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African Journal of Biotechnology Vol. 6 (7), pp. 853-860, 2 April 2007
Available online at http://www.academicjournals.org/AJB
ISSN 1684–5315 © 2007 Academic Journals
Full Length Research Paper
Genetic diversity of Tamarindus indica populations:
Any clues on the origin from its current distribution?
Boukary Ousmane DIALLO1*, Hélène, I. JOLY2, Doyle McKEY3, Martine HOSSAERT-
McKEY3 and Marie Hélène CHEVALLIER2
1 DPF/INERA/CNRST 03 BP 7047 Ouagadougou, Burkina Faso.
2CIRAD-Forêts TA 10 / C Campus International de Baillarguet 34398 Montpellier Cedex 5 France.
3CEFE/CNRS 1919 Route de Mende, 34293 Montpellier Cedex 5 France.
Accepted 6 October, 2006
Tamarindus indica is a domesticated species of high economic value for the Sahel region. Despite this
importance, very few data is available on its diversity as well as its structure leading to controversial
discussions on its origin. Thus it is questionable whether the knowledge of its genetic diversity and
organisation may help in identifying the area of its origin. We have studied 10 populations using markers
RAPDs with the seeds collected from Asia (India and Thailand), Africa (Burkina Faso, Senegal, Kenya and
Tanzania), from three islands (Madagascar, Réunion and Guadeloupe). The results showed that T. indica
has a high intra population genetic variability with a higher value obtained in the population from
Cameroon. This high intra-population variability did not allow us to determinate the origin of the species.
However, if we take into account the paleontological and anthropological results, we can assume that T.
indica has an African origin.
Key words: Tamarindus indica, RAPDs markers, genetic diversity origin.
INTRODUCTION
Human migrations and/or exchanges have promoted the
worldwide distribution of numerous plant species, such as
wheats, rice, maize, potato, and citrus (Brush et al.,
1995; Hamon et al., 1999; Nicolisi et al., 2000). Selection
has produced specialized phenotypes with genetic differ-
entiation between cultivated forms and their wild rela-
tives. The organization of the genetic diversity of these
widespread domesticated plants has been studied and
their centres of origin identified in many cases (FAO,
1996). For tree species of lesser current economic impor-
tance little information is available and doubts remain
regarding the area of origin of some widespread species.
This knowledge is of importance for developing efficient
in situ conservation strategies (Chevallier 1999).
*Corresponding authors E-mail: ousboukdiallo@yahoo.fr.
Tamarindus indica is widely distributed in dry parts of
Africa and tropical Asia and has been recently introduced
into South America, as well as into the Antilles
(Guadeloupe) and the Indian Ocean (La Réunion). The
periods of these introductions are still unknown and the
precise origin of this species is still a subject of
controversy (Lefévre, 1971; El-Siddig et al., 1999; Grollier
et al., 1998) claimed that it originated in Africa and was
introduced into India at an early date, whereas Wunderlin
(1998) and Poupon and Chauvin (1983) assumed that its
origin is in Asia particularly in India, because of its
appelation "Tamar hindi” which when translated means
“Indian date" and owing to the fact that Marco Polo in his
writings quotes its presence into 1298 and the boudhists
sources make noted it 650 years AV J.C.
The spatial distribution of genetic diversity could yield
clues to resolve these uncertainties. Introduction events
are often associated with a population bottleneck, which
should reduce genetic diversity (Citation). The short time
elapsed since introductions should not have been suffici-
854 Afr. J. Biotechnol.
ent for mutation to counter such reduction in diversity
(Citation). The diversity in an area of introduction should
thus be a subset of initial diversity (Citation).
The main objectives of this study were i) to estimate the
genetic diversity of T. indica and ii) to evaluate whether
patterns in the organization of this diversity can provide
clues as to the area of origin of the species. In order to
address these questions we assessed the genetic
diversity of 10 populations of T. indica, distributed over its
range with Random Amplified Polymorphic DNA (RAPD)
markers, which have proved useful for estimating genetic
diversity especially for previously unstudied taxa when
DNA sequence information are not available (Williams et
al., 1990). RAPD analysis has been widely used in asse-
ssing the genetic diversity of populations of a large
number of trees, including tropical species (Bekessy et
al., 2002; Cardoso et al., 1998; Dawson and Powell,
1999; Degen et al., 2001; Gillies et al., 1997; Heaton et
al., 1999; Lee et al., 2002; Lowe et al., 2000).
