JBI347.

fm Page 141 Tuesday, April 25, 2000 1:27 PM

Journal of Biogeography, 27, 141–157

Structure, floristic composition and natural

Blackwell Science, Ltd

regeneration in the forests of Cat Tien National

Park, Vietnam: an analysis of the successional
trends
L. Blanc*, G. Maury-Lechon and J.-P. Pascal Laboratoire de Biométrie et Biologie Evolutive,
UMR CNRS 5558, Université Claude Bernard, Lyon 1, 43 Bd du 11 novembre 1918,
69622 Villeurbanne Cedex—France. E-mail: gemaml@biomserv.univ-lyon1.fr

Abstract
The forests in Cat Tien National Park, appear as a mosaic of different communities, distinct
from each other with respect to their floristic and structural parameters. The objectives
of this study are (1) to characterize the different formations occurring in the lowland part
and (2) to identify the main successional trends in the area. Understanding forest succession
is important for silviculture and restoration of forests and land rehabilitation, as adequate
information on the ecological role of local species in the functioning of the forests is not
available in Vietnam. Five plots (1 ha each) were established in the lowland part of Cat
Tien National Park, where all the trees ≥ 10 cm d.b.h. (diameter at breast height) were
located, measured and identified. A systematic sampling was made to assess the regener-
ation. Three plots (A, C and D) can be considered as secondary forests on the basis of
their structural parameters. Plots A and C are dominated by Lagerstrœmia calyculata and
plot D by Dipterocarpus alatus. The other two plots can be regarded as mature forests.
Plot B corresponds to a semideciduous formation dominated by Lagerstrœmia calyculata
and Fabaceae species, and plot E to an evergreen one dominated by dipterocarp species.
The floristic composition of plots A and C will change in the future because dominant
canopy species are rare or absent in regeneration. A correspondence analysis performed
on the number of trees per species shows two kinds of successional trends: one from A
to B on shallow and drier soils, and another from C to E on deeper and wetter soils.

Keywords
Secondary forests, semideciduous forests, Vietnam, diversity, reciprocal ordination,
Lagerstrœmia calyculata, Dipterocarpaceae.

Nomenclature: Leconte (1905–52) and Ho (1991–93).
INTRODUCTION
Vietnamese forests, especially the lowland dense forests in
the southern part of the country, suffered severe damage
during the last century (Durand, 1994). The causes were
mainly land clearing for farming, collection of forest products
by the local population, over-logging and use of herbicides
during the last war that degraded a large number of forests,
especially mangroves (Tschirley, 1969; Orians & Pfeiffer,
1970; Boffey, 1971; Norman, 1974). At present, only 17%
*Correspondence: Laboratoire de Biométrie et Biologie Evolutive, UMR CNRS
5558, Université Claude Bernard, Lyon 1, 43 Bd du 11 novembre 1918,
69622 Villeurbanne Cedex–France, E-mail: gemaml@biomserv.univ-lyon1.fr
(5.5 million hectares) of Vietnam’s total area is under forest
cover (Collins et al., 1991) whereas about 44% was forested
in 1944. Depletion of forest is considered to be one of the
most important environmental threats in Vietnam, chal-
lenging the economic development of the rural areas. It is
estimated that more than 6 million hectares have to be
reforested in Vietnam (Mai, 1983) and hence there is an
urgent need for rehabilitation of land and restoration of
forests as in many other Asian countries (Lamb, 1994).
Instead of planting exotic species, future programmes
should emphasize the use of local species as mentioned in
the National Plan for Environment and Sustainable Develop-
ment (SRV et al., 1991). The aim is to recreate an ecosystem
which would satisfy the basic forestry needs of rural popula-
tions. The priority therefore is to study forest ecosystems in
order to understand their dynamics, especially the processes

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142 L. Blanc, G. Maury-Lechon and J.-P. Pascal
of secondary succession. This basic knowledge is necessary
for reforestation programmes as well as to manage the exist-
ing secondary forests.
Very little quantitative data is available for the south viet-
namese lowlands forests. Some studies have given a general
floristic description of these forests (Maurand, 1943; Trung,
1966; Schmid, 1974) and of similar formations (Vidal, 1960
for Laos; Rollet, 1972 for Cambodia). However, there is no
information on descriptive parameters such as density, basal
area, floristic richness, diversity, etc., and very little data is
available on the dynamics of these forests (Rollet, 1960).
According to Trung (1966) and Schmid (1974), natural lowland
forest is a closed deciduous tropical forest dominated by L.
calyculata associated with Dipterocarpaceae (D. turbinatus,
D. dyeri) and Fabaceae. They estimated that Lagerstrœmia-
dominated stands belong to ‘climax forests’ whereas Rollet
(1960) regarded them as secondary forests. He considered
this formation as the probable result of shifting cultivation.
It is then necessary to clearly describe the stages of the forests
to distinguish secondary from mature forests. Corlett (1994)
defined secondary forest as forest regenerating after complete
clearing. In this study, we consider, as Brown & Lugo (1990),
‘secondary forests as those formed as a consequence of
human impact’. When structural and floristic parameters of
the forests do not reflect past degradations, we call it mature
forests, as Hartshorn (1980).
These works include the Nam Cat Tien area now clas-
sified as Cat Tien National Park where the current studies
were carried out. Satellite data analyses (De Cauwer & De
Wulf, 1994) showed that the forests of Cat Tien National
Park appear like a mosaic of forest formations more or less
dominated by Lagerstrœmia species, confirming the field
observations of Vandekerkhove et al. (1993).
The objectives of this study are to (1) describe the main
forest types of Cat Tien National Park with structural and
floristic parameters; (2) focus on the distinction between
secondary and mature forests; (3) study the floristic links
between these types of forests.
MATERIALS AND METHODS
Study area
Nam Cat Tien Reserve Forest is located in the southern part
of Vietnam, approximately 130 km north-east of Hochiminh
City (Dong Nai Province, Fig. 1). It was created in 1978 and
became a National Park in 1992. It lies at the foot of the
central Vietnamese highlands (11°20′50″–11°32′13″ N and
107°11′13″–107°28′20″ E) and covers an area of 40,000
hectares. The few scientific studies made in the park were
limited to inventories of flora and fauna (Schmid, 1974;
Boulbet, 1960; Vandekerkhove et al., 1993; De Cauwer
& De Wulf, 1994).
The park is subjected to a tropical monsoon climate with
two distinct seasons: a rainy season from April to November
and a dry season from December to March. The mean annual
rainfall is 2450 mm. August and September are the most
rainy months in the year. The temperature amplitude is very
low, varying from 24 to 29 °C, and the mean annual temper-
ature is 25.4 °C.
Two major topographical areas can be distinguished
(Fig. 1). (a) The eastern part, delimited by Dong Nai river, is
a low and flat area with altitude not exceeding 150 m. All
the plots studied were established in this area. (b) The west-
ern part is a hilly area with altitude ranging from 150 to
375 m. These two parts are separated by a large swampy
area with lakes at elevations below 130 m.
The bedrock is mainly basalt with stony outcrops. Very
fertile, black ferrallitic soils developed on this bedrock. They
mostly occur in the low eastern part of the park. In the other
parts, impoverished and poorly drained grey soils developed
on schist. Along the Dong Nai river, the soil is made of dif-
ferent layers of sediments.
Data collection
Field work was carried out from January 1995 to November
1996. Five plots were established in the forest mosaic, each
one different from the other with respect to their floristic
and structural parameters. Four plots (A, B, C and E) were
on basaltic soils and the fifth (plot D) on schist. A detailed
vegetation map of this area is not available except for the
satellite data analysis of De Cauwer & De Wulf (1994)
which has, however, not been fully verified in the field. Plots
were selected from aerial photos and after discussion with
Vietnamese foresters and researchers. Each plot covers 1 ha,
divided into small quadrats of 10 × 10 m. All the trees with
a diameter ≥ 10 cm d.b.h. (d.b.h.: diameter at breast height
or at 1.3 m) were numbered, measured, plotted on a map
and identified. The trees were classified into two groups
(depending on the potential maximum height of the species):
canopy and emergent species more than 25 m (G1) and under-
storey species less than 25 m (G2).
Regeneration was studied by systematic sampling to com-
pare floristic composition with adult trees. Sub-quadrats (4 ×
4 m) were established in the middle of alternate 10 × 10 m
quadrats, accounting for 8% of the plot area. All saplings ≥
2 m high with d.b.h. < 10 cm were counted, identified and
classified under G1 or G2. Species which can’t reach 10 cm
diameter even at maturity were grouped under G3. Trees
were considered as adult when they attained the critical size
to produce fruits. If not they belong to natural regeneration.
For canopy species (G1), we considered that trees are adult
when they are ≥ 20 cm d.b.h. For species belonging to G2,
all trees ≥ 10 cm d.b.h. were considered to be adult.
Soil humidity was measured in plots A, B, C and E (four
borings for each plot and four soil samples per boring). The
ratio between the weight of the sample before and after
drying (48 h in an oven at 100°C) was taken as the humid-
ity rate. The soil was measured down to 2 m depth or down
to the mother rock if found before (twenty-five borings for
each plot).
A herbarium was created with specimens of trees and
saplings collected systematically from the plots. Except for
a few rare species, all the other specimens were identified up
to the species level.
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Forest typology in Cat Tien National Park 143

