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Quantitative morphological descriptors confirm traditionally classified morphotypes of Tamarindus indica L. fruits

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This study used quantitative descriptors to assess morphological variation of traditionally classified tamarind fruits and its relation to ecological conditions. Tamarind trees were sampled spanning locally recognized fruit morphotypes within three ecological zones. Twelve morphological descriptors were measured on 3000 fruits and seeds. Univariate and canonical discrimiant analyses performed on morphological descriptors revealed significant differences and confirmed morphotypes distinction as perceived by local people. Nevertheless, the variance components analysis showed substantial variations within morphotypes, suggesting a significant heterogeneity within fruits traditionally classified as belonging to the same morphotypes. To get a more powerful morphological discrimination, quantitative descriptors should hence be combined with locally perceived qualitative traits (pulp taste and colour). Observed variations were significantly correlated with ecological factors. Fruits’ and seeds’ size and mass tended to increase with humidity and decline with aridity. Results also indicated that fruit mass is a good predicator of pulp yield, although its predicting power differed among morphotypes. Outputs from the variance component analysis suggested that pending further genetic studies, germplasm collection should be done by sampling a moderate number of trees per morphotype, to ensure capturing a wide range of genetic diversity. The observed extensive variation has provided with relevant information for further improvement programs. KeywordsEcological zones–Folk classification–Morphological variation– Tamarindus indica –West Africa
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RESEARCH ARTICLE
Quantitative morphological descriptors confirm
traditionally classified morphotypes of Tamarindus
indica L. fruits
Belarmain Fandohan Achille Ephrem Assogbadjo
Romain Gle
`le
`Kakaı
¨Tina Kyndt
Brice Sinsin
Received: 18 December 2009 / Accepted: 17 May 2010 / Published online: 9 June 2010
ÓSpringer Science+Business Media B.V. 2010
Abstract This study used quantitative descriptors
to assess morphological variation of traditionally
classified tamarind fruits and its relation to ecological
conditions. Tamarind trees were sampled spanning
locally recognized fruit morphotypes within three
ecological zones. Twelve morphological descriptors
were measured on 3000 fruits and seeds. Univariate
and canonical discrimiant analyses performed on
morphological descriptors revealed significant differ-
ences and confirmed morphotypes distinction as
perceived by local people. Nevertheless, the variance
components analysis showed substantial variations
within morphotypes, suggesting a significant hetero-
geneity within fruits traditionally classified as belong-
ing to the same morphotypes. To get a more powerful
morphological discrimination, quantitative descrip-
tors should hence be combined with locally perceived
qualitative traits (pulp taste and colour). Observed
variations were significantly correlated with ecolog-
ical factors. Fruits’ and seeds’ size and mass tended
to increase with humidity and decline with aridity.
Results also indicated that fruit mass is a good
predicator of pulp yield, although its predicting
power differed among morphotypes. Outputs from
the variance component analysis suggested that
pending further genetic studies, germplasm collection
should be done by sampling a moderate number of
trees per morphotype, to ensure capturing a wide
range of genetic diversity. The observed extensive
variation has provided with relevant information for
further improvement programs.
Keywords Ecological zones Folk classification
Morphological variation Tamarindus indica
West Africa
Introduction
The domestication of indigenous fruits and nuts for
the diversification of subsistence agriculture is play-
ing a big role in the achievement of the Millennium
Development Goal, trying to combat poverty and
hunger and mitigate environmental degradation in
developing countries (Leakey et al. 2007). Studies on
the biological variability of indigenous fruit tree
species, their propagation using cheap and simple
methods appropriate for rural development projects,
and their suitability for domestication have been
progressively increased in West Africa over the last
decades (Leakey et al. 2000). In some cases, a
participatory approach to cultivar development was
implemented with success (Leakey et al. 2003). In the
B. Fandohan (&)A. E. Assogbadjo
R. Gle
`le
`Kakaı
¨B. Sinsin
Faculty of Agronomic Sciences, University of Abomey-
Calavi, 01 BP 526 Cotonou, Benin
e-mail: bfandohan@gmail.com
T. Kyndt
Faculty of Bioscience Engineering, Ghent University,
Coupure Links 653, 9000 Ghent, Belgium
123
Genet Resour Crop Evol (2011) 58:299–309
DOI 10.1007/s10722-010-9575-3
fulfilment of cultivars development for priority tree
species, two key elements are (1) the identification of
‘plus trees’’ in natural populations and (2) their
propagation by vegetative techniques (Leakey and
Page 2006). Prior to ‘‘plus trees’’ identification,
quantitative characterization of fruit, nut and kernel
variation (Leakey et al. 2005a), variation in nutritive
value and other food properties (Leakey et al. 2005b)
have to be studied and an understanding of the
interactions between different traits for multi-trait
selection is needed (Leakey 2005). Tamarind, Tam-
arindus indica L. (Leguminosae: Caesalpinioideae),
is a semi-evergreen multipurpose tree typical of
savannah ecosystems, featuring prominently in ripar-
ian habitats (Fandohan et al. 2010). The tamarind tree
has an important role in local economies, supple-
ments the local diet, and is used in traditional and
modern therapies (El-Siddig et al. 2006). Its pulp is
much appreciated in condiments, used to make juice
and is a good source of proteins, fats and carbohy-
drates that could be used to alleviate malnutrition in
children (El-Siddig et al. 2006). In efforts to enhance
the species’ genetic conservation and utilization it has
been recently identified as one of the top ten
agroforestry tree species to be prioritized for future
crop diversification programs and development in
sub-Saharan Africa (Eyog Matig et al. 2002).