MATERIALS AND METHODS
Knowledge on T. Indica
T. indica L. is a semi-evergreen tree which can reach up to 20 m in
height. It is a member of the family Fabaceae (Leguminosae), which
is the third largest family of flowering plants with 700 genera and
approximately 17,000 species (Chant, 1993). It belongs to the
subfamily Caesalpinioideae, which can be divided into five to nine
tribes or groups of genera based on the morphological characters.
In the Bentham’s classification (Pettigrew and Watson, 1977),
tamarind belongs to the tribe of the Amherstieae Benth. According
to Léonard (1957), the Amherstieae comprises 25 genera in total,
21 in tropical Africa, two in tropical America and two in Asia. There
are only three relic Afro-Asiatic genera within the group, namely
Humboldtia, Tamarindus and Amherstia. Both Tamarindus and
Amherstia are more derived, with zygomorphic and showy flowers
and stamen filaments connate in a sheath, but have many
differences in the floral structure, leaves, fruits and seeds. These
genera are isolated from each other and from the main part of the
tribe, centred in western Africa (Polhill and Raven, 1981).
Tamarindus is said to have some resemblance to Heterostemon
Desf. from the upper Amazon region of South America. No other
tree bearing any resemblance to tamarind has been reported in
other countries (Dassanayake and Clayton, 1999). The genus
Tamarindus contains only one species (monotypic genus). It is a
diploid species with a chromosome number of x = 12 and 2n = 24
(Purseglove, 1987; El-Siddig et al., 1999).
Plant material and DNA extraction
Seeds were collected from 10 trees (Guadeloupe) to 30 trees
(Kodiena, Burkina Faso) in eight natural or naturalised plantations
and two planted stands of T. indica covering a large part of the
current range of the species (Table 1). For the populations of
Cameroon, India and Senegal the number of trees was not
specified by the seed centres.
The seeds were allowed to germinate in polycarbonate boxes
containing sand and young seedlings were kept in an incubator at
37°C. Total DNA was extracted from 150 mg of foliar tissues from
fifteen-day-old seedlings following the protocol of Bousquet et al.
(1990). The integrity of the DNA was estimated on agarose gels
and the quantity was determined using a fluorimeter DNA was then
diluted in sterile water to a concentration of 5 ng/µl for use in
amplification reactions.
RAPD analyses
Fifteen primers were screened to identify suitable primers for RAPD
analysis, using five individual samples from different provenances.
Eight primers able to detect distinct, clearly resolved and
polymorphic amplified products were selected for further analysis.
PCR was carried out in a final volume of 25 µl containing 10 mM
Tris (pH 8.0), 50 mM KCl, 2 mM MgCl2, 0.1 mM of each dNTP, 0.56
µM of primer, and 1 unit of Taq DNA polymerase. Amplification of
RAPDs was performed on a Perkin-Elmer thermocycler using the
following PCR conditions: an initial denaturation step at 94°C for 4
min, followed by 45 cycles each at 94°C for 1 min, 36°C for 1 min,
72°C for 2 min and a final extension step at 72°C for 2 min.
Negative controls, in which DNA was omitted, were included in
each run. Amplification products were separated by electrophoresis
on 1.8% agarose gels run in 1X TBE (Tris-Borate-EDTA) buffer,
stained with ethidium bromide, visualised and photographed under
UV light.
Scoring of bands and statistical analysis
Amplified bands were scored present (1) or absent (0) regardless of
band intensities. Ambiguous data were scored as missing data. We
analysed the data on three levels of comparison: among
individuals, among populations and among areas.
