Six Decades of Natural Vegetation Studies

in Indonesia

Kuswata Kartawinata

INTRODUCTION

Vegetation is defined as the mosaic of plant communities in the landscape

and the natural vegetation refers to the vegetation which exists in the
landscape unaffected by man (Kchler, 1967). Taylor and White (1958)
acknowledged that vegetation studies may be made purely for scientific
reasons, but many of them are for definite purposes to obtain knowledge to
aid mans control of environment, such as land classification for the use in
agriculture, forestry, and grazing and for the latter two cases knowledge of
successional changes is of importance. It should be noted that land use centers
around the growth of plants, such as forest plants, pasture plants, and crops,
hence land classification should necessarily consider factors affecting plant
growth, i.e., climate, geology, geomorphology, soil and vegetation. Full
understanding of these factors and their inter-relationships will enable land use
planner to determine the best use of the land resources in relation to
sustainable development.
Indonesia extends over 5,000 km from West to East and its land area
comprises a diversity of dryland and swamp vegetation. Biological investigations,
including vegetation studies, in Indonesia have barely scratched the surface,
although they date back to the beginning of the eighteenth century. A historical
account of the research undertakings prior to 1945 is summarized in Science
and Scientists in Netherlands Indies (Honig and Verdoorn, 1945) and later in
Chronica Naturae, volume 106 (6) in 1950. Vegetation and plant ecological
investigations were concerned primarily with floristic and faunistic exploration
and inventory, descriptive accounts based on visual observations, natural his-
tory, and to a lesser extent, with ecological factors. Most of these were
published in Tropische Natuur, Natuurkundig Tijdschrift voor Nederlandsch

95
Indie, Treubia, Bulletin du Jardin Botanique de Buitenzorg, Annales du Jardin
Botanique de Buitenzorg.
The state of vegetation research up to 1980 was reported by Kartawinata
(1990) who evaluated available references on vegetation and plant ecology in
Indonesia revealing that this field have not been much studied. Much of plant
ecological information in various literatures such as the series of books on
Ecology of Indonesia (e.g., MacKinnon et al., 1996 and Whitten et al., 1984)
have been based on investigations carried out in Malaysia. The studies have
been largely concentrated on forests, particularly the lowland dipterocarp
forests. Much of the information for phytogeographic region Malesia (Brunei,
Indonesia, Malaysia, Philippines, Papua New Guinea and Timor Leste) have
been synthesized by Whitmore (1984) in his Tropical Rain Forest of the Far
East. Vegetation data on Papua New Guinea (e.g., Paijmans, 1976; Gressitt,
1982; Johns, 1985, 1987a & b; Brouns, 1987; Grubb and Stevens, 1985) provide
information on vegetation, biogeography and ecology applicable to Papua.
The following account is a one-person brief review of the state of research
on natural vegetation and plant ecological studies in Indonesia during the last
six decades. It is unavoidably biased and in no way constitutes an exhaustive
and critical review and is more like a bibliography, but I hope I have identified
the state of the development of the studies that can provide information on
what have been done and data sources for further analysis and synthesis on
vegetation in particular.

VEGETATION AND OTHER PLANT ECOLOGICAL STUDIES

Botanical studies in Indonesia date back to the pre-Linean time when

Rumphius initiated the work on Herbarium Amboinense in 1662 (Steenis, 1950).
Vegetation studies and plant ecological' investigations, however, have barely
started and only during the last three decades the work on the subject by
resident and visiting specialists has increased substantially.

Vegetation mapping
Published vegetation maps covering Indonesia include the Atlas van Tropisch
Nederland (Steenis, 1938), Vegetation Map of Malaysia (Steenis, 1958),
Vegetation Map of Malesia (Whitmore, 1984b) and Map of Forest of Indonesia
(Hannibal, 1950 cited in Dilmy and Kostermans, 1958). The latter was based
on Steenis' 1935 map and the results of the field strip surveys made by the
Forest Service. Some of the surveying strips were maintained as permanent
plots until early 1970s when regrettably the logging activities encroached the
area.
As a byproduct of the intensified forest inventory of commercial timber trees
throughout the country, complemented with aerial photographs, to meet the
increasing needs of logging concession, the Directorate of Forestry Planning or
Direktorat Bina Program Kehutanan (1980) published a Map of Forest Stands of

96
Indonesia (scale 1: 2,750,000), To date the mapping activities have been
continued by the Badan Planologi Kehutanan (Agency for Forestry Planning)
Meijer and Withington (in Withington, 1981) prepared a map of natural
vegetation and of human occupancy of Sumatra based on LANDSAT images
and used Steeniss 1958 map as a base map. BIOTROP and the "Institute de
la Carte International du Tapis Vegetal" published vegetation and bioclimatic
maps of Sumatra (scale 1: 1,000,000), based on satellite imageries and field
surveys' data (Lamounier et al., 1983; 1986a,b). Other maps published recently
include the "Land Use Map" (scale l :200,000) (Tata Guna Tanah, 1980) and
the Original Vegetation Types of the Sundaic and Wallacean Sub-regions
(MacKinnon and MacKinnon, 1986). The latter was used by Whitmore (1984b)
as one of the bases for his Vegetation Map of Malesia. In recent years various
NGOs , projects and private companies have published vegetation and land-
use maps of limited areas, such as those for Leuser Ecosystem by the Leuser
Ecosystem Management and the copper and gold mining concession of the
Freeport Indonesia.
Large-scaled vegetation maps often complement the management plans of
various national parks, nature reserves and other protected areas, prepared by
Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam (the Directorate-
General of Nature Conservation and Forest Protection) in cooperation with
various institutions such as FAO, CII (Conservation International Indonesia),
TNC (The Nature Conservancy), WCS (Wildlife Conservation Society) and WWF
(World Wide Fund for Nature) Indonesia Program. The floristic composition of
various vegetation types, however, is often lacking, except for the list of
common species. The most detailed large-scaled vegetation map is that of the
Ujung Kulon National Park (scale I: 75000) and each vegetation type is
complemented with floristic enumeration and character species (Hommel, 1987).
The RePPProT (Regional Planning Program for Transmigration) produced
accurate land use, land suitability and land capability maps (scale 1: 250,000).
As I checked some of the maps on the ground, I found that they are good
indicators of vegetation types, but unfortunately floristic indicators are lacking.
The maps could constitute a good base for further detailed vegetation studies.
Vegetation maps on wetlands are contained in survey reports prepared by
Wetland International Indonesia Program.
The vegetation maps become quickly obsolete in just a few years even
months, due to the rapid destruction of millions of hectares of natural
vegetation, especially lowland forests, throughout the country. It is attributed
to logging, development projects (e.g., tree and crop plantations, and
transmigration), smallholder conversion including shifting cultivation, and fire
loss.

Vegetation types
The work of Steenis (1935) on "Maleische vegetatieschetsen" is by far the
most complete, qualitative, and detailed account of the vegetation of Indonesia
before the 2nd World War. It includes also various ecological factors responsible

97
for the patterning of the vegetation, correlating the climate, soil, geology, water
availability and soil with vegetation types. Steenis (1957a) used habitat factors
as the basis for constructing the outline of vegetation of Malesia, and it is
adopted also by Whitmore (1984). It has been incorporated into the plant
geography of Java (Steenis 1965) and further refined and applied to Indonesia
(Kartawinata, 1989); the latest version is attached as Table 1. Earlier,
Kartawinata (1978b) prepared also the ecological zones of Indonesia and later
he made a forest classification scheme (Kartawinata, 1981a). Steenis (1965,
1972) provided qualitative descriptions of vegetation types of Java and
Kartawinata ( 1994a) has published a brief account on forest types of
Indonesia, while the vegetation of Indonesia is currently in preparation
(Kartawinata, in prep.)
The vegetation of Indonesia is composed of plant formations which are
related to soil types, soil water condition and elevation and are dominated by
tropical forest (Table 1). Climatically these formations are located within the
everwet and monsoon (seasonally dry) regions. The everwet region,
dominated by the rain forest, occupies areas with per-humid climate, where the
index Q or the ratio between dry and wet periods (Schmidt and Ferguson's,
1951) ranges between 0 and 33.3 percent, thus having only slight dry season.
The monsoon region, dominated by monsoon forest, covers areas with Q
ranges between 33.3 and 300 percent, showing markedly and seasonally dry
period, in which water situation becomes seasonally and seriously limiting to
plants.

Lowland vegetation studies

The progress of the vegetation studies up to 1958 was reported by Dilmy
and Kostermans (1958). During the last three decades vegetation research
along with botanical explorations and other botanical studies have been
undertaken throughout the country but mostly in Kalimantan, Sumatra and
Sulawesi. Table 2 shows the list of 90 sites where quantitative phytosociological
studies using 223 plots ranging in size from 0.04 15 ha to enumerate trees
with DBH (Diameter at Breast Height) 10 cm reported in various publications
and unpublished data. The studies include botanical explorations, enumeration
of tree species and ecology of the effects of human activities on the lowland
forests. Tree species enumeration within plots was aimed at gathering
information on floristic composition of various vegetation types in relation to
habitat conditions and indication of species diversity in the explored areas and
ecology of the effects of human activities on the lowland forests. They were
conducted in dryland forests, peat swamp forests, freshwater swamp forests,
heath forests and mangrove forests. So far no records on the investigation of
limestone forests and forests on ultrabasic soils, which are otherwise
ecologically interesting.
I believe that there are others whose data are unpublished or published in
journals and reports that are not available to me. The quantitative study by
Yamada (1975) in the montane forest of Mt. Pangrango, West Java, was