Although of high local economic importance, our
knowledge of the morphometric and ecological diver-
sity of tamarind is still limited outside of Asia. Works
on Asian tamarind populations have revealed a
considerable phenotypic and genotypic variation and
allowed the selection of superior trees based on pulp
mass, pulp taste and fruit length (El-Siddig et al. 2006).
Other studies have addressed its domestication poten-
tial in Africa and provided data on biochemical
analyses (Soloviev et al. 2004), the comparison of
the genetic diversity of African, Asian and South-
American populations as indicator of the species native
area, breeding systems and pollination-related issues
(Diallo et al.2007,2008). To our knowledge, no study
has documented (1) indigenous perception of qualita-
tive or size group morphological variation within the
species and (2) quantitative morphological and genetic
structuring to test whether locally perceived variations
and preferences are either genetically or ecologically
determined. Such bottom-up approach may help to
identify and characterize ‘‘plus trees’’, locate ecolog-
ical conditions allowing the species to better express its
potential (i.e. fruit size and yield, pulp productivity,
pulp taste, etc.), identify links between traits and is
crucial to make improvement strategies realistic. The
current study aims at (1) matching the quantitative
assessment of tamarind fruits traits with the folk
classification based on local knowledge, (2) analysing
its relationship with ecological conditions and,
(3) analysing the implications for further improvement
programs. Thus, the following questions were
addressed: Do quantitative descriptors confirm folk
classification of tamarind morphotypes? Which eco-
logical factors drive the pattern of morphological
variability in T. indica?
Materials and methods
Study area
The study was conducted in Benin (West Africa).
Three different ecological zones were targeted based
on tamarind distribution range: the Sudanian zone
(9°450–12°250N), the Sudano-Guinean zone (7°300
9°450N) and the sub-humid Guineo-Congolian zone
(6°250–7°300N). The Guineo-Congolian zone is the
wettest with a bimodal rain regime whilst the
Sudanian zone is the driest with nearly a 7 month
drought period. The vegetation is made of grassland
and thickets and some relic rain forests in the Guineo-
Congolian zone. In the Sudano-Guinean zone, the
vegetation is dominated by Isoberlinia spp. wood-
lands whereas in the Sudanian zone, the vegetation is
characterised by Combretum spp. and Acacia spp.
tree savannas (White 1983). Table 1summarizes the
ecological characteristics of the three study sites.
Data collection
Tamarind individuals were sampled in the Sudanian,
the Sudano-Guinean and the Guineo-Congolian eco-
logical zones of Benin. Within each zone, trees were
sampled where local people had experience and
knowledge on tamarind tree. Ethnobotanical surveys
were carried out on the local perception of the
morphological variation in tamarind fruits. The
survey revealed that local people distinguish ten
morphotypes (Table 2), a morphotype being a group
of tamarinds sharing some qualitative fruit traits as
perceived by interviewees. Two experienced women
300 Genet Resour Crop Evol (2011) 58:299–309
123
very familiar to the described morphotypes chosen
with the help of local leaders in each study site were
asked to participate in the selection of tamarind
individuals to be sampled for fruit morphological
traits description. Five trees were sampled per
morphotype: 25 individuals in the Sudanian zone,
15 in the Sudano-Guinean zone and 10 in the Guineo-
Congolian zone. The variability in the number of
samples per zone was due to the fact that not all
morphotypes were found in all zones. From each
selected tree, 30 samples of both fruits and seeds
were collected for measurement following the proto-
col described by Leakey et al. (2000). We measured
twelve morphological descriptors on fruits (length,
width, thickness, number of seeds, fresh mass, dry
mass, pulp mass, and the ratio pulp mass/fruit mass)
and seeds (length, width, thickness and mass). To
improve accuracy, fruit’s width and thickness were
measured at the first, the second and the third quarter
of each fruit and the arithmetic means were consid-
ered as the fruit’s width and thickness. Similar
descriptors have already been used in other studies
like IBPGR (1980), Leakey et al. (2000), (2005a,b),
El-Siddig et al. (2006).