Ressemblances/differences among individuals
The Sokal and Michener index of similarity S (Sokal and Sneath,
1963) was calculated between all individuals, using the software
NTSYS-PC (Rohlf, 1993). The value of this index varies between 0
(two individuals not having any common marker) to 1 (two
individuals showing identical banding patterns). The matrix of
similarities allowed us to calculate the matrix of genetic distances
(1-S). A neighbor-joining (NJ) dendrogram was then constructed
from these distances using the software DARwin 3.6 (Perrier et al.,
1999). We performed a first analysis in order to quantify genetic
variability within the potential native range of T. indica, and then
added planted individuals and individuals from populations of recent
introduction.
Quantification and comparison of population diversity
The Shannon index (HS=- pi ln pi) and Nei’s index (H=n(1- pi2)/n-
1) where pi (in both indices) is the frequency of the ith RAPD band,
as well as the percentage of polymorphic loci (P) at the 0.99
criterion were calculated for each population using Popgen 1.32
(Yeh and Boyle, 1997). Calculation of P is a heuristic approximation
for these dominant markers. It is not known whether bands at a
particular level correspond to a single locus. Furthermore, only two
“alleles” are possible, ‘present’ and ‘absent’. The population
genetic structure FST was computed using AFLP-SURV 1.0
(Vekemans, 2002) following Lynch and Milligan (1994) instead of
ST normally applied during AFLP data analysis.
Comparison among areas
To examine differences among areas (East Africa, West Africa,
Cameroon, Réunion and Guadeloupe) we grouped individuals with-
Diallo et al. 855
Table 1. T. indica samples studied for RAPD analysis. N denotes the sample size for seeds and
Nb denotes the sample size of female donor trees of each population.
Country Population name Population type Nb trees N seeds
Burkina Faso Kodiena Natural 30 18
Cameroon Maroua Natural - 16
Kenya Gédé Natural 25 16
Senegal Pamene Natural - 18
Tanzania M’tandika Natural 22 18
Guadeloupe Mahaudière Planted 10 21
India Dehra Dun Natural - 24
Madagascar Anarafaly Natural 15 22
Réunion Etang sale Natural 16 21
Thailand Bangkok Planted 10 12
in each area as an entity and calculated distances of Nei using the
software AFLP-SURV 1.0 (Vekemans, 2002). Robustness of the
nodes was evaluated by bootstrap (2000 replicates). We used the
Neighbor and Consense procedures of the Phylip95 software
package (Felsenstein, 1993) to construct the dendrogram and infer
boostrap confidence on tree branches.
RESULTS
Intra-population genetic diversity
The eight primers used to screen RAPD diversity of T.
indica populations generated 58 polymorphic amplifica-
tion fragments across the whole sample. (Table 2) lists
the polymorphic primers, their sequences and the num-
ber of polymorphic markers found in the 10 populations
(Table 3). We obtained 5 to 10 fragments per amplifica-
tion with an average of 7.25. Marker 1 is fixed in Guade-
loupe, polymorphic in Thailand and India and completely
absent from the other populations (Figure 1). The
percenttage of polymorphic loci ranged from 39.7% for
Thailand to 77.6% for Cameroon with a mean of 69.3 for
the African populations. The average within-population
diversity was 0.28 and 0.31, respectively, over all popula-
tions and the African populations.
Inter-population genetic diversity
The dendrogram of individuals from populations of the
potential native range (Figure 2) exhibits three groups:
the first group includes the eastern African populations
along with the populations from Madagascar and from
India, the second group comprises the western African
populations and the individuals from Cameroon form a
third group (Figure 2). The same analysis carried out with
only the five African populations, groups the Came-
roonian individuals in a cluster with those from West
Africa; Kenyan and Tanzanian individuals form two other
groups (Figure 3). The among-populations, genetic
diversity was 0.09 for these five populations with FST =
Table 2. Polymorphic primers and number of polymorphic
markers obtained.
