Journal of Vegetation Science 6: 205-210, 1995

IAVS; Opulus Press Uppsala. Printed in Sweden

- Restoration of natural vegetation in degraded Imperata cylindrica grassland -

205

Restoration of natural vegetation in degraded Imperata cylindrica

grassland: understorey development in forest plantations
Kuusipalo, Jussi1*, djers, Gran1, Jafarsidik, Yusuf1, Otsamo, Antti1, Tuomela, Kari1
& Vuokko, Risto2
Reforestation and Tropical Forest Management Project c/o Reforestation Technology Center, P.O. Box 65 (Jl. Sei
Ulin 28 B), 70711 Banjarbaru, Kal-Sel, Indonesia; 2Enso Forest Development Oy Ltd., Kuparintie 47, FIN-55100
Imatra, Finland; *Correspondence: Jl. P. Suriansyah 56, 70711 Banjarbaru, Kal-Sel, Indonesia; Fax + 62 511 93222

Abstract. Reclamation of former, degraded forest lands occupied by Imperata cylindrica is one of the crucial environmental and forestry issues in the humid tropics, notably Southeast
Asia. We suggest that it is possible to gradually restore the
original natural forest cover with the help of a sacrifice fallow
crop of fast-growing exotic tree species. Recently, a set of
suitable fast-growing plantation tree species has been identified and stand establishment methods developed for this purpose. We assessed the regeneration of natural vegetation in
stands of different plantation tree species and evaluated the
ecological impact of species composition in the plantation
understorey. PCA ordination, regression analysis and analysis
of covariance were applied at different stages of the study. We
found a marked vegetational resemblance between stands
dominated by Acacia mangium: they had the highest number
of indigenous trees in their understorey, whereas stands of
other plantation trees supported more diverse grass and herb
vegetation. A high proportion of evergreen woody vegetation
reduces the risk of fire and grass competition and enhances
secondary succession towards natural forest.
Keywords: Acacia mangium; Natural regeneration; Reforestation; Restoration ecology; Forest management.
Nomenclature: George (ed.) (1981); Anon. (1982).

Introduction
Along with the increasing population pressure and
need of new agricultural land, natural forests of southeast Asia are being decimated. During 1978 - 1988, the
deforestation rate has been 16 000 km2/yr. However, the
forest area is actually diminishing twice as fast as the
permanent agricultural land is increasing (Anon. 1992).
The difference may be accounted for by unsustainable
land use practices, including repeated logging of valuable timber particularly Dipterocarpaceae trees, which
often predominate the upper canopy layers (Kostermans
1992). Logging is usuall followed by illegal cutting of
the residual timber, and subsequent shifting cultivation

with a short fallow period (see e.g. Eussen & Wirjahardja

1973; Dela Cruz 1986 and references therein).
Abandoned shifting-cultivation areas are almost always invaded by one of the most noxious colonizers of
the degraded humid tropical forest soils, Imperata cylindrica (alang-alang), a rhizomatous perennial pantropical
grass which occurs over a wide range of soil conditions
and can take advantage of the alternation of moist and
not too long and severe dry seasons, typical for Southeast Asia. Due to its high growth rate (8 - 20 t/ha/yr),
I. cylindrica is a strong competitor for water, light and
nutrients, it is also known to have allelopathic effects on
tree seeds and other plants (Eussen & Wirjahardja 1973;
Soerianegara 1980; Ohta 1992). Natural regeneration of
tree vegetation on alang-alang grasslands is therefore
retarded or impossible. Regular fires over vast areas
covered with dry Imperata shoots hamper any such succession (Eussen & Wirjahardja 1973; Dela Cruz 1986).
The largest alang-alang grasslands are found in the
interior of the island of Borneo, on former dipterocarp
forest soils. A frequently cited, though not accurate
figure for Indonesia alone is 200 000 km2 with an annual
increase of 150 000 ha (Soerjani 1970; Anon. 1990).
Skerman & Riveros (1990) estimated that the total area
of Imperata grasslands throughout the tropics, natural
grasslands included, is 2 000 000 km2.
I. cylindrica is known as a nuisance weed in agriculture. It suppresses the growth of annual and perennial
crops, and most farmers have little interest in reclamation of alang-alang lands for arable crops but prefer
slash-and-burn cultivation in primary or secondary forests instead. Large-scale animal husbandry is virtually
absent in Southeast Asia; moreover, the fodder value of
I. cylindrica is poor, except for a short period in the
beginning of the rainy season (Soerjani 1970). All in all,
Southeast Asian alang-alang grasslands are largely considered unproductive wastelands. For the benefit of
local population and environment another type of land
use would be more valuable (Townson 1991).