To estimate pulp mass, fruits were oven-dried at
65°C for 48 h to obtain the dry weight. Dried fruits
were broken and the content extracted (pulp ?
seeds ?fibers). The pulp was removed by soaking
the content in water. The residu (seeds and fibers) was
oven-dried at 65°C for 48 h. This protocol was
successfully used before for the baobab tree
(Assogbadjo et al. 2005).
Overall, 18,000 individual values were recorded for
1500 fruits and 1500 seeds from the 50 analyzed trees.
Monthly climatic data (rainfall, relative humidity,
minimum and maximum temperatures and insolation)
and number of dry months per year for over 30 years
(1978–2008) were obtained for each study site within
ecological zones from Hijmans et al. (2004).
Analysis
The pulp mass (wp) in each fruit was computed using
the following formula:
wp ¼wPiwRið1Þ
where, wp is the pulp content of a given fruit; wP
i
is
the dried mass of the fruit (i); wR
i
is the total dried
mass of seeds, fibers and husk of the fruit (i).
Univariate analyses of variance and Student–
Newman–Keuls (SNK) tests were used to describe
the morphotypes and identify the discriminative
descriptors. Then, Least Square Means of fruits and
seeds descriptors were estimated and a Canonical
Discriminant Analysis (using the Mahalanobis dis-
tance) was performed to reveal links between the
descriptors and plot distances between morphotypes.
This multivariate analysis is a relevant and powerful
test to distinguish between entities that fall into
natural groupings i.e. morphological or ecological
groups (Lowe et al. 2004).
Afterwards, the within and between morphotypes
variability was evaluated using Variance Component
Analysis (Goodnight 1978). To examine the influence
of ecological conditions on tamarind fruit and seed
traits, a Principal Component Analysis (PCA) was
performed only on the quantitative descriptors. The
PCA factor scores were correlated with the climatic
index of Mangenot (1951), the minimum and the
maximum temperatures and the insolation using a
Pearson correlation.
The climatic index of Mangenot (I
M
) was com-
puted for each sampled site as follows:
IM¼
P
100 þMSþUx
nS þ500
Un ð2Þ
where P: mean annual rainfall (mm), M
S
: mean rainfall
of dry months (i.e. months with rainfall less than
Table 1 Characteristics of the three study sites (adapted from Hijmans et al. 2004)
Guineo-Congolian Sudano-Guinean Sudanian
Average rainfall (mm/year) 1200 1200 675
Temperature range (°C) 18–33 20–36 17.1–42.1
Relative humidity range (%) 30–98 31–98 18–99
Climate type Sub-humid Sub-humid with tendency to dry Sudanian dry
Soil type Ferralitic without concretions Ferralitic with concretions Ferruginous on sedimentary rocks
Genet Resour Crop Evol (2011) 58:299–309 301
123
50 mm, nS: number of dry months, Ux: maximum of
annual relative humidity (%), Un: minimum of annual
relative humidity (%). A higher Mangenot index
indicates wetter ecological conditions.
As the pulp is the principal trait of commercial
importance we also carried out a linear regression to
identify predictors of pulp yield per fruit and to test if
the predicting power of the explanatory variables
differs between morphotypes. We built a linear
regression for pulp mass per fruit, with eight indepen-
dent variables measured on fruits (length, width,
thickness, and mass) and seeds (length, width, thick-
ness and mass). Pearson’s correlation was performed
between the independent variables to test multicollin-
earity. Since there were significant strong correlations
between pairs of variables (r[0.60, P\0.001) only
one independent variable (fruit mass) was finally used
in the regression model. We insert traditional morpho-
type in the model as a dummy variable (see Kutner
et al. 2005). The model tested was: pulp mass =
b
0
?b
1
(fruit mass) ?b
2
(traditional morphotype) ?
e.b
0
indicates the intercept, b
1
and b
2
the partial
regression slopes and ethe unexplained error associ-
ated to the model. The residuals normality plot, the
residual vs. fitted plot and the residuals vs. leverage
plots with Cook distance were used to diagnose
the regressions models (Quinn and Keough 2005).
Data were processed under SAS version9.1 (SAS Inc.
2003).