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seminal and followed eventually by Kartawinata et al. (1981) and Riswan
(1982) in dipterocarp forest of East Kalimantan. To date there has been no
attempt to do computer analysis of floristics and synthesize all data on national
scale. The data in various plots as listed in Table 2 can be synthesized in that
fashion, but different methods and plot sizes used in the sampling may
present problems.
Data that include all plants and trees with DBH 10 cm, such as those from
Karimun Jawa (Djarwaningsih et al., 2003), Nusakambangan (Partomihardjo and
Prawiroatmodjo, 2003) and Nusa Tenggara Barat (Yusuf 1996), and those
collected with plotless sampling method ( e.g., Abdulhadi, 1978; Kartawinata,
1980; Kartawinata et al., 1985; Mackie 1986; Setiadi, 2005) are not included in
Table 2. In addition many reports and articles based on qualitative observations
on vegetation usually made during botanical explorations (e.g., Hidayat and
Wiriadinata, 2003; Iwatsuki et al., 1980; Kalkman, 1963; Kalkman and Vink,
1970; Kartawinata, 1965; Kostermans, 1960; Kostermans, 1965; Kostermans,
and Kartawinata, 1961; Mogea, 2002; Noteboom, 1987; Rugayah et al., 2002,
2003; Wiriadinata, 1996) and ethnobotanical studies (e.g., Purwanto and
Fanani, 2003), are interesting and important (see also Flora Malesiana Bulletin
1950-2004).
Most of the above studies have been carried out by the Lembaga Biologi
Nasional (National Institute of Biology), now Pusat Penelitian Biologi (Research
Center for Biology) of LIPI (Lembaga Ilmu Pengetahuan Indonesia,
Indonesian Institute of Sciences), the results of which have been reported in
the series of Technical Reports 1970-2004 or appeared in the institutes journals
Reinwardtia (a peer-reviewed international journal) and Berita Biologi as well
as in Ekologi Indonesia, a journal published by the Himpunan Ekologi
Indonesia (Indonesian Ecological Society) and Indonesian Journal of Biology of
the Indonesian Biological Society. Occasionally they appeared in international
journals. Unfortunately the data presentation in and editing of the recent
Technical Reports are poor that make data contained therein often difficult to
synthesize.
As indicated above Direktorat Bina Program Kehutanan (now known as the
Agency of Forestry Planning) of the Ministry of Forestry since mid-1960s has
intensified forest inventories throughout the country and has produced an
enormous amount of data, but unfortunately the collection and treatment of
data were done as such that made the data and information unreliable,
including the species identification. However, there are some good results of
inventories undertaken prior to mid 1960s such as that presented by Dilmy
(1965) on peat swamp forests of Sampit, Central Kalimantan. Pusat Penelitian
Hutan dan Konservasi (Center for Forest and Conservation Research) of Badan
Penelitian dan Pengembangan Kehutanan (Forestry Research and Development
Agency, FORDA) has done similar work emphasizing commercial timbers and
other aspects of forestry in various parts of the country (e.g. Jafarsidik 1980).
The results are usually reported in the Forest Research Bulletin published by
the institute. One should also refer to the work of Goor and Kartasubrata

99
(1982) who wrote the Indonesian Forestry Abstracts of Dutch literatures until
about 1960. Forestry Faculties and Biology Departments of various universities
did undertake vegetation studies and the results have appeared in currently
mushrooming biological journals (in Indonesian) published by various
universities. Unfortunately the quality of most papers and editing of local
journals need a great deal of improvement. Consequently poor articles do not
attract international scientists who often considered such articles as grey
literature or even thrash, hence they do not enter international circulation.
PTFI or P.T. Freeport Indonesia (1997) launched a biggest biodiversity
study ever undertaken in Papua and even in Indonesia in its mining concession
area. The purpose of this study was to document the current status of
biodiversity in the PTFI Contract of Work Area adjacent to the western side of
the Lorentz National Park covering approximately 3,000 km2, from the Grasberg
mine at the altitude of about 4,000 m to the coastal area at sea level to provide
baseline data for long-term management of flora and fauna in the region. The
study involved international team of scientists from the Indonesian Institute of
Sciences (LIPI), University Cendrawasih in Manokwari (UNCEN), Bishop
Museum (Hawaii), Western Australian Museum (Perth), Smithsonian Institution
(Washington, D.C.), and several other research organizations in Indonesia. The
study shows that the montane, subalpine, alpine and nival zones are rich in
biodiversity, including ecological diversity, organismal diversity, and genetic
diversity. The lowland zone (0 650 m above sea level) was subdivided into 7
subzones, comprising beaches, tidal swamps, meander belts, peat swamps,
alluvial fans, alluvial valleys, and dissected terraces within which 250
communities were identified.
Pusat Penelitian Biology has established permanent sample plots of primary
forests in 17 stations in Java, Kalimantan and Sumatra (Mirmanto et. al., 2005;
Suzuki et al., 1998). Badan Penelitian dan Pengembangan Kehutanan has
established several Balai Penelitian Hutan (Forestry Research Institute) in
various provinces and each has and manages a set of permanent sample plots
in forest areas within its jurisdiction. CIFOR (Center for International Forestry
Research) has established 24 permanent sample plots of one-hectare each,
emphasizing trees with DBH 20 cm at Seturan, Malinau, East Kalimantan
(Sist et al., 2002). Similarly Cirad-Fort and PT INHUTANI has a series of
permanent sample plots at Berau, East Kalimantan (Sist and Saridan 1999).
Harvard University has established a field station at the Gunung Palung National
Park, West Kalimantan (Leighton and Darnaedi, 1996) managing many
permanent sample plots. Similarly Cambridge University in collaboration with
FORDA has a field station with many permanent sample plots at Barito Ulu,
Central Kalimantan (Chivers, 1992). Permanent plots to monitor successional
changes in Krakatau were established by Oxford University and Pusat Penelitian
Biologi (Whittaker et al., 1995; 1998) Permanent plots have been used to
facilitate long-term studies and monitoring on vegetation, forest dynamics,
nutrient cycling, phenology, habitat preferences, ethnobotany, ecology of
species, growth studies, effects of logging, silviculture, and carbon sequestration.

100
They can be used also as sites for collecting samples to test medicinal values
of species such as anticancer and anti-AIDS testing of plants and relate them to
the ecology of the species (Soejarto 1996), although this has not been done in
Indonesia. Figure 7 shows an example of the value of a lowland forest in two
one-hectare permanent plots in Jambi in terms of category of use of non-
timber forest product (Mirmanto et al., 2005).
Various foreign universities, especially those from United States, Japan and
United Kingdom, have been taking part in undertaking botanical studies and the
results may be found in various international journals. International conservation
NGOs operating in Indonesia, while they need vegetation data for their programs,
have paid little attention to vegetation studies.
Several studies on peat swamp forests have been undertaken (e.g.,
Anderson, 1976; Mirmanto et al., 1993, 2003; Siregar, 2002). Many papers
discussing peat swamp forest ecology and other related aspects from
Kalimantan and Sumatra are presented in a proceedings of symposium on
tropical peatlands (Iwakuma et al., 2000) and on land management and
biodiversity (Osaki et al. 2002).
Mangroves on mud deposits are the most well studied vegetation, but those
on sand deposits and coral reefs received little attention, hence the information
is limited (see Budiman et al., 1986; Kartawinata and Waluyo, 1977). Their
floristic composition and structure are well known although they vary from one
place to another, contingent upon habitat conditions. Steenis (1957b) synthesized
available qualitative data and published the result as an introduction to
Rhizophoraceae in Flora Malesiana. More quantitative ecological data are
contained in several volumes of proceedings of national seminars since 1978
(e.g., Soemodihardjo et al., 1979, 1984; Soerianegara et al., 1986), review
(Soemodihardjo, 1985), research report (Ogino and Chihara, 1988) and other
separate papers published elsewhere. A good treatise on mangrove restoration
in Sumatra is presented by Soemodihardjo et al. (1996). Noor et al. (1999)
made a good account on mangrove vegetation and field guide to flora.

Montane vegetation studies

Floristic enumeration of montane vegetation has received little attention. In
Java, Steenis (1972) synthesized available information on vegetation and
ecology. Quantitative assessment was carried out in Mt.Gede-Pangrango by
Meijer (1959), Yamada (1975, 1976a,b, 1977, 1990) and Abdulhadi et al.
(1998), in Mt. Slamet by Abdulhadi (1978), in Mt. Halimun by Simbolon and
Mirmanto (1997), Mirmanto and Simbolon (1998), Rahayoe et al. (2004),
Yoneda et al. (2001) among others. Simbolon and Mirmanto (1997) did
vegetation analyses along altitudinal gradients and confirmed quantitatively the
currently accepted altitudinal montane vegetation zonation of lower montane,
upper montane, lower subalpine and subalpine forests of Steenis (1972).
Unpublished report of a UNESCO training course on vegetation studies in 1976
gave good data on the forest of Mt. Gede. There have been a number of
vegetation studies carried out by undergraduate and graduate students in this

101
mountain, but the results on floristic composition of some of them are
unreliable. Outside Java, quantitative floristic assessment has been limited to
East Kalimantan (Bratawinata, 1986), Sumatra (Ohsawa et al., 1985), Maluku
(Edwards et al., 1990) and Papua (Hope et al., 1976). Bratawinata shows the
altitudinal variation of primary forest in the Apo Kayan and identified six
vegetation types. Here Dipterocarpaceae is still dominant. Ohsawa et al. (1985)
studied the altitudinal zonation and identified three forest zones between 1750
and 2950 m altitude and a scrub zone between 2950 and 3250 m. In the
Manusela National Park, Seram, Maluku, Edwards et al. (1990) studied the
altitudinal zonation of the rain forest and divided the vegetation and soil into
five zones, i.e., lowland, lower montane, montane, subalpine and alpine. The
only comprehensive study of high altitude vegetation was carried out by Hope
et al. (1976) in the Sudirman Range, Papua, particularly around the
permanently snow-capped Mt. Jaya. Not less than 24 community types
occurring on varied habitats could be identified, ranging from upper montane
forest to alpine tundra. They represent typical vegetation on the highland of
Irian Jaya and perhaps of New Guinea. Mangen (1933) discussed also the
ecology and vegetation of Mt. Trikora.
PTFI (1997) identified in the montane zone over 80 communities which can
be divided into three main formations, based on structure and floristic, i.e., the
lower montane forest formation at 650-1500 m, the mid montane forest
formation at 1500-2800 m, and the upper montane forest formation at 2800-
3200. The Subalpine Zone occurs from 3,200-4,170 m above sea level, and is
divided into two subzones: the lower subalpine subzone from 3,200-3,650 m
and the upper subalpine subzone from 3,650-4,170 m. The Alpine Zone occurs
from 4,170-4,585 m, and the Nival Zone occurs above 4,585 m. Within each
zone and subzone, numerous plant communities can be defined based on
structural formations, dominant species, and successional stage. These occur
along a number of complex environmental gradients.

Patterns of species richness and vegetation

Species richness
The species richness varies a great deal, as indicated also by MacKinnon
(1982) for various forest types. Species richness is a good indicator of species
diversity which is comparable to index of diversity (Gentry, 1988). In some
instances, however, the low number of species may not necessarily indicate low
species richness or low diversity, but may be attributed to inadequacy of field
identification as well as to different techniques of establishing plots, where data
on density and number of species from vertical projection of the plots are
higher than those from plots laid down on ground surface. In some cases
identification uses mainly vernacular names without complete set of voucher
specimens; the results are definitely unreliable, since only few species have
consistent vernacular names.

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 450
WEST MALESIA EAST MALESIA SOUTH MALESIA
400 386

350
NUMBER OF SPECIES

300

250 239

203
200
147 155
150 125

100 79

50 37
10 8 12 10 9 3 1
0
Sum Mal Kal Sar Sab Brun Phi Sul Mlk Pap PNG Jav Bal NT Tot
COUNTRY/ISLAND/REGION

Figure 1. Number of species and distribution of Dipterocarpaceae in Malesia (Brunei,

Indonesia, Malaysia, Philippines, Papua New Guinea and Timor Leste)
showing that the majority of species occur in West Malesia. Bal Bali; Brun
Brunei; Jav Java; Kal Kalimantan; Mal Malaysia Semenanjung;Mlk
Maluku; NT East and West Nusa Tenggara; Pap Paua; Phi Philippines;
PNG Papua New Guinea; Sab Sabah; Sar Sarawak; Sul Sulawesi; Sum
Sumatra; Tot Total.