Name of
primer Sequence of primer
(5’ to 3’)
No. of polymorphic
markers
OPA-A09 GGGTAACGCC 10
OPA-B06 TGCTCTGCCC 7
OPA-K06 CACCTTTCCC 6
OPA-K17 CCCAGCTGTG 9
OPA-R15 GGACAACGAG 6
OPA-W01 CTCAGTGTCC 8
OPA-X01 CTGGGCACGA 7
OPA-Y01 GTGGCATCTC 5
Total 58
Table 3. Estimates of genetic diversity for T. indica.
Population P HS H
Burkina Faso 65.5 0.325 (0.281) 0.216 (0.199)
Cameroon 77.6 0.429 (0.262) 0.290 (0.186)
Guadeloupe 63.8 0.312 (0.283) 0.207 (0.198)
India 72.4 0.345 (0.268) 0.227 (0.188)
Kenya 75.9 0.401 (0.271) 0.270 (0.193)
Madagascar 60.3 0.326 (0.295) 0.221 (0.208)
Réunion 67.2 0.370 (0.289) 0.251 (0.205)
Senegal 60.3 0.334 (0.299) 0.227 (0.211)
Tanzania 67.2 0.351 (0.287) 0.236 (0.205)
Thailand 39.7 0.219 (0.287) 0.148 (0.200)
100.0 0.521 (0.180) 0.350 (0.144)
P, percentage of polymorphic loci; HS, mean Shannon diversity
index; H, Nei’s diversity index.
Values in parentheses denote standard deviations.
0.23, highly significant (P < 0.001).
856 Afr. J. Biotechnol.
Tanzania Senegal Burkina Faso
Guadeloupe Cameroon Madagascar
OPA-K06
750 pb
Thaïland Kenya India
OPA-K06
310 pbe
Figure 1. RAPD profiles obtained after amplification of 9 populations of Tamarindus indica with OPA-KO6 after
separation on a 1.8% agarosse gel. The size marker included was 1 Kb DNA Ladder.
Figure 2. Dendrogram of the populations from native range of T. indica, generated by neighbor joining cluster analysis of Sokal
and Michener distances. Bf: Burkina Faso; Cm: Cameroon; Id: India; Ky: Kenya; Md: Madagascar; Sn: Senegal; Tz: Tanzania.
The individuals from the potential native range that
were grouped into three clusters were analyzed again
with the individuals of the population from Thailand,
which are planted, and the individuals from Guadeloupe
and Réunion, which are from naturalised populations
resulting from recent introductions. The dendrogram
(Figure 4) shows that individuals from the Réunion are
grouped with the western African individuals with a
statistically strong bootstrap value (72%); the individuals
from Thailand and Guadeloupe form a cluster which is
closest to those from Cameroon but with a lower boot-
strap value (44%); the individuals from eastern Africa,
Madagascar and India are the most different from all the
others (bootstrap value of 68%). Thus, a factorial analysis
Diallo et al. 857
Figure 3. Dendrogram of the African populations generated by neighbour joining cluster analysis of Sokal and
Michener distances. Bf: Burkina Faso; Cm: Cameroon; Ky: Kenya; Sn: Senegal; Tz: Tanzania.
East
West
Réunion
Cameroon
Thailand
Guadeloupe
72
44
63
68
Figure 4. Dendrogram of the various areas considered, constructed
using neighbour joining method. Bootstrap values are indicated for
each corresponding node (2000 replicates) East: Kenya, Tanzania,
Mada-gascar and India; West: Burkina Faso, Senegal.
of correspondence (AFC) (Figure 5) showed the same
structuring of populations in addition to predominant
markers associated with each group.
DISCUSSION
Presumed origin of T. indica
The high intra population variability from the populations
of the presumed origins of T. Indica do not allow for
confirmation of the geographical origin of the species
between Africa, Madagascar and India, as the sampling
was small in Asia and Madagascar. In fact, Polhill and
Raven (1981) who studied the centre of diversity of some
Leguminosae tribes measured by the percentage of the
total number of genera that are endemic to each region
showed that out of the 25 genera of Amhertieae (Tam-
arindus tribe), 23 are endemic to Africa and Madagascar,
and only two to Asia and America, respectively. However,
our findings reinforce paleontological observations cond-
ucted in the tertiary sediments in the south and centre-
north of Tanzania respectively, that revealed the presen-
ce of the pollen related to five genera of Caesalpinioideae
(Brachystegia; Cassia-type didymobotrya; Cassia-type
italica; Isoberlinia and Jubernardia-type paniculata) (Vin-
cens et al., 2003; Herendeen and Jacobs, 2000).
