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Acta Biologica Malaysiana (2013) 2(3): 85-94
http://dx.doi.org/10.7593/abm/2.3.85
Composition and Diversity of Plant Seedlings and Saplings at Early
Secondary Succession of Fallow Lands in Sabal, Sarawak
Karyati • Isa B. Ipor • Ismail Jusoh • Mohd. Effendi Wasli • Idris Abu Seman
Received: 04 April 2013 / Accepted: 15 December 2013
© Acta Biologica Malaysiana 2013
Abstract Seedlings and saplings represent the
juvenile stage of plant life and their presence can
reflect the future forests regeneration. However,
still less information is available on the
composition and diversity of seedlings and
saplings under secondary forests at Sarawak,
especially in fallow lands after shifting cultivation.
In this study, the composition and diversity of
plant seedlings and saplings in secondary forests at
various age stands was conducted in order to
obtain basic information on species under
succession of secondary forests after shifting
cultivation. A survey was carried out in four stages
of fallows land such as 3 years
Karyati
Faculty of Forestry, University of Mulawarman,
Kampus Gunung Kelua,
Samarinda, East
Kalimantan, Indonesia, 75119.
Ipor I. B., Jusoh I., Wasli M. E.
Faculty of Resource Science and Technology,
Universiti Malaysia Sarawak, 94300, Kota
Samarahan, Sarawak, Malaysia.
Seman I. A.
Ganoderma and Diseases Research of Oil Palm Unit,
Malaysian Palm Oil Board (MPOB), Bandar Baru
Bangi, 43000 Kajang, Selangor, Malaysia.
Karyati (
)
Faculty of Forestry, University of Mulawarman,
Kampus Gunung Kelua,
Samarinda, East
Kalimantan, Indonesia, 75119.
Email: karyati.hanapi@yahoo.com
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of fallows lands (hereafter called Temuda I),
5 years old secondary forest (hereafter called
Temuda II), 10 years old secondary forest
(hereafter called Belukar I), and 20 years old
secondary forest (hereafter called Belukar II)
in Sabal area, Sarawak. Twenty five plots
with the size of 20 m × 20 m were established
in each study sites and all plant seedlings and
saplings within the plot were enumerated and
identified. The results showed that Temuda I
and Temuda II were mostly dominated by
pioneer species such as Melastoma
malabathricum L., Ficus aurata Miq.,
Ploiarium alternifolium Melchior, Dillenia
spp. and Macaranga spp. At Belukar II,
significant changes in terms of species
composition was obvious where plant species
such as Artocarpus sarawakensis Jarrett,
Artocarpus integer (Thunb.) Merr., and
Palaquium decurrens H.J. Lam were among
the most common species in this study site.
Among all the study sites, species diversity of
Belukar I was the highest based on the indices
of diversity (3.12), evenness (0.90), and
richness (7.68). By understanding the
composition and diversity of plant
regeneration at early stages of secondary
succession on fallow lands, such information
will be useful for biodiversity conservation,
and social and economic values for future
forest.
Keywords Floristic composition • diversity •
seedlings • saplings • secondary succession •
fallow lands
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Introduction
Secondary forests cover more than 600 million ha
of the land area in the tropics in which, accounts
for about 40% of the total forest area with rates of
formation are about 9 million ha year-1 (Brown &
Lugo 1990). FAO (1996) estimated that the area of
secondary forest in 1990 in Asia to be 87.5 million
ha, while the figures for Latin America and Africa
were 165 and 90 million ha, respectively. Such
situation in the tropical region suggested that
future goods and services such as timber resources,
environmental services, biodiversity conservation,
and forest products that society obtains from
tropical forests will increasingly have to come
from secondary forests, or from some other kind of
anthropogenically-induced forest (De Jong et al.
2001). For the case of Sarawak, Malaysia, the land
use pressure on primary forests to provide
ecological services are at stake due to the needs for
various activities ranging from commercial
activities such as timber logging to shifting
cultivation by subsistence farmers. Such activities
has purpose being rapidly reduced due to
combination of various activities such as logging
as well as shifting cultivation and are being
replaced by the secondary forests of lower stature
and altered species composition (Jomo et al. 2004;
Primack & Hall 1992). The human disturbance
could bring negative effects to forest and cause the
decline of species diversity and simplicity of plant
community structure (Dianpei et al. 2004).
Swidden fallows provide rotating habitats
for successional species in a primary-secondary
forests matrix thus enhancing biodiversity. Due to
limited forest destruction and rapid re-growth, the
watershed and soil properties of this primarysecondary forest landscape are almost the same as
to the land under primary forest (Chokkalingam et
al. 2001). Plant species composition, diversity, and
growth during the fallow period after shifting
cultivation are resulted from complex interactions
among a number of conditions and factors which
occur before and during the fallow period such as
degree of disturbance, historical factors, land
management, tree composition and seed sources in
soils or from the surrounding forests, soil fertility,
and climate conditions (Awang Noor et al. 2008;
Kendawang et al. 2007; Van Do et al. 2010).
The plant seedlings, usually the most
transitory of life-history stages, provide
opportunities to explore novelties, as well as life
cycle continuum feature and vulnerability which
are responsible for the plant species population and
community dynamics (Leck et al. 2008).