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Kuusipalo, J. et al.

Development of alang-alang grassland into tropical

(preferably dipterocarp) rain forest is difficult due to
annual fires and the lack of standing trees and a soil seed
bank. Despite years of trial plantings, no feasible largescale method has been found for restoring the original
vegetation. Planting of dipterocarp or other indigenous
tree seedlings in the middle of grassland, even after
complete cultivation and regular spraying with herbicides, has had very limited success. A more suitable
method is to establish a shade cover crop of fast-growing plantation trees to gradually suppress the grass
(Otsamo et al. in press). A special cover crop, like the
rubber tree, is too expensive to be used on a large scale.
The most promising plantation trees to suppress
I. cylindrica include Australasian Acacia spp., especially A. mangium (Anon. 1983).
The rationale for the hypothesis that natural forest
vegetation can be reclaimed with the help of a sacrifice
fallow crop of fast-growing plantation trees is that (a)
physical properties and nutrient status of the top soil are
improved; (b) cover tree crop favours woody undergrowth at the expense of more light-demanding grassdominated vegetation because light, instead of nutrients, becomes a limiting factor for the understorey, and
(c) biological diversity increases due to the more heterogeneous and limited light availability, created by the
crowns of plantation trees (Tilman 1982).
Our hypotheses were that (1) the diversity of the
natural primary or secondary trees and (2) improvement
of soil conditions, should be highest under trees forming
the densest canopies, which most effectively suppress
grasses and herbs.
Study area
The Riam Kiwa plantation area is located in South
Kalimantan, Indonesia, 3 30' S, 115 E, at 100 - 200 m
a.s.l. The topography is undulating with red-yellow
podzolic soils (ultisols), deeply weathered and heavily
textured, and partially replaced by oxisols (Burnham
1984). The pH of non-forested grassland is 3.9 - 5.0,
total nitrogen 0.025 - 0.33 % and clay content 54 - 58 %.
Rainfall is ca. 2200 mm/yr, with a dry season from May
to September, with only 20 % of the annual total.
Imperata cylindrica grasslands cover the area since
World War II. In 1983, the Reforestation and Natural
Forest Management Project, part of the Indonesia-Finland Forestry Programme, established a trial area in
Riam Kiwa in order to study methods of reforestation.
Today, 60 % of the 1000-ha trial area has been reforested. Since 1983, more than 100 species with various
provenances have been tested; 10 - 15 species were found
promising for grassland reforestation and different plantation methods have been tested (Otsamo et al. in press).

Sampling and statistical analysis

17 compartments out of the total of 40 were sampled randomly for the inventory. The compartments
measured from 0.21 - 4.51 ha. 13 compartments were
plantations with one or two species planted 3 - 6 yr
before the inventory; one was a 0.5-ha natural stand of
Vitex pubescens; three, 0.5 ha each, represented the original Imperata grasslands (see Table 1).
The inventory was carried out systematically along
the planting lines - 2 m apart, including all occurring
vascular plant species, representing the life forms trees,
shrubs, climbers, herbs, grasses and ferns. The Vitex
pubescens stand and Imperata sites were sampled similarly. Species occurrence was expressed as frequency.
Due to the varying size of the compartments, the
effect of sample area on the number of species was
checked with curve-fitting regression analysis.
Floristic relations between the compartment understories were described through Principal Component
Analysis, based on the Jaccard similarity index (Jongman
et al. 1987). Compartments were grouped for analysis of
covariance according to the results of the PCA.
The two hypotheses mentioned in the Introduction
were tested with Covariance Analysis.

Results
151 vascular plant species, 89 trees, 19 shrubs, 13
climbers, 11 herbs, 9 grasses and 10 ferns, were found
growing spontaneously in the 5 to 6-yr old plantations.
Complete data are available upon request. Tree species
with a frequency > 50 % in the plantations include
Glochidion capitatum, Vitex pubescens, Alstonia angustiloba, Buchanania arborescens and Ficus grossularioides; the most frequent herbaceous species were Imperata cylindrica and Eupatorium pallescens. Both species
produce much above-ground biomass, particularly under open canopies. 18 of the tree species found in the
plantations, belonging to 11 families, were also found in
the Imperata cylindrica grasslands. They are apparently
adapted to the frequent occurrence of fires. Grasses
accompanying the dominant I. cylindrica include Saccharum spontaneum, Bromus insignis and Scleria spp.
Species-area relationships
The number of species appeared to be independent
of compartment size (Y = 27.12 + 5.262 log X; r2 = 0.14).
However, the number of tree species (Fig. 1) and the
arcsin-transformed ratio: tree species / total number of
all vascular plant species (Fig. 2) were significantly
related to log compartment size.