According to Polhill and Raven (1981) three of these
genera were encountered in the south of central Africa in
Guineocongolese forest. However, none of these studies
mentioned the Tamarindus genus suggesting a very late
migration to this zone.
Genetic diversity and origin of newly introduced
populations
In general, T. indica displays a high genetic diversity (H =
0.38), the value of this species is higher than values
obtained with isoenzymes for tropical rain forest species
(H = 0.11) (Hamrick and Loveless, 1986) and for
Australian Acacia species (0.07 to 0.20) (Moran et al.,
1989a,b; Coates,1988), or for coniferous species (H =
0.27) (Miton, 1983). In turn, the diversity value of T. indi-
ca is close to the values reported for Fadherbia albida in
tropical arid zones (Joly et al., 1992; Harris et al., 1997)
and Prunus africana (Dawson and Powell,1999). Results
obtained in this study are congruent with published stud-
ies on other African tree species which indicate a high
level of among populations diversity (Lowe et al., 2000),
858 Afr. J. Biotechnol.
Plan 1/2
- 0 .8
0
0. 8
- 0 . 8 0 0 .8
a xe 1
Guadeloupe Cam e roun Madagascar Thailande Kenya
Inde Tanzanie Sénégal Burkina Réunion
Plan 1/3
-0.8
0
0.8
-0.8 0 0.8
axe 1
axe 2
Guadeloupe Cameroun Madagascar Thailande Kenya
Inde Tanzanie Sénégal Burkina Réunion
Figure 5. Analyzes Factorial correspondences (AFC) on the individuals (provenances) in
plans 1/2 and 1/3.
and a high differentiation between East African and West
African populations on Prunus africana (Dawson and Powell, 1999), Faidherbia albida (Harris et al., 1997) and
Acacia senegal (Unpublished data Dolmia).
A partial analysis performed on populations coming
from the presumed natural distribution of the species
(Africa, Madagascar, India) revealed its structure per
geographic great region. The partial analysis including
only African populations shows a high differentiation bet-
ween populations from West and East of the continent.
Such differentiation has already been found with other
species like F. albida (Joly et al., 1992; Rendell, 1998),
Ivirginia gabonensis and I. wombolu (Lowe et al., 2000).
This differentiation may presumably stem from the role
that the Rift valley might have played in stopping gene
flow between the two regions of the continent both in
terms of seed dispersion and those induced by human
exchanges and migration. Moreover, it should be noted
that the population from Cameroon displayed the highest
genetic diversity value (H = 0.77). This was also found in
bush mango (Lowe et al., 2000). The high variability in
the Cameroonian population can be explained by two
plausible reasons, namely Maroua had been a com-
mercial point between Sub-saharan Africa and India via
the Mahgreb using the Sahara road as emphasized by
Omer-Cooper et al. (1968), and Cameroon could have
been a primary center from which the species migrated
toward the other regions of the continent with favorable
conditions for its growth.
The two genetically close West African populations pre-
sented high intra population diversity that contrast with
findings of Chevallier et al. (1994) on Senegal popula-
tions of A. senegal. However, the contradictory results
may be linked to the commercial exchanges in the sub-
region favoured either by Dioula merchants of nowadays
Mali or via the massive displacements of human cara-
vans during transhumance, nomadism and wars. In fact,
Omer-Cooper et al. (1968) noted that in the 19th century
Fulani (Peulh) conducted many Jihads in the region. The
best known example is that of Utsman Dan Fodio in the
State of Gobir whose forefathers came from Fouta Toro
in actual Senegal. After the fall of the empire, an impor-
tant part of his fellows went to the northern part of
Burkina Faso to create the Liptako. During this second
migration, seeds originating from Senegal may have
been introduced via Gobir State. Afterward, hybridization
of the populations might have happened with the local
populations. The second hypothesis is that it might be
only one population that never differentiated, because
they are geographically isolated. That did not allow the
drift effect to impact on populations.