Acta Biologica Malaysiana (2013) 2(3): 85-94
Intraspecific
differences
in
sapling
abundances as characterized by the coefficient
of skewness are the potentially useful tool for
predicting future trends in vegetation
population change (Grime & Hillier 2000). To
understand the mechanisms of secondary
forest succession, time since abandonment
has to be considered as a compound factor
integrating variables of community structure
(Van Breugel et al. 2006). Many studies have
been conducted on the floristic and structure
of trees with a DBH > 5 and 10 cm in the
tropical forest of Malaya and Borneo Island
(Adam & Ibrahim 1992; Faridah-Hanum
1999; Faridah-Hanum et al. 1999, 2008; Ipor
et al. 1999; Kartawinata et al. 1981; Nizam et
al. 2006; Proctor et al. 1983; Soepadmo 1987;
Sukardjo et al. 1990; Yamakura et al. 1986).
However, there is still limited information
available on the plant floristic composition as
well as diversity of seedling and saplings in
various ages of secondary forests in Sarawak.
This study was conducted in order to
determine the composition and diversity of
plant seedlings and saplings in secondary
forests after shifting cultivation at various
fallow periods in Sabal.
Materials and Methods
Study Sites
The study was carried out at sites with four
stages of fallow period namely lands with
fallow period of 3, 5, 10, and 20 years
(hereafter called Temuda I (01°04'35.6''N
110°58'49.7''E), Temuda II (01°04'43.3''N
110°59'02.0''E), Belukar I (01°03'55.9''N
110°55'51.4''E), and Belukar II (01°03'55.9''N
110°55'51.4''E), respectively) in Sabal, Sri
Aman, Sarawak, East Malaysia (Figure 1).
The study plots at Sabal were located
approximately 110 km South East of Kuching
along the Kuching-Sri Aman Road and 5 to
15 km from the Sabal Agroforestry Center.
All study sites are formerly shifting
cultivation land for upland rice farming with
almost similar land use history (fallow
cropping rotation). The original vegetation at
Sabal site is classified as lowland mixed
dipterocarp forest with heath forest (kerangas)
(Kendawang et al. 2007; Whitmore 1975).
The soils of the study site are derived from
non-calcareous sedimentary rocks consisting
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Acta Biologica Malaysiana (2013) 2(3): 85-94
87
N
Legend :
N
To
Simunjan
SF
BRUNEI
DARUSSALAM
= main road
= study site
= Sabal Agroforestry Center
= village (kampung)
= secondary forest
SABAH
EAST
MALAYSIA
SARAWAK
Sabal
KALIMANTAN
INDONESIA
Temuda II
(5 yr old SF)
Sabal Kruin
Sabal Agroforestry
Center
Kuching-Sri Aman Road
Belukar II
(20 yr old SF)
Temuda I
(3 yr fallow period)
Sabal Kruin
Baru
Abok
Belukar I
(10 yr old SF)
2 km
Figure 1 – Map of the study area
of fine and whitish sandstone during the mid
Tertiary period (Butt 1983). Most of the soils are
classified
into
Oxyaquic
or
Spodic
Quartzipsamments at Sabal site based on the
USDA classification system (Soil Survey Staff
1994). According to the climatic data were
collected from Sri Aman Station, which is located
nearest to the study area, the area received an
average of 3,491 mm year-1 of rainfall, 26.6oC of
monthly temperature, and 85.1% of relative
humidity during the past 20 years (1992-2011).
According to the Schmidt-Ferguson classification
system (1951), the area is characterized as zone A
with Q (Quotient) of 0.013 where very humid area
with vegetation of tropical rain forest (Karyati et
al. 2012).
Data Analysis
The dominant species of forest community
were determined by the summed dominance
ratio (SDR) of species. To calculate the SDR
of a particular species within the plots, the
following formulas were used (Krebs 1999;
Mueller-Dombois & Ellenberg 1974):
RF = ____Frequency of species______ x 100
Total of frequencies of all species
Rd = Number of individual of a species x 100
Total number of individuals
SDR
Data Collection
The surveys of Temuda I, Temuda II, Belukar I,
and Belukar II were conducted from January 2010
to January 2011. Twenty five sub plots of 20 m ×
20 m were established from every study sites,
enabling sampling and data collection of the main
study to be carried out in a systematic manner. All
plant seedlings and saplings with diameter at
breast height (DBH) of less than 5 cm within the
plot
were
enumerated
and
identified.
Nomenclature was checked in the flora records of
the study area (Anderson 1980; Ashton 1988; Jawa
& Chai 2007; Soepadmo et al. 1996, 2002, 2004,
2007, 2011; Soepadmo & Saw 2000; Soepadmo &
Wong 1995). The habitat condition and all species
of each community were recorded.
RF Rd
2
where, RF is relative frequency and Rd is
relative density.
The floristic similarity of species
composition among different communities
was evaluated using Sorensen similarity index
(ISS) (Fachrul 2007; Misra 1992) and defined
as:
ISS
2C
A B
where, A = number of species found within
site A, B = number of species found within
site B, and C = number of species common to
both A and B.
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Acta Biologica Malaysiana (2013) 2(3): 85-94
Four diversity indices were used to
measure species diversity of standing tree in each
community, such as, the Shannon-Wiener’s index
(H') (Ludwig & Reynolds 1988; Magurran 1988;
Shannon & Weaver 1949) (diversity index), the
Simpson’s index (Ds) (Odum 2005; Simpson 1949)
(ecological dominance index), Pielou’s index (J')
(Ludwig & Reynolds 1988, Pielou 1975)
(community evenness index), Margalef’s index (R)
(Ludwig & Reynolds 1988; Margalef 1958)
(species richness index).
s
n
n
H ' i ln i
N
i 1 N
n
Ds i
i 1 N
s
J'
R
2
H'
ln( S )
S 1
ln n
As stated here, ni = number of individuals of the ith species, N = total number of all the individuals
in a unit area, and S = number of species in each
plot.