Figure 1 Map of the Nam Cat Tien area (Cat Tien National Park) and location of the five 1 ha plots.

Data analysis where S is the species richness of the population, Sobs the
number of species observed in the sample, n the sample size
As a first approach, Jaccard’s Index of similarity was used to
and ni the abundance of species i in the sample.
compare vegetation types. I = 100*c/(ux + uy + c) where c is
Hurlbert’s species richness curve gives the expected number
the number of species common to both plots X and Y; and
of species E(Sx) in a sample of x individuals selected at
Ux and Uy, the number of species found only in plots X and
random (without replacement) from a plot containing n
Y, respectively.
individuals and S species (Hurlbert, 1971). The species
Differences in species diversity in the forest stands were
richness of different plots for a given number of species can
assessed using two indices, derived from shannon-Wiener
thus be compared.
species diversity index and Simpson’s (Gimaret-Carpentier
et al., 1998). Their estimators are:
n – ni
S  x 
E(Sx) = ∑ 1 – ----------------
Sobs
Ŝ
ni ni
 --- ni ( ni – 1 )
ˆ ′ = –∑ --- + Ŝ----------
– 1- and D̂ = 1 − λ with λ =
∑ --------------------
-
H n-log -
2 n  2n n(n – 1)
-
i=1 n
i=1 i=1 x
n 3
n
with a theoretical variance Var(D̂) = 4 ∑---n-i
2
--n- –λ An index derived from the Importance Value Index (Curtis
i=1 & McIntosh, 1950) was used to study the floristic structure.

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144 L. Blanc, G. Maury-Lechon and J.-P. Pascal
Figure 2 Relationship between soil humidity and depth (mean and
confidence interval) in plots A, B, C and E.
Like Pascal & Pelissier (1995), we also consider IVIi as the
sum of the relative density Di and relative basal area Gi for
species i in one plot. The sum of all the species in one plot is
equal to 200.
IVIi = Di + Gi,
Di = (ni/n)·100,
Gi = (gi/g)·100.
Correspondence analysis and reciprocal scaling were per-
formed with ADE.4 software (Thioulouse et al., 1997).
Reciprocal scaling was performed from a correspondence
analysis of the species (lines) × diameter class (columns)
table. This statistical technique allows a joint display of spe-
cies and diameter classes. From the coordinates of diameter
classes ordered on a factorial axis seen as a gradient, the
conditional mean and variance of species can be calculated.
Variance of the means is maximized. The conditional means and
variances of diameter classes can be obtained reciprocally
(Thioulouse & Chessel, 1992). The range of diameter classes
for each species can thus be derived.
RESULTS
Soil property
The soils of plots A, B, C and E are developed on basalt.
They all have good chemical properties (P. Deturck, pers.
comm.) but differ physically. The soil is shallow, less than
50 cm deep, in A and B, and deeper, around 1 m, in C and E
(Fig. 2). Using Bonferroni Test, significant differences are
found between A and C (P = 0.0027), A and E (P = 0.0008),
B and C (P = 0.0011) and B and E (P = 0.0003). Soil humid-
ity at the end of the dry season is also significantly different
between A and C (P = 0.0001), B and C (P < 0.0001) and B
and E (P = 0.004). Root system is superficial in shallow soils
inducing more frequent tree falls resulting in opening the
canopy. Plot D has poor grey soil developed on schist, poorly
drained and flooded during several months in the year.
Stand structure
The diameter class distributions for all the trees with a d.b.h.
≥ 10 cm in each plot are shown in Fig. 3. The distribution
pattern of plots B and E is a negative exponential distribution
or reverse ‘J-shaped’ curve, typical of mature forests (Rollet,
1978). Compared to this, the distribution patterns of plots
A, C and D indicate a disturbed forest. Plots A and C show also
a negative exponential distribution but with a high frequency
of trees in medium diameter class for A and a very high fre-
quency of trees in the first diameter class as well as in the last
(diameter ≥ 110 cm) for plot C. Diametric distribution for plot
D is very different from the four others with the highest fre-
quencies for the medium class, which is characteristic of stands
with poor regeneration (Knight, 1975). Density and basal area
are given in Table 1. Density in plot D is much lower (195 trees/ ha)
than in the other four plots (389–540 t/ ha). This plot appears
like a very disturbed forest. The basal area of plots A and C is
high: 69.41 and 54.89 m2/ha, respectively, whereas it is around
31 m2/ha for the other three plots. In plot C, the twenty
trees belonging to the last diameter class (3.7% of the total
number of trees) account for 71.4% of the total basal area.
Figure 3 Diameter class distribution of trees
≥ 10 cm d.b.h. for the five plots.
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Forest typology in Cat Tien National Park 145