Genetic differentiation of La Réunion from West Afri-
ca, and genetic differentiation of Guadeloupe from
Thailand populations
Results obtained after analysis of populations from the
natural distribution and areas of recent introductions sug-
gest that the population of Réunion Island comes from
Diallo et al. 859
West Africa. With the discovery of the Island in 1500 by
Tristan Da Cunta, the possibilities for Vanilla and sugar
cane production led to a need for labour, therefore the
first French migrants recruited cheaper labour on the
western coasts. These workers may have transported
with them seeds of tamarind tree. However, material
could have been introduced by Arab merchants who follo-
wed the Spanish explorers of the island since medieval
times and may have transported seeds to the island.
Even though not inhabited, the island was the way to the
Indies and may have served as a stop point for boats
coming from the western African coasts because the
reports relating to the company of Indies (1711) ment-
ioned the presence of this species on the island (Lacou-
ture).http//perso.wanadoo.fr/daniel.lacouture/divers/dossi
er/nature/culture/nature020/
From this analysis it was also evident that the popu-
lation of Guadeloupe comes from Thailand. These two
populations present also a low genetic diversity compa-
red to the populations from the natural distribution area of
the species. This may be due to the fact that the planted
trees came from seeds collected on a low number of
individuals. The differentiation between the two popula-
tions of East Africa (Tanzania and Kenya) is probably due
to either natural and/or ethnic barriers (i.e. wars and
plundering without occupying the conquered lands) that
stopped human movements and indirectly the exchanges
of vegetal material.
In conclusion, the intra population variation observed,
as well as the low sample number of Asian populations
did not allow for clarity on the origin of T. indica. How-
ever, the observed genetic diversity within the African
populations shows that there are no risks of genetic ero-
sion in the short term. Molecular analyses based chloro-
plastic DNA maternally inherited characteristics may help
to understand the mode of dissemination of T. indica.
ACKNOWLEDGEMENTS
The field and laboratory works of this study were spon-
sored by the International Foundation of Science (IFS),
CIRAD-Forêts and the French cooperation mission.
REFERENCES
Bekessy SA, Allnutt TR, Premoli AC, Lara A, Ennos RA, Burgman MA,
Cortes M, Newton AC (2002) Genetic variation in the vulnerable and
endemic Monkey Puzzle tree, detected using RAPDs. Heredity 88:
243-249.
Bousquet J, Simon L, Lalonde M (1990) DNA amplification from
vegetative and sexual tissues of trees using polymerase chain
reaction. Can. J. Forest Res. 20: 254-257.
Brush S, Kesseli R, Ortega R, Cisnero P, Zimmerer K, Quiros C (1995).
Potato diversity in the Andean Center of crop domestication.
Conservation Biol. 9: 1189-1198.
Cardoso MA, Provan J, Powell W. Ferreira PCG, De Oliveira DE (1998).
High genetic differentiation among remnant populations of the
endangered Caesalpinia echinata Lam. (Leguminosae-
Caesalpinioideae). Mol. Ecol. 7: 601-608.
860 Afr. J. Biotechnol.
Chant SR (1993) Fabales. In Flowering Plants of the World. Editor
Heywood VH, Batsford BT. Ltd.London.
Chevallier MH (1999). Diversité génétique des arbres forestiirs In : Vers
une approche régionale des ressources génétiques forestières en
Afrique Sub-Saharienne eds. Ouédraogo AS., & Boffa JM. Pp IPGRI,
FAO, Rome, Italie. 299: 124-137.
Chevallier MH, Brizard JP, Diallo I, Leblanc JM (1994). La diversité
génétique dans le complexe Acacia senegal. Bois et Forêts des
Tropiques 240: 5-12.