The category of plant species diversity
was adapted from Odum (2005), while
classification of plant species according to
ecological dominance and community evenness
was adapted from Krebs (1999). Odum (2005)
classified the Shannon-Wiener index (H') in a
community into three diversity categories: H' < 1 =
low diversity, 1 < H' < 3 = intermediate diversity,
and H' > 3 = high diversity. On the basis of
ecological dominance (Ds) in a community the
species are grouped into three categories: 0.00 <
Ds < 0.30 = low dominance, 0.30 < Ds < 0.60 =
intermediate dominance, 0.60 < Ds < 1.00 = high
dominance (Krebs, 1999). The species may be
grouped into three categories of community
evenness (J'): J' < 0.4 = low evenness, 0.4 < J' <
0.6 = intermediate evenness, and 0.6 < J' < 1 =
high evenness (Krebs, 1999). The mean values of
H', Ds, J', and R for each site were compared with
one-way analysis of variance (ANOVA) by
Tukey’s tests. All statistical tests were conducted
using SPSS version 18 for Windows (SPSS Inc.,
2012).
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Results and Discussion
Floristic Composition
The survey on various ages of secondary
forests showed significant variation with their
plant density, species composition, and
diversity. The number of plant seedlings and
saplings decreased in secondary forests with
increasing fallow period. Density of the plant
seedlings and saplings (DBH of < 5 cm) was
considerably high in Temuda I (3332
individuals per hectare), Temuda II (3149
individuals per hectare), Belukar I (3092
individuals per hectare), and Belukar II (2352
individuals per hectare) as shown in Table 1.
Table 2 presented relative frequency (RF),
relative density (Rd), and summed dominance
ratio (SDR) of ten most common plant species
among the seedlings and saplings in each
study site. According to density and SDR, the
plant seedlings and saplings in Temuda I and
Temuda II were dominated by light
demanding and fast growing species, such as
M. malabathricum, P. alternifolium, and F.
aurata as well as Dillenia spp. and
Macaranga spp. Dillenia suffruticosa Martelli
was also common species in both Belukar I
and Belukar II. The other common species of
Belukar I were Syzygium arcuatinervum
(Merr.) Craven & Briffin, Diospyros siamang
Bakh., Agrostistachys longifolia Benth. ex
Hook. f., Macaranga caladifolia Becc., and
Whiteodendron moultonianum (W.W.Sm.)
Steenis. Belukar II was dominated by P.
decurrens, Nephelium cuspidatum Blume,
Antidesma neurocarpum Miq., and Syzygium
polyanthum Walp. as well as Artocarpus spp.
Seedlings and saplings of M.
malabathricum was the most dominant
species at the early stage of secondary
succession period till 5 years after land
abandonment. In degraded old fields in
Peninsular Malaysia, stands that were
dominated by Melastoma in the early stages
then became occupied by other species after
4-8 years (Kochummen & Ng 1977). During
the early fallow period after shifting
cultivation (less than about three years),
Kemunting (Melastoma polyanthum) was
dominant with higher frequency and density
in the Mujong River area, Sarawak (Tanaka et
al. 2007). In burned plots of East
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Acta Biologica Malaysiana (2013) 2(3): 85-94
89
Table 1 – Ten most common species of plant seedlings and saplings (DBH of < 5 cm) in terms of density in 1 hectare of
each study site
No.
Species
Family
Temuda I
Temuda II Belukar I
Belukar II
1
Agrostistachys longifolia
Euphorbiaceae
59 (6)
Benth. Ex Hook. F.
2
Alstonia spatulata Blume
Apocynaceae
111 (9)
3
Antidesma neurocarpum
Euphorbiaceae
65 (7)
Miq.
4
Artocarpus integer (Thunb.)
Moraceae
203 (2)
Merr.
5
Artocarpus sarawakensis
Moraceae
422 (1)
Jarrett
6
Cratoxylum arborescens
Clusiaceae
120 (10)
Blume.
7
Cratoxylum glaucum Korth.
Clusiaceae
147 (6)
101 (10)
8
Dillenia pulchella Gilg
Dilleniaceae
141 (7)
140 (6)
62 (8)
9
Dillenia suffruticosa Martelli
Dilleniaceae
150 (5)
179 (3)
91 (1)
94 (5)
10
Diospyros siamang Bakh.
Ebenaceae
55 (10)
11
Euodia glabra (Bl.) Bl.
Rutaceae
139 (8)
127 (7)
12
Ficus aurata Miq.
Moraceae
171 (3)
173 (4)
13
Goniothalamus andersonii J.
Annonaceae
49 (10)
Sincl.
14
Gonystylus costalis Airy
Thymelaeaceae
114 (8)
Shaw
15
Hopea kerangasensis P.S.
Dipterocarpaceae
58 (7)
Ashton
16
Leea indica (Burm.f.) Merr.
Ampelidaceae
57 (8)
17
Lepisanthes sp.
Sapindaceae
56 (9)
18
Macaranga beccariana
Euphorbiaceae
201 (2)
Merr.
19
Macaranga caladifolia Becc.
Euphorbiaceae
64 (3)
20
Macaranga igantean Mull.
Euphorbiaceae
135 (9)
Arg.
21
Macaranga trichocarpa
Euphorbiaceae
168 (5)
Mull. Arg.
22
Melastoma malabathricum
Melastomataceae
292 (1)
409 (1)
L.