The rain forest is prevalent over most part of Indonesia. In this forest there
are considerable floristic variations which are related to differences in soils and
topography Broader variations are also reflected by the horizontal zonation
(related to geologic history) and vertical zonation (related to elevation). The
main types of the tropical forest of Indonesia include the lowland rain and
monsoon forests, lower montane forest, upper montane forest, subalpine
forest, heath forest, limestone forest, forest over ultrabasic rocks, beach forest,
mangrove forest, peat swamp forest and fresh water swamp forest.
The rain forest has the richest flora and shows also a much greater local
endemism than elsewhere. These conditions are likely attributed to a greater
stability of the climate than anywhere else in the humid tropical region
(Whitmore, 1984a). The flora belongs to the phytogeographic region of
Malesia. The Malesian flora differs distinctly from that of the adjacent regions.
Steenis (1950) has shown that the flora contains about 300 endemic genera
and the floristic boundary is very sharp, which is marked by distinct
demarcation knots. Whitmore (1984) has indicated that strong knots coincide
with the boundaries of the major forest types, and where the knot is weakest,
such as eastward in the Pacific, there is no such distinct boundaries of the
forest types.

103
 500
441: Samboja, E. Kalimantan
450

400
NUMBER OF SPECIES

350

300

250 221: Malinau, E. Kalimantan

200 182: Batang Gadis, N. Sumatra

150
87: Lore Lindu, N. Sulawesi 104: Mt. Gede, W. Java
100
71: Mt. Halimun, W. Java
50
72: Wanonii Island , S.E. Sulawesi
0
0 1 2 3 4 5 6
AREA(HA)

Figure 2.Species-area curves of various lowland and montane forests. The sources of
data are Kartawinata et al. (2004) for Batang Gadis; Kartawinata et al.
(unpublished) for Malinau and Mt. Gede; Purwaningsih & Yusuf (2005) for
Lore Lindu; and Simbolon & Mirmanto (1997) for Mt. Halimun.

Although smaller in extent than that of tropical America and Africa the
Malesian forest is exceedingly rich in species, richer than either one of them
(Whitmore, 1984a). The number of flowering plant species is estimated to be
about 25,000, or ten percent of the world flora, of which 3000 - 4000 species
are Orchidaceae (Steenis, 1971) and the majority of them occur in Indonesia.
Dipterocarpaceae is one of the large woody families and contains 386 species
(Ashton, 1984) which form the upper canopy and dominate the rain forest of
West Malesia. The role of Dipterocarpaceae, however, is less important towards
the east (Ashton, 1984). Other large woody families include Moraceae
(especially Ficus) and Myrtaceae (especially Eugenia), each with about 500
species and Ericaceae (Rhododendron and Vaccinium) with 740 species
(Whitmore, 1984). The richness of the flora is attributed to some trees that
provide environment to smaller trees and plants to grow. The richness of the
flora, for example, is clearly indicated by the great number of species growing
together and the number increases as the area increases (see Figure 2).

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Table 1. An outline of vegetation type of Indonesia

CLIMATE ELEVATION WATER SOIL VEGETATION TYPE

STATUS

I. Everwet Littoral (< 0 m) Saline water 1. Vegetation on saline water

Q<60 <0m Saline water Coral and sand; Wet 1.1. Littoral vegation
(Type A, B, <0m Saline water Alluvial; Wet 1.2. Mangrove forest
C); Lowland: Stagnant fresh Alluvial; Wet 2. Freshwater swamp vegetation
Annual 0 (750) 1000 m water
rainfall: Stagnant fresh Alluvial; Wet 2.1. Fresh water herbaceous vegetation
100-1700 mm water
Stagnant fresh Alluvial; Wet 2.2. Freshwater sago swamp vegetation
water
Stagnant fresh Alluvial; Wet 2.3. Fresh water swamp forest
water
Stagnant fresh Alluvial; Wet 2.4. Riparian swamp forest
water
Stagnant peat 3. Peat swamp vegetation
water
Stagnant peat Organosol; Wet 3.1. Peat swamp forest
water
Stagnant peat Organosol; Wet 3.2. Herbaceous peat swamp vegetation
water
Dryland 4. Dryland vegetation
Dryland Regosol; Dry 4.1. Sandy and/or rocky beach herb
vegetation
Dryland Regosol, Red Yellow 4.2. Sandy and/or rocky coastal forest
Podsolic; Dry.
Dryland Red Yellow Podsolic, 4.3. Lowland dipterocarp forest
Latosol; Dry
Dryland Red Yellow Podsolic, 4.4. Lowland non-dipterocarp forest
Latosol; Dry
Dryland Podsol; Dry 4.5. Kerangas (heath) forest
Dryland Renzina; Latosol; Dry 4.6. Limestone forest
Dryland Soils with high Mg, Fe, 4.7. Forest on ultrabasic rocks
and Si content; Dry
Highland/mountain: 5.. Montane vegetation
(750) 1000-4500 m Dryland 5.1. Montane forest
(750) 1000-2500 m Dryland Red Yellow Podsolic, 5.1.1. Lower montane forest
La-tosol, Andosol,
Lithosol; Dry
(1500) 2500-3300 m Dryland Red Yellow Podsolic, 5.1.2. Upper montane forest
La-tosol, Andosol,
Lithosol; Dry
2400-3800 m Dryland Organosol, Lithosol; 5.1.3. Lower sub-alpine forest
Dry
3800-4100 m Dryland Lithosol; Dry 5.1.4. Upper sub-alpine forest
Dryland 5.2. Sub-
alpine shrub and herb vegetation
3300-3800 m Dryland Lithosol, Limestone, 5.2.1. Forest-edge grasslland shrubland
Dry

3200-3700 m Dryland Lithosol; Dry 5.2.2. Tussock grassland with tree ferns

3300-4100 m Dryland Lithosol; Dry 5.2.3. Tussock grassland

3800 m Dryland Rocks; Dry 5.2.4. Lichen vegetation
3800-4100 m Dryland Limestone, Lithosol; 5.2.5. Moss vegetation
Dry
> 3800 m Dryland Lithosol, Rocks; Dry 5.2.6. Rock overhanging vegetation

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Table 1. continued

WATER
CLIMATE ELEVATION SOIL VEGETATION TYPE
STATUS
Stagnant peat/ fresh 5.3. Subalpine swamp vegetation
water
3300-4000 m Stagnant peat/ fresh Organosol; Wet 5.3.1. Grass bog
water
3400-4100 m Stagnant peat/ fresh Organosol; Wet 5.3.2. Herbaceous bog
water
3900-4200 m Stagnant peat/ fresh Lithosol, Calcite 5.3.3. Shrub vegetation
water deposit; Wet
Dryland 5.4. Alpine vegetation
4100-4200 m Dryland Lithosol; Dry 5.4.1. Alpine short grassland
4000-4500 m Dryland Lithosol; Dry 5.4.2. Alpine tussock grassland
4200-4600 m Dryland Morain (Lithosol); Dry 5.4.3. Alpine dry tundra
>4200 m Dryland Limestone MoraIin; 5.4.4. Alpine moist tundra
Moist
> 4200 m Dryland Morain (Lithosol), 5.4.5. Alpine heath moss & algal
Organosol; Moist vegetation
>4200 m Dryland Lithosol, Limestone, 5.4.6. Alpine heath dwarf shrub
Dry vegetation
4300-4500 m Snow Rocks, Lithosol; Moist 5.5. Nival vegetation
Littoral (< 0 m ) Saline water 6. Vegetation on saline water
II. Seasonally < 0 m Saline water Coral and sand; Wet 6.1. Monsoon littoral vegetation
Short to Long < 0 m Saline water Alluvial; Wet 6.2. Monsoon mangrove forest
Dry Monsoon
Lowland: Stagnant fresh water 7. Freshwater swamp monsoon vegetation
Q = < 61 - 300
0 (750) 1000 Stagnant fresh water Alluvial; Wet 7.1. Fresh water monsoon herb vegetation
(Type D, E, F)
m Stagnant fresh water Alluvial; Wet 7.2. Fresh water monsoon swamp forest
Annual rainfall:
700-2900 mm. Dryland 8. Dryland vegetation
Dryland Regosol; Dry 8.1. Sandy and/or rocky beach monsoon
herb vegetation
Dryland Regosol; Red Yellow 8.2. Sandy and/or rocky coastal monsoon
Podsolic, Latosol; Dry forest
Dryland Podsol; Renzina, 8.3. Lowland seasonal deciduous monsoon
Latosol; Dry forest
Dryland Podsol; Renzina, 8.4. Lowland evergreen monsoon forest
Latosol; Dry
Dryland Podsol; Renzina, 8.5. Monsoon scrub
Latosol; Dry
Dryland Podsol; Renzina, 8.6. Monsoon savanna
Latosol; Dry
Dryland Podsol; Renzina, 8.7. Monsoon grassland
Latosol; Dry
Highland/mount Dryland 9. Montane vegetation
ain: Dryland 9.1. Montane forest
(750) 1000- Dryland Red Yellow Podsolic, 9.1.1. Lower montane monsoon forest
3000 m Latosol, Andosol, Dry
Dryland Red Yellow Podsolic, 9.1.2. Uper montane monsoon forest
Latosol, Andosol, Dry
Dryland Latosol, Andosol, Dry 9.1.3. Sub-alpine monsoon forest
Dryland Lithosol, Dry 10. Sub-alpine monsoon scrub
Dryland Lithosol; Dry 11. Sub-alpine monsoon non-grass herb
vegetation
Dryland Lithosol; Dry 12. Sub-alpine monsoon grassland