Coates DJ (1988) Genetic diversity and population genetic structure in
rare Chittering grass wattle, Acacia anomala Court. Aust. J. Bot. 36:
273-286.
Dawson IK, Powell W (1999). Genetic variation in the Afromontane tree
Prunus africana, an endangered medicinal species. Mol. Ecol. 8: 151-
156.
Dassanayake MD, Clayton WD (eds.) (1999). Revised handbook to the
flora of Ceylon, Vols XIII Amerind Publ. New Delhi.
Degen B, Caron H, Bandou E, Maggia L, Chevallier MH, Leveau A,
Kremer A (2001). Fine-scale spatial genetic structure of eight tropical
tree species as analysed by RAPDs. Heredity 87: 497-507.
El-Siddig K, Ebert G, Ludders P (1999). Tamarind (Tamarindus indica
L.): a review on a multipurpose tree with promising future in the
Sudan. J. Appl. Botany 73: 202-205.
FAO (1996). Conservation des ressources génétiques dans
l’aménagement des forêts tropicales. Principes et concepts. Etude
FAO, forêts 107. FAO, Rome. p. 101.
Felsenstein J (1993). PHYLIP (Phylogeny Inference Package) version
3.5c. Distributed by the author. Department of Genetics., University of
Washington, Seattle.
Gillies ACM, Cornelius JP, Newton AC., Navarro C, Hernandez M,
Wilson J (1997). Genetic variation in Costa Rican populations of the
tropical timber species Cedrela odorata L., assessed using RAPDs.
Mol. Ecol. 6: 1133-1145.
Grollier C, Debien C, Dornier M, Reynes M (1998). Principales
caractéristiques et voies de valorisation du tamarin. Fruits 53: 271-
280.
Hamon P, Seguin M, Perrier X, Glaszmann JC (Eds) (1999). Genetic
diversity of tropical cultivated plants. Science Publishers Inc.Cirad
,Montpellier (France)
Hamrick Jl, Loveless MD (1986). Isozymes variation in tropical trees :
procedures and preliminary results. Biotropica 18: 201-207.
Harris SA, Fagg CW, Barnes RD (1997). Isozyme variation in
Faidherbia albida (Leguminosae, Mimosoideae). Plant Syst. Evol
207: 119-132.
Heaton HJ, Whitkus R, Gomez-Pompa A (1999). Extreme ecological
and phenotypic differences in the tropical tree chicozapote (Manilkara
zapota (L.) Royen P) are not matched by genetic divergence: a
random amplified polymorphic DNA (RAPD) analysis. Mol. Ecol. 8:
627-632.
Herendeen PS & Jacobs BF (2000). Fossil legumes from the Middle
Eocene (46.0 Ma) Mahenge flora of Singida, Tanzania Am ( r ). J.
Botany 87: 1358-1366.
http//perso.wanadoo.fr/daniel.lacouture/divers/dossier/nature/culture/nat
ure020/
Joly H, Zeh-Nlo M, Danthu P, Aygalent C (1992). The genetics of
Acacia albida (syn. Faidherbia albida) in Faidherbia albida in west
African semi-arid tropics : proceedings of a workshop, Apr. 1991.
Niamey, Niger (Vandenbeldt, R J., ed.). Patachncheru, A.P. 502324,
India : International Crops Research Institute for the Semi-Arid tropics
and Nairobie, Kenya : International Centre for research in
Agroforestry. 22-26
Lowe A.J, Gillies ACM, Wilson J, Dawson IK (2000). Conservation
genetics of bush mango from central/west Africa: implications from
random amplified polymorphic DNA analysis Mol. Ecol. JUL 2000 9:
831-841
Lee CT, Wickneswari R, Mahani MC, Zakri AH (2002). Effect of
selective logging on the genetic diversity of Scaphium macropodum.
Biol. Conser. 104: 107-118.
Lefèvre JC, (1971). Revue de la littérature sur tamarinier. Fruits 26:
687-695.
Lowe AJ, Gillies ACM, Wilson J, Dawson I (2000). Conservation
genetics of bush mango from central/west Africa: implications from
random amplified polymorphic DNA analysis. Mol. Ecol. 9: 831-841.