23
Nephelium cuspidatum
Sapindaceae
121 (4)
Blume
24
Palaquium decurrens H.J.
Sapotaceae
180 (3)
Lam
25
Ploiarium alternifolium
Theaceae
152 (4)
220 (2)
Melchior.
26
Shorea faguetiana Heim
Dipterocarpaceae
60 (5)
27
Shorea pinanga Scheff.
Dipterocarpaceae
56 (9)
28
Syzygium arcuatinervum
Myrtaceae
66 (2)
(Merr.) Craven & Briffin
29
Syzygium polyanthum Walp.
Myrtaceae
67 (6)
30
Whiteodendron
Myrtaceae
61 (4)
moultonianum (W.W.Sm.)
Steenis
Total
1648
1742
627
1319
Total per hectare
3332
3149
3092
2352
Number of families
39
38
55
46
Number of genera
74
72
140
86
Number of species
97
93
220
106
The figures in parentheses represent the ranking in terms of density per hectare. (1) represent species with the highest density.
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Acta Biologica Malaysiana (2013) 2(3): 85-94
Table 2 – Ten most common species of plant seedlings and saplings (DBH of < 5 cm) in terms of summed dominance ratio
(SDR) in 1 hectare of each study site.
No.
Species
Family
RF (%)
Rd (%)
IVi
SDR
A. Temuda I
1
Melastoma malabathricum L.
Melastomataceae
4.49
8.76
13.26
6.63
2
Ficus aurata Miq.
Moraceae
4.31
5.13
9.44
4.72
3
Ploiarium alternifolium Melchior.
Theaceae
3.56
4.56
8.12
4.06
4
Dillenia pulchella Gilg
Dilleniaceae
3.00
4.23
7.23
3.61
5
Euodia glabra (Bl.) Bl.
Rutaceae
3.00
4.17
7.17
3.58
6
Cratoxylum glaucum Korth.
Clusiaceae
2.62
4.41
7.03
3.52
7
Macaranga beccariana Merr.
Euphorbiaceae
0.94
6.03
6.97
3.48
8
Macaranga gigantea Mull. Arg.
Euphorbiaceae
2.81
4.05
6.86
3.43
9
Adinandra dumosa Jack
Theaceae
3.37
3.12
6.49
3.25
10
Macaranga havilandii Airy Shaw
Euphorbiaceae
2.81
3.60
6.41
3.21
B. Temuda II
1
Melastoma malabathricum L.
2
Ficus aurata Miq.
3
Ploiarium alternifolium Melchior.
4
Macaranga trichocarpa Mull. Arg.
5
Dillenia suffruticosa Martelli
6
Vitex pubescens Vahl.
7
Euodia glabra (Bl.) Bl.
8
Dillenia pulchella Gilg
9
Leea indica (Burm.f.) Merr.
10
Gonystylus costalis Airy Shaw
C. Belukar I
1
Dillenia suffruticosa Martelli
2
Syzygium arcuatinervum (Merr.)
Craven & Briffin
3
Endospermum diadenum (Miq.)
Airy Shaw
4
Diospyros siamang Bakh.
5
Agrostistachys longifolia Benth. ex
Hook. f.
6
Hopea kerangasensis P.S. Ashton
7
Macaranga caladifolia Becc.
8
Whiteodendron moultonianum
(W.W.Sm.) Steenis
9
Hopea dryobalanoides Miq.
10
Santiria rubiginosa Blume
D. Belukar II
1
Artocarpus sarawakensis Jarrett
2
Artocarpus integer (Thunb.) Merr.
3
Palaquium decurrens H.J. Lam
4
Nephelium cuspidatum Blume
5
Antidesma neurocarpum Miq.
6
Syzygium polyanthum Walp.
7
Dillenia suffruticosa Martelli
8
Xylopia ferruginea Baill.
9
Lepisanthes sp.
10
Dillenia pulchella Gilg
Melastomataceae
Moraceae
Theaceae
Euphorbiaceae
Dilleniaceae
Verbenaceae
Rutaceae
Dilleniaceae
Ampelidaceae
Thymelaeaceae
3.93
4.72
2.55
3.73
2.95
4.52
3.34
1.96
3.34
2.36
12.99
5.49
6.99
5.34
5.68
3.14
4.03
4.45
2.70
3.62
16.92
10.21
9.54
9.07
8.63
7.66
7.37
6.41
6.04
5.98
8.46
5.10
4.77
4.53
4.32
3.83
3.69
3.21
3.02
2.99
Dilleniaceae
Myrtaceae
2.19
1.04
2.94
2.13
5.13
3.18
2.56
1.59
Euphorbiaceae
1.56
1.58
3.15
1.57
Ebenaceae
Euphorbiaceae
1.35
1.25
1.78
1.88
3.13
3.12
1.57
1.56
Dipterocarpaceae
Euphorbiaceae
Myrtaceae
1.25
0.94
1.04
1.84
2.07
1.94
3.09
3.01
2.98
1.55
1.50
1.49
Dipterocarpaceae
Burseraceae
1.35
1.25
1.52
1.62
2.87
2.87
1.44
1.43
Moraceae
Moraceae
Sapotaceae
Sapindaceae
Euphorbiaceae
Myrtaceae
Dilleniaceae
Annonaceae
Sapindaceae
Dilleniaceae
17.94
8.63
7.65
5.14
2.76
2.85
4.00
2.04
2.38
2.64
0.95
3.09
3.33
1.90
3.09
2.85
1.66
3.09
2.38
1.66
18.89
11.72
10.98
7.04
5.85
5.70
5.66
5.13
4.76
4.30
9.45
5.86
5.49
3.52
2.93
2.85
2.83
2.56
2.38
2.15
RF = relative frequency, Rd = relative density, IVi = importance value index, and SDR = summed dominance ratio.