Table 1 Density, basal area, percentage of deciduous species and Table 3 Jaccard’s index of similarity for the five plots (trees with
trees, diversity indices and species richness for all the trees ≥ 10 cm d.b.h. d.b.h. ≥ 10 cm).
in the five plots.
Plots A B C D E
Plots A B C D E
A — 32.3 26.8 0 32.5
Density (trees/ ha) 419 389 540 195 469 B — 22.3 2.7 27.8
Basal area (m2/ ha) 69.41 31.71 54.89 29.30 31.33 C — 0 33.3
Deciduous species (%) 26 35 13 11 10 D — 4.2
Deciduous trees (%) 47 27 17 1 6 E —
Simpson’s index 0.84 0.92 0.97 0.51 0.96
Shannon’s index 4.15 4.64 5.62 1.98 5.41
Species richness 70 57 91 18 81
dominated by species of the understorey (G2), except for
Dipterocarpaceae and Combretaceae whose species all
Table 2 Number of species of the main families in the five plots. belong to G1 (see Appendix). Some differences can be
observed in the five plots. Dipterocarpaceae is absent in
Family Nb species A B C D E plots A and C. Fabaceae has six species in plots A and B, but
only three in plot C and one in plots D and E. In contrast,
Rubiaceae 15 7 5 6 — 7 species richness is higher in plots C and E for Lauraceae,
Lauraceae 15 4 2 9 — 6 Euphorbiaceae and Meliaceae. Floristic composition in plot
Fabaceae 12 6 6 3 1 1 D is very different from the other four. It contains four spe-
Euphorbiaceae 11 4 4 8 2 7 cies of Dipterocarpaceae but most of the main families are
Meliaceae 11 3 2 8 — 5
absent (Table 2 and Appendix). Jaccard’s Index of similarity
Anacardiaceae 11 4 3 3 — 5
Ebenaceae 9 4 3 8 — 4
is very low (Table 3) between this plot and B or E, and nil
Annonaceae 9 5 5 7 — 5 for A and C. None of the species is common to all the five
Clusiaceae 8 5 2 2 2 4 plots due to the highly specific composition of plot D. How-
Myrtaceae 6 — 1 1 3 2 ever, sixteen species are common to plots A, B, C and E, four
Verbenaceae 5 1 2 1 1 4 of them represented by a large number of trees: Lagerstrœmia
Sterculiaceae 5 3 2 3 — 2 calyculata (197, Lythraceae), Diospyros longebracteata (151,
Dipterocarpaceae 5 — 2 — 4 3 Ebenaceae), Ochrocarpus siamensis (87, Clusiaceae) and
Oleaceae 4 2 2 3 — 2 Alphonsea philastreana (69, Annonaceae). Jaccard’s Index
Moraceae 4 3 2 4 — 1 ranges for these four plots from 22.3 to 33.3 (Table 3). It is
Combretaceae 4 1 3 1 — 1
highest (over 30%) between A and B, A and E and C and E.
The percentage of deciduous species (and trees) is higher in
plots A and B than in the other three plots (Table 1). Sub-
Floristic composition
stantial difference was also observed in the species richness
In the five plots, trees with d.b.h. ≥ 10 cm belong to 176 which varies from eighteen species in plot D to ninety-one
species. Rubiaceae and Lauraceae have the highest number species in plot C (Table 1). The shape of the Hurlbert’s spe-
of species (Table 2). The most abundant families are mainly cies richness curve (Fig. 4) is similar for plots B, C and E.

Figure 4 Hurlbert’s species richness curve

for the five plots.

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146 L. Blanc, G. Maury-Lechon and J.-P. Pascal
Figure 5 Percentage of Importance Value Index {IVI = Di ( ) + Gi (h)} for the main G1 (j) and G2 (
The slope is steeper for plot A and shallow for plot D. Plots
dominated by evergreen species (C and E) have a higher
species richness than plots dominated by deciduous species
(A and B). However, plot D, although evergreen, has a low
species richness. The same trend is observed for species
diversity (Table 1). For the two pairs of plots, disturbed forests
are richer. But plot A is less diverse than plot B, whereas
diversity in plot C is slightly higher than in plot E in terms of
Simpson’s and Shannon’s indices. Plots A, C and D are strongly
dominated by one species (Fig. 5). IVI of Lagerstrœmia
calyculata is 53% and 39.9%, respectively, in A and C and IVI
of Dipterocarpus alatus accounts for 75.1% in D. The pattern
of species importance is quite different for B and E where quite
a few species have similar IVI values. In B, L. calyculata and
Fabaceae species like Afzelia xylocarpa and Dalbergia mammosa
are the most dominant G1 species mixed with Anogeissus
acuminata and Lagerstrœmia ovalifolia. Plot E contains a mixture
of L. calyculata and Dipterocarpus turbinatus. Dipterocarpus
alatus and Afzelia xylocarpa have only two trees in the plot
but with very big diameter. It should be noted that both, B and
E, are dominated by Cleistanthus sumatranus and Diospyros
longebracteata in the understorey. Another important feature
is that L. calyculata is dominant in all the plots except in D.
However, its density is higher in plot A than in the others
where it is represented by few trees but with larger diameter.
) species (IVI ≥ 2) in the five plots.
Floristic relationships
To study the floristic relationships, a between-plots cor-
respondence analysis was performed on the number of trees
per species and per subplot (20 × 20 m). Only species with
more than five trees in the four plots taken together were
selected. Plot D was not included in the analysis due to its
very distinct floristic composition. The results are given in
Fig. 6. Several points can be formulated.
The floristic composition of the plots A and C is very dis-
tinct and there is no specific floristic link between these two
plots (Fig. 6a,b).
Plots E and B are close on axes 1 and 2 because they share
two very common species: Cleistanthus sumatranus and
Diopsyros longebracteata (Fig. 6b). However, most of their
subplots have a distinct floristic composition and are separ-
ated on axis 3 (Fig. 6c).
There is a clear floristic link between plots C and E with a
succession of species present only or mainly in these two
plots (Fig. 6b).
From Fig. 6(b) (axes 1 and 2), it is not possible to deter-
mine the floristic links between A on one side and B or E on
the other side.
This can be analysed through the third axis (Fig. 6c). The
canopy or emergent species (G1) are positioned between A
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Forest typology in Cat Tien National Park 147