106
Lowland rainforest
The lowland rainforest is the most extensive one occurring in Indonesia and
is the most luxurious vegetation compared to other types of forest in the world.
Variation occurs within this forest and the most obvious one is the big
difference between the western and eastern parts of Indonesia. The floristic
composition of various dryland forests in the country vary as indicated in Table
2, illustrating the variation of species richness and density in various forests.
There is a tendency that the forests in islands east of the Wallace line is poorer
than in the west. The distribution of Dipterocarpaceae follows this pattern
where out of 386 species only 26 species occur in eastern Indonesia and none
in the seasonally dry areas of East Nusa Tenggara (Ashton, 1984) (Figure 1).
In the western part (Sunda shelf) the family Dipterocarpaceae especially the
genera Anisoptera, Dipterocarpus, Dryobalanops, Parashorea and Shorea are
dominating the large trees of the emergent layer. Out of 12 genera and about
470 species of Dipterocarpaceae in Asia, 10 genera and about 350 species are
found in the Sunda shelf (Ashton, 1984). The forest in this part is also
characterized by a rich ground-layer palm flora of shade and moisture loving
genera, such as Iguanura, Pinanga, Areca, Nenga and Rhopalobaste (Whitmore,
1984a). In East Malesia the role of Dipterocarpaceae in the rainforest
ecosystem is less important and locally dominant. The important species here
include Araucaria cunninghamii, A. hunsteini and Agathis labillardieri and there
are more conifer species typical to the montane forest, such as Dacrydium
elatum, D. novoguineense, Papuacedrus spp., Phyllocladus hypophyllus and
Podocarpus papuanus (Whitmore, 1984a). Agathis, although it occurs also in the
West Malesia, the number of species or at least the number of forms is larger to
the east of the Wallace line. Pinus, on the other hand, occurs only in the west.
Variation in species richness reflects habitat conditions and on coarse level
floristic differences exist between forest formations on contrasting soil types,
such as between peat swamp, heath and lowland evergreen rain forests while
at finer level floristic differences reflect different topographic preferences within
a formation likely reflecting differences in soil (Whitmore, 1989a). Siregar
(2001) has shown the relationship of number of species with topography in
dryland and peat swamp forests and Miyamoto et al. (2003) in heath forest.
Whitmore (1989b) stressed that the richest tropical rain forest have more
than 200 species of tree with DBH 10 cm per hectare and such plots are
found in Asia although a few have been recorded from tropical America. In
Indonesia all such plots are in Kalimantan, notably plots at Malinau, Wanariset
Samboja and Lempake (Table 2). The richest plot for Indonesia is the plot of
Malinau 1, where in one hectare 221 tree species with density of 759 trees
per hectare were recorded. Whitmore (1989b) further suggested that more
counts are needed and such undertaking is straight forward and can be carried
out by students so that geographical picture can be established to test the
theory of Pleistocene Refugia explaining the cause of patterns of species
richness. Species richness is also a good indicator of species diversity and is
comparable to but simpler than the index of diversity (Gentry, 1988). Lembaga

107
Biologi Nasional was on the right tract then when in mid-1970s it initiated and
integrated the vegetation analysis project into its overall biological research
program and was later incorporated into the Indonesian MAB (Man and
Biosphere) Program to strengthen the integrated research program on human
impacts on tropical rain forest ecosystem, whose results have been reported
elsewhere (.e.g., Kartawinata, 1984; Kartawinata et al., 1981; Kartawinata and
Vayda, 1984).

A. Batu Karang, Central Kalimantan (Data from Riswan B. Sungai Barang, E. Kalimantan (Data from Bratawinata (1986).
et al. 2002). Plot size at each altitude: 30 x 50 m Plot size at each altitude: 4 (40 x 40 m)

800 838
647 680 900 785 785
700 607 800 719
648 654
600 700
460 600
500
500
400
400
300 300
179 157 157
200 200
78 117 106
54 57 51 63 100
100
0
0
770 930 990 1350 1560 1920
300 500 700 900
ALTITUDE (M)
ALTITUDE (M)
Density (Trees/ha) Number of Species. Density (Trees/Ha) Number of Species

C. Mt. Gede, West Java (Data from Yamada 1977)

Plot size at each altitude: 20 x 20 m D. Lore Lindu National Park , Central Sulawesi (Data
from Purwaningsih & Yusuf 2005). Plot size at each
160 152
altitude: 30x50 m
140 570 567
600
120 110
500
100 89
400 323
80
300
60 44
200
40
100 30 47 41
20 10 8 7 9
0
0
500 750 1000
2300 2600 2800 3000
ALTITUDE (M) ALTITUDE (M)
Density (Trees/Ha) Number of Species Density (Trees/Ha) Number of Species

Figure 3. Number of species and tree density along altitudinal gradient at various sites.

To date there have been very few enumerations of all plant species in
tropical forest plots in the world as it is very time consuming (Whitmore, 1989b)
although the results will be of scientific value and reflect more accurate species
richness. Such count was carried out in a one-hectare plot of lower montane
forest at 1500 m altitude at Mt. Gede by Meijer (1959) who recorded 333
species phanerogams and ferns, comprising 78 species of trees, 40 of shrubs,
30 of climbers, 10 of creepers, 100 of epiphytes and 73 of terrestrial herbs.
Figure 2 shows examples of species area curves for several lowland and
montane forests indicating the lowland forest curves rise steeply even up to an area
of 4 hectare at Samboja forest. In such species-rich lowland forests it is almost
impossible to define the point of inflection, while in the montane forest the point of
inflection is reached at an area of less than one hectare indicating the beginning
of the decrease of diversity. They are less so in the poorer forests in eastern
Indonesia (such as in Lore Lindu) and in the montane forests in West Java.

108
Table 2. Density and number of species of trees with dbh 10 cm in selected plots of
different forest types in Java, Kalimantan, Maluku, Papua, Sumatra, and Sulawesi .
N.P. = National Park; N.R. = Nature Reserve.

Mean
Plot size Number of
Locality Alt. (m) Density Reference
(ha) Species
(Trees/ha)
1 2 3 4 5 6
DRYLAND FOREST
East Kalimantan
Malinau 1 100 1.0 759 225 Kartawinata et al. (in prep.)
Malinau 2 100 2 x 1.0 413 240 Yusuf (2003)
Malinau 3 100 4 x 1.0 606 404 Samsoedin (in prep. )
Berau <100 3x4.0 521 538 Sist & Saridan (1999)
Wanariset Samboja 1 < 100 10.5 567 562 Kartawinata (upublished)
Wanariset Samboja 2 <100 1.5 541 239 Kartawinata et al. (1981)
Lempake <100 1.6 445 209 Riswan, S. (1987)
Sungai Barang
Site 1 700-770 4 x 0.16 719 179 Bratawinata (1986)
Site 2 850-930 4 x 0.16 838 78 Bratawinata (1986)
Site 3 930-990 4 x 0.16 785 157 Bratawinata (1986)
Site 4 1100-1350 4 x 0.16 648 117 Bratawinata (1986)
Site 5 1350-1560 4 x 0.16 785 156 Bratawinata (1986)
Site 6 1780-1920 4 x 0.16 654 106 Bratawinata (1986)

West Kalimantan
Muara Kendawangan N.R.
Hill forest 30 0.3 770 33 Purwaningsih & Amir (2001)
Central Kalimantan
Wanariset Sangai 15 584 72 1298 Wilkie et al. (2004)
Bukit Karung 300-900 4 x 0.15 598 225 Riswan et al. (2002)
North Sumatra
Gunung Leuser N.P.
Ketambe 1 450-670 1.6 538 116 Abdulhadi et al. (1989)
Ketambe 2 350-450 1.6 420 94 Abdulhadi (1991)
Ketambe 3 350-450 1.6 475 127 Abdulhadi et al. (1991)
Batang Gadis N.P.
Aek Nangali 660 1 583 182 Kartawinata et al. (2004)

Riau
Bukit Tiga Puluh N.P.
Buikit Lawang 297 1.0 453 216 Polosokan (2001)

North Sulawesi
Toraut 1 408 109 Whitmore & Sidiyasa (1986)
Lore Lindu N.P.
Plot 1 500 0.3 323 30 Purwaningsih & Yusuf (2005)
Plot 2 750 0.3 570 47 Purwaningsih & Yusuf (2005)
Plot 3 1000 0.3 567 41 Purwaningsih & Yusuf (2005)
B.N. Wartabone N.P.
Dudepo 180 0.4 610 64 Rahayoe et al. (1996)
Tanganga 1 200 0.2 410 31 Rahayoe et al. (1996)
Tanganga 2 200 0.2 480 21 Rahayoe et al. (1996)
Gunung Kabela 1 150 0.26 323 46 Polosokan & Siregar (2001)
Gunung Kabela 2 250 0.2 430 45 Polosokan & Siregar (2001)
Gunung Kabela 3 300 0.2 485 33 Polosokan & Siregar (2001)
Gunung Kabela 4 400 0.2 575 61 Polosokan & Siregar (2001)

Central Sulawesi
Sausu 45 - 85 0.4 770 68 Sidiyasa (1995)

South Sulawesi
Wotu 0.4 725 62 Sidiyasa (1988)

109
Table 2. continued

Mean
Plot size Number of
Locality Alt. (m) Density Reference
(ha) Species
(Trees/ha)
1 2 3 4 5 6
South East Sulawesi
Wawonii Island
Lampeapi < 300m 11 x 0.09 568 91 Rahayoe et al. (2004b)
Lansinowo 1 100 0.5 436 50 Purwaningsih (2003)
Lansinowo 2 300 0.5 492 40 Purwaningsih (2003)
Lansinowo 3 500 0.3 813 9 Purwaningsih (2003)
North Buton W.R.
Gunung Wani 1 300 0.5 462 60 Mansur & Wardi (2004)
Gunung Wani 2 400 0.5 442 49 Mansur & Wardi (2004)
Soloi 100 0.5 554 60 Mansur (2003)

Maluku
Halmahera 630 0.5 742 76 Whitmore et al. (1987)
Papua
Wamena: Wanduga 2800 0.5 528 28 Partomihardjo & Supardiyono (1993)
Wamena:Tengon 1600 0.15 813 38 Partomihardjo (1991)
Wamena: Kurulu 1600-2350 0.7 564 76 Partomihardjo (1991)
Yapen Tengah N.R. 600-1200 14 x 0.1 799 235 Partomihardjo (2001)
Supiori N.R. 1- Slope 320 0.5 1024 93 Siregar (2001)
Supiori N.R. 2- Flat ridge 320 0.5 1010 77 Siregar (2001)

Central Java
Karimun Jawa N.P.
Plot 1 100 m 0.4 738 68 Yusuf et al. (2004)
Plot 2 200 m 0.4 725 71 Yusuf et al. (2004
Plot 3 300 m 0.4 866 56 Yusuf et al. (2004

West Java
Gunung Halimun N.P.
Purwabakti: 2nd forest 900 1.0 441 69 Yusuf (2004)
Citalahab: 2nd forest 1000-1200 0.7 395 51 Rahayoe (1996)
Citorek: Plot Group 1 905-1127 5 x 0.1 530 56 Mirmanto & Simbolon (1998)
Plot Group 2 761-893 5 x 0.1 384 61 Mirmanto & Simbolon (1998)
Plot Group 3 784-939 2 x 0.1 106 26 Mirmanto & Simbolon (1998)
Cikaniki 850-1500 26 x 0.09 601 73 Simbolon & Mirmanto (1997)
Cikelat 1000-1600 21 x 0.09 624 80 Simbolon & Mirmanto (1997)
Cisarua 1100-1500 13 x 0.09 408 85 Simbolon & Mirmanto (1997)
Cisangku 1050-1800 13 x 0.09 671 84 Simbolon & Mirmanto (1997)
Gunung Gede-Pangrango
N.P. 1500-1900 4.0 889 93 Abdulhadi et al. (1998)
Cibodas 1 1600 1.0 427 57 Yamada (1975)
Cibodas 2 1700 0.1 450 19 Yamada (1977)
Cibodas 3 1900 0.1 560 15 Yamada (1977)
Cibodas 4 2100 0.1 840 14 Yamada (1977)
Cibodas 5 2300 0.1 1100 11 Yamada (1977)
Cibodas 6 2400 0.4 1516 10 Yamada (1976)
Cibodas 7 2600 0.04 2225 8 Yamada (1977)
Cibodas 8 2800 0.04 1100 7 Yamada (1977)
Cibodas 9 3000 0.04 3829 9 Yamada (1977)
Cibodas 10
PEAT SWAMP FOREST
Sumatra
Teluk Kiambang < 100 0.2 752 34 Anderson (1976)
Muara Tolam < 100 0.2 705 46 Anderson (1976)
Sungai Siak Kecil < 100 0.2 831 50 Anderson (1976)
Bengkalis 1 < 100 0.24 584 33 Siregar (2002)
Bengkalis 2 <100 0.24 801 28 Siregar (2002)