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Acta Biologica Malaysiana (2013) 2(3): 85-94
Kalimantan, the occurrence of M. malabathricum,
Eupatorium inulaefolium, Ficus sp., and Vitex
pinnata L. strongly increase with the age of
regeneration (last burned 3 years, 4 years, and 9
years previously), but were rarely found in the
secondary forest after fire burning (Yassir et al.
2010). Melastoma is one of the characteristic
species of Adinandra-belukar communities which
grow in very low-nutrient soils in South-East Asia
(Turner 1991). Several studies had reported the
similar result on the abundance of Melatoma and
Macaranga spp. at the secondary forests in
Sarawak, East Malaysia (Ipor & Tawan 2004), in
East Kalimantan, Indonesia (Slik et al. 2003), and
in Mindanao, Philippine (Weidelt & Banaag
1982).
The dominance of the fast-growing
pioneer trees species were not exist in Belukar II.
Although Dillenia spp. and Macaranga spp. were
still encountered in Belukar II, they were not as
abundant in Temuda I, Temuda II, and Belukar I.
The occurrence of pioneer species, such as D.
suffruticosa and M. caladifolia were still common
in Belukar I. In Belukar II, pioneer species were
not dominant based on density and SDR, while A.
sarawakensis, A. integer, and P. decurrens were
dominant in this Belukar. Sorensen’s index (Cs) is
regarded as one of the most effective presence or
absence similarity measures (Magurran 2004;
Southwood & Henderson 2000). The similarity
index of association Temuda I and Temuda II was
the highest (64.21%), followed by Temuda II and
Belukar II (56.28%), Temuda I and Belukar II
(49.26%), Belukar I and Belukar II (40.49%),
Temuda II and Belukar I (37.70%), and Temuda I
and Belukar I (37.22%) (Table 3). The
development and changes of species composition
of plant seedlings and saplings after slash and burn
process was mostly influenced by secondary
succession process and fallow age in abandoned
lands. The result showed that during early stage
secondary succession of fallow lands after shifting
cultivation, the floristic composition was
dominated and obtained by many common and
similar species in Temuda I, Temuda II, and
Belukar I. However, Belukar II showed relatively
different species composition among all study
sites. This showed that species composition at
abandoned lands after burning begin to change
after 20 years of abandonment. Several species of
Dipterocarpaceae were also recorded, including
Hopea dryobalanoides Miq., Hopea kerangasensis
P.S. Ashton, Shorea faguetiana Heim, and Shorea
pinanga Scheff. in Belukar I. In Belukar II, the
density of plant seedlings and saplings was less
91
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
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73
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76
77
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80
81
82
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105
106
than those recorded in Belukar I. Late pioneer
species and secondary species, such as A.
neurocarpum, N. cuspidatum, S. polyanthum,
Xylopia ferruginea Baill., and Lepisanthes sp.
were common in this site.
Floristic Diversity
The diversity indices of plant seedlings and
saplings in various ages of secondary forests
are presented in Table 4. The ShannonWiener diversity indices (H') of all study sites
were categorized as ‘intermediate to high
diversities’. This was due to high density of
plant seedlings and saplings recorded in every
study sites. High species diversity indicates a
highly complex community, for a greater
variety of species allows for a larger array of
species interactions (Brower et al. 1990). The
diversity index in Belukar I was significantly
higher than Temuda I, Temuda II, and
Belukar II perhaps due to the high number of
families, number of genera, and number of
species were recorded in Belukar I compared
to the other sites as shown in Table 1.
The ecological dominance (Ds value)
of all studied forests was categorized as ‘low
dominance’. The Ds value of Belukar II was
the highest among the four studied sites
(0.17). This value suggested that a few or
almost no plant species were dominant in
every study sites. The nearly zero values
correspond to low diverse or more
homogeneous plant ecosystem. The Ds values
is an expression of how many equally
abundant species would have a diversity equal
to that in the observed collection (Brower et
al. 1990). The dominance index of Belukar I
was significantly lower than the other study
sites. It was probably due to the number of
individuals of every species and mostly
related to diversity index. All study sites had
‘high evenness index (J')’. This indicated that
every species was distributed evenly within
the plant community. As mentioned for H'
and Ds, the values of J' and R (species
richness index) showed no significant
different in three sites of Temuda I, Temuda
II, and Belukar II. The J' and R values of
Belukar I were significantly higher compared
to the other studied sites. The high values of J'
and R may effected by a large number of
number of individuals and number of species
observed in the study sites.
The results showed that species
1
2
3
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Acta Biologica Malaysiana (2013) 2(3): 85-94
Table 3 – Sorensen similarity index (ISS) of plant seedlings and saplings (DBH of < 5 cm) in the study sites.
Type
Temuda I
Temuda II
Belukar I
Belukar II
Temuda I
64.21
37.22
49.26
Temuda II
Belukar I
Belukar II
37.70
56.28
40.49
-
Sorensen similarity index was computed for the entire study plots (1 ha).
Table 4 – Diversity indices of plant seedlings and saplings (DBH of < 5 cm) in the study sites.
No.
