

Ped

icel

indu

men

tum

Pube

scen

tglab

rous

Glabrou

sGlabrou

sglab

rescen

tVelvety

pube

scen

tGlabrou

sVelvety

pube

scen

tGlabrescent

Velvety

pube

scen

tGlabrou

s

Flower

bud

width

(mm)

(6.7)7.4

8.7(-9

.5)

(6.9)7.4

8.8(-9

.6)

7.3

9.2

(4.5)5.2

6.2(- 7

.5)

611

45.5

(4)5.4

8(-9

)3.2

4.5

(6.2)8

9.4

Sep

alindu

men

tum

Pube

scen

tmargins

andap

ex

Glabrou

sexcept

margins

Ciliate

marginan

dap

ex

Pube

scen

tGlabrou

s,ciliate

margin

Velvety

pube

scen

tPube

scen

tmargins

andap

ex

Ciliate

marginan

dap

ex

Glabrou

s,bu

tciliate

margin

Stamen

filamen

tindu

men

tum

Pube

scen

tat

leastat

1/3

ofitslength

Pube

scen

tat

leastat

2/3

ofitslength

Pube

scen

tat

leastat

2/3

ofitslength

Pube

scen

tat

leastat

2/3

ofitslength

Glabrou

sPube

scen

tat

leastat

2/3

ofitslength

Pube

scen

tat

leastat

2/3

ofitslength

Pube

scen

tat

leastat

2/3

ofitslength

Pube

scen

tat

leastat

2/3

ofitslength

Ova

ryindu

men

tum

Glabrou

sto

pube

scen

tin

margins

Glabrou

sGlabrou

sDen

sely

villou

sGlabrou

sDen

sely

villou

sPube

scen

tglab

rescen

tFew

hairs

alon

gsu

tures

Glabrou

s



In DA2, D. ogea, D. pilosa, D. soyauxii, D. thuriferaand D. pynaertii were included (Fig. 19). The first fourappear clearly separated from each other; D. pynaertiiis 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 mis-classified as D. klainei and one of D. thurifera thatwas misclassified as D. pynaertii. In DA2, some pre-dicted classifications were erroneous. One OTU of

D. ogea was misclassified as D. pynaertii, one OTUof D. pilosa as D. pynaertii, two OTUs of D. pynaertiias D. ogea and two OTUs of D. thurifera asD. pynaertii (Table 3).

DISCUSSION

Hutchinson & Dalziel (1928) published the first workthat included keys for the eight accepted speciesfound in West Tropical Africa; they used the numberof lateral nerves as a significant character in species

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.



delimitation, which we found in our study to be oflittle value (Table 1). They also used the number ofpairs of leaflets, but as shown in Table 1 these valuesare normally overlapping between species. The ovaryindumentum and shape of leaflets were also used.We found that these characters are quite variablewithin a species; for example, in D. thurifera theleaflets are reported to have different sizes andshapes on depending of the tree age (Lonard, 1950).Baker (1930) used the same characters for speciesdelimitation.Lonard (1950) used almost the same characters in

species delimitation as previous authors, but he alsointroduced two new characters that have been foundrelevant to our study: pedicel length (Fig. 8) and the

length of petiolules in some species (although he didnot state which leaflet was measured).Keay (1958) established the guidelines for the cur-

rently accepted species limits. These were followedlargely by Aubrville (1968, 1970), who recognizednine species and one variety. Both authors used somecharacters that we consider too variable betweenspecies, such as number of leaflets, leaflet shape,ovary indumentum and flower colour. However, theyalso used some characters that appear to be signifi-cant, e.g. pedicel length (Fig. 8), size and indumen-tum of sepals (Figs 10, 11) and indumentum on thelower 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

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

DA1 DA2

Root 1 Root 2 Root 1 Root 2

Leaf length -0.64914 0.52263 -0.79948 -0.44039Largest leaflet width -0.24551 -0.30511 1.08569 0.35456Largest leaflet petiolule length 1.11395 0.00474 0.90353 -0.89617Apical leaflet width -0.04235 0.51413 -0.62804 0.56807Inflorescence lateral branch lenght 0.49774 -0.10209 -0.00955 0.35357Pedicel length 0.36537 -0.05699 0.19058 -0.74487Receptacle width 0.54468 0.06549 0.28071 -0.35115Sepal length -0.03427 -0.49918 0.61141 0.78235Sepal hair length 0.16673 -0.27217 -0.00421 -0.17748Lateral petal length -0.16856 0.05097 -0.52099 0.20015Stamen filament union length -0.11682 -0.71725 0.43291 0.00132Anther length 0.37161 -0.27287 0.15073 0.32697Ovary stipe length 0.25591 0.46709 -0.10173 0.32651Pod length 0.63612 0.05870 -0.34432 0.56595Eigen values 6.34754 1.05295 3.54284 2.25548Total cumulative proportion 0.76864 0.89615 0.49164 0.80463

