Biodiversity and Conservation of Tropical Peat Swamp Forests

Author(s): Mary Rose C. Posa, Lahiru S. Wijedasa and Richard T. Corlett

Source: BioScience, Vol. 61, No. 1 (January 2011), pp. 49-57
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 Articles

Biodiversity and Conservation of

Tropical Peat Swamp Forests
Mary Rose C. Posa, Lahiru S. Wijedasa, and Richard T. Corlett

Tropical peat swamp forest is a unique ecosystem that is most extensive in Southeast Asia, where it is under enormous threat from logging, fire,
and land conversion. Recent research has shown this ecosystem’s significance as a global carbon store, but its value for biodiversity remains poorly
understood. We review the current status and biological knowledge of tropical peat swamp forests, as well as the impacts of human disturbances. We
demonstrate that these forests have distinct floral compositions, provide habitat for a considerable proportion of the region’s fauna, and are important
for the conservation of threatened taxa, particularly specialized freshwater fishes. However, we estimate that only 36% of the historical peat swamp
forest area remains, with only 9% currently in designated protected areas. Given that peat swamp forests are more vulnerable to synergies between
human disturbances than other forest ecosystems, their protection and restoration are conservation priorities that require urgent action.

Keywords: peat, Southeast Asia, blackwater, wetland, threatened

P eatlands are ecosystems characterized by the 

accumulation of partially decayed organic matter, called
peat, which is formed from plant debris under waterlogged
conditions (Andriesse 1988). In the tropics, peat and peaty
soils (histosols) form in a variety of conditions, but the
greatest peat depths—and thus carbon stores—occur in
peat swamp forests situated at low altitudes in the river
valley basins, watersheds, and subcoastal areas of Southeast
Asia. Until recently, the inaccessibility of peat swamp forests
and the belief that they support lower species diversity than
dryland rainforests meant that they received relatively little
attention from scientists (Prentice and Parish 1990, Yule
2008). Countries with extensive peat swamp forests have
tended to regard them as wastelands that must be converted
to more productive land use (e.g., Rijksen and Peerson 1991).
Thus, they remain poorly understood, and their importance
is underappreciated. Vast tracts of peat swamp forest have
already been degraded, and remaining areas are quickly
disappearing as a result of logging, fire, and conversion to
agriculture and industry (Yule 2008, Hansen et al. 2009).
In the last decade, there has been increased interest in the
role of peatlands in the global carbon cycle because of the large
amount of carbon they store and can potentially release to the
atmosphere (55 gigatons is estimated for Indonesia alone; Page
et al. 2004, Jaenicke et al. 2008). The realization that carbon
dioxide emissions from the clearance and burning of peat
swamp forests in Southeast Asia may comprise as much as 3%
of total global anthropogenic emissions (Ballhorn et al. 2009,
van der Werf et al. 2009) has sparked an explosion of interest in
their past, present, and future roles as carbon stores. However,
the biological implications arising from the loss and degrada-
tion of peat swamp forests are not yet fully understood, and
proper assessment is hampered by the lack of baseline data
on their flora and fauna. In this article, we summarize current
knowledge on (a) the development and biogeography of peat
swamp forests in Southeast Asia, (b) their current geographic
extent and conservation status, (c) peat swamp forest biodiver-
sity, and (d) threats and their impact on flora and fauna. We
restrict our review to the biogeographical region of Southeast
Asia and exclude the island of New Guinea because of the
paucity of information regarding the apparently extensive
peat swamp forests there. Our aim is to galvanize research and
emphasize the urgent need to afford more protection to this
neglected yet extremely valuable ecosystem.
Formation and biogeography of tropical peat swamps
In the tropical lowlands of Southeast Asia, peat soil is
formed from woody plant debris under high-rainfall and
high-temperature conditions, in contrast with temperate and
boreal regions where peat originates mainly from mosses and
herbs (Andriesse 1988, Chimner and Ewel 2005). In areas with
poor drainage, such as basins and valleys, peat can accumulate
over long periods of time until it is above the groundwater
level. This process eventually creates dome-shaped “ombrog-
enous” raised bogs that are fed only by rainwater and possess
their own perched water table, with the peat acting as a reser-
voir holding water by capillary forces (Andriesse 1988). Some
highly developed domes have peat depths of up to 20 meters
(m) and their centers may be dry (Anderson 1983). The bogs
are deficient in nutrients (oligotrophic) because of the lack
of mineral input, and the leaching of organic compounds
causes the water to become extremely acidic (pH 4 or less).
Ombrogenous peat swamps can also develop inland on flat or
gently convex areas between rivers in places with year-round
rainfall. Conversely, “topogenous” peat forms in areas with a
high water table where nutrients are available from seasonal