Diversity indices
Temuda I (n=25)
Temuda II (n=25)
Belukar I (n=25)
Shannon-Wiener diversity
a
a
1
2.41 (+0.07)
2.43 (+0.09)
3.12 (+0.14)b
index (H')
2
Simpson dominance index (Ds)
0.13 (+0.01)ab
0.14 (+0.02)b
0.07 (+0.01)a
a
a
3
Pielou evenness index (J')
0.86 (+0.01)
0.82 (+0.01)
0.90 (+0.01)b
a
a
4
Margalef species richness (R)
3.66 (+0.29)
4.03 (+0.25)
7.68 (+0.72)b
Belukar II (n=25)
2.28 (+0.10)a
0.17 (+0.03)b
0.82 (+0.03)a
3.78 (+0.24)a
Calculation was done according to the 20 m × 20 m subplots. Values are average and standard error in parentheses. Different letters in each
line indicate a significant different at 5% level by Tukey's test among different ages of secondary forests.
diversity indices of plant seedlings-saplings varied
widely among the four study sites. These three
indices increased as fallow periods increased then
at 20 years old secondary forest, these indices
showed decreasing value (Table 4). The highest
values of H', J', and R were recorded in Belukar I.
In contrast, the lowest evenness index was also
observed in this studied site. It may be due to past
intermediate disturbance in this site as compared to
Temuda I, Temuda II) and Belukar II. The results
showed that, as the H', J', and R increased, the Ds
decreased. Diversity will be greatest at
intermediate disturbance frequencies because the
landscape includes patches of a great variety of
ages supporting a wide mix of species (Wright
1999). A forest is most rich in species when at an
intermediate state of recovery from disturbance, or
when disturbance is at an intermediate intensity or
frequency, because it will then contains both
pioneer and climax species (Whitmore 1993).
The development and changes of floristic
composition and diversity of plant seedlings and
saplings during early stages of secondary
succession process was mostly influenced by
secondary succession process and fallow period in
various ages of secondary forests after slash and
burn process. The floristic composition may affect
to values of H', Ds, J, and R. Information on the
composition and diversity of seedlings and
saplings are useful for predicting future trends in
the vegetation succession, especially on secondary
succession of fallow lands. By understanding the
composition and diversity of plant regeneration at
early stages of secondary succession on fallow
lands, such information will be useful for
biodiversity conservation, and social and economic
values for future forest.
Acknowledgement We acknowledge to Malaysian
Palm Oil Board (MPOB) for supporting the funding
of this research project. We thank all support staff at
Faculty of Resource Science and Technology, En.
Hidir Marzuki, En. Sekudan Tedong, En. Salim Arip,
and En. Muhd Najib Fardos for their field assistance
and companionship during this survey.
References
Adam, J.H. & Ibrahim, K. (1992). An Enumeration of
One Hectare of Lowland Dipterocarp
Forest at Danum Valley Field Centre,
Lahad
Datu,
Sabah,
Malaysia.
Rehabilitation of Tropical Rainforest
Ecosystems : Research and Development
Priorities, pp. 43-56.
Anderson, J.A.R. (1980). A Check List of the Trees
of Sarawak. Malaysia: Forest Department
Sarawak.
Ashton, P.S. (1988). Manual of the Non-Dipterocarp
Trees of Sarawak, Vol. II. Kuala Lumpur:
Dewan Bahasa dan Pustaka.
Awang Noor, A.G., Faridah Hanum, I. & Tuan
Marina, T.I. (2008). Relationship between
Economic Value and Species Diversity of
Timber Resources in a Hill Forest in
Peninsular Malaysia.
Journal of
Sustainable Development, 1(2), 17-26.
Brower, J.E., Zar, J.H. & Von Ende, C.N. (1990).
Field and Laboratory Methods for General
Ecology, 3rd Ed. USA : Wm. C. Brown
Publishers. 237 pp.
Brown, S. & Lugo, A.E. (1982). The Storage and
Production of Organic Matter in Tropical
Forest and Their Role in the Global Carbon
Cycle. Biotropica, 14, 161-187.
Butt, G. (1983). Semi-Detailed Site Evaluation of
Sabal Forest Reserve Northern Half.
Forest Research Report No. S.S.2.
Kuching: Soil Survey Unit, Research
1
Acta Biologica Malaysiana (2013) 2(3): 85-94
93
2
3
Section, Forest Department, Sarawak.
Chokkalingam, U., Smith, J. & De Jong, W. (2001). A
Conceptual Framework for the Assessment of
Tropical Secondary Forest Dynamics and
Sustainable Development Potential in Asia.
Journal of Tropical Forest Science, 13(4), 577600.
De Jong, W., Chokkalingam, U. & Smith, J. (2001).
Tropical Secondary Forests in Asia: Introduction
and Synthesis. Journal of Tropical Forest
Science, 13(4), 563-576.
Dianpei, W., Shuyi, J., Feipeng, C. & Shaolin, P. (2004).
Composition and Characteristics of Natural
Secondary Forests in Shenzhen, South China.
Forestry Studies in China, 6(2), 6-11.
Fachrul, M. F. (2007). Metode Sampling Bioekologi.
Jakarta: Bumi Aksara.
Faridah-Hanum, I. (1999).
Plant Diversity and
Conservation Value of Ayer Hitam Forest,
Selangor, Peninsular Malaysia. Pertanika Journal
of Tropical Agriculture Science, 22(2), 73-83.
Faridah-Hanum, I., Simin, M. & Awang Noor, A.G.
(1999). Tree Species Diversity and Economic
Value of a Watershed Forest in Ulu Muda Forest
Reserve, Kedah. Pertanika Journal of Tropical
Agriculture Science, 22(1), 63-68.