Results
Quantitative morphological assessment
of traditionally classified tamarind morphotypes
Table 3shows the mean values recorded for quan-
titative fruit and seed descriptors in the 10 identified
morphotypes of T. indica. Mean fruit traits (length,
width, thickness, number of seed, fresh mass, dry
mass, pulp mass, ratio length/width and ratio pulp/
fruit) and mean seed traits (length, width, thickness,
mass and ratio length/width) significantly differed
between morphotypes (P\0.0001; Wilks’ Lambda
C0.124). The lowest fresh and dry fruit mass and
pulp mass, seed length, seed width and seed mass
were recorded for fruits from morphotype A, whilst
fruits from morphotype Bshowed the lowest fruit
thickness. Fruits from morphotype Cportrayed the
Table 2 Folk classification of tamarind morphotypes
Morphotypes Fruit shape Pulp colour Pulp taste Seed shape Seed colour Seed brightness Seed roughness
Curved Straight Brown–
yellowish
Brown-
darkish
Sour Intermediate Sweet Irregular Bowl-
shaped
Quadrangular Brown Brown-
darkish
Black Non-
brilliant
Brilliant Rough Polished
A?- ? - ?- -? -- ?--? - ?-
B?- ? - ?- -- -? ?- -? - -?
C?- - ? -- ?- -? -? -- ? -?
D?- - ? -- ?- -? -? -- ? ?-
E?- - ? -- ?? -- -? -- ? ?-
F?- - ? -? -- -? -? -- ? ?-
G?- - ? -? -- ?- -? -? - ?-
H?- - ? -- ?- ?- -- ?- ? ?-
I-? ? - ?- -? -- ?- -? - ?-
J-? - ? -- ?- -? -? -- ? ?-
Locally recognized morphotypes are indicated as (?) present or (-) absent
302 Genet Resour Crop Evol (2011) 58:299–309
123
highest ratio pulp/fruit and seed thickness whereas
fruits from morphotype Eexhibited the lowest
length. Fruits from morphotype Fshowed the
highest values in width, fresh and dry mass and
pulp mass while fruits from morphotype Gshowed
the highest values in length and thickness, seed
length, seed thickness and seed mass. Fruits from
morphotype Hshowed the lowest values in width,
number of seeds per fruit and seed thickness
whereas fruits from morphotype Jshowed the
highest values for number of seeds per fruit but
the lowest values for the ratio pulp/fruit.
The multivariate canonical discriminant analysis
on fruit and seed descriptors using the Mahalanobis
distance calculation confirmed the morphotypes as
discriminated by local people (P\0.0001; Wilks’
Lambda =0.82).
The canonical discriminant analysis performed on
the ten morphotypes showed that the first two axes
explained 82% of the observed variation. These axes
were thus used to describe the relationships between
the investigated descriptors and traditional morpho-
types. The correlation between the axes and the used
descriptors is shown in Table 4. The first axis
showed a strong and positive link with and between
the fruit length, width, thickness, fresh mass, dry
mass pulp mass and number of seeds per fruit and
the seed length, seed width and seed mass. This axis
was negatively correlated with the ratio pulp/fruit.
Figure 1shows the projection of the individuals
from the ten morphotypes onto axes 1 and 2. From
this plot and Table 4it can be deduced that
overall, morphotypes Fand G(located in the upper
positive part of the axis 1) outclassed the others for
most of the quantitative descriptors but showed low
values for the ratio pulp/fruit. In contrast, the
other morphotypes had high values for the ratio
pulp/fruit.
Despite the significant differences among morpho-
types suggested by the canonical discriminant anal-
ysis, the variance components analysis revealed that
the variation within morphotypes is higher than that
between them for all fruit and seed descriptors except
seed length (Table 5). In general, 47 to 95% of the
morphological variation was present within morpho-
types. Nevertheless, important amounts of between
morphotypes variations were detected for fruit mass,
pulp mass, seed length, seed width and seed mass (24
to 53%).