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flooding or underlying mineral soils. Extensive areas of peat
swamp vegetation occur on the island of New Guinea but
there is no record that they have a domed surface or are cor-
related with the forest types found in the rest of Southeast
Asia (Marshall and Beehler 2007). Similar forested bogs occur
in other tropical regions where appropriate topography and
climate coincide (e.g., Troxler 2007, Lähteenoja et al. 2009),
but little is known about their extent or biology.
The fossil record confirms the presence of angiosperm-
dominated peat swamp forests in Southeast Asia since the
Early Miocene (> 20 million years ago), and suggests that
these were floristically similar to contemporary forests by
the Middle Miocene, when they probably reached their
maximum extent and depth (Morley 2000, Belkin et al.
2009, Petersen et al. 2009). There are too few paleoecologi-
cal records from the Quaternary to assess the extent of peat
swamp forest, but a variety of evidence suggests that ever-
green rainforests remained extensive during the last glacial
maximum (Cannon et al. 2009), and dated cores show that
peat survived at inland sites, even when it was not actively
accumulating (Anshari et al. 2004, Page et al. 2004). In
contrast, dates from modern peat swamp forests in basins
behind the coast suggest that these formed only in the last
6000 years, after the stabilization of global sea levels. Similar
coastal peatlands presumably formed at previous sea-level
stillstands, although these must have been restricted in
extent during periods when sea levels fell below the margins
of the Sunda Shelf. Peat swamp forests in Southeast Asia are
thus collectively ancient but individually ephemeral, and
their total areal extent has probably fluctuated widely.
Current extent and protection status
Approximately 60% of the currently known tropical peat-
lands occur in Southeast Asia, mostly in Indonesia, Malaysia,
Brunei, and Thailand, and to a lesser extent in the Philippines
and Vietnam (table 1; Rieley et al. 1996, Joosten 2004). Peat-
lands can be found throughout the tropics (Rieley et al. 1996,
Andriesse 1988), where many types of wetlands are capable
of forming peat soils, but other forested peat deposits do
not reach the extent and depth of those in Southeast Asia
(e.g., Chimner and Ewel 2005, Troxler 2007, Lähteenoja
et al. 2009). There is still no consensus on the precise extent
and condition of tropical peatlands, as accurate delineation
of peat soils without ground truthing is difficult and many
areas have already been lost or degraded (Joosten 2004).
Estimates of the maximum predisturbance peatland area in
Southeast Asia range from 20 million to 30 million hectares
(ha) (Rieley et al. 1996). Using published estimates from
various sources, we calculate that a maximum of only 36%
of the historical peat swamp forest area remains (table 1,
figure 1). Even this number is likely to be an overestima-
tion—our calculation does not consider degradation, largely
from logging. It is evident from satellite images that very
little peat swamp forest remains undisturbed, but there has
been no regionwide analysis of the rates of clearance and
degradation. Forest loss has accelerated in recent decades
and has been exacerbated by fires, particularly in El Niño
years. For the whole of Borneo, the average annual defor-
estation rate for 2002–2005 was estimated as 2.2% for peat
swamp forest, compared with 1.8% for lowland dipterocarp
forest (Langner et al. 2007). Forest loss in the lowlands of
Sumatra and Kalimantan, the two Indonesian provinces
containing the largest areas of peat swamp forest, accounted
for more than 70% of forest clearing in the country from
1990–2005, resulting in a staggering 41% loss in total area in
just 15 years (Hansen et al. 2009).
We estimate that only around 9% of the region’s peat
swamp forests are currently in areas designated in the
United Nations Environment Programme World Conser-
vation Monitoring Centre and the International Union