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

DANo. ofOTUs

Correct predictedclassifications (%)

Incorrect predictedclassifications P

D. alsteeniana 1 12 91.7 1 OTU = D. klainei 0.23529D. klainei 1 14 100 0.27451D. pynaertii 1 11 100 0.21569D. thurifera 1 14 92.9 1 OTU = D. pynaertii 0.27451D. ogea 2 15 93.3 1 OTU = D. pynaertii 0.30612D. pilosa 2 5 80 1 OTU = D. pynaertii 0.10204D. pynaertii 2 11 81.8 2 OTU = D. ogea 0.22449D. soyauxii 2 4 100 0.08163D. thurifera 2 14 85.7 2 OTU = D. pynaertii 0.28571

OTU, operational taxonomic unit.



species delimitation: petiole indumentum, petiolewidth, number and position of glands on the lowersurface of the leaflet and presence or absence ofglands at the insertion of each pair of leaflets.

Daniellia alsteeniana is the most southern speciesof the genus. Our analyses show that D. alsteenianaand D. klainei overlap to some extent. The former iseasily distinguished from the majority of Danielliaspecies by its comparatively long lateral branches inthe inflorescence (Fig. 7) and the presence of onegland on the lower surface on the midrib 13 cm fromthe petiolule. These two characters are shared withD. klainei. Most specimens of D. alsteeniana havesome pubescence on the petiolules, rachis, leafletsand inflorescence, which allows differentiation fromD. klainei, a completely glabrous species (Table 1).Additionally, D. alsteeniana has larger sepals andpetals (Figs 10, 12); the leaflets are usually larger inD. klainei and in this species we frequently foundpetiolules thickened by glands. Daniellia alsteenianafilaments are slightly pubescent for 1/3 of their lengthin contrast with D. klainei, the filaments of which aredensely pubescent for 2/3 of their length.In our morphometric study, we found that D.

pynaertii is the species with most overlap with theother Daniellia species included in the analyses(Table 3). This is probably a result of the great vari-ability in leaflet texture and size found in D.pynaertii, which nevertheless is usually readily dis-tinguished from other Daniellia species by means ofits pubescent midribs (Table 1). Daniellia pynaertiishares this feature with D. pilosa; the former isdistinguished by the presence of a pair of glands atthe insertion of each pair of leaflets and its usuallysimple, paniculate inflorescence, whereas, in D.pilosa, rachis glands have not been found and theinflorescence is usually a compound panicle.Material of D. soyauxii in herbaria is limited. This

species is poorly known and its classification willremain a problem until more collections are obtained.In our previous work (Estrella et al., 2007) severalcharacters allowed us to present D. pilosa as a dis-tinct species closely related to D. soyauxii. Daniellapilosa has a pubescent midrib on the lower surface (insome specimens a few leaflets were glabrous) and thesepals are velvety pubescent, whereas in D. soyauxiithey have only a ciliate margin and a tuft of hairs onthe top. Other features concern flower size or numberof lateral branches in the inflorescence (912 in D.pilosa vs. 56 in D. soyauxii) which also distinguishthese two species (Table 1).

Daniellia thurifera is one of the species with great-est variability in leaflet shape, size and texture.Lonard (1950: 100) said that, according to fieldobservations by Melville, this species had small leaf-lets in older trees. Daniella thurifera is the only

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 thecharacters presented in Table 1, allows the differen-tiation of this species.

Daniellia ogea is widely dispersed in West and WestCentral Africa. This is the most problematic specieswith respect to its delimitation from D. thurifera.Daniellia ogea also shows great variability in leafletshape, size and texture (as a result of the differencebetween young and mature leaflets). Daniellia ogea isdistinguished from the remaining Daniellia speciesby its usually glabrous leaflets, a ring of hairs aroundthe glands at the insertion of each pair of leaflets andflowers with densely pubescent sepals and a denselyvillous ovary.In view of these data, we conclude that nine species

should be recognized in the genus Daniellia. We alsoreflect the need for future explorations to improvethe species description and delimitation within thisAfrican genus.