- Restoration of natural vegetation in degraded Imperata cylindrica grassland -

207

Table 1. Data on the 17 plantation compartments and natural

stands, with year of planting, tree species, area (ha), number
of tree species, NT, and other vascular plant species, VT.
No. Yr

Species

Area NT VT

A
B
C
D
E
F
G
H
I
J
K
L
M
N
15
16
17

Acacia mangium + Swietenia macrophylla

Paraserianthes falcataria
A. mangium
A. mangium
Gmelina arborea
A. mangium
A. mangium
Vitex pubescens (natural)
A. mangium + Shorea spp.
A. mangium
P. falcataria
Species trial
Eucalyptus deglupta
A. mangium + P. falcataria
Natural Imperata cylindrica (slope)
Natural I. cylindrica (valley)
Natural I. cylindrica (crest)

0.91
2.75
0.84
1.98
0.25
4.51
2.57
0.20
1.41
0.50
1.27
2.31
0.37
0.85
0.50
0.50
0.50

1987
1987
1986
1986
1991
1986
1986
1986
1986
1986
1986
1986
1987
-

13
16
10
29
6
24
35
4
16
16
25
14
3
13
4
9
10

1
21
6
5
18
3
2
8
6
22
31
3
6
26
10
12
7

Fig. 1. Regression of the number of tree species and the area of

the compartment. Regression coefficient is significant (p =
0.01).

Principal component analysis

The approach was considered justified since the
total number of species appeared to be statistically independent of sample area.
The results are shown in Fig. 3. No verbal interpretations are given to the principal components; instead, our
interest was in the configuration of different plantation
compartments in the floristic ordination. Three groups
were recognized.
1. Most of the pure and one-mixed Acacia mangium
compartments: D, F, G, I and J, are grouped together,
whereas the A. mangium compartment C, trial compartment L - where A. mangium has currently taken over as
a predominant species - and one of the Paraserianthes
falcataria trials (K) are in the vicinity, grouped together
with a mixed compartment of A. mangium and S.
macrophylla (A) and a P. falcataria compartment (B).
2. This group consists of a Eucalyptus deglupta stand
(M) and a mixed stand of A. mangium and P. falcataria
(N).
3. This group comprises the pure Gmelina arborea
stand (E) and a natural Vitex pubescens stand (H9 which
are clearly separated from the others. The Vitex and
Gmelina stands resemble each other in their very low
numbers of understorey species. In this group A. mangium
is missing.

Fig. 2. Regression of the ratio: number of tree species / total

number of vascular plant species in the area of the compartment. Regression coefficient is significant (p = 0.01).

Analysis of covariance
The arcsin- transformed percentage of the number of
tree species out of the total number of all understorey
vascular plant species was compared in the three
floristically different subsets of samples, pure Acacia
mangium stands, mixed A. mangium stands and stands

Fig. 3. Diagram of the PCA of compartments A - N (see Table

1) for the first two principal components. Variance explained
by Axis 1 = 16.9 %, by Axis 2 = 14.5 %.

208

Kuusipalo, J. et al.

where A. mangium was not planted. Due to the dependency of the dependent variable on the size of the sample
plot (see Figs. 1 and 2), compartment size was used as a
covariate in the analysis (Table 2).
The result is quite straightforward. Both stand type
and compartment size have a reasonably significant
effect with a statistical risk level of about 1/20 on the
ratio between the number of tree species and the number
of all species. Multiple a posteriori comparison of means
(LSD) reveals that both pure and mixed A. mangium
stands differ significantly from all other stand types but
not from each other. In other words, the proportion of
tree species is significantly higher in pure or mixed A.
mangium stands than in the rest of the stands, where
grasses and other herbaceous vegetation prevail.