Faridah-Hanum, I., Philip, L. & Awang Noor, A.G. (2008).
Sampling Species Diversity in a Malaysian Rain
Forest: The Case of a Logged-Over Forest.
Pakistan Journal of Botany, 40(4), 1729-1733.
FAO. (1996). Forest Resources Assessment 1990. Survey
of Tropical Forest Cover and Study of Change
Processes. Rome, Italy: FAO Forestry Paper
130. 152 pp.
Grime, J.P. & Hillier, S.H. (2000). The Contribution of
Seedling Regeneration to the Structure and
Dynamics of Plant Communities and Larger
Units of Landscape. In: M. Fenner (Ed.), Seeds:
The Ecology of Regeneration in Plant
Communities, 2nd Ed. (pp. 349-364). United
Kingdom: CABI Publishing.
Ipor, I.B. & Tawan, C.S. (2004). Succession of Fallows
after Shifting Cultivation in Sungai Sarawak
Basin, Kuching, Sarawak. In: E.M. Donoghue,
G.L. Benson, & J.L. Chamberlain (Eds.),
Sustainable Production of Wood and Non-Wood
Products: Proceedings of IUFRO Division 5
Research Groups 5.11 and 5.12 (pp. 19-28).
Ipor, I.B., Tawan, C.S., Ismail, J. & Bojo, O. (1999).
Floristic Compositions and Structures of Forest
at Bario Highlands, Sarawak. ASEAN Review of
Biodiversity and Environmental Conservation
(ARBEC). pp. 1-15.
Jawa R. & Chai, P.K. (2007). A New Check List of the
Trees of Sarawak. Sarawak, Malaysia: Lee
Miing Press Sdn. Bhd.
Jomo, K.S., Chang, Y.T. & Khoo, K.J. (2004). Deforesting
Malaysia: The Political, Social Ecology of
Agricultural Expansion and Commercial
Logging. United Kingdom: Zed Books Ltd.
Kartawinata, K., Abdulhadi, R. & Partomihardjo, T.
(1981). Composition and Structure of a Lowland
Dipterocarp Forest at Wanariset, East
Kalimantan. The Malaysian Forester, 44(2 &3),
397-406.
Karyati, Ipor, I.B., Jusoh, I. & Wasli, M.E. (2012).
Suitability
of
Plant
Species
for
Agroforestry Program at Sri Aman,
Sarawak. In: F.B. Ahmad, S. Muid, I.B.
Ipor, R. Zainudin, M.E. Wasli, M. Kalu, &
Z.B. Assim (Eds.),
Taxonomy and
Ecology: Beyond Classical Approaches
(pp. 203-213). Malaysia: Universiti
Malaysia Sarawak.
Kendawang, J.J., Ninomiya, I ., Tanaka, K., Ozawa,
T., Hattori, D., Tanaka, S. & Sakurai, K.
(2007). Effects of Burning Strength in
Shifting Cultivation on the Early Stage of
Secondary Succession in Sarawak,
Malaysia. Tropics, 16, 309-321.
Kochummen, K.M. & Ng, F.S.P. (1977). Natural
Plant Succession after Farming in Kepong.
Malaysian Forester, 40, 61-78.
Krebs, C.J. (1999). Ecological Methodology, 2nd Ed.
USA:
Addison-Welsey
Educational
Publishers, Inc.
Leck, M.A., Simpson, R.L. & Parker, V.T. (2008)
Why Seedlings?. In: M.A. Leck, V.T.
Parker, & R.L. Simpson (Eds.), Seedling
Ecology and Evolution (pp. 3-13). New
Yok : Cambridge University Press.
Ludwig, J. A. & Reynolds, J.F. (1988). Statistical
Ecology: A Primer on Methods and
Computing. USA: John Wiley & Sons,
Inc.
Magurran, A.E. (1988). Ecological Diversity and Its
Measurement.
Princeton, New Jersey:
Princeton University Press.
Magurran, A.E. (2004). Measuring Biological
Diversity.
Oxford, United Kingdom:
Blackwell Science.
Margalef, R. (1958). Information Theory in Ecology.
International Journal of General Systems,
3, 36-71.
Misra, K.C. (1992). Manual of Plant Ecology. New
Delhi: Raju Primlani.
Muller-Dombois, D. & Ellenberg, H. (1974). Aims
and Methods of Vegetation Ecology. New
York: John Wiley & Sons.
Nizam, M.S., Norziana, J., Sahibin, A.R. & Latiff, A.
(2006). Edaphic Relationships Among
Tree Species in the National Park at
Merapoh, Pahang, Malaysia.
Jurnal
Biosains, 17(2), 37-53.
Odum, P.E. (2005). Fundamentals of Ecology, 5th
Ed. Belmont, CA: Thomson Brooks/Cole.
Pielou, E.C. (1975). Ecological Diversity. New
York: John Wiley & Sons Inc.
Primack, R. B. & Hall, P. (1992). Biodiversity and
Forest Change in Malaysian Borneo.
Bioscience, 42, 829-837.
Proctors J., Anderson, J.M., Chai, P. & Vallack, H.W.
(1983).
Ecological Studies in Four
Contrasting Rain Forests in Gunung Mulu
National Park, Sarawak. I.
Forest
Environment, Structure and Floristics.
Journal of Ecology, 71, 237-260.
Shannon, C.E. & Weaver, W. (1949). The
Mathematical Theory of Communication.
Urbana, IL: University of Illinois Press.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
56
57
94
Simpson, E.H. (1949). Measurement of Diversity. Nature
(London), 163, 688.