Figure 6 Correspondence analysis performed on the number of trees per species (species with more than five trees in the four plots) and
per subplot (20 × 20 m). The figure shows the results of the between-plot analysis for (a) the subplots; (b) the species on axes 1 and 2; and
(c) the species on axes 1 and 3. Axes 1 and 2 account for 41.5% and 35.3%, respectively, of the total variance of the scatterplot. The observed
dispersion of the centres of gravity of the four plots is highly significant. No higher value was found in 1000 simulations. Species with a high
relative contribution are underlined for the first axis and in bold for the second axis. Species code are given in the Appendix.

and B. They are Afzelia xylocarpa and Dalbergia mammosa,
which are two typical species of B, Tetrameles nudiflora
which is also in C, Beilschmiedia micranthopsis also found
in E and Pterospermum diversifolium also in C and E. Only
Vitex pinnata doesn’t belong to A but to B and E. Other
species belong to the understorey (G2) and occur in all the
four plots: Alphonsea philastreana, Streblus taxoides, Diospyros
longebracteata, Polyalthia luensis, Garcinia nigrolineata,
Linociera pierrei, Antidesma velutinosum and Madhuca pierrei.
Nephelium melliferum and Cleistanthus sumatranus occur in
three plots (A, B and E). Garcinia vilersiana and Dehasia caesia
are the only species found in A and E. The distribution of

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148 L. Blanc, G. Maury-Lechon and J.-P. Pascal

Figure 7 Ordination by reciprocal scaling after a correpondence analysis performed on the number of trees in eleven diameter classes for forty-
four species. The dot (proportional to the number of trees) and the associated line represent the conditional mean and variance, respectively, for
(a) the species of the 5 plots and (b) the diameter classes, on the first axis (inertia = 28.9%). (c) show the diametric class distribution of
Lagerstrœmia calyculata in plot A and Dipterocarpus alatus in plot D. Species code are given in the Appendix.

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Forest typology in Cat Tien National Park 149

Table 4 Estimated density (and confidence interval) of saplings from the systematic sampling (≥ 2 m and < 10 cm d.b.h.) and species richness in
the five plots.

Plots A B C D E

Density
Estimated number 4275 6338 8150 2850 6175
of saplings (per ha) (3646 –4904) (5480 –7195) (7450 –8885) (2393 –3307) (5372– 6978)
Species richness
Total number of species 63 57 111 29 82
% G3 species 52 42 35 59 35
% G3 saplings 60.8 41 36.2 85.5 55.1

the species on axes 1 and 3 thus shows a clear floristic link Table 5 Percentage of deciduous species and species richness of
between plot A and plot B. mature trees and natural regeneration of G1 and G2 species in the
five plots. The trees of G1 species in the first diameter class
(10 ≤ cl1 < 20 cm) are not considered as adult trees and are added to
Stand ordination the data from the systematic sampling only when they were present
in the subquadrats (4 × 4 m). Two trees with d.b.h. < 20 cm were
A correspondence analysis was performed on the number of
added in plots A and B, 3 in plots C and E and none in the plot D.
trees in each diameter class for all species with more than two
individuals. The first axis distinguishes the first two diameter Plots A B C D E
classes from the others. When Lagerstrœmia calyculata of
plot A (152 trees) and Dipterocarpus alatus of plot D Percentage of deciduous trees
(135 trees) are not included in the analysis, the diameter classes G1 adult trees 92 83 53 1 43
are more spread out on the first factorial axis. From the G1 regeneration 50 52 20 1 24
results, a reciprocal scaling was made (Fig. 7) as proposed G2 adult trees 4 3 12 12 0
by Thioulouse & Chessel (1992). (i) Plot A is composed G2 regeneration 2 0 3 8 2
Species richness
of two groups of species. Four species, Afzelia xylocarpa,
G1 adult trees 16 22 15 9 19
Tetrameles nudiflora, Haldina cordifolia and Bursera serrata G2 adult trees 48 33 70 7 57
are only in the higher diameter classes (≥ 50 cm diameter). Total 64 55 85 16 76
Moreover, only few trees of Lagerstrœmia calyculata are in G1 regeneration 5 12 9 5 11
the first three diameter classes (Fig. 7c). These species show G2 regeneration 26 21 66 4 43
very poor regeneration. In contrast, Dalbergia mammosa Total 31 33 75 9 54
and Pterospermum diversifolium are restricted to the smallest
diameter classes indicating an obvious change in the future
floristic composition in this plot. (ii) A discontinuity is also
observed in plot C. Lagerstrœmia calyculata is restricted
to the large diameters (out of a total of twenty-seven trees, larger tree (diameter 200.6 cm). Thus in plots B and E, the
only three belong to the first diameter class and twenty to canopy and emergent species that have attained large diameters
the last one). The majority of species is in the smallest classes. are in general seen to be regenerating and the floristic com-
But for Lagerstrœmia calyculata, this plot appears like a position is therefore unlikely to change.
regenerating area. (iii) Plot D has few small diameter trees
of Dipterocarpus alatus (Fig. 7c). The dominance of this
Natural regeneration
species will dramatically decrease to the benefit of non-
dominant species like Garcinia benthami and Parinari Sapling density (≥ 2 m high and < 10 cm d.b.h.) was estimated
annamensis. (iv) Plots B and E show a more uniform dis- from the fifty subquadrats sampled in each plot. Concerning
tribution with several species present in a large array of density, the same conclusions can be drawn (Table 4) as for
diametric classes including the smallest ones: Pterospermum trees with d.b.h. ≥ 10 cm. It is significantly lower in plots D
diversifolium, Dalbergia mammosa, Sindora siamensis, and higher in plot C. In all the plots, saplings are dominated
Anogeissus acuminata, Lagerstrœmia ovalifolia in B, Dipte- by understorey species (G3) both in the number of species and
rocarpus turbinatus, Lagerstrœmia ovalifolia and Pterosper- individuals (Table 4). In plot D, Colona auriculata (Tiliaceae)
mum diversifolium in E. Only Afzelia xylocarpa in B and accounts for 70.2% of the total number of individuals.
Lagerstrœmia calyculata in E do not have trees in the smallest The comparison of floristic parameters between adult
diameter classes and might disappear in these two plots. In trees and natural regeneration is shown Table 5 and Fig. 8.
B, six trees of Lagerstrœmia calyculata were counted in the The canopy of plots A and B is almost deciduous whereas
first two diameter classes. Three are on one side of the plot the canopy of plots C and E is semideciduous (Table 5). The
which is a part of a large gap and a fourth is a sprout of a frequency of deciduous trees in regeneration is about half

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150 L. Blanc, G. Maury-Lechon and J.-P. Pascal

Figure 8 Simpson’s index of species diversity

for G1 + G2 species (Nx) and for only G1
(G1x) for mature trees (s) and regeneration
(j) for each plot (x = A, B, C, D or E).