Table 1. Estimates of major peat swamp forest areas (in square kilometers) in Southeast Asia.
Region Initial area Remaining area Percentage remaining Protected area Percentage protected

Indonesia
 Sumatra 82,525 25,622 31.1 7212 8.7
  Kalimantan 67,876 31,606 46.6 7632 11.2
 Sulawesi 3115 18 0.6 300 9.6
Malaysia
  Peninsular Malaysia 9845 2492 25.3 444 4.5
 Sabah and Sarawak 17,460 6328 36.2 984 5.6
Brunei 1040 873 83.9 218 21.0
Thailand 680 304 44.7 206 30.3
Southeast Asia total 182,541 67,243 36.8 16,995 9.3

Note: To determine which protected areas are found on peat, maps from the World Database of Protected Areas were overlaid with the World
Wildlife Fund Ecoregions of the World map. Areas listed as “proposed” were excluded. For national parks in Indonesia that encompass more than
one ecoregion type, we consulted other references to extract areas over peat where possible. Updated information for peat areas in the Philippines
and Vietnam were not found. Sources available from authors upon request.

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Figure 1. Extent of peat swamp forest (dark areas) in 2004 for Peninsular Malaysia, Sumatra, and Borneo. Peat swamp
forest cover was determined using the Global Landcover Product (ESA GlobCover Project, led by MEDIAS-France), along
with validation of vegetation type using the following soil maps: 1968 Reconnaisance Soil Map of Peninsular Malaysia
(Ministry of Agriculture and Fisheries, Malaysia); 1968 Soil Map of Sarawak (Land and Survey Department, Sarawak,
Directorate of National Mapping, Malaysia); 1974 The Soils of Sabah (Directorate of National Mapping, Malaysia); and
2000 Soil Resource Atlas of Indonesia (Departemen Pertanian, Badan Penelitian dan Pengembangan Pertanian Pusat
Penelitian Tanah Dan Agroklimat).

for Conservation of Nature (IUCN) World Database of
Protected Areas (table 1). The conservation status of many
of these sites is unclear, as they have not been assigned
IUCN protected area categories, and therefore the primary
management objective is not known. Nevertheless, even the
declaration of an area as a national park does not guarantee
long-term conservation, as significant forest clearance, log-
ging, and hunting occur within park boundaries in many
parts of Southeast Asia, particularly in Indonesia (Curran
et al. 2004, Corlett 2009, Gaveau et al. 2009). Such degrada-
tion is especially detrimental to peat swamp forests, as it
increases susceptibility to fire during droughts (as discussed
below).
Peat swamp biodiversity
Tropical peat swamp forests remain inadequately under-
stood scientifically, which is alarming given the vast areas
that have been lost or degraded. The paucity of informa-
tion is in part the result of early views that collecting trips
to peat swamp forests were a waste of time because of
their low animal diversity and abundance. This, coupled
with the extremely difficult logistics posed by swampy
conditions, has until recently discouraged biologists from
survey work. In contrast, foresters and soil scientists have
appraised peat swamp forests for their timber concession
and agriculture potential since the mid-20th century. It
has been asserted in several papers that peat swamps har-
bor significant and highly specialized biodiversity (e.g.,
Phillips 1990, Page et al. 1997, Yule 2008), but this has yet
to be comprehensively assessed. We compiled the avail-
able species lists for angiosperms and vertebrates, and
estimated the numbers of species in Southeast Asia that
have been observed in peat swamp forests, and those that
are restricted to or strongly associated with (e.g., recorded
more often or occur in higher densities than other habi-
tats) this ecosystem (table 2). These data must be treated
with caution, since many areas are still unsurveyed and the
complete distributions of many species are not yet known.
However, the major patterns indicated by our data are
likely to be robust to future additions and corrections.

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Table 2. Number of species recorded from peat swamp forests of Southeast Asia.
Total number of species Plants Mammals Birds Reptiles Amphibians Freshwater fish

Recorded from PSF 1524 123 268 75 27 219

Restricted to PSF 172 0 0 0 0 80
Species strongly associated with PSF — 6 5 1 3 —

PSF, peat swamp forests.