110
Table 2. continued

Mean
Plot size Number of
Locality Alt. (m) Density Reference
(ha) Species
(Trees/ha)
1 2 3 4 5 6
West Kalimantan
Sungai Durian < 100 0.2 1006 39 Anderson (1976)
Sungai Sebangan < 100 0.2 1200 50 Anderson (1976)
Sungai Pinyuh & Mandor 30 -60 1.6 342 119 Mirmanto et al. (1993)

FRESH WATER SWAMP

FOREST
East Kalimantan
Sesayap > 100 1.0 400 76 Suhardjono & Wiriadinata (1984)
West Kalimantan
Kendawangan N.R.
Freshwater swamp 15 0.3 590 17 Purwaningsish & Amir (2001)
Semi-swamp 20 0.3 390 54 Purwaningsish & Amir (2001)

KERANGAS (HEATH)
FOREST
East Kalimantan
Samboja 1 < 100 0.5 750 24 Riswan (1987b)
Samboja 2 < 100 0.5 554 14 Riswan (1987b)

In East Kalimantan dipterocarps species are prevalent in the lowland

forests, except in freshwater swamp forests. In a permanent plot of 10.5
hectare lowland dipterocarp forest at Wanariset in East Kalimantan, 406 species
of trees (DBH > 10 cm), 178 genera and 57 families were recorded, with a
density of 534 trees/ha, a mean basal area of 29 sq, m/ha (Partomihardjo et al.,
1984). The dipterocarp species are the largest trees and dominate the forest
and the number of species amounts to 30 (7.4 %) with the total basal area of
79.5 sq.m (26 %). As indicated in Table 2 above the one-hectare plot at
Malinau 1, where in one hectare 221 tree species with density of 759 trees
per hectare is the richest forest in the country, followed by the plot at Samboja
2 where in 1.6-hectare plot 239 species with density of 541 per hectare are
recorded. Table 2 and Figure 2 show that the plot at Malinau has higher
density and number of species than elsewhere. The forest at Lempake,
Samarinda, differs in its tree species composition and structural characteristics
than that of Wanariset. It contains less dipterocarps, amounting to only 13
species in a plot of 1.6 ha with density of 27 trees per hectare and basal area of
10 sq. m per hectare (Riswan, 1987a).
In 25 plots of primary forest at the Gunung Leuser National Park, North
Sumatra, 332 tree species, 179 genera, and 68 families were recorded
(Abdulhadi et al., 1984) . The forest is less diverse than those at Lempake and
Wanariset. Dipterocarps are less prominent where only 12 species with a
density of 23 trees/ha are recorded. Meliaceae, however, is richer having 22
species with a mean density of 74 trees/ha, second only to Euphorbiaceae. The
forest has a low stature with a mean height of 15 -23 m depending on the
topography. Another forest type in North Sumatra may be reflected by data

111
from the Batang Gadis National Park, where the number of species and tree
density was 182 and 583, respectively (Table 2 and Figure 2) (Kartawinata et
al., 2004).

Fresh water swamp vegetation

The type of vegetation developed on freshwater swamps varies considerably
with the wide variation in its soils. In some areas the natural vegetation is
dominated by grasses, sedges and other herbaceous species, in others it is
palm or pandan-dominated forest or forest with structure and composition
similar to lowland rain forest. The freshwater vegetation, occurs in all islands of
Indonesia in areas where climate ranges from seasonally dry to very wet, in the
lowlands and the highlands. The greatest extent is the freshwater swamp forest
occurring in the lowlands of Sumatra, Kalimantan and Papua. The freshwater
swamp occurs on permanently or seasonally flooded mineral soils in vast
coastal alluvial plains, deltas and meander belts . Belts of freshwater swamp
forest of up to 5 km wide can be found along big rivers usually adjacent to peat
swamp forests. Endert (1927), Steenis (1935, 1957, 1965), Paijmans (1976),
MacKinnon et al. (1996) and Whitten et al. (1984, 1987) described freshwater
swamp vegetation in various parts of the country. Freshwater vegetation may
comprise freshwater swamp forest, sago swamp forest and herbaceous
freshwater swamp vegetation. Variations of floristic composition and species
richness occur within each of these vegetation types.
So far quantitative studies were undertaken in East Kalimantan (Suhardjono
and H. Wiriadinata, 1984) and Papua (PTFPI 1997). The floristic composition
shows a great diversity which is related to the great range of habitat conditions,
and the flora is usually not different, at family and generic level, from that of
the dry lowland forest occurring in the same region (Endert, 1927; Whitmore,
1984a). Some species tend to form pure stands, such as Metroxylon sagu and
Campnosperma brevipetiolatum in New Guinea. Other three species common to
this forest include those of the genera Alstonia, Barringtonia, Campnosperma,
Dillenia, Eugenia, Shorea, Mangifera, Neesia, and Pholidocarpus

Peat swamp forest

The peat swamp forest is the swamp forest occurring in oligotrophic waters,
that make the peat formation possible. The peat layer may extend to the depth
of 20m. It is found extensively in Sumatra and Kalimantan. Species richness in
peat swamp forest is lower than that in dryland forest. Variation in tree density,
height and floristic composition in response to habitat changes especially the
depth of the peat deposit leads to the concentric forest zonation, with the
poorest floristic composition and lowest stature located in the center. Study of
Mirmanto et al. (2003) revealed that the distance from river may be used as a
suitable parameter for investigating plant community distribution in peat swamp
forest, even if peat thickness differ greatly.
Although not all types are present, Anderson (1976) noted the sequence of
forest types like in Sarawak occurs also in Riau, West Kalimantan and Central

112
Kalimantan: (1) the Gonystylus-Dactylocladus-Neoscortechinia association (mixed
swamp forest), (2) the Shorea albida-Gonystylus-Stemonurus association, (3) the
Shorea albida consociation, (4) the Shorea albida-Litsea-Parastemon association,
(5) the Tristania-Parastemon-Palaquium association, and (6) the Combreto-
carpus-Dactylocladus association. This pattern is related to the structure of the
swamps and the fertility of the peat swamp soils. Anderson noted that the flora
of the peat swamp forests in Kalimantan is richer than in Sumatra. It has been
shown that there are differences and similarities in number of species and
structural characteristics of the peat swamps in Sumatra and Kalimantan. In
Sumatra the number of species per unit area along the catenary sequence is
constant, although floristic composition varies from one locality to another,
while in Kalimantan the number of species in the Mixed Swamp forest is
significantly higher than in the padang. The tree density increases across the
catena, but in Kalimantan the density is higher in each forest type than in each
equivalent forest type in Sumatra. The total basal area across the catena, does
not show a significant reduction, and the total biomass is assumed to follow the
same pattern also. The mean basal area and the girth, however, decrease
across the catena and these are more pronounced in Kalimantan than in
Sumatra. Most of the dominant species in Mixed Swamp Forest are common to
both Sumatra and Kalimantan. These include Alstonia pneumatophora,
Lophopetalum multinervium, Campnosperma coriacea, Gonystylus bancanus,
Palaquium burckii, Tetramerista glabra, Durio carinatus, Koompassia
malaccensis, Parastemon urophyllum, Dyera lowii, Mezzettia leptopoda, Shorea
platycarpa, S. teysmanniana and S. uliginosa.
50

45 Transition area: Modified

boundary between Montane Myrsine affinis
40 and Subalpine Zones at 2300 m
Schima wallichii
35
RELATIVE DOMINANCE (%)

30
Polyosma illicifolia
25 Podocarpus imbricatus

20 Achronychia
punctata
Eurya obovata
15

10 Symplocos
sessilifolia
5

0
1600 1700 1900 2100 2300 2400 2600 2800 3000
-5
ALTITUDE (M)
MONTANE ZONE SUBALPINE ZONE Steenis Zonation
MONTANE ZONE SUBALPINE ZONE Modified Zonation

Figure 4. Species dominance along the altitudinal gradient at the Gunung Gede-
Pangrango National Park with modified boundary between Montane Zone
and Subalpine Zones at 2300 m altitude (Data after Yamada, 1977).

113
Kerangas forest
Few quantitative studies, carried out in kerangas or heath forests
(Kartawinata, 1979; Miyamoto et al., 2003; Purwaningsih and Amir, 2001;
Riswan, 1982; 1987b), showed poorer species composition and higher tree
density, especially in the padang type, compared to those of lowland
dipterocarp forest. Kerangas, in which peat deposit is underlain by acid white
sandstone that form podsol soils, showed also concentric zonation like in peat
forest mentioned above (Kartawinata, 1979). Bratawinata (1986) noted kerangas
forest in the highland dominated by Agathis borneensis. Miyamoto et al. (2003)
showed different tree species with small-scale variation of humus depth and
topography in a heath forest in Central Kalimantan. Various types of kerangas
occurring in Kalimantan and Sumatra were described qualitatively by various
authors and have been summarized by Kartawinata (1978a), who at the time
could not find any record of kerangas in eastern Indonesia. Only in recent years its
occurrence in Papua was recorded (Kartawinata and Widjaja 1988; PTFI 1997).
Earlier reports from Indonesia (Kartawinata 1978a) and Borneo (Whitmore,
1984a) stressed that kerangas forest occurs on podsol soils which are poor in
nutrients and very acid. It differs from the lowland dryland forests in floristic
composition, structure and physiognomy. The trees are shorter and smaller in
size, generally with sclerophyllous leaves. Myrtaceae is prominent in this forest,
particularly Tristania, but also Eugenia, Whiteodendron moultanianum and
Baeckia fruteascens. Other prominent species include Cratoxylum glaucum, C.
arborescens, Combretocarpus rotundatus, Casuarina nobillis, Cotylelobium
burckii, C. malaynum, C. melanoxylon, Dacrydium elatum, D. subulatum,
Dactylocladus stenostachys, Ilex cymosa, I. hypoglauca , Shorea balangeran,
S.coriacea, and S. havilandii (Kartawinata, 1979; Riswan, 1982, 1987b). Many
species occurring in kerangas forests, for example many of Dipterocarpaceae,
are also found in lowland dipterocarp forest; and there are also many species
common to both kerangas and peat swamp forests and some to both
kerangas and upper montane forests (Whitmore, 1984a).