Table 6 Floristic composition of regeneration

Plots A B C D E Total of G1 species in the five plots.

Dalbergia mammosa Pierre 1* 2 3

Lagerstrœmia calyculata Kurz. 3* 2 1 6
Garcinia benthami Pierre 4 1 1 6
HCT822 1 1 2
Vitex quinata Williams 1 1 2
Firmania simplex (L.) W. F. Wight 1 1
Mangifera foetida Lour. 1 2** 1
Terminalia calamansanai (Bl.) Rolfe 2* 2
Xylia xylocarpa (Roxb.) Taubert 1 1
Vatica odorata (Griff.) Sym. subsp odorata 3* 3
Lagerstrœmia ovalifolia Teijsin. & Binn. 2 1* 1 4
Garcinia ferrea Pierre 5 2 1 8
Disoxylum loureirii Pierre 3 2 5
Vitex pinnata L. 4 2 6
Cryptocarya ferrea Bl. 1 1
Aphanamixis polystachia J. N. Parker 1* 1
Disoxylum cauliflorum Hiern 1 1
Hibiscus macrophyllus Roxb. ex Horbem 1* 1
Neolamarckia cadamba (Roxb.) Bosser 1 3* 4
Anisoptera costata Korth. 1 1
Callophyllum calaba var. bracte. (Wight) Stevens 1 1
Dipterocarpus alatus Roxb. 2 2
Crypteronia paniculata Bl. var affinis (pl.) Beus 4 4
Beilschmiedia micranthopsis Kost. 1 1
Melanorrhea usitata Wall. 1 1
Pterospermum diversifolium Bl. 1 1
Dipterocarpus turbinatus Gaertn. f. 2 2

*Represents one tree with diameter ≥10 cm (see legend table 5).

that of adult trees in these four plots. Adult trees and regen-
eration of G2 species are almost evergreen. Plot D is ever-
green. The species richness decreases from plot A to B and
from C to E for adult trees (Table 5), but this decrease is
only due to G2 species as species richness of G1 shows an
increase. The same trend is observed for regeneration. Species
diversity of G1 adult trees is significantly lower than that of
their regeneration for plots A, C and D (Fig. 8). For plots B and
E, Simpson’s index values for adult trees and regeneration
are high and not significantly different. In all the five plots,
there is no significant difference (χ2 = 0.20; 8ddl; NS)
between G1 species number represented only by adult trees, or
only in regeneration, or by both adult trees and regeneration.
Table 6 shows the number of saplings measured for regen-
eration in each plot, together with the nonadult (G1) trees
with ≥ 10 cm d.b.h. Regeneration of Lagerstrœmia calyculata

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JBI347.fm Page 151 Tuesday, April 25, 2000 1:27 PM
Figure 9 Stand dominated by Lagerstrœmia
calyculata with bright grey bark (photo L.
Blanc).
is not abundant compared to the number of adult trees in
each of the four plots. It shows very poor regeneration in
the understorey, confirming the results given in Fig. 7(c). We
can conclude that in all the plots where this species is domin-
ant, it tends to disappear (C and E) or decrease drastically
(A and B). In A and C, regeneration is too poor for a clear
trend to be observed (Table 6). In D, the dominant species,
Dipterocarpus alatus, has poor regeneration in contrast to
Crypteronia paniculata. In B, some of the more important
species like Dalbergia mammosa, Terminalia calamansanai,
Vatica odorata, Lagerstrœmia ovalifolia and Vitex pinnata
regenerate well. In E, Mangifera fœtida, Vitex pinnata,
Neolamarckia cadamba and Dipterocarpus turbinatus show
good regeneration. Five regenerating species are common to
both B and E.
DISCUSSION
A dynamic view of forests in southern Vietnam is necessary
because primary forests disappeared almost completely and
were replaced by a mosaic of forest formations disturbed in
varying degrees of several kinds of degradations in the past.
Satellite data analyses (De Cauwer & De Wulf, 1994) showed
that the forests of Cat Tien National Park appear like a mosaic
of forest formations more or less dominated by Lagerstrœmia
species. De Cauwer & De Wulf (1994) distinguished three
forest formations in the lowland area: primary, secondary
and mixed forests but floristic composition are not precise.
Primary forests are found mainly in the middle of the lowland
area, i.e. in the most inaccessible part of the park, and the
other two types are localized along the river.
Our results give some elements to establish a better
typology of the formations in Cat Tien. The mature or
nonmature stage of the plots was analysed from floristic and
structural parameters and from differences in the floristic
© Blackwell Science Ltd 2000, Journal of Biogeography, 27, 141–157
Forest typology in Cat Tien National Park 151
composition between trees and regeneration. In plot A,
diameter class distribution of all the species shows a high
frequency of medium diameters (Fig. 3) which is a reflection
of the distribution of the very dominant species, Lager-
strœmia calyculata (Fig. 7c). This plot appears like a ‘pure
stand of Lagerstrœmia’ (Fig. 9) as defined by Rollet (1960).
The floristic composition of the plot is not stable and will
change in the future with the progressive disappearance of
Lagerstrœmia calyculata and all the other main species
Tetrameles nudiflora, Haldina cordifolia, Afzelia xylocarpa
(deciduous) and Bursera serrata (evergreen) which have no
or poor regeneration. On the other hand, there will be an
increase in Dalbergia mammosa and Garcinia benthami, and
also a slight increase in the total number of evergreen spe-
cies. Plot C appears as a regenerating plot with a diameter
class distribution characterized by a high number of trees
with small diameters (below 20 cm), which have not yet
reached the adult size (Fig. 3). This could be one reason to
explain the relatively small number of regenerating stems.
Lagerstrœmia calyculata, which is dominant among the big
trees, has poor regeneration and will become less dominant.
The regeneration, with a high number of species, indicates
that the stand will evolve towards a more evergreen com-
position. Thus, these two plots are both characterized by the
high dominance of L. calyculata, inducing a low diversity
of G1 species and a high basal area. They have a nonstable
floristic composition and should be regarded as secondary
forests or secondary stages of succession. These two plots
differ however, by their diametric distribution. Plot A has a
high frequency of medium diameters whereas plot C is strongly
dominated by big L. calyculata trees. Data available do not
support the hypothesis that C is a more degraded stand than
A. Furthemore, no successional trends are observed between
A and C in the correspondence analysis. In contrast, plots B
and E can be considered as mature forests from their diameter
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152 L. Blanc, G. Maury-Lechon and J.-P. Pascal