Source: Data compiled from various sources available from authors by request.

The small number of nonfish species confined to peat
swamp forest is surprising for an ancient tropical ecosystem
with a large total area spread across several biodiversity hot
spots. The lack of endemic terrestrial vertebrates probably
largely reflects the relatively ephemeral existence of individual
peat swamp forest areas that we highlighted above, although
it is not possible to rule out an impact from a reduced total
areal extent during low Pleistocene sea levels. When new
peat swamp forest patches develop in a nonpeat landscape,
species must be acquired from surrounding ecosystems. For
the flora, at least, the extensive and relatively permanent
heath forests that occur on coarse, white-sand soils with
trees adapted to water stress and nutrient-poor conditions
appear to have been the major source of species, along with
topogenous freshwater swamp forests that, although also
locally ephemeral, are well connected through river systems.
Other taxa have apparently been acquired from a variety
of other forest types. Peat swamp forest endemism is most
likely to have evolved in organisms that (a) directly inhabit
the environmentally rigorous microhabitats provided by
the peat and the associated waters, and (b) could track the
occurrence of these habitats across the Sunda Shelf during
glacial cycles. Conversely, canopy-living species or species
that are very poorly dispersed are unlikely to have the need
or opportunity to evolve specialized adaptations to this habi-
tat. Although the peat swamp forest specialists are clearly of
greatest conservation concern, the astonishingly high rates of
deforestation in lowland Sumatra and Kalimantan in recent
decades mean that peat swamp forests have now become
important refuges for many species that can use a wide range
of other forest types. In the following sections we expand on
what information is known of peat swamp flora and fauna.
Flora.  The extreme chemical and hydrological conditions
of peat swamp forests apparently restrict local and regional
tree diversities. However, tropical peat swamp forests have
the highest floral diversity compared with all other peatland
types in the world, the majority of which are found in the
Northern Hemisphere and are dominated by Sphagnum moss
or grasses. There are between 30 and 122 tree species greater
than 10 centimeters in diameter recorded in 1-ha plots of
peat swamp forest in Southeast Asia, a number lower than
most dryland rainforests in the same region (100 to 290
species) but similar to heath forests (29 to 129 species)
(Purwaningsih and Yusuf 2000, Corlett 2009). Adaptations
to acidic, nutrient-deficient soils are required for species
to grow in the highly oligotrophic heath forests, which
are particularly extensive in the coastal regions of Borneo.
Adaptations to high and fluctuating water levels, such as
pneumatophores, stilt roots, and buttresses, are required for
the relatively nutrient-rich freshwater swamp forests. Peat
swamp forest, on the other hand, is unique in requiring
both sets of adaptations at the same time. In our literature
review, around 11% of plants recorded from peat swamp
forests are reported only from that habitat (table 2). The
majority (63%) of these restricted species are trees, followed
by epiphytes and climbers (16%). The high proportion of
restricted epiphytes is surprising, since the canopy of peat
swamp forest does not provide a unique environment,
but the result may simply reflect the undercollection of
epiphytes in the taller, denser forests of dryland sites.
Most tree families and many genera of lowland evergreen
dipterocarp rainforest are found in peat swamp forest, but
the species composition is distinctive, except on very shallow
peats (Anderson 1983). Anderson (1963) was the first
to undertake rigorous ecological characterization of peat
swamp forest flora, describing a catenary sequence of forest
“phasic communities” in concentric zones on ombrogenous
domes in Sarawak and Brunei. These range from mixed
swamp forest, similar in species composition to dryland
rainforests, on the more fertile periphery of the dome, to
stunted vegetation with xeromorphic features in the dry
center. Other peat swamp forests across Southeast Asia have
a similar concentric sequence of forest types, but they dif-
fer from one another in species composition and structure
(Rieley et al. 1996), varying in response to the local gradi-
ents in hydrology, nutrient availability, depth of peat (Page
et al. 1999), and presumably, the availability of species within
dispersal distance. Thus, there is no single peat swamp for-
est vegetation, and this ecological and floristic heterogeneity
suggests that each swamp is distinct (Corner 1978). For
example, on Borneo, the peat swamp forests of Sarawak and
Brunei are dominated mainly by the endemic dipterocarp
Shorea albida, but in Kalimantan, this species is found only
in the northwest. In southern Kalimantan, the congener
Shorea balangeran can be found in pure stands and may play
the same role as S. albida in these forests. This broad-scale
heterogeneity in floristic composition means that extensive
protection of multiple sites will be needed to encompass the
full range of variation of this ecosystem.