Table 3 Means and standard errors of quantitative morphological descriptors of fruits and seeds of the ten locally identified tamarind morphotypes
Fruit length
(cm)
Fruit width
(cm)
Fruit
thickness
(cm)
Fruit fresh
mass (g)
Fruit dry
mass (g)
Pulp
mass (g)
Ratio- pulp/
fruit
Number of
seeds per fruit
Seed
length (cm)
Seed
width (cm)
Seed
thickness (cm)
Seed mass (g)
Morphotypes
A12.58 ±0.69e 1.86 ±0.02 h 1.47 ±0.01f 11.15 ±0.26 h 10.29 ±0.21 h 3.92 ±0.11i 0.35 ±0.01b 8.16 ±0.13d 1.08 ±0.01i 0.87 ±0.02 g 0.55 ±0.003d 0.45 ±0.004 g
B15.01 ±0.41c 1.97 ±0.02f 1.4 ±0.02 g 15.28 ±0.46e 12.78 ±0.34e 5.26 ±0.26e 0.37 ±0.02a 8.80 ±0.28c 1.2 ±0.01 g 1.06 ±0.01d 0.55 ±0.01d 0.62 ±0.02d
C12.28 ±0.27f 1.88 ±0.03 g 1.61 ±0.03d 12.47 ±0.66 g 10.94 ±0.53 g 4.86 ±0.34f 0.38 ±0.01a 7.50 ±0.24e 1.32 ±0.01e 0.98 ±0.02e 0.68 ±0.01a 0.63 ±0.009d
D14.76 ±0.36d 2.32 ±0.03d 1.74 ±0.02b 18.40 ±0.56d 17.68 ±0.55d 7.14 ±0.32c 0.38 ±0.01a 8.88 ±0.34c 1.39 ±0.01d 1.07 ±0.02d 0.57 ±0.009c 0.74 ±0.01c
E11.76 ±0.22 g 1.73 ±0.02i 1.54 ±0.04e 14.19 ±0.83f 12.40 ±0.64f 4.79 ±0.24 g 0.35 ±0.005b 7.49 ±0.18e 1.09 ±0.02i 0.88 ±0.01 g 0.54 ±0.006e 0.46 ±0.01 g
F25.95 ±0.71b 3.88 ±0.03a 2.3 ±0.02a 57.9 ±1.85a 46.55 ±1.47a 17.00 ±0.55a 0.29 ±0.004c 10.57 ±0.35b 1.65 ±0.11b 1.25 ±0.01b 0.58 ±0.006c 1.08 ±0.01b
G28.24 ±0.92a 3.06 ±0.04b 1.76 ±0.02b 39.8 ±1.82b 31.27 ±1.36b 12.18 ±0.78b 0.31 ±0.02c 8.9 ±0.32c 1.75 ±0.02a 1.54 ±0.01a 0.57 ±0.007c 1.21 ±0.03a
H12.5 ±037e 1.53 ±0.07j 1.67 ±0.02c 15.22 ±0.84e 12.81 ±0.57e 4.22 ±0.27 h 0.3 ±0.01c 5.32 ±0.19f 1.24 ±0.01f 1.1 ±0.02c 0.53 ±0.01e 0.56 ±0.01e
I12.4 ±0.27e 2.0 ±0.02e 1.58 ±0.03d 12.32 ±0.31 g 10.47 ±0.26 g 4.16 ±0.18 h 0.34 ±0.01b 7.9 ±0.19d 1.15 ±0.02 h 0.88 ±0.01 g 0.57 ±0.007c 0.50 ±0.02f
J15.62 ±0.3c 2.61 ±0.03c 1.68 ±0.02c 21.99 ±0.35c 18.04 ±0.28c 5.73 ±0.24d 0.26 ±0.007d 10.9 ±0.19a 1.52 ±0.02c 0.92 ±0.02f 0.61 ±0.007b 0.75 ±0.01c
Means followed by the same letter within a column are not significantly different at P\0.05 (Student Newman and Keuls test)
Genet Resour Crop Evol (2011) 58:299–309 303
123
Influence of ecological conditions
on the quantitative descriptors of tamarind
fruits and seeds
The Principal Component Analysis performed on
morphological traits showed that the first two axes
explained 63% of the variation. Table 6shows the
correlation between the axes and quantitative descrip-
tors. The first axis shows a positive link between
some fruit traits (length, width, thickness, fresh mass,
dry mass and pulp mass) and seed traits (the length,
width and mass). The second axis was correlated with
seed thickness only. Moreover, the first axis was
found significantly and positively correlated with the
climatic index (I
M
) of Mangenot whereas it was
negatively correlated with the maximum temperature
(T
max
) and insolation (Ins) (Table 6). This means that
the fruit traits (length, width, thickness, fresh mass,
dry mass and pulp mass) and the seed traits (length,
width and mass) increase with higher I
M
but decline
with higher maximum temperature and insolation.
The other relationships were not significant. Overall,
it can be deduced that fruits from wetter zones (i.e.
the Guineo-Congolian zone) generally had greater
fruit and seed size and mass, whilst fruits from drier
zones (i.e. the Sudanian zone) showed more thin-
shaped and lightweight fruits and seeds.
Modelling pulp yield per fruit
Regression equations were used to build predictive
models for pulp yield (the principal trait of commer-
cial importance) based on fruit mass (Table 7). There
were highly significant and strong relationships
between fruit mass and pulp mass (R
2
=0.795).