Lynch M, Milligan BG (1994). Analysis of population genetic structure
with RAPD markers. Mol. Ecol. 3: 91-99.
Mitton JB (1983). Conifers. In plant Genetics and Breeding . Part B.
(eds SD Tanskley, TJ Orton) Elsevier. Amsterdam. pp. 423-441.
Moran GF, Muona Od, Bell JC (1989a). Acacia mangium : a tropical
forest tree of the coastal lowland with low genetic diversity. Evolution
43: 231-235.
Moran GF, Muona O, Bell JC (1989b). Breeding system and genetic
diversity of Acacia auriculiformis and A. crassicarpa. Biotropica
21: 250-256.
Nicolosi E, Deng ZN, Gentile A, La Malfa S, Continella G, Tribulato E
(2000). Citrus phylogeny and genetic origin of important species as
investigated by molecular markers. Theor. Appl. Genet. 100: 1155-
1166.
Perrier X, Flori A, Bonnot F (1999). Les méthodes d'analyse des
données, p. 43-76, In P. Hamon, M. Séguin, X. Perrier JC
Glaszmann, eds. Diversité génétique des plantes tropicales cultivées.
Cirad, éditions Repères, Montpellier.
Pettigrew CJ, Watson L (1977). On the classification of
Caesalpinioideae. Taxon 26: 57-64.
Polhill RM, Raven PH (1981). Biogeography of leguminosae. In
Advances in legume systematics, Part 1. pp 27-34. eds. Polhill R.M.
& Raven P.H. Royal Botanic Gardens, Kew. p. 425.
Poupon J, Chauvin G (1983). Les arbres de la Martinique, p. 256, In
ONF, ed. Direction régionale pour la Martinique.
Purseglove JW (1987). Tropical crops. Dicotyledons, Longman Science
and Technology: 204-206.
Léonard J (1957). Genera des Cynometreae et des Amherstieae
africaines (Leguminosae-Caesalpinioideae). Mémoire Académie
Royale Belgique 30: 1-314.
Omer-Cooper JD, Ayandele EA, Afigbo AE, Gavin RJ (1968). The
growth of African Civilisation. The making of modern Africa. Vol 1:The
Nineteenth century to the partition. Printed in Singapore by New Art
Printing Co. (pte) LtD ISBN 0582 60247-5
Rendell S (1998). Population genetic structure of Faidherbia albida
(Del) A. Chev. (Leguminosae, Mimosoideae). Thesis of Biological
Sciences, University of Oxford. The British Library, Document supply
centre. Boston Spa, Wetherby West Yorkshire, United Kingdom.
LS23 7BQ. P219.
Rohlf FJ (1993). NTSYS-pc: numerical taxonomy and multivariate analysis
system. Version 1.80 Setauket, N.Y. Exeter Software, c1993. 2
computer disks 1 manual (loose-leaf); 23 cm.
Sokal RR, Sneath PHA (1963). Principles of Numerical Taxonomy.
Freeman, San Francisco.
Vekemans X (2002). AFLP-SURV version 1.0. Distributed by the author.
Laboratoire de Génétique et Ecologie Végétale, Université de
Bruxelles, Belgium.
Vincens A, Williamson D, Thevenon F, Taieb M, Buchet G, Decobert M,
Thouveny N (2003). Pollen-based vegetation changes in southern
Tanzania during the last 4200 years: climate change and/or human
impact Palaeogeography palaeoclimatology palaeoecology 198: 321-
334.
Williams JGK, Kubelik AR, Livak KJ, Rafalski JA, Tingey SV (1990).
DNA Polymorphisms Amplified by Arbitrary Primers Are Useful as
Genetic-Markers. Nucleic Acids Research 18: 6531-6535.
Wunderlin RP (1998). Guide to the vascular plants of Florida Gainesville
: University Press of Florida, p. 806.
Yeh FC, Boyle TJB (1997). Popgene Version 1.32. Microsoft Windows-
based software for population genetic analysis.pp. 135