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Fauna.  The low nutrient content of ombrotrophic peats
results in a relatively low primary productivity compared
with other tropical forest formations (Bruenig and Droste
1995). All else being equal, this would result in a lower
diversity and abundance of fauna, although only a few stud-
ies have made direct comparisons with different forest eco-
systems (e.g., Gaither 1994). There have been no systematic
studies of animal distribution by forest phasic community,
but anecdotal evidence suggests greater wildlife abundance
in the mixed swamp forest at the margins of peat domes
(Phillips 1990, Page et al. 1997). Compiling available data
from faunal surveys, we found that 23% to 32% of all species
of mammals and birds in Peninsular Malaysia and Borneo
have been recorded from peat swamp habitats (figure 2).
A similar percentage of freshwater fish were represented in
peat swamps in Peninsular Malaysia (33%), but not Borneo
(10%). This most likely reflects the lack of comprehensive
sampling in the latter. The proportions are somewhat lower
for snakes (7% to 18%) and amphibians (19% to 23%),
Figure 2. Percentage of fauna in Peninsular Malaysia and
Borneo recorded from peat swamp forest.
which may reflect their more intimate contact with the
extreme substrate and water conditions. On the whole, these
results show that peat swamp forests provide a habitat for a
considerable proportion of the region’s fauna.
Aside from fish, no vertebrate species can be considered
entirely dependent on peat swamp habitats (table 2). How-
ever, peat swamp forests support a substantial number of
rare, specialized, and threatened species. We found that 45%
of mammals and 33% of birds recorded in peat swamp for-
ests had an IUCN Red List status of near threatened, vulner-
able, or endangered (figure 3). Eight species of threatened
freshwater turtle have been recorded, indicating that peat
swamps are an important habitat for this highly endangered
group. Also, because peat swamp forests are usually found
in an ecological mosaic with other forest ecosystems, they
can provide resources for species with wide ranges and
may be a sanctuary for species whose habitats have been
altered by human activities (see Gaither 1994, Page et al.
1997, Singleton and van Schaik 2001, Struebig et al. 2006).
For instance, in Sarawak, all maternity colonies of the large
flying fox (Pteropus vampyrus natunae) are found in remote
and inaccessible areas in mangrove and peat swamp forest
(Gumal 2004).
Peat swamps are also important for the conservation
of a number of endangered primate species. The richest
habitats for orangutans are high-quality swamp forests and
lowland alluvial forests (Russon et al. 2001). In Gunung
Palung National Park in western Kalimantan, primary peat
forest had a higher density of Bornean orangutan (Pongo
pygmaeus) nests (49% more) and individuals (31% more)
than lowland forest (Johnson et al. 2005). The Seban-
gau catchment in central Kalimantan supports the largest
single orangutan population in Borneo (Morrogh-Bernard
et al. 2003). Peat swamp forests are also important for the

Figure 3. Proportions of threatened fauna found in peat swamp forests. Numbers of species and IUCN Red List categories
are given in each slice. CR, critically endangered; DD, data deficient; EN, endangered; LC, least concern; NT, near
threatened; VU, vulnerable.