However, fruit mass was a stronger predictor of pulp
mass for morphotypes J,D,C, I and A(i.e. higher
estimated regression slopes) than for morphotypes F,
G, E, H and B(0.32 \b
1
\0.47 versus 0.07 \b
1
\0.3; P\0.0001; Table 7).
Discussion
This paper quantifies variation in traditional morpho-
types of tamarind and provides basic knowledge on
the range of variation of several quantitative mor-
phological descriptors within and between locally
identified morphotypes, across ecologically different
sites. Ten morphotypes were recorded using folk
taxonomy. This is consistent with previous studies on
tamarind in India, Thailand and Philippines were
eight to fifty cultivars are differentiated based on fruit
size and degree of sweetness (El-Siddig et al.
2006).These morphotypes may have resulted from
complex genetic inter-crossing processes, but local
people link the differences in pulp taste to habitat
types. For instance, they affirm that sweet fruits are
found in gallery forest while sour fruits in savannah
lands (observations from an ongoing survey).
The quantitative morphological analyses on fruits
and seeds of the 10 identified morphotypes confirmed
the traditional discrimination to be effective. From
the results we can conclude that fruits having a
greater size and mass have a lower pulp/fruit ratio,
despite a significant increase in pulp mass, in general.
This may indicate that for superior morphotypes, the
increase in pulp mass is lower than that of the
remaining part of the fruits (seeds and oaks mass).
The correlations were less evident for the number of
seeds per fruit nonetheless. In fact, the number of
seeds per fruit seemed to result from a trade-off
between fruit length and seed size (e.g. fruits having
greater or lower length may contain either lower or
higher number of seeds depending on seeds size;
personal observation).
Table 4 Correlation between quantitative morphological
descriptors of tamarind fruit and seed and canonical discrimi-
nant axes
Morphological
descriptors
Axis1 (0.66;
P\0.0001)
Axis 2 (0.16;
P\0.0001)
Fruit length 0.989 -0.005
Fruit width 0.907 0.296
Fruit thickness 0.751 0.476
Fruit fresh mass 0.934 0.336
Fruit dry mass 0.928 0.356
Pulp mass 0.937 0.290
Ratio-pulp/fruit -0.417 -0.278
Number of seeds per fruit 0.524 0.268
Seed length 0.913 -0.047
Seed width 0.898 -0.269
Seed thickness 0.159 -0.147
Seed mass 0.975 -0.069
In brackets are the percentages of variation explained by the
axes and statistical significances
304 Genet Resour Crop Evol (2011) 58:299–309
123
Despite the confirmation of the traditional dis-
crimination, the statistical analysis revealed that most
of the variability of morphological traits of fruits and
seeds is present within the morphotypes. This
suggests a significant heterogeneity within fruits
traditionally classified as belonging to the same
morphotypes. To get a more powerful morphological
discrimination, quantitative descriptors should hence
be combined with locally perceived qualitative traits
(pulp taste and color). The very extensive variation
found irrespective of the descriptors is consistent with
previous studies on tamarind (El-Siddig et al. 2006)
Fig. 1 Canonical
discriminant analysis to
reveal differences between
morphotypes and links
between descriptors–
legend: A,B,C,D,E,F,G,
H,Iand Jare the different
morphotypes–can1 =first
canonical axis,
can2 =second canonical
axis
Genet Resour Crop Evol (2011) 58:299–309 305
123
and other indigenous fruit trees such as Detarium
microcarpum Guill. and Perr. (Kouyate
´and Van
Damme 2002), Irvingia gabonensis (Aubry-Lecomte
ex O’Rorke) Baill. ex Lanen and Dacryodes edulis
(G. Don) H.J. Lam (Leakey et al. 2004), Adansonia
digitata L. (Assogbadjo et al. 2006,2008,2009;
Kyndt et al. 2009), Vitellaria paradoxa C.F. Gaertn.
(Sanou et al. 2006) and Canarium indicum L.
(Leakey et al. 2008).
Variation in fruit size and in number of seeds per
fruit was found to be significantly affected by cross
pollination and resource availability for T. indica
(Thimmaraju et al. 1989; Diallo et al. 2008).
According to these authors, self-incompatibility in
self-pollinated flowers and resources limitation
(which imposes a sorting by tamarind trees) may
reduce fruit size and the number of seeds per fruit.