Monsoon forest
Very little quantitative information on the monsoon (seasonal) forest and
available information show that it is generally poorer in species as exemplified
by the forest in Ruteng (Setiadi, 2005) and Sumbawa, where in a series of plots
from 50 m to 900 m totaling 0.64 ha contained only 170 species of trees and
shrubs (Yusuf 1996). Based on phytosociological assessment using the Braun-
Blanquet method, Meijer-Drees (1951) developed a scheme on distribution,
ecology and silvicutural possibilities of the trees and shrubs from the savanna-
forest region in eastern Sumbawa and Timor. He identified 20 communities
that are grouped into four alliances. Monk et al. (1997) synthesized available
information for East and West Nusa Tenggara. The monsoon vegetation of
Indonesia may be divided into deciduous forest, dry evergreen forest and
savanna. Typical species in these vegetation types include Acacia leucophloea,
A. tomentosa, Aegle marmelos, Albizia chinensis, A. lebbekoides, Azadircahta

114
indica, Borassus flabellifer, Butea monosperma, Caesalpinia digyna Cassia
fistula, Corypha elata, Dalbergia latifolia, Dischrostachys cinerea, Feronia
limonia, Garuga floribunda, Homalium tomentosum, Lannea grandis, Melia
azedarach, Schleichera oleosa, Schoutenia ovata, Stereospermum suaveolens,
Streblus asper, Tamarindus indica, Tectona grandis, Tetrameles nudiflora, and
Zizphus rotundifolia (Meijer-Drees, 1951; Steenis, 1957; Whitmore, 1984a). In
Papua and Nusa Tenggara, the Australian elements are very strong, species
such as Eucalyptus alba, Melaleuca leucadendron and Santalum album (Steenis,
1957) are abundant.

Limestone forest and forest on ultrabasic rocks

Little is known about the limestone forest which occur throughout the
country but constitutes only a fraction of the total forest area in Indonesia. In
Malaysia limestone forest is rich in endemic species, which could be so also in
Indonesia. Similarly we practically know very little about the forest on ultrabasic
rocks, which cover relatively much smaller area compared to the lowland
rainforest. It occurs on very specialized habitat in Kalimantan, Sulawesi and
Papua. Soils under this forest derived from serpentinites and have a high iron,
magnesium and a low silica content and are also characterized by high
concentrations of phytotoxic elements, especially nickel, cobalt and chromium
(Burnham in Whitmore, 1984a). The forest cover varies from shrubby open
vegetation to high and dense forest and so does the floristic composition that could
be very distinctive or similar to that of forest over other soils (Whitmore, 1984a)

Montane and subalpine forests

There is a tendency that in the montane and subalpine forests the number
of species decrease with altitude while tree density increases (Bratawinata,
1986; Edwards et al., 1990; Ohsawa et al., 1985; Yamada, 1977, 1990), but is
not the case in lowland forest (Figure 3). The species-area curves in the
montane forests on Mount Gede and Mount Halimun (Figure 2) flatten out
rather quickly where the minimum area can be attained at an area less than
one hectare as shown by the point of inflection indicating the beginning of the
decrease of diversity. Edwards et al. (1990) noted that tree species diversity in
the montane forests in Manusela National Park, Seram, Maluku, is lower than
that in many rain forests in the Far East, including Borneo and Papua, but
comparable to that recorded by Whitmore et al. (1987) in other part of Seram.
The species-area curves in nine plots flattened out rather quickly also and are
comparable to that of Mt. Halimun.
Yamada (1977) shows at Mt. Gede and Mt. Pangrango that Schima
wallichii and Achronychia punctata are the dominant tree species in the
montane forest at the altitudes of 1600-2400 m and Myrsine affinis and Eurya
obovata in the subalpine forest at altitudes of 2400-3000 m with distinct
abundance and dominance of Vaccinium varingiaefoliumm at the summit at
altitude of 3000 m (Figure 4). It can be noted that the altitude of 2300 m is
the meeting point of dominant species of upper montane forest and sub-alpine

115
forest, hence the elevation of 2300 may be suggested as the boundary beween
montane zone and subalpine zone differing from the boundary at altitude of
2400 m previously defined floristically by Steenis (1972). The subalpine forest is
known also as elfin forest (Steenis, 1972), referring to forest with low canopy,
high density, crooked and gnarled trees because of substratum, in which the
common species include those of the genera Astronia, Eurya, Leptospermum,
Myrsine, Pittosporum, Rhododendron, Schefflera, Turpinia, Vaccinium, etc.
Vaccinium and Rhododendron (Ericaceae) are the most prominent, hence the
forest is often called also ericoid forest. The elfin forest often carries mosses
(e.g. Aerobryum and Usnea) also and is, therefore, known as mossy forest. On
much higher elevation the mosses may be found even on the ground, including
peat moss or Sphagnum. Such moss-covered ground are favorable habitat for
Nertera, Corybas and Utricularia.
On Mt. Kerinci in Sumatra, Ohsawa et al. (1985) divided the forest zone
above 1700 m into (i) a subtropical zone below 2400 m dominated by Schima
wallichii and Eugenia spp. and (ii) a warm temperate zone above 2400 m
dominated by Lithocarpus sp or other trees of Symplocaceae and Myrsinaceae.
This division is also characterized by changes of structural and floristic features
at about 2400 m, thus confirming the boundary of van Steenis montane zone
and subalpine zone. Lamounier (1990) performed cluster analysis on flora
distribution in Sumatra and the results revealed six floristic sectors in the
lowland and three in the mountains, leading to a need for a more precise
definition of lowland areas while agreeing with Steenis zonation in the higher
elevation. Contrary to Steenis zonation scheme in Jawa, PTFPI (1997) found
different zonation as follows: (i) the lower montane forest formation (650-1500
m), (ii) the mid montane forest formation (1500-2800 m), (iii) the upper
montane forest formation (2800-3200 m), (iv) the Subalpine Zone (3,200-4,170
m) with two subzones (the lower subalpine subzone at 3,200-3,650 m and the
upper subalpine subzone from 3,650-4,170 m), (v) the Alpine Zone (4,170-
4,585 m) and (vi) the Nival Zone (> 4,585 m). In the Mt Jaya area. PTFPI
(1997) noted that the lower montane subzone is a transition zone between
lowland forest and mid-montane forest, and is characterized by the presence of
Castanopsis and Lithocarpus species. The mid montane forest is the main
montane forest, and is characterized by the presence of Nothofagus species
and montane families such as Lauraceae and Elaeocarapaceae. The upper
montane zone is a transition zone to the subalpine forest at higher altitudes and
is characterized by species of southern conifers and members of the family
Ericaceae. From various studies mentioned above it is noted that the number of
tree species does not always decrease with increasing altitudes, but species
composition, prevalent species and structural characteristics do change.

Sub-alpine and alpine herbaceous vegetation

The diversity of vegetation at higher altitudes covering subalpine, alpine
and nival zones are greatest in Papua (see Table 1). To date PT Freeport
recognized approximately 35 communities in the subalpine, alpine and nival

116
zones, while Hope et al. (1976) identified at least 19 community types
occurring on varied habitats, ranging from grass communities to tundra. In
other islands the variability is less and the characteristic species are recorded in
Steenis (1962, 1972). In Table 1 they are grouped into Subalpine shrub and
herb vegetation, Subalpine swamp vegetation, and Alpine vegetation. They are
rich in grass communities occupying different habitats. These grasslands occur
in the mountains with altitudes greater than 3000 m and may cover pure grass
communities, open scrubs and savanna, but the grass species characterize
these communities and constitute factors determining the structure and
selection of other species that can grow among them. The subalpine and alpine
herbaceous vegetation are also rich in algal and moss communities.
The timber line or the upper elevation limit for tree growth occurs in high
mountains, such as in Papua. Hope et al. (1976) noted that this elevation limit
varies from one place to another spreading also to open areas or places whose
snow cover just melted at lower elevation, hence alpine vegetation can not be
necessarily defined as all vegetation above certain elevation; the highest timber
line is at 4170 m altitude on the north slope of Meren Valley on Carstenz Range.
Above this limit the vegetation is characterized by the presence of Senecio a
species of Asteraceae comparable to Anaphalis in high mountains of Java.
Common species include Coprosma brassii, Rhododendron culminicolum and
Styphelia suaveolens.

Vegetation dynamics
The dynamics of vegetation received a greatest emphasis in the ecological
studies. The classic studies of primary succession on Krakatau of 1883 eruption
were summarized by Richard (1952, 1996). The anniversary of the eruption has
stimulated the organization of interdisciplinary symposium (Sastrapradja et al.
1985) and new studies which have produced a large number of publications
providing new evidence on succession and on island biogeography (e.g., Bush
et al., 1995; Kartawinata et al., 1985; Partomihardjo, 1995; Partomihardjo et
al., 1992, 1993; Richards and Wiriadinata, 1985; Tagawa, 1985, 2005;Tagawa et
al., 1985; Thornton, 1992, 1996; Whittaker, 2004; Whittaker et al., 1984, 1989,
1995, 1999, 2000; etc.). The forests thus far developed during the last 100
years are still in successional stage. The return to conditions similar to a
primary forest will take a very long time and probably will never be reached
since sources of propagules in Java and Sumatra are declining. It has been
shown that the succession on the islands is a complex process, hierarchically
structured by interrelationship between plants and animals and affected by
volcanic disturbance and climatic unpredictability (Whittaker, 2004).
Studies on successions in naturally formed gaps have been just initiated in
lowland and montane forests (Partomihardjo et al., 1987; Srijanto, 1987,
Mackie et al., 1987). The gaps in primary forest constitute 16 - 28 % with size
mostly of less than 500 sq. m and regeneration within them consists mostly of
primary forest species, while in secondary forest the gaps constitute 13 % with
size range of 18 - 186 sq.m. Gaps created by traditional timber extraction by