Slik, J.W.F., Keßler, P.J.A. & Van Welzen, P.C. (2003).
Macaranga
and
Mallotus
Species
(Euphorbiaceae) as Indicators for Disturbance in
the Mixed Lowland Dipterocarp Forest of East
Kalimantan (Indonesia). Ecological Indicators, 2,
311-324.
Soepadmo, E. (1987). Structure, Above Ground Biomass
and Floristic Composition of Forest Formation at
Gunung Janing Barat, Ulu Endau, Johore,
Malaysia. Malayan Nature Journal, 41, 275-290.
Soepadmo, E. & Saw, L.G. (2000). Tree Flora of Sabah
and Sarawak. Vol. Three. Kuala Lumpur: Forest
Research Institute Malaysia (FRIM), Sabah
Forestry Department and Sarawak Forestry
Department.
Soepadmo, E. & Wong, K.M. (1995). Tree Flora of Sabah
and Sarawak. Vol. One. Kuala Lumpur: Forest
Research Institute Malaysia (FRIM), Sabah
Forestry Department and Sarawak Forestry
Department.
Soepadmo, E., Wong, K.M. & Saw, L.G. (1996). Tree
Flora of Sabah and Sarawak. Vol. Two. Kuala
Lumpur: Forest Research Institute Malaysia
(FRIM), Sabah Forestry Department and
Sarawak Forestry Department.
Soepadmo, E., Saw, L.G. & Chung, R.C.K. (2002). Tree
Flora of Sabah and Sarawak. Vol. Four. Kuala
Lumpur: Forest Research Institute Malaysia
(FRIM), Sabah Forestry Department and
Sarawak Forestry Department.
Soepadmo, E., Saw, L.G. & Chung, R.C.K. (2004). Tree
Flora of Sabah and Sarawak. Vol. Five. Kuala
Lumpur: Forest Research Institute Malaysia
(FRIM), Sabah Forestry Department and
Sarawak Forestry Department.
Soepadmo, E., Saw, L.G., Chung, R.C.K. & Kiew, R.
(2007). Tree Flora of Sabah and Sarawak. Vol.
Six. Kuala Lumpur: Forest Research Institute
Malaysia (FRIM), Sabah Forestry Department
and Sarawak Forestry Department.
Soepadmo, E., Saw, L.G., Chung, R.C.K. & Kiew, R.
(2011). Tree Flora of Sabah and Sarawak. Vol.
Seven. Kuala Lumpur: Forest Research Institute
Malaysia (FRIM), Sabah Forestry Department
and Sarawak Forestry Department.
Soil Survey Staff. (1994). Keys to Soil Taxonomy. 6th Ed.
Washington, DC: U.S. Department of
Agriculture & Soil Conservation Service.
Southwood, T.R.E. & Henderson, P.A. (2000). Ecological
Methods. Oxford, United Kingdom: Blackwell
Science.
Acta Biologica Malaysiana (2013) 2(3): 85-94
Sukardjo, S., Hagihara, A., Yamakura, T. & Ogawa
H. (1990). Floristic Composition of a
Tropical Rain Forest in Indonesia Borneo.
Bulletin of the Nagoya University Forests,
10, 1-44.
Tanaka, S., Wasli, M.E., Seman, L, Lee, A.,
Kendawang, J.J., Sakurai, K. & Morooka,
Y. (2007). Ecological Study on Site
Selection for Shifting Cultivation by the
Iban of Sarawak, Malaysia. A Case Study
in the Mujong River Area. Tropics, 16 (4),
357-371.
Turner, I.M. (1991). Effects of Shade and Fertilizer
Addition on the Seedlings of Two Tropical
Woody
Pioneer
Species.
Tropical
Ecological Monographs, 57, 189-214.
Van Breugel, M., Martinez-Ramos, M. & Bongers, F.
(2006). Community Dynamics during Early
Secondary Succession in Mexican Tropical
Rain Forests. Journal of Tropical Ecology,
22, 663-674.
Van Do, T., Osawa, A. & Thang, N.T. (2010).
Recovery Process of a Mountain Forest
after Shifting Cultivation in Northwestern
Vietnam. Forest Ecology and Management,
259, 1650-1659.
Weidelt, H.J. & Banaag, V.S. (1982). Aspects of
Management and Silviculture of Philippine
Dipterocarp Forests. German: German
Agency for Technical Cooperation.
Whitmore, T.C. (1975). Tropical Rain Forests of the
Far East. United Kingdom: Oxford
University Press.
Whitmore, T.C. (1993). An Introduction to Tropical
Rain Forests.
Great Britain: Oxford
University Press.
Wright, S.J. (1999). Plant Diversity in Tropical
Forests. In: F.I. Pugnaire & F. Valladares
(Eds.), Handbook of Functional Plant
Ecology (pp. 449-471). New York: Marcel
Dekker Inc.
Yamakura, T., Hagihara, A., Sukardjo, S. & Ogawa,
H.
(1986). Tree Size in a Mature
Dipterocarp Forest Stand in Sebulu, East
Kalimantan, Indonesia. Tonan Ajia Kenkyu
(Southeast Asian Studies), 23(4), 452-478.
Yassir, I., Van der Kamp, J. & Buurman, P. (2010).
Secondary Succession After Fire in
Imperata Grasslands of East Kalimantan,
Indonesia. 19th World Congress of Soil
Science, Soil Solutions for a Changing
World 1-6 August 2010. Brisbane,
Australia.
Ismail Jusoh