The partial link with resource limitations is mirrored
by the pattern of correlations found in this study
between some ecological factors and some pinpointed
morphological traits. For instance, the positive link
between fruit traits (length, width, thickness, fresh
mass, dry mass and pulp mass) and seed traits (the
length, width and mass) on one hand and ecological
factors such as the climatic index of Mangenot on the
other hand, and their opposite link with maximum
temperature and insolation suggest that fruit and seed
size and mass of tamarind trees tend to increase with
humidity (i.e. higher climatic index) and decline with
aridity (i.e. higher maximum temperature).
Phenotypic plasticity was found in several species
of tropical and temperate trees for many traits,
usually in response to changes in ecological condi-
tions (Heaton et al. 1999) and biogeographic history
of individual species (Schlichting and Pigliucci
1998). Nevertheless, ecological differences may only
partly explain the observed variations, the remaining
part being driven by genetic variation.
The high variability indicates great potential for
further improvement through the development of
cultivars from elite trees using horticultural tech-
niques (Leakey et al. 2008). Speedy benefits may be
obtained by selecting superior morphotypes and
propagating such stocks as clones (El-Siddig et al.
2006). Since morphotypes Fand G(see Table 2for
their specific characteristics) portrayed the highest
values for most of the investigated descriptors,
especially pulp mass per fruit, they may be of
particular interest if improvement programs are to be
implemented with the purpose of improving pulp
yield per fruit. As they showed greatest seed mass,
Table 5 Results of the variance components estimation procedure (in percentage) on tamarind fruit and seed traits
Variance
component
Fruit
length
Fruit
width
Fruit
thickness
Fruit
fresh
mass
Fruit
dry
mass
Pulp
mass
Ratio-
pulp/
fruit
Number of
seeds per fruit
Seed
length
Seed
width
Seed
thickness
Seed
mass
Between
morphotypes
17.13 18.28 0.42 26.28 25.46 24.36 15.9 5.34 53.09 43.9 2.7 46.69
Within
morphotypes
82.87 81.72 99.58 73.72 74.54 75.64 84.1 94.66 46.91 56.1 97.3 53.31
Error 5.71 0.03 0.01 14.29 9.89 3.06 0.01 2.69 0.005 0.01 0.002 0.004
Table 6 Correlation between quantitative morphological
traits, ecological factors and PCA factors
Axis1 Axis 2
Morphological descriptors
Fruit length 0.874 -0.320
Fruit width 0.828 0.155
Fruit thickness 0.684 0.384
Fruit mass 0.960 -0.020
Fruit dry mass 0.959 -0.043
Fruit mass 0.894 -0.084
Ratio-pulp/fruit -0.094 -0.145
Number of seeds per fruit 0.402 -0.257
Seed length 0.843 0.256
Seed width 0.692 -0.375
Seed thickness 0.181 0.649
Seed mass 0.887 0.155
Ecological factors
T
min
0.309 -0.016
T
max
-0.706 *** -0.027
Ins -0.590 *** -0.177
I
M
0.761 *** 0.153
Significance: *** P\0.001
306 Genet Resour Crop Evol (2011) 58:299–309
123
they may also portray higher germination, seedling
growth and survival performances (Khan 2004).
Thus, they are potential candidates to be used as
root stock onto which cultivars can be grafted. Other
morphotypes like C,D,E,Hand J(see Table 2for
their specific characteristics) showed intermediate or
low fruits and seeds size and mass but high pulp/fruit
ratio and sweet pulp and hence may also be of great
interest as far as improving the ratio pulp/fruit and
pulp taste is concerned.
The relatively strong relationships between fruit
mass and pulp mass suggested by the predictive
models indicate that selection for pulp can be based
on fruit mass. The variability of the relationship
between fruit mass and pulp mass confirms the
differences between morphotypes and may have been
driven by both ecological and genetic variation. Thus,
further use of the obtained models should be made
with respect to the morphotypes.
Practical conservation measures are to be taken to
preserve genetic diversity and maintain multiple
specimens. This study indicates that based on the
quantitative descriptors, most of the variation is held
within morphotypes. Nevertheless, the between mor-
photype variation was found to relatively high,
particularly for fruit mass, pulp mass and seed mass.
In addition, the perceived qualitative variation may
be genetically determined and should not be
neglected. Thus, pending further genetic finger-
printings, one possible strategy for germplasm col-
lection may consist of sampling a moderate number
of trees within all the morphotypes. This may ensure
capturing a wide range of variation. Because
phenotypic variability results from both ecological
and genetic effects, studies of genetic diversity and
gene flow among ecological zones are needed to
explain all the observed variation prior to effective
germplasm collection, ‘‘plus trees’’ selection and
propagation in traditional agroforestry systems.