117
the natives are comparable to natural treefall gaps in size and the density of
regenerating seedlings. Heavy wind storm in 1984 has created large gaps in the
montane forest of Mt. Gede and Srijanto (1987) found negative correlation
between gap-size and the seedling and sapling densities of major species,
Castanopsis argentea, Lithocarpus pseudomoluccus, Schima wallichii and
Villebrunea rubescens. Gaps resulted from logging activities, however, are much
bigger, amounting to more than 30 % and a gap can be as big as one hectare
(Abdulhadi et al., 1987). In such a big gap regeneration consists mostly of
secondary forest species. From a long-term 2-ha plot in Mt. Gede Abdulhadi et al.
(1998) reported zero mortality in 22 species and 10-100 % in 58 species with
total density of 29 trees/ha as well as an increase of species by 14 in 17 years.
Studies on regeneration after human disturbance deal with the effects of
logging and shifting cultivation. Substantial data have been collected from
studies on logged-over forests conducted by the Forest Research Institute. The
studies are concerned primarily with the regeneration of dipterocarps, and the
regeneration is identified by commercial species group not by species, hence it
is difficult to make floristic interpretation. Most current papers (e.g., Cannon et
al., 1994; Okimuri and Matius, 2000; Ruslim et al., 2000; Sist et al., 2003)
present detailed effects of logging and indicate also the value of logged-over
forests for conservation. Nieuwstadt et al. (2001) described the negative
ecological consequences of salvaged logging in the burned forests commonly
practiced in various places. Environmental effects of mechanized logging in
Indonesian forests have been summarized in various articles (Kartawinata,
1981b; Kartawinata et al., 1989, 2001). It is shown that on the average 50 %
of the residual trees are damaged, regeneration is dominated by secondary
forest species, and there is a sign of genetic erosion due to the creaming of
extraction of the best trees (dipterocarps particularly) as well as loss of species.
In recent years, no successional study has been undertaken in the logged-over
forests in Indonesia. The recovery of logged-over forest was studied by Priyatna
et al. (in prep) who reported that after 23 years there has been no shift in most
important species and families, while the tree density and canopy are still
developing and estimated to reach complete recovery in about 60 years.
Earlier, Kramer (1926, 1933) did an experimental tree cutting in montane forest
of Mt. Gede, West Java, to determine the regeneration in relation to gap sizes
that may suitable for selective logging, although his findings never taken up by
the Ministry of Forestry for formulating the Indonesian Selection Cutting System
to be employed in the forest exploitation since 1968.
Richards (1996) claimed that the investigation on succession after clear
cutting and burning of lowland dipterocarp forest and kerangas forest in East
Kalimantan (Riswan, 1982; Riswan and Kartawinata, 1988, 1989, 1991) was the
only experimental successional studies ever done in the world. The results
show that seedlings were the main component in the vegetation recovery
process in dipterocarp forest in contrast to vegetative resprouts in kerangas.
Different regimes of disturbances lead to differences in species composition, but
in kerangas forest composition is similar. The pattern of secondary succession is

118
critically determined by the successional process during the first six-month
period. Burning altered the soil properties in dipterocarp forest but soil fertility
returned to the original level within 1.5 years, but in kerangas forest. they are
otherwise. The studies show also that tree species differ in their requirements
for germination and for seedling establishment and species may be classified
into species with shade-tolerant seedlings and species with seedlings that
require full illumination created by large gaps. This lead to the species
grouping into the climax, primary, or shade-tolerant species or known also as
dryad for the former and the pioneer, secondary, shade-intolerance species or
nomad for the latter (Riswan, 1982; Whitmore, 1984a). Related to both
primary and secondary successions is the availability of soil seedbanks, but this
area has not been much investigated. Whittaker et al. (1995) studied the
surface and buried seed banks on Krakatau to characterize the nature and
variability of the seedbanks and their implication for the sterilization hypothesis.
Riswan (1982), although indirectly, referred the role of seedbanks in the
regenerartion of dipterocarp forests after clear-cutting and burning and Brearley
et al., (2004) studied floristic and soil changes by comparing old secondary and
primary forests in Barito Ulu, Central Kalimantan. Soedjito (1985) studied the
succession and nutrient dynamics after shifting cultivation and later he
investigated the nutrient dynamics in relations to the root systems of secondary
forest species (Soedjito, 1990).
Secondary successions on cleared peat swamp forests based on existing
reports were summarized by Kostermans (1960). Based on his qualitative
observations in various parts of Indonesia, Kostermans (1962) further
acknowledged that secondary successions are of a very complex nature,
differing from one place to another and formulated factors determining floristic
composition in the stages of succession. In the mean time Dilmy (1962)
emphasized the effects of fire used by early man on the vegetation of the
humid tropics, resulting in the formation of Imperata grasslands and
subsequent successions. Riswan and Abdulhadi (1992) indicated that slash and
burn cultivation resulted in drastic decrease of species and repeated burning led
to the formation of Imperata grassland within just a few years after the clear-
cutting of primary forest. Systematic examination on succession after burning
has not been much investigated and so far nothing is more revealing than the
experiments done in East Kalimantan mentioned above (Riswan, 1982; Riswan
and Kartawinata, 1988, 1989, 1991). Various impacts of fire on vegetation
have been described in various papers (e.g. Dennis et al., 2001; Kartawinata,
1993; Simbolon et al., 2003; Wirawan, 1993; see also papers in Guhardja,
2000 and Osaki, 2003)
Practices of traditional shifting cultivation by Kenyah Dayak do not lead to
permanent forest destruction (Mackie, 1986; Mackie et al., 1986). If left
fallowed for more than 100 years the old field communities resulted there from
would acquire structure and composition of primary forest. However, the rate of
return to primary forest composition is dependent upon the proximity of seed
sources. Studies on a series of different ages of fallowed slash-and-burn

119
agriculture in East Kalimantan (Okimuri and Matius in Guhardja et al., 2001)
show that the pioneer Macaranga spp. were dominant during the first 15 years
and dipterocarp species and other primary forest species replaced Macaranga
after 30 years and only after 70 years dipterocarp species became dominant
although the forest lacks emergent trees. The recovery is very slow and
estimated to take 148-478 years for a young secondary forest to reach maturity
(Riswan, 1982; Riswan and Kartawinata,1988; Riswan et al., 1985).

PRIMARY
FOREST (Climax)

Clear-cut logging

SWIDENED LAND Selective logging

YOUNG SECONDARY FOREST

< 1 YEAR
(Grasses, herbs & shrubs)
EXOTIC OR
NATIVE
SPECIES
PLANTATION
FALLOWED YOUNG SECONDARY FOREST
SWIDENED LAND > 1 YEAR
(Shrubs & young trees)

LOGGED-OVER
RF FOREST
(Primary forest
ANR intercalated with
secondary forest)
E
IMPERATA
GRASSLAND
EXOTIC OR
NATIVE SPECIES
OLD SECONDARY PLANTATION
FOREST > 30 YEARS
RF (Trees)

IMPERATA GRASSLAND E &S

(Fire climax)
ANR

C FOREST OF FAST
GROWING
COMMERCIAL
SPECIES
EXOTIC AND/OR NATIVE
CLIMAX FOREST
TREE SPECIES PLANTATION

Figure 5. Natural succession after clear-cutting and selective logging of tropical

forests and possible intervention for further development through
Accelerated Natural Regeneration (ANR), Conversion (C), Enrichment
Planting (R), Enrichment Planting (E) and Enrichment Planting and Selection
(E & S)

120
 Indonesia has extensive areas of post-extraction secondary forests and
degraded lands arising from intensive exploitation of forest resources in recent
decades. Using the area of forests resulting from selective logging practices as
an estimate, in year 2000, post-extraction secondary forests covered about 23
million ha, or about 55% of the total concession area. Kartawinata et al.
(2001) analyzed the underlying causes of transformation of primary to
secondary forests and degraded lands, including policy and regulations in
forestry and forest resources, poor enforcement of regulations, and the lack of
recognition of timber exploitation rights for local communities. The government
is committed to promoting participation of local communities in managing
forests. Recent policy changes for ameliorating some of the degrading factors
have resulted in increased pressure on post-extraction secondary forests due to
rampant illegal logging and use claimed by local communities and land
speculators. While the largest proportion of post-extraction secondary forests
has been maintained as part of the permanent forest estate, substantial areas
have been converted for swidden agriculture, industrial tree and estate crop
plantations and transmigration areas. Local community involvement and an
understanding of the underlying degradation pressures would be imperative for
the effective rehabilitation and sustainable management of post-extraction
Rehabilitation programs should be an integral part of development programs
in the country, although the rate of deforestation is outpacing that of
rehabilitation. Papers on rehabilitation techniques and constraints in forest
management are presented in Guhardja et al. (2000). Methods and use of
secondary forest species in rehabilitation has been suggested (Kartawinata,
1994b, c). Assisted natural regeneration, a method of reforestation which
exploits the natural processes of vegetation recovery, is a potentially rapid,
efficient, and cost-effective means of reforestation. The scheme is presented in
Figure 5, showing different possible pathways beside natural successions after
clear-cutting of a primary forest, in which community-based reforestation
strategies may be exercised. The rehabilitation of deforested lands requires
urgent and serious attention in order to (1) outpace the rate of deforestation,
(2) restore biological diversity. (3) diversify the products and increase the
productivity of deforested lands, (4) provide socio-economic benefits to both
government and rural communities, and (5) supply raw material to the wood
industry so that the pressure on remaining primary forests can be reduced and
even eliminated. While there is sufficient knowledge and experience that can be
used as a basis for trial programs, further research is needed in particular
concerning the use of economically valuable fast-growing species of seral
communities (Kartawinata, 1994c). Rehabilitation may be also undertaken by
applying agroforestry approach (Kartawinata and Satjapradja, 1983) involving
local communities.

Other ecological aspects

The Ecology of Indonesia Series on Sumatra (Whitten et al., 1984), Sulawesi
(Whitten et al., 1987), Kalimantan (MacKinnon et al., 1996), Nusa Tenggara

121
(Monk et al., 1997) and Java and Bali (Whitten et al., 1996) have been recently
published. They are the synthesis of available information and contain good
accounts on plant ecology. The Ecology of Sulawesi includes also keys to plant
species of mangroves and estuaries, aquatic macrophytes and tree ferns. The
Rainforest Ecosystems of East Kalimantan (Guhardja et al., 2000) contains a
rich information in diverse articles by different authors on the results of 15-year
cooperative studies between Indonesia and Japan concerning the El Nio,
drought, fire and human impacts, addressing the current conditions of the
rainforest ecosystems and the regeneration techniques that can be used in
developing sustainable forest management.
160

140
Hal-2

120
Total litterfall (g/m2/mo)

Hal-3
100

80 Lahei-HF

60
Lahei-PSF

40

Barito Ulu
20

0
Jun Jul Agu Sep Oct Nov Dec Jan Feb Mar Apr May Jun
Month
Figure 6. Montly litter production in montane forest of Mt. Halimun, West Java (Hal 2
& Hal 3) and heath forest at Laei (Lahei HF), peat swamp forest at Lahei
(Lahei PSF) and lowlandland forest at Barito Ulu, Central Kalimantan (After
Mirmanto et al., 2005)
40

30
No. of species (%)

20

10

0
Fruits Vegetables Spices Rattan Medicinal Poison Firewood Home
plants plants industry
and art
Figure 7. Percentage of number of species according to category of plant use in two
one-hectare permanent plots in Jambi (After Mirmanto, 2005)