ACKNOWLEDGEMENTS

The authors wish to thank the staff of the citedherbaria for their support in our visit and/or loan ofselected material, and also J. L. Castillo and A.Martn for their technical support. We are indebted toJ. J. Wieringa for his help and B. Mackinder for heradvice and critical review of the manuscript. Thiswork was financed by the Flora of Equatorial Guineaproject (CGL 2006-01223). M. de la Estrella wasfunded by a Universidad Complutense de Madridpre-doctoral grant and visited BR under FPVIEuropean-funded Integrated Infrastructure Initiativegrant SYNTHESYS, BE-TAF 2142 project.

REFERENCES

Aubrville A. 1968. Flore du Gabon 15: Lgumineuses Caesalpinoides. Paris: Musum National dHistoireNaturelle.

Aubrville A. 1970. Flore du Cameroun 9: Lgumineuses Caesalpinoides. Paris: Musum National dHistoireNaturelle.

Baker EG. 1930. The Leguminosae of Tropical Africa. Ostend:Unitas Press.

Bennett JJ. 1854. Description of the Bungo, or FrankincenseTree of Sierra Leone. Pharmaceutical Journal and Trans-actions 14: 252253.

Bruneau A, Forest F, Herendeen PS, Klitgaard BB,Lewis GP. 2001. Phylogenetic relationships in the Caesal-pinioideae (Leguminosae) as inferred from chloroplast trnLintron sequences. Systematic Botany 26: 487514.

Cowan RS, Polhill RM. 1981. Detarieae. In: Polhill RM,Raven PH, eds. Advances in Legume systematics. Part 1.Kew: Royal Botanic Gardens, 117134.



Estrella M, Aedo C, Velayos M. 2007. Daniellia pilosa (J.Lonard) Estrella, comb. & stat. nov. (Leguminosae).Annales Botanici Fennici 44: 149150.

Hutchinson J, Dalziel JM. 1928. Flora of West TropicalAfrica 1(2). London: The Crown Agents for the Colonies.

Keay RWJ. 1954. Revision of the Flora of West TropicalAfrica V. Kew Bulletin 4: 487492.

Keay RWJ. 1958. Flora of West Tropical Africa 1(2), 2nd ed.London: The Crown agents for Oversea Governments andAdministrations.

Lonard J. 1949. Notulae Systematicae IV (Caesalpinaceae-Amherstieae africanae americanaeque). Bulletin du JardinBotanique de ltat 19: 383408.

Lonard J, 1950. tude botanique des copaliers du CongoBelge. Publications de l Institut National pour ltudeAgronnomique du Congo Belge, Srie Scientifique 45: 1158.

Mackinder B. 2005. Detarieae. In: Lewis G, Schrire B, Mac-kinder B & Lock M, eds. Legumes of the World. Kew: RoyalBotanic Gardens, 69109.

Rico ML, Bachman S. 2006. A taxonomic revision ofAcaciella (Leguminosae, Mimosoideae). Anales del JardnBotnico de Madrid 63: 189244.

Sneath PH, Sokal RR. 1973. Numerical taxonomy. SanFrancisco: W.H. Freeman.

APPENDIXLIST OF SPECIMENS MEASURED AND USED FOR THE MORPHOLOGICAL AND NUMERICAL STUDIES

Species 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 deKabwele, 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 HollandCAMEROON. 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).



APPENDIX Continued

Species Collections studied

Daniellia oliveri (Rolfe) Hutch. & DalzielBURKINA 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 AFRICANREPUBLIC. 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, fortFathala, 26.i.1994, Madsen, Goudiaby & Traor 3052 (AAU). SENEGAMBIA. Without locality, Heudelot 364 (K).WITHOUT LOCALITY. Mann 978 (K).

Daniellia pilosa (J.Lonard) EstrellaGABON. Nyanga: Mayumba (Mayoumba), 24.v.1915, Le Testu 2062 (LISC). Ogoou-Lolo: rgion de la fort desAbeilles, 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, CEBexplotation, 23.viii.1992, Wieringa & van de Poll 1462 (WAG); fort des Abeilles, 13 km SE of the confluence ofGongu-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 deCoq, 3.xii.1946, Lonard 1090 (BR); Forestier Central, Dundusana, xii 1913, Mortehan 931 (BR); Eala, route deCoq, 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) RolfeGABON. Estuaire: Munda, Sibange Farm, 2.vi.1880, Soyaux 90 (K). Moyen-Ogoou: 26 km ENE of Lambarn, 6 kmENE 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 ofKakata, 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, Nongowachiefdom, 20.xii.1965, Samai 261 (K). Western Area: road to Kent from main road aroun the Peninsula, 6.xii.1963,Morton 197 (K).



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