Discussion
Reforestation of Imperata cylindrica grasslands is
extremely difficult due to compact and nutrient-deficient soil, hydrologic instability, large variation in surface temperatures of the soil, grass competition and
allelopathy, and high fire susceptibility of the grass
(Soerianegara 1980; Ohta 1992; Parrotta 1993). Natural
succession processes are prevented by annual fires,
which, together with other conditions listed above, destroy plant propagules and seedlings, as well as fungal
root symbionts.
Problems of reforestation have already been studied
intensively in the Indonesian-Finnish trial area of Riam
Kiwa for more than ten years. More than 100 species
have been tested, including fast-growing exotic plantation species already mentioned above, but also easily
propagated indigenous trees have been planted and evaluated in the area, including dipterocarps such as Shorea
spp., Hopea spp. and Vatica spp., and others, e.g. Durio
spp., Peronema canescens, Artocarpus spp., Macaranga
spp. and Agathis borneensis.
Several exotic species of pioneer tree character have
shown promising reforestation results in terms of growth
as well as in suppressing I. cylindrica. However, a
successful establishment of these species requires rather
heavy site preparation, which turns over the rhizomes of
I. cylindrica and improves the rooting of tree seedlings
(Otsamo et al. in press). On the other hand, reforestation
with native climax species directly on grassland has
proved impossible or at least very laborious and expensive (e.g. Appanah & Weinland 1993). Low survival of
these most probably results from shallow rooting unable to penetrate into compact soil, and from the lack of
adequate fungal root symbionts (e.g. ectomycorrhizae
of dipterocarps). Rehabilitation of original vegetation
seems feasible only if environmental conditions are im-

proved with the help of tree plantations; native species

or agricultural crops can be interplanted later within the
rows of plantation trees (Parrotta 1992; Lugo 1993).
This is an important option for land rehabilitation programmes (e.g. Anon. 1991).
An interesting phenomenon is the rapid spontaneous
emergence of indigenous trees and other plant species
under the plantations of exotics. The propagules of these
plants must have been dispersed into the plantations by
animal vectors or by wind from the narrow strips of
natural vegetation along the creeks in the trial area or
from the larger forests in the mountains several km
away. The plants found in the plantations include trees,
shrubs, climbers, herbs, ferns and grasses, and they may
be pioneer, secondary or primary forest species.
The plantations create microclimate and soil conditions which improve the germination and survival of
species unable to germinate in the open or within
I. cylindrica grass cover. Most of the tree species spread
naturally to the plantation area are fruit-bearing plants,
with animal vectors, notably birds and bats as the principal means of dispersal. This suggests that plantation
canopy promotes the regeneration process by providing
roosting habitats for seed-dispersing animals. Others
have light-weight seeds dispersed easily by wind over a
long distance. Plants characterized by a heavy-weight
seed with no animal vectors, such as most of the dipterocarps, are absent in the spontaneous undergrowth
(Whitmore 1972-1978). Our results thus give further
support to the findings from Central America presented
e.g. by Lugo et al. (1993) and Parrotta (1993).
The majority of the tree species dispersed spontaneously to the plantations belong to the pioneer or secondary forest species characterized by effective means of
rapid dispersal. However, there were also 17 tree species typical of primary forests, including Buchanania
arborescens with a frequency of 57 %, Diospyros buxifolia, Litsea firma, Dacryodes rostrata, Gluta wallichii,
Sandoricum borneense and Stergulia rubiginosa. These
species occur frequently in the nearby patches of natural
forest vegetation still persisting on hilltops and deep
gullies along the brooks and rivers (Whitmore 19721978).
The proportion of trees was largest in pure or mixed
stands of Acacia mangium. This tree is known as a very
effective species in suppressing light-demanding grass
and herb vegetation, mainly due to its dense crown,
which also creates a moister and cooler microclimate,
and abundant litter fall (Anon. 1983; Otsamo et al. in
press). It is evergreen even during a prolonged dry
season in climatic conditions typical of humid tropics
(Anon. 1983). A. mangium is a nitrogen-fixing legume;
nitrogen-rich litter increases the amount of organic input into the soil maintaining its biological activity and