122
 Plant-animal interactions received great interests of various specialists
(e.g., Rijksen, 1978; Whitten, 1980; Curran and Leighton, 2000; Curran and
Webb, 2000; see also articles in Guhardja, 2000). Although the studies
emphasized the ecology of primates, they contain good accounts on structure
and composition of vegetation, phenology and seed dispersals. Current
information on production ecology is very limited and studies have been carried
out mainly in lowland forests of Sumatra and Kalimantan (e.g., Rahayoe et al.,
2004a,c; Schaik et al., 1985; Yamakura et al., 1986) and montane forests of
Java (Yamada, 1976; Rahayoe et al., 2004a). Litter production of the lowland
forest of Sumatra is estimated to be 7.6-11.5 ton.ha-1.year-1 and that of
Kalimantan 7.2 ton.ha-1.year-1 while that of the montane forest 5.9 ton.ha-
1
.year-1. The above ground biomass of a lowland forest of east Kalimantan is
estimated to be 509 ton.ha-1. Figure 6 shows monthly litterfall productions in 5
stations representing montane forests, heath forest, peat swamp forest and
dryland forest, based on 2-4 years observations (Mirmanto et al., 2005). It is
apparent that heath forest and peat swamp forest have similar production
patterns and both are lower than montane forest and dryland forest in the
lowland. Pattern in the montane forest varies even in the same general area
perhaps attributed to different floristic composition. The litter production in
montane forest of Mt. Halimun was 7.0-8.2 ton.ha-1.year-1 (Rahayoe et al.,
2004a) while on Mt. Pangrango (Yamada, 1976) was 5.96 ton.ha-1.year-1.
Sukardjo (1995) recorded litterfall in a mangrove of East Kalimantan, while
Alrasyid (1988) reported that in a mangrove forest in West Kalimantan the litter
production was 10.5 ton.ha-1.year-1 and organic carbon release 2.02 ton.ha-
1
.year-1. Litter production and nutrient studies in burned forests are presented
in various papers in Guhardja et al. (2000). Mirmantos (2000) study in peat
swamp forest of Tanjung Putting National Park reveals that the N, P, and K
content in litter is lower than in fresh leaves and that the translocation of N and
P is very high which is suspected to be an inhibitory factor of plant growth in
this forest. Few studies by Bruijnzeel (1983, 1984, 1985), Riswan (1982) and
Soedjito (1985, 1990) deal with some aspects of nutrient cycling
Long-term ecological studies of peat swamp, fresh water, kerangas
lowland and montane forests, have been carried out at the Gunung Palung
National Park, West Kalimantan by Harvard University group (see Leighton and
Darnaedi, 1996). The studies cover such aspects as vegetation structure and
composition, regeneration, phenology, and plant-animal interaction as the
bases for forest management. Much of the palaeoecological studies have been
carried out by Flenley and his students and the results are incorporated into his
book on "The equatorial rain forest: a geological history " (Flenley, 1979).

PROBLEMS AND CONSTRAINS

Data on timber species resulted from forest inventory undertaken by

Badan Planologi Kehutanan are enormous, but unfortunately timber species
identification is mainly based on vernacular names, rendered them scientifically

123
less valuable, otherwise they would constitute good data base for formulating
vegetation or at least forest types. To do vegetation analysis needs good
knowledge of plant identification and identification using vernacular names not
complemented with voucher specimens is unreliable; vernacular names are
generally not consistent and only few species have consistent vernacular
names. The results are definitely unreliable some time anecdotal and amusing.
Such practices can be found in various reports produced by forestry agencies,
consulting firms, and universities as well as student theses. Collection of
voucher specimens are mandatory in this regard, but it presents problems also
as good (fertile or sterile) herbarium specimens are sometimes difficult to
collect, thus relying only on fallen leaves. This is one of the constraints in
securing good data on tree enumeration which has been a nagging problem.
Only the Herbarium Bogoriense, Pusat Penelitian Biologi, LIPI, has the
capability to identify voucher specimens. Various agencies and universities in
the country has to rely on the services provided by the Herbarium Bogoriense,
where again its capability is constrained by limited number of trained staff able
to do the work. Regional herbaria with limited capabilities are available in
various universities and Balai Penelitian Kehutanan.
Trained plant ecologists able to undertake vegetation studies are few. It is
not uncommon also to find plant ecologists who are not able to do identification
and know plant species well and for identification they rely on the work of
parataxonomists or technicians and accepted the results uncritically. Efforts to
require young plant ecologists to master identification techniques and
familiarize themselves with species of major families prior to their assignment
as plan ecologists have been attempted. Teaching dendrology, plant taxonomy
and field botany in various universities are not adequate to make students able
to do identification and undertake acceptable vegetation studies.
Going through various publications of Indonesian scientists and theses of
graduate students, I am sad to find that most of the present-day researchers
and graduate students are not well-read; they seem to be disinclined to consult
both past and present literatures critical for their studies. Such reputable and
important pre-war journals published by research institutions and scientific
organizations in Bogor as Tectona, Tropische Natuur, Mededelingen Boshbouw-
proefstation, Bulletin du Jardin Botanique de Buitenzorg and Annales Bogoriensis,
etc., and even the Indonesian Forestry Abstract, summarizing Dutch literatures
up to 1960, are also rarely consulted. Furthermore, I noticed also that the
CIFOR library that houses important state-of-the-art international journals and
other publications on plant sciences, forestry, agroforestry ecology, conservation,
etc. are hardly visited by non-CIFOR scientists, graduate students and university
lecturers.
Currently biological journals (in Indonesian) published by various universities
are mushrooming. Unfortunately the quality of most papers and editing of local
journals need a great deal of improvement. Consequently poor articles do not
attract international scientists who often considered such articles as grey
literature or even thrash, hence they do not enter international circulation.

124
 CONCLUSION AND RECOMMENDATIONS

It is encouraging to record that the number of studies and publications on

vegetation science and related plant ecology has increased significantly since I
made an evaluation many years ago (Kartawinata, 1990), although it is not
pleasing to see that most of them were done by scientists of the Herbarium
Bogoriense, Pusat Penelitian Biologi, LIPI. In terms of quality many of the
reports and publications are not as good as those published in international
journals, hence they need improvement. Data presentation and editing are poor
that make further analyses and synthesis difficult. Only a few Indonesian
ecologists have been able to publish their articles in peered-reviewed
international journals.
So far research has been focused mainly on vegetation and dynamics of the
lowland rain forests of Sumatra and Kalimantan. Furthermore most of the
studies have been carried out in protected areas, such as National Parks and
few only undertaken in non-protected areas, including production and
conversion forests, which cover the largest proportion of primary forest areas
in the country and are vulnerable to destruction and even extinction.
Lack of trained personnel is one of the factors of the slow progress on the
studies of vegetation and ecology in Indonesia and advanced training is
necessary in this respect. Recruitment of undergraduate and graduate students
to work with senior plant ecologists in research institutions could help solving
this problem.
In order to ease the dependence of identification of plant specimens
necessary for botanical research and vegetation studies on the services
provided by the Herbarium Bogoriense, establishment of new regional herbaria
in various research institutions and universities should be encouraged and the
existing regional herbaria should be strengthened. In line with this, taxonomists
should be persuaded to write island flora and/or regional flora.
Opportunities for undertaking various facet of research on natural
vegetation of Indonesia is wide open. The followings are several aspects
recommended for future studies (see also Ashton, 1978; Whitmore, 1978):
1. Revising UNESCO vegetation map of Malesia with special reference to
Indonesia, mapping all ecosystem types, compiling available vegetation
maps and identification which ecosystem types have not been represented
in the protected area system of Indonesia.
2. More studies on floristic composition and structure of various plant
communities in both protecrted and non-protected areas are needed and
such undertaking is straight forward which can be even carried out by
students so that geographical picture can be established not only to test
scientific theory explaining the cause of patterns of species richness but also
for utilization, management and conservation of biodiversity. Standardized
methods are required for such studies in order to facilitate comparison and
synthesis on regional and national scales.

125
3. Studies on floristic composition and structure of various plant communities
along altitudinal gradients, complemented with soil and climatic data are
necessary to understand the relationships between vegetation and
environmental factors.
4. Research on fresh water and peat swamp forests, limestone forests, forests
on ultrabasic soils, seasonal forests, montane forests outside Mt. Gede-
Pangrango and Mt. Halimun, savannas, grasslands and littoral vegetation
should be encouraged as to date information and data about these plant
communities are not available.
5. Very high priority should also be given to studies of secondary forest which
is expanding very rapidly and yet remains poorly investigated.
6. Attention should be given also to research on all aspect of ethnobotany in
various vegetation types.
7. Studies on lianas and epiphytes should be promoted further since to date
they have been neglected.
8. There is an urgent need to develop rapid assessment of plant diversity
because current methods of plant species inventory require a great deal of
expertise and time. Expert on tropical plants is limited, resulting in slow
progress of inventories. In this regard training of parataxonomists will help
a great deal.
9. Intensified plant monographic studies, publication of local flora, island flora
and regional flora as well as plant collecting are urgently needed in all
island outside Java especially in Sulawesi, Maluku, Nusa Tenggara and
Papua. Availability of mono-graphs of various taxa as well as local flora,
island flora and regional flora will help identification.
10. Long-term synecological studies of pattern and process in permanent plots
in different forest types, coupled with meteorological stations, are necessary for:
a. Studies of growth rate covering periodic measurement of girth of all
mapped trees, seedling recruitment and mortality in representative
samples, and continuous phenological recording of all numbered
individuals.
b. Studies of dynamics in relation to canopy srtructure in both natural forest
and manipulated experimental forest plots:
i. Distribution of canopy gaps in space and time.
ii. Understorey composition, canopy species regeneration, understorey
dynamics as affected by canopy structure.
iii. The relationship between gap size on gap phase succession.
iv. Effects of gap size on soils.
v. Seed dormancy, germination, seedling competition, etc. in gap
succession.
vi. Investigation on diaspores whether they arrive after the formation of
a gap or are there and are released from dormancy.
c. Study on the ecology of succession:
i. Investigation on changes in floristic composition, structure and habitat
conditions through time.

126
 ii. Discovering whether secondary forest pioneer species colonize from
seeddormant in the soil or from seeds invading the clearing after its
creation.
iii. Study the ecology of good-quality and fast-growing secondary
species that have economic potentials useful for practical application.
iv. Ecological research on restoration and rehabilitation, applying prin-
ciples of succesion to provide bases for better implementation of
restoration and rehabilitation of deforested and degraded lands,
which in Indonesia cover millions of hectares. The scheme outlined in
Figure 5 may be followed in the implementation, in which techniques
and use of economically valuable fast growing seral species
suggested by Kartawinata (1994b, c), application of agroforestry
approaches and participation of local communities can be promoted.
11. Autecological studies, including phenology, reproductive ecology, fruiting
biology, seed ecology, seedling ecology, ecophysiology, ecology of exotic
invaders into tropical forests, and allelopathy in mixed forests (see Ashton
1978) as well as plant-soil relationships and production ecology to
increase basic understanding of mineral and water relations and nutrient
cycling in rainforest should be initiated.

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