deciduous trees). Plot E occurs on deeper soil (like C) and

Figure 10 Large tree of Lagertsrœmia calyculata (photo L. Blanc).
distribution, basal area, and also because most of the G1
species have reached large diameters and regenerated well.
Both these plots occur on basaltic soils, but with different
depths and humidity. Plot B, which is on shallow soil (like
A) is dominated by L. calyculata and species of Fabaceae. It
can be described as a semideciduous stand (with 27% of
is an almost evergreen forest dominated by dipterocarp
species, mainly Dipterocarpus turbinatus, and a few L.
calyculata. This evergreen nature will be accentuated in the
future as L. calyculata is not regenerating.
The correspondence analysis based on the floristic com-
position clearly shows a floristic link between A and B on one
hand, and C and E on the other, confirming the distribution
of the two pairs of plots on two different soil conditions.
Both Jaccard’s index and correspondence analysis show that
plots B and E share a pool of common species which seem to
be not affected by the edaphic differences. Others species,
more sensitive to the depth and the humidity of these soils,
make the difference in the floristic composition and pheno-
logical nature (deciduous/evergreen) between these two forma-
tions. Thus from our results, we can conclude that (i) two
mature formations are present in the studied area of Cat
Tien: one semideciduous, dominated by Lagerstrœmia caly-
culata and Fabaceae species (plot B) and another, almost
evergreen, dominated by dipterocarp species (plot E); (ii)
two successional trends can be discerned from A to B and
from C to E; (iii) as suggested by Jaccard’s index of similarity
and the pure evergreen nature of the stand, plot D appears
to be a third formation dominated by Dipterocarpus alatus
whose successional relationships are not easy to predict.
Lagerstrœmia calyculata seems to be a key species in the
succession. We agree with Rollet (1960) who considered
Lagerstrœmia-dominated stands to be secondary forests.
This species can be considered as a secondary species with a
long life span and large diameters (Fig. 10), not typical of
mature forests. Its regeneration is very poor and limited to
large openings. It appears to be a very good competitive spe-
cies able to regenerate on denuded areas: along roads and
on land abandoned after cultivation. Human disturbances
have mostly affected dipterocarps for resin and Fabaceae for
their wood, thus reducing the number of trees of these
species and creating favourable conditions for regeneration of
Lagerstrœmia calyculata. In plot B, Lagerstrœmia calyculata
is still maintained. One hypothesis is that treefalls are more
frequent on this shallow soil, due to poor rooting, creating
gaps which allow the regeneration of this secondary species
and its occurrence as rare individuals in mature forests.

Table 7 Comparison of mean density and basal area in some tropical forests of Asia.

Minimum Mean density Mean basal

Location gbh (cm) (ha-1) area (m2.ha-1) References

Uppangala, India 30 625 39.7 Pascal & Pelissier (1995)

Danum Valley, Sabah 30 470 26.6 Newbery et al. (1992)
Pasoh, Malaysia 31.4 530 25.2 Kochumen, Lafrankie & Manokaran (1990),
Manokaran & Lafrankie (1990)
Sungei-Menyala, Malysia 31.4 493 32.4 Manokaran & Kochumen (1987)
Mulu, Sarawak 31.4 739 57.0 Proctor et al. (1983)
Cat Tien Vietnam, Plot E 31.4 469 31.3 Present study
Cat Tien Vietnam, Plot B 31.4 389 31.7 Present study
Xishuangbanna, China 31.4 386 30 Cao et al. (1996)

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JBI347.fm Page 153 Tuesday, April 25, 2000 1:27 PM
Rollet (1960) observed that Lagerstrœmia and Fabaceae
forest type has maximum stability on rocky black basaltic
slopes. This edaphic condition favours treefalls. The main-
tenance of this forest type could be the result of this fre-
quent natural disturbance. Table 7 shows that the density
and basal area of the two mature formations (plots E, ever-
green, and B, semideciduous) are quite comparable to other
forests in Asia.
The next step will be to focus on the biology of the main
species involved in the successional processes and coloniza-
tion of the open areas. As this region is in urgent need of
land rehabilitation, it is necessary to obtain this data at the
earliest.
ACKNOWLEDGMENTS
The authors are grateful to the Programme Environnement,
Vie et Sociétés, of CNRS for supporting the project and to
the board of Cat Tien National Park for allowing the
research work. They also thank Nguyen Manh Sum,
Nguyen Tran Vy, Luu Huong Truong, Laurence Curtet,
Bruno Dutreve and students from the Department of Bot-
any and Ecology, University of Ho Chi Minh City for field
assistance; Prof. Le Cong Kiet, Vu Van Bien, Nguyen Thien
Thich for their help in species identification; Drs My Hanh
Diep and Pol Deturck for soils description and analyses and
S. Dolédec for data analyses. The authors thank also two
anonymous reviewers for useful critiques.
L. Blanc received a grant from the Ministère de l’Enseignement
Supérieur et de la Recherche and additional financial support
from the Rhône-Alpes Region.
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BIOSKETCHES
Lilian Blanc is a tropical forest ecologist. He spent 2 years in
Vietnam (1995–6) on fieldwork for his Ph.D. thesis. He
graduated in July 1998.
Géma Maury-Lechon and Jean-Pierre Pascal are researchers
at CNRS (Centre National de la Rechereche Scientifique) and
have worked in Asia for more than 20 years. Géma Maury-
Lechon is a dipterocarp specialist (phylogeny and forest dynamics)
and has collaborated mainly with Malaysia and Vietnam. She
is the leader of a reforestation project in Cat Tien National Park.
Jean-Pierre Pascal is head of the research team ‘nodelisation
and dynamics of tropical rain forest at the laboratory UMR
CNRS 5558. For 10 years he was the director of the French
Institute of Pondichery (India).

Appendix IVI values (in %) of tree species in the five plots established in Cat Tien National Park. The unidentified species are named after
their reference number in the herbarium (HCT xxx).