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conservation of other primates, such as proboscis monkeys
(Nasalis larvatus), the Bornean banded langur (Presbytis
chrysomelas) (Phillips 1990) and the four species endemic to
Siberut island, Hylobates klossi, Presbytis potenziani, Macaca
siberu, and Simias concolor (Quinten et al. 2009). A num-
ber of endangered felids also make use of swamp forests,
including the flat-headed cat (Prionailurus planiceps), Sunda
clouded leopard (Neofelis diardi), and marbled cat (Pardofe-
lis marmorata) (Cheyne et al. 2009). Finally, peat swamp
forest is the favored habitat of the endangered false gharial
(Tomistoma schlegelii) (Bezuijen et al. 2001).
Among the faunal groups, fish exhibit the highest ende-
micity to peat swamps (table 2). Work in Peninsular Malay-
sia has shown that the blackwaters of peat swamps are not
species poor or low in biomass, and up to 33% of the known
freshwater fish species are associated with peat swamps
(figure 2; Ng et al. 1994, Kottelat et al. 2006). Peat swamps
also harbor a number of miniature fishes, including Paedo-
cypris progenetica, the smallest known vertebrate (Kottelat
et al. 2006). Many of these fishes were discovered only in
the last 20 years and many more await formal description.
Of the 219 fish species identified in our data collation of
fauna recorded from peat swamps, 80 species are restricted
to this ecosystem, 31 of which are point endemic species
found only in single locations. Kottelat and colleagues
(2006) suggested that the conditions in peat swamps have
favored the evolution of these specialized fish species,
and that each patch of peat swamp could contain its own
suite of endemic species. However, even the most rapid
presumed rate of speciation could not have produced
local-endemic fish faunas within the few thousand years
that appear to have been available at most sites. Thus,
this question requires further study and the sampling of
adjacent but unconnected peat swamp forests and the con-
struction of calibrated molecular phylogenies. Compre-
hensive studies of peat swamp fish have been carried out in
only a few regions: Thailand (Vidthayanon 2002); Jambi,
central Sumatra (Tan and Kottelat 2009); north Selangor,
Peninsular Malaysia (Ng et al. 1992); and Pulau Bintan,
Riau Archipelago (Tan and Tan 1994). Unfortunately, the
ongoing destruction of peat swamp forests means that
such research will soon be impossible. Given the extremely
restricted ranges of many of these species, their long-term
survival is unlikely unless urgent conservation action is
taken. For instance, the critically endangered Betta perse-
phone, Betta miniopinna, and Betta spilotogena are listed
on the IUCN Red List as highly threatened by extinction as
a result of declines in area of occupancy, extent of occur-
rence, and quality of habitat. At the moment, there are 17
species of peat swamp fish on the IUCN Red List classified
as vulnerable, endangered, or critically endangered, 12
of which are point endemics. The threat statuses of the
majority of this ecosystem’s specialized fishes have yet to
be assessed and they must be incorporated in future con-
servation, as they are the most distinctive element of the
peat swamp forest vertebrate fauna.
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Invertebrates of peat swamp forests are less well known—
surveys are usually short, not comprehensive, and do not
identify organisms down to the species level. There do
not seem to be any insects specific to or characteristic
of peat swamp forests, and the fauna has been described
as impoverished (e.g., Ng et al. 1992, Abang and Hill
2006). However, among aquatic invertebrates, a few rotifer
(Chittapun et al. 2007) and decapod crustacean species
(Ng et al. 1992, Wowor et al. 2009) have been found to be
restricted to blackwater habitats. The distribution of fresh-
water Macrobrachium prawns found in acidic peat habitats
is surprisingly wide for species adapted to such a specialized
habitat and merits further study (Wowor et al. 2009).
Threats and their impacts
Perhaps more than any other forest ecosystem, tropical peat
swamp forest is susceptible to the synergistic effects of mul-
tiple human disturbances because of the balance that exists
among vegetation, peat, and hydrology. Positive feedback
loops among deforestation, drainage, and fire mean that
peatlands are easily degraded once the balance is upset.
In following sections we discuss the impacts of the major
threats to this ecosystem.
Logging.  Tropical peat swamp forests contain a number of
valuable timber species, sometimes at high densities, and
have thus been intensively exploited all over Southeast Asia.
There is very little information on the long-term effects
of logging on the flora and fauna of peat swamp forests;
however, evidence from forestry assessments indicates that
selective logging causes changes in forest structure and
composition (Bruenig and Droste 1995, Hadisuparto 1996).
For instance, logging disturbance leads to a lack of regenera-
tion by ramin trees (Gonystylus bancanus), the most valu-
able timber species in peat swamps, which is now listed as
vulnerable on the IUCN Red List. Early logging operations
were possible only through the use of manual labor, and logs
were moved by railway, but the mechanization of methods
has led to more intensive extraction and greater damage to
the residual forest (up to 50%; Rashid and Ibrahim 1994).
Sustainable harvest of timber, which is rarely practiced,
would require strict standards of long felling cycles and high
diameter limits, as well as total protection of the more frag-
ile phasic communities (Bruenig and Droste 1995). Canals
cut for floating out logs—often also used in illegal logging
operations—are particularly damaging because they change
hydrological conditions, making peat swamp forest more
vulnerable to fire (see below).
Even where hydrology is not compromised, impacts of log-
ging on fauna in peat swamp forest are similar to those reported
in other forest types. Logging was found to reduce orangutan
densities by 21% to 22% when compared with unlogged
peat swamp forest, with lower densities in more recently
disturbed areas (Felton et al. 2003, Morrogh-Bernard et al.
2003, Johnson et al. 2005). It has been postulated that this
trend was the result of changes in habitat quality that include