- Restoration of natural vegetation in degraded Imperata cylindrica grassland improving the nutrient status (Sanchez & Miller 1986).
Phosphorus uptake is efficient due to the symbiosis with
compatible vesicular-arbuscular (VA) mycorrhizal fungi
(Malloch et al. 1980). All in all, A. mangium seems to
favour shade-tolerant, relatively nutrient- and moisturedemanding tree species at the expense of light-demanding herbs and grasses in its understorey.
Inside the Eucalyptus deglupta plantation the light
intensity was seemingly much higher than in A. mangium
plantations of the same age, and the amount of I.
cylindrica grass consequently much larger. Poor shading and vigorous competition by grass may have inhibited the germination and survival of the indigenous plant
seedlings. Plantations of Paraserianthes falcataria provided better shading than E. deglupta, and I. cylindrica
had been mostly replaced by other plants. However, the
stand was light enough to allow the weed Eupatorium
pallescens to become dominant, followed by Clibadium
spp. as the second commonest shrub. The shrubby understorey of P. falcataria is also pyrogenic, making the
stand very susceptible to fire during the dry season.
Some of the promising tree species were perhaps
underrated in this study due to a limited sample of
stands; e.g. Gmelina arborea was represented just by
one recently established compartment, although it is
known to have a good capability to suppress the grass
(Otsamo et al. in press).
Successful plantations seem to be able to act as
initial steps in the secondary succession of I. cylindrica
grasslands back to natural forests by allowing many
indigenous forest tree species to colonize the site within
a relatively short period of time. Acacia mangium is a
particularly suitable species because it turns the resource supply ratio into a more favourable one for
woody perennial undergrowth: light becomes a more
limiting resource than nitrogen, moisture and other soil
resources. Grasses and herbs cannot take advantage
from improved soil conditions due to their generally
higher light demand compared to woody perennials
which, in turn, can only establish on an improved soil
(Tilman 1982).
The more woody, evergreen undergrowth, the lesser
susceptibility to fire damage. This is a rule of thumb
which has, for a long time, been appreciated by foresters
and forest scientists working in Imperata areas, as well
as elsewhere in the tropics (Lugo et al. 1993; Parrotta
1993). A. mangium seems to provide an alternative
which creates suitable conditions for an abundance of
understorey trees. The results of this study support
suggestions made by Parrotta (1993), i.e. that forest
plantations on degraded tropical lands can be used not
only for production of wood and other forest commodities but also for soil improvement and acceleration of
secondary forest succession. Species and provenance

209

Table 2. Analysis of covariance and a posteriori comparison

of means (LSD) of the Riam Kiwa data. Legend to variables:
ASNREL (Dependent variable): Ratio between number of tree species
and total number of vascular plant species; AREA (Covariate): Area
of compartment (ha); EXTRA (Independent class variable):
1. Stands without Acacia mangium (n = 5)
2. Pure A. mangium stands (n = 5)
3. Mixed A. mangium stands (n = 4)
Analysis of covariance
Source

Sum-of squares

Extra
Area
Error

0.445
0.284
0.571

DF

2
1
10

Mean-square

0.223
0.284
0.057

F-ratio

3.901
4.976

0.056
0.050

Fishers least-significant difference test

1. Stands without A. mangium
2. Pure A. mangium stands
3. Mixed A. mangium stands

1.000
0.045
0.031

1.000
0.867

1.000

selection, together with plantation design and silvicultural

practices, play a key role in realizing these objectives
(Otsamo et al. in press). Nitrogen-fixing species such as
Albizia lebbek used by Parrotta (1993) in Costa Rica, or
Acacia mangium in the present study, can have a dramatic effect on soil fertility through their production of
decomposable, nutrient-rich litter and turnover of fine
roots and nodules.
The influence of the proximity of natural forests on
seed recruitment into plantations poses a special problem when using fast-growing tree plantations as foster
ecosystems (Parrotta 1993). Understanding of seed ecology, including dispersal characteristics and habitat requirements for germination and seedling development
is of particular importance in restoration plantation design. Many important species, e.g. most of the dipterocarp
trees predominating Southeast Asian moist forests have
heavy, seeds and, subsequently, spread very slowly into
areas where mother trees have completely disappeared
(Appanah & Weinland 1993). Planting of dipterocarps
and other primary forest species characterized by a slow
dispersal within the shelter of plantation tree canopies
and spontaneously grown natural trees would, therefore,
speed up the restoration process considerably.

Acknowledgements. We thank Mr. A.P.S. Sagala and two

anonymous referees for their invaluable comments, and Dr.
John A. Parrotta for sending us literature we needed for the
completion of the study at our remote base. The study was
carried out under the auspices of the Reforestation and Tropical Forest Management Project financed by the Finnish International Development Agency (FINNIDA) and implemented
as a joint effort of the Ministry of Forestry, Indonesia, and
Enso Forest Development Ltd., Finland.

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Kuusipalo, J. et al.

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Received 1 July 1994;

Revision received 3 November 1994;
Accepted 14 November 1994.