Plots

Family Species Code G A B C D E

Anacardiaceae Allospondias lakonensis (Pierre) Stapf. Allak 2 0.34 1.52 0.36

Bouea poilanei Evr. Boupoi 1 0.97
HCT575 Mangifera Hct575 2 0.27
Mangifera dongnaiensis Pierre Mandon 1 0.53
Mangifera duperreana Pierre Mandup 1 1.15
Mangifera fœtida Lour. Manfoe 1 0.59 3.41
HCT103 Hct103 1 0.42
Melanorrhoea usitata Wall. Melusi 1 0.26 0.25
Semecarpus graciliflora Evr. & Tard. Semgra 2 0.25
Semecarpus reticulata Lec. Semret 2 0.23
Spondias pinnata (Koenig & L. f.) Kurz Spopin 1 2.03
Annonaceae Alphonsea philastreana (Pierre) Fin. & Gagn. Alpphi 2 4.83 7.09 3.63 2.60
Cananga latifolia (Hook. f. & Thoms.) Fin. & Gagn. Canlat 1 0.41 3.48
HCT821 Polyalthia Hct821 2 0.41
HCT899 Polyalthia Hct899 2 0.91 0.20
Mitrephora thorelii Pierre Mittho 2 0.25 0.45 1.51 3.26
Polyalthia corticosa (Pierre) Fin. & Gagn. Polcor 2 1.66
Polyalthia luensis (Pierre) Fin. & Gagn. Pollue 2 0.54 0.69 0.25 0.53
Sageraea elliptica (A. DC.) Hook. & Thoms. Sagell 2 0.25 4.60 1.08
Xylopia vielana Pierre ex Fin. & Gagn. Xylvie 2 0.57
Apocynaceae Alstonia scholaris (L.) R. Br. Alssch 1 0.79 2.04 0.24
Kibatalia laurifolia (Ridl.) Woods Kiblau 2 1.26 0.24
HCT050 Hct050 2 0.36
Aquifoliaceae Ilex cochinchinensis (Lour.) Loesen. Ilecoc 2 0.25 7.60 2.73
Bignoniaceae Markhamia stipulata var. pierrei (Dop) Sant. Marsti 1 0.45 0.40
Stereospermum colais (Dillw.) Mabb. Stecol 1 2.02 1.31
Bombacaceae Bombax insignis Wall. Bomins 2 0.32
Bursera serrata Wall. ex Colebr. Burser 1 5.31 2.77
Celastraceae Lophopetalum wightianum Arn. Lopwig 1 14.80
Clusiaceae Callophyllum calaba var. bracteatum (Wight) Stevens Calcal 1 1.96
Garcinia benthami Pierre Garben 1 2.78
Garcinia ferrea Pierre Garfer 1 0.28
Garcinia nigrolineata Pl. ex T. And. Garnig 2 2.10 1.04 0.40 2.13
Garcinia vilersiana Pierre Garvil 2 0.78 0.79

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Forest typology in Cat Tien National Park 155

Appendix continued

Plots

Family Species Code G A B C D E

HCT397 Garcinia Hct397 2 0.51

HCT535 Garcinia Hct535 2 0.55
Ochrocarpus siamensis T. Anders. Ochsia 2 17.13 3.26 2.22 4.55
Combretaceae Anogeissus acuminata (DC.) Guill. & Perr. Anoacu 1 9.94
Terminalia calamansanai (Bl.) Rolfe Tercal 1 0.57 4.23
Terminalia corticosa Pierre ex Lan. Tercor 1 1.45 1.39
Terminalia pierrei Gagn. Terpie 1 1.48
Crypteroniaceae Crypteronia paniculata Bl. var. affinis (Pl.) Beus. Crypan 1 1.78
Cycadaceae Cycas circinalis L. Cyccir 2 0.25 0.61 3.11
Datiscaceae Tetrameles nudiflora R. Br. Tetnud 1 2.19 1.27 0.43
Dilleniaceae Dillenia hookeri Pierre Dilhoo 2 0.20
Dipterocarpaceae Anisoptera costata Korth. Anicos 1 2.14
Dipterocarpus alatus Roxb. Dipala 1 150.2 6.83
Dipterocarpus turbinatus Gaertn. f. Diptur 1 10.70
Hopea odoata Roxb. Hopodo 1 0.43 1.64 2.16
Vatica odoata (Griff.) Sym. subsp odoata Vatodo 1 2.60 0.59
Ebenaceae Diospyros castanea (Craib.) Fletcher Diocas 2 0.30 1.88
Diospyros filipendula Pierre ex Lec. Diofil 2 0.28 2.34
Diospyros frutescens Bl. Diofru 2 7.92
Diospyros hasseltii Zoll. Diohas 2 2.09 0.20
Diospyros longebracteata Lec. Diolon 2 6.54 25.50 1.19 20.11
Diospyros longipedicellata Lec. Diolog 2 0.59 2.98
Diospyros maritima Bl. Diamar 2 2.90 0.52
Diospyros nitida Merr. Dionit 2 3.96 6.59
Diospyros rubra Lec. Diorub 2 0.41 6.17
Elaeocarpaceae Elaeocarpus petiolatus (Jack.) Wall ex Kurz. Elapet 2 0.22
Elaeocarpus stipularis Bl. Elasti 2 1.15
Euphorbiaceae Antidesma velutinosum Bl. Antvel 2 0.50 0.47 0.21 0.52
Aporusa dioica (Roxb.) Muell.-Arg. Apodio 2 0.21
Aporusa planchoniana H. Baill. ex Muell. Apopla 2 0.25 0.61 1.01
Aporusa yunnanensis (Pax. & Hoffm.) Metc. Apoyun 2 0.69
Baccaurea ramiflora Lour. Bacram 2 2.12 1.78
Cleistanthus sumatranus (Miq.) Muell.-Arg. Clesum 2 0.25 22.16 11.71
Erismanthus sinensis Oliv. Erisin 2 3.76 2.61
Exccaria oppositifolia Griff. Excopp 2 0.23 1.97
HCT773 Hct773 2 0.61
HCT822 Hct822 1 2.00 4.15 0.54
Suregada multiflora (Juss.) H. Baill. Surmul 2 0.25 0.81 0.54
Fabaceae Adenanthera pavonina L. Adepav 2 0.25 0.25
Afzelia xylocarpa (Kurz.) Craib. Afzxyl 1 6.68 19.69 1.55 5.53
Albizia lucidior (Steud.) J. Niels. Albluc 1 1.15
Albizia vialenea Pierre Albvia 2 0.55
Dalbergia mammosa Pierre Dalmam 1 1.65 8.59
Milletia diptera Gagn. Mildip 2 0.50
Milletia nigrescens Gagn. Milnig 2 2.26
Ormosia tonkinensis Gagn. Ormton 2 0.57
Peltophorum dasyrrachis var. tonkinensis (Pierre)
K. & S. S. Larsen Peldas 2 0.26
Pterocarpus macrocarpus Kurz. Ptemac 2 0.25
Scolopia spinosa (Roxb.) Warb. Scospi 2 0.24
Sindora siamensis Teysm. ex Miq. var. siamensis Sinsia 1 1.76
Xylia xylocarpa (Roxb.) Taubert Xylxyl 1 0.90
Hypericaceae Cratoxylon cochinchinensis (Lour.) Bl. Cracoc 2 0.87 0.21
Icacynaceae Gonocaryum lobbianum (Miers.) Kurz. Gonlob 2 0.42 0.25
Irvingiaceae Irvingia malayana Oliv. ex Benn. Irvmal 1 1.55
Lauraceae Beilschmiedia foveolata Kost. Beifov 2 0.50
Beilschmiedia laotica Kost. Beilao 2 1.38