Conclusion
This study has highlighted preliminary required
information for tamarind further improvement based
on natural individuals. It demonstrates opportunities
to select wild ‘‘plus trees’’ for pulp production to
meet the needs of traditional and modern markets.
The developed predictive models could allow
researchers and policy makers in partnership with
local people to make quantitative assessment of the
pulp yield potential of tamarind trees established in
traditional agroforestry systems. However, further
endeavours on phenotypic and genetic diversity in the
species are required and much larger populations
should be examined to build effective management
strategies for its genetic resources. If the domestica-
tion of tamarind is to be implemented, more
comprehensive researches extend to other sub-Saharan
countries (Burkina Faso, Mali, Togo, Ghana, etc.)
will be needed. Since traits for nutritional values
should be taken into account in selection processes,
further evaluations of organoleptic traits are also
needed. Studies on provenance variation in germina-
tion and seedling growth dynamics are as well
required to identify best provenances to be used as
Table 7 Linear regression model for T. indica pulp yield per fruit
Morphotypes Model FAdjusted R
2
P
APulp mass =0.322 (Fruit mass) 26.38 0.795 \0.0001
BPulp mass =-0.198 ?0.07 (Fruit mass)
CPulp mass =0.416 (Fruit mass)
DPulp mass =0.462 (Fruit mass)
EPulp mass =0.883 ?0.275 (Fruit mass)
FPulp mass =0.280 (Fruit mass)
GPulp mass =0.238 (Fruit mass)
HPulp mass =1.700 ?0.166 (Fruit mass)
IPulp mass =0.375 (Fruit mass)
JPulp mass =-4.700 ?0.464 (Fruit mass)
The independent variables are fruit mass and morphotype (here used as a dummy variable)
Genet Resour Crop Evol (2011) 58:299–309 307
123
rootstocks on which selected cultivars will be grafted.
As far as the aforementioned further research steps
are concerned, African research programs can benefit
from the experience and results of Asian research
teams on the species.
Acknowledgments This work is supported by Domestication
and Development of Baobab and Tamarind (DADOBAT-EU
funded project). We are particularly grateful to Orou G. Gaoue
and Jean TC Codjia for guidance and comments on an earlier
version. We are indebted to local women for their substantial
contribution to this work especially during morphotypes
identification. Elisabeth Agadja, Alice Bonou and Gbaguidi
Marie-Ange are gratefully acknowledged for their advice and
warm-hearted help without which we could not have made
efficient surveys. We are indebted to anonymous reviewers
whose comments have improved this paper.
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... So, it is recommended that selection of elite trees should not be based only on morphological traits, but also nutritive traits should be given priority. Many recent studies have documented variation in fruit traits in fruit tree species (Abasse et al., 2011;Assogbadjo et al., 2011;Fandohan et al., 2011;Gouwakinnou et al., 2011). In Bangladesh, very little information is available on morphological and nutritional traits for many wild fruits species. ...
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SUMMARY This study obtained quantitative data on fruit and nut traits from two indigenous fruit trees in West Africa (Irvingia gabonensis and Dacryodes edulis), which have led to the identification of trees meeting ideotypes based on multiple morphological, quality and food property traits desirable in putative cultivars. The same data also indicates changes in population structure that provide pointers to the level of domestication already achieved by subsistence farmers. D. edulis represents 21-57% of all fruit trees in farmers' fields and plays an important part in the economy of rural communities. An investigation of the socio-economic and biophysical constraints to indigenous tree cultivation found that indigenous fruits could play an even greater role in the rural economy of West and Central Africa. The opportunity to build on this through further domestication of these species is considerable, especially as retailers recognise customer preferences for certain D. edulis fruit traits, although at present the wholesale market does not. This project was linked to a larger participatory tree domestication programme within ICRAF's 2 wider agroforestry programme with traditionally valuable indigenous trees. Together these projects provided insights into the value of domesticating indigenous fruit trees, which are of strategic importance to poverty alleviation and sustainable development worldwide.
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1. Introduction 2. Estimation 3. Hypothesis testing 4. Graphical exploration of data 5. Correlation and regression 6. Multiple regression and correlation 7. Design and power analysis 8. Comparing groups or treatments - analysis of variance 9. Multifactor analysis of variance 10. Randomized blocks and simple repeated measures: unreplicated two-factor designs 11. Split plot and repeated measures designs: partly nested anovas 12. Analysis of covariance 13. Generalized linear models and logistic regression 14. Analyzing frequencies 15. Introduction to multivariate analyses 16. Multivariate analysis of variance and discriminant analysis 17. Principal components and correspondence analysis 18. Multidimensional scaling and cluster analysis 19. Presentation of results.
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