altered tree size distribution, reduced availability of large
fruit crops, and greater extent of canopy gaps (Felton
et al. 2003). Other primate species and large mammals may
be similarly negatively affected by these changes, but this
question remains unstudied. A short study demonstrated
that the abundance and species diversity of ground-active
small mammals increased after selective logging (Doody et al.
1997), which may indicate they are favored by disturbance.
Moreover, 80% of bird species in pristine peat swamp forest
were also observed in selectively logged areas (Page et al.
1997). More rigorous research is needed to determine the
specific impacts of logging on peat swamp forest fauna.
Fire.  Tropical peat swamp forests are more vulnerable to
destruction by fire than any other forest type because the soil
substrate itself is extremely flammable when dry (Langner
et al. 2007, Langner and Siegert 2009). On Borneo, 73% and
55% of the forests affected by fire in 2002 and 2005, respec-
tively, were on peat (Langner et al. 2007). Transects in Tan-
jung Puting National Park peat swamp forest surveyed after
fires from 1997–1998 had 13% to 69% fewer species than
before, compared with a 7% to 31% loss of species in low-
land dipterocarp forests in Sungai Wain Nature Reserve and
Kutai National Park (Yeager et al. 2003). The general effects
of fire on peat swamp forest vegetation are similar to those
in other forest types—burned forest has lower canopy cover,
decreased species richness, and reduced tree and sapling
density compared with unburned forest (Yeager et al. 2003).
However, because of peat’s high combustibility, fires can
burn both above and below the surface, destroying vegeta-
tive structures underground, as well as the seed bank (Page
et al. 2009). Deep peat fires can smolder below the surface
for months and are difficult to extinguish. These subsurface
fires can also cause the collapse of overlying material, creat-
ing additional tree mortality.
During extreme droughts, often associated in Southeast
Asia with strong El Niño events, peat swamps are extremely
susceptible to burning, as evidenced by recent large wildfires
that have destroyed hundreds of thousands of hectares on
Borneo (Langner and Siegert 2009, Page et al. 2009). Previous
disturbance greatly exacerbates the severity of fire damage.
For instance, logging creates gaps in the canopy that change
forest microclimates and increase the temperature of the peat
surface, whereas any drainage will cause the water table to
drop below critical levels (i.e., 0.5 meters to 1 meter below the
surface) and can result in shrinkage and irreversible drying of
the peat (Andriesse 1988, Wosten et al. 2006). An assessment
of the effects of the 1982–1983 El Niño event in east Kalim-
antan found that 97% of the swamp forest plots that were
previously disturbed had been burnt (Goldammer 2007).
Areas that have been logged and then burned multiple times
have dramatically lower stem densities and diversities rela-
tive to areas that have burned only once and had little or no
logging history (Yeager et al. 2003, Page et al. 2009). Peat forest
that has been subjected to one fire has a high probability of
burning again because of the accumulation of unburned dead
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Articles
biomass and the presence of fire-prone regrowth vegetation
that can act as fuel for the next fire (Siegert et al. 2001, Page
et al. 2009). Analyses of fires in central Kalimantan showed
that repeated fires occur mostly in nonforest secondary veg-
etation, and that fire incidence is becoming decoupled from
El Niño events (Page et al. 2009). Greater burn intensity and
frequency causes the vegetation to become dominated by
low-growing herbaceous plants, with few or no trees (Page
et al. 2009). Peat combustion also reduces surface elevation;
modeling predicts that where both vegetation and peat is
burned, flooding will increase and impede plant regeneration
(Wosten et al. 2006, Page et al. 2009).
Land conversion.  With much of the available arable land on
mineral soils already under cultivation, peat swamps are one
of the last frontiers for production of food, pulp, and biofuels.