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Appendix continued

Plots

Family Species Code G A B C D E

Beilschmiedia micranthopsis Kost. Beimic 1 0.93 4.58 0.80

Cinnamomum iners Reinw. Cinine 2 0.77 0.50
Cryptocarya ferrea Bl. Cryfer 1 0.30
Cryptocarya obovata R. Br. Cryobo 2 0.21 1.28
Cryptocarya ochracea Lec. Cryoch 1 0.71
Dehaasia caesia Bl. Dehcae 2 1.63 1.17
Dehaasia poilanei Liouhe. Dehpoi 2 0.82
HCT093 Hct093 2 0.27
HCT439 Dehaasia Hct439 2 0.95
HCT846 Hct846 2 0.44
HCT879 Hct879 2 0.20 0.24
Persea mollis (W. W. Sm.) Kost. Permol 1 1.58
Phoebe pallida Nees. Phopal 2 0.43
Lecythidaceae Barringtonia acutangula (L.) Gaertn. Baracu 2 0.27
Barringtonia pauciflora King. Barpau 2 0.25
Careya arborea Roxb. Cararb 1 0.67
Lythraceae Lagerstrœmia calyculata Kurz. Lagcal 1 106.1 30.30 79.86 14.22
Lagerstrœmia duperreana Pierre ex Gagn. Lagdup 1 0.40
Lagerstrœmia ovalifolia Teijsin. & Binn. Lagova 1 0.94 7.56 1.32 2.81
Malvaceae Hibiscus macrophyllus Roxb. ex Hornem Hibmac 1 1.96
Meliaceae Aglaia hoaensis Pierre Aglhoa 2 0.43 0.29
Aglaia pirifera Glance Aglpir 2 0.22
Aphanamixis polystachia J. N. Parker Aphpol 1 2.13
Disoxylum arborescens Miq. Disarb 1 0.21
Disoxylum binectariferum Hook. f. Disbin 2 0.79 0.64
Disoxylum cauliflorum Hiern Discau 1 0.22 0.76
Disoxylum loureirii Pierre Dislou 1 0.25 0.68
HCT097 Disoxylum Hct097 2 0.20 0.29
Disoxylum tpongense Pierre Distpo 2 1.26
HCT569 Hct569 2 9.39
Walsura robusta Roxb. Walrob 2 0.25 0.42 1.06
Moraceae Artocarpus nitida ssp. lignanensis (Merr.) Jarr. Artnit 2 0.48
Artocarpus rigida ssp. asperulus (Gagn.) Jarr. Artrig 2 0.52 0.21
HCT102 Hct102 1 0.60 0.32 0.46
Streblus taxoides (Heyne) Kurz Strtax 2 0.25 2.58 0.21 1.02
Myristicaceae Knema lenta Warb. Knelen 2 7.21 0.51
Myrtaceae Cleistocalyx nigrans Gagn. Merr. & Perry. Clenig 2 0.81
HCT211 Syzygium Hct211 2 2.00
Syzygium grandis Wight Syzgra 1 0.37
Syzygium jambos (L.) Alston. Syzjam 2 1.23 2.54
HCT115 Hct115 2 1.15
Syzygium zeylanicum (L.) DC. Syzzey 2 3.75
Oleaceae Linociera microstigma Gagn. Linmic 2 0.27 0.31
Linociera pierrei Gagn. Linpie 2 2.61 0.58 0.85 2.90
HCT551 Linociera Hct551 2 0.41 7.50
Oleaceae Olea macrophylla Gagn. Olemac 2 0.40
Proteaceae Heliciopsis terminalis (Kurz) Sleumer Helter 2 0.24
Rhizophoraceae Carallia brachiata (Lour.) Merr. Carbra 2 0.55 1.22
Chryso balanaceae Parinari annamensis Hance Parann 1 12.51 2.77
Rubiaceae Aidia pycnantha (Drake) Tirv. Aidpyc 2 0.29
Haldina cordifolia (Roxb.) Ridsd. Halcor 1 10.64
HCT436 Hct436 2 0.28 0.57
HCT525 Hct525 2 0.25 0.41
HCT543 Hct543 2 0.31 0.25
HCT572 Hct572 2 0.28
HCT814 Randia Hct814 2 0.20
HCT860 Hct860 2 0.26

© Blackwell Science Ltd 2000, Journal of Biogeography, 27, 141–157

JBI347.fm Page 157 Tuesday, April 25, 2000 1:27 PM

Forest typology in Cat Tien National Park 157

Appendix continued

Plots

Family Species Code G A B C D E

Hymenodictyon orixense (Roxb.) Malb. Hymori 2 0.62

Neolamarckia cadamba (Roxb.) Bosser. Neocad 1 0.25 1.33
HCT924 Randia Hct924 2 0.23
Rothmania encodon (K. Schum.) Brem. Rotenc 2 0.27 0.41 0.42 3.49
Rothmania vietnamensis Tirv. Rotvie 1 2.70 0.34
Tarenna disperma (Hook. f.) Pit. Tardis 2 0.21 0.51
HCT391 Tarenna Hct391 2 0.25 0.26
Rutaceae Acronychia pedunculata (L.) Miq. Acrped 2 4.22
Murraya paniculata (L.) Jack. Murpan 2 1.58 0.30
Sapindaceae Arytera littoralis Bl. Arylit 2 1.27 0.24
Nephelium melliferum Gagn. Nepmel 2 1.28 3.15 0.98
Xerospermum noronhianum (Bl.) Bl. Xernor 2 0.25 1.92 1.49 10.70
Sapotaceae Madhuca pierrei (Williams) H. J. Lam. Madpie 2 1.01 1.76 0.91 1.04
Palaquium obovatum (Griff.) Engleter var. obovatum Palobo 1 3.41
Xantolis dongnaiensis (Dub.) Aubr. Xandon 2 1.88 0.36 1.05
Staphyleaceae HCT178 Hct178 2 0.20
Sterculiaceae Pterospermum diversifolium Bl. Ptediv 1 1.27 3.19 1.20 3.06
Pterospermum grewiaefolium Pierre Ptegre 2 0.79
Sterculia foetida L. Stefoe 1 1.46
Sterculia lanceolata Cav. Stelan 2 0.37 1.83 0.64
Sterculia tonkinensis A. DC. Steton 2 0.22
Symplocaceae Symplocos longifolia Fletcher Symlon 2 0.52 0.36
Theaceae Camelia fleuryi (Chev.) Sealy. Camfle 2 0.62
Tiliaceae Grewia paniculata Roxb. ex DC. Grepan 2 0.61 1.58 1.08
Verbenaceae HCT617 Vitex Hct617 2 0.24
Vitex pinnata L. Vitpin 1 0.93 5.10
Vitex quinata Williams Vitqui 1 0.25 0.93 0.36
HCT925 Vitex Hct925 2 0.71
Vitex tripinnata (Lour.) Merr. Vittri 2 9.95 2.45

© Blackwell Science Ltd 2000, Journal of Biogeography, 27, 141–157