Peat is considered a “problem” soil because its physical
and chemical properties make the cultivation of many
agricultural crops difficult (Andriesse 1988). Large-scale
land conversion of peat for agricultural purposes requires
clearance, drainage, fertilizer application, and liming to
increase the pH and boost microbial activity. Construction
of canal systems for drainage and water-table management
is often the first step. Drainage has numerous negative
consequences, chief of which is the greater decomposition
of organic matter by microbial oxidation. Decomposition
eventually leads to lowering of the peat surface (subsidence)
and, if underlying acid sulfate soil is exposed, acidification
(Wosten et al. 1997). Oxidation releases carbon dioxide and
other greenhouse gases, significantly contributing to climate
change (Couwenberg et al. 2009, van der Werf et al. 2009). To
exacerbate matters, both corporations and smallholders use
fire as a cheap method of clearing land, and unless efforts are
made to control the area burned, fires can easily spread and
run out of control (Page et al. 2009). Many attempts to grow
crops on peat were performed without proper knowledge of
the soil's unique nature and have resulted in failure and the
degradation of vast areas of peatlands (e.g., Ex-Mega Rice
Project in central Kalimantan; see Page et al. 2009); these
areas then become prone to fire. However, there have also
been at least short-term successes with tree crops, such as
Acacia crassicarpa (grown for pulp) and oil palm (Elaeis
guineensis). These successes have encouraged the current
pressure on peat swamps, and development continues
despite repeated calls for a moratorium (Pearce 2007,
Koh et al. 2009). It has yet to be seen whether carbon
credit–offset schemes such as REDD (reducing emissions
from deforestation and forest degradation) will convince
governments to protect and restore tropical peatlands.
Conclusions
Until very recently, tropical peat swamp forests were
a neglected ecosystem with low conservation priority.
Well-justified concerns about their increasingly impor-
tant role in global anthropogenic carbon dioxide emis-
sions have attracted much attention, but have not led to
Articles
a commensurate growth in the understanding of their
biodiversity value. We reviewed the available information
and demonstrated that peat swamp forests are immensely
important for the conservation of freshwater fish diver-
sity and are of clear conservation significance for a wide
range of plants and nonfish vertebrates. Further research
on previously neglected invertebrate groups is likely to
reveal additional peat swamp forest–dependent species,
particularly among taxa that come into intimate contact
with the peat and associated waters. Research is urgently
needed into the biodiversity and ecology of the remain-
ing peat swamp forests in Southeast Asia and those in
the Indonesian province of Papua, which are likely to
face the same threats in the longer term. Given the large
areas of damaged but not totally destroyed peatlands in
the region, there is also an urgent need for research into
the possibilities for restoration. Decisionmakers must be
vm of the fragility of the ecosystems and the negative
consequences of peat degradation at local, regional, and
global scales. Finally, protection cannot wait for research:
We already know enough to conclude that the conversion
of deep peats into marginally profitable plantation agri-
culture makes no environmental, economic, or political
sense. The future of tropical peat swamp forests depends
on urgent conservation and policy action to safeguard
their existence.
Acknowledgments
The authors thank Sue Page, Aljosja Hooijer, Heok Hui Tan,
and Robert Morley for useful discussions on tropical peat
swamp forests, and three anonymous reviewers for their
comments on the manuscript. MRCP was supported by the
Singapore-Delft Water Alliance peatland research program
(R 264-001-004-272).
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Richard T. Corlett is a professor, in the Department of Biological Sciences at the
National University of Singapore. Lahiru S. Wijedasa is a senior arborist at the
Botany Centre at Singapore Botanic Gardens.