Wetlands (2013) 33:257267

DOI 10.1007/s13157-013-0378-0

ARTICLE

Using Conceptual Models to Understand Ecosystem

Function and Impacts of Human Activities in Tropical
Peat-swamp Forests
Mark E. Harrison

Received: 27 August 2012 / Accepted: 10 January 2013 / Published online: 25 January 2013
# Society of Wetland Scientists 2013

Abstract Increased resource investment in conservation is

generating greater appreciation of the importance of ecological monitoring programmes to assess the effectiveness of
conservation interventions in achieving their stated goals. A
key component for developing such a programme is an
appropriate conceptual model of ecosystem function and
the effects of human activities on this. Tropical peatswamp forests are a particularly important ecosystem for
reducing carbon emissions and biodiversity conservation.
This is leading to increased investment in their protection
and, consequently, an increased need for effective ecological
monitoring programmes and conceptual models of ecosystem function on which to base these. Here, I adapt previous
conceptual models of ecosystem function developed for
terrestrial forests and other wetland ecosystems, to create a
habitat-specific model for tropical peat-swamp forests. This
provides a tool to guide thinking in developing ecological
monitoring studies in this habitat and understanding (i)
ecosystem processes and function, (ii) the impact of human
activities on these, and (iii) subsequent impacts on ecosystem services. This is relevant for monitoring the effectiveness of conservation interventions with varying goals in
tropical peat-swamp forest, including emission reductions;
and highlights considerations relevant for conservation
M. E. Harrison (*)
Department of Geography, University of Leicester,
University Road,
Leicester( LE1 7RH, UK
e-mail: mharrison@outrop.com
M. E. Harrison
Orangutan Tropical Peatland Project, Jl. Semeru 91,
Palangka Raya 73112, Central Kalimantan, Indonesia
M. E. Harrison
Centre for the International Cooperation in Sustainable
Management of Tropical Peatlands, University of Palangka Raya,
Palangka Raya 73112, Central Kalimantan, Indonesia

management and ecological monitoring in other wetland

habitats.
Keywords Conceptual model . Ecological monitoring .
Ecosystem services . Peat-swamp forest . Anthropogenic
threats . Management interventions

Introduction
Preventing biodiversity loss and curtailing global warming
are two of the most pressing challenges facing humanity
today. Reducing deforestation of tropical wet and dryland
forests is considered one of the cheapest and most effective
ways to mitigate these threats, owing to these forests high
levels of biodiversity and carbon storage, and the interrelatedness of these threats (Whitmore and Sayer 1992;
IPCC 2007; Stern 2007). Consequently, there is international interest in ensuring that the development of mechanisms
for reducing carbon emissions through forest protectioni.e.,
Reduced Emissions from Deforestation and Degradation
(REDD)also leads to positive impacts on biodiversity and
local forest communities (CCBA 2008; Venter et al. 2009a;
Harvey et al. 2010; projects implementing measures to
achieve these co-benefits are termed REDD+ projects).
Other economic incentives for habitat protection focussed
primarily on biodiversity, such as biodiversity offsetting/banking, are also beginning to receive interest in the tropics and
have potential to achieve a similar suite of co-benefits (e.g.,
Yaap et al. 2010; Kumaraswamy and Udayakumar 2011).
Although cheap compared to other methods, reducing forest loss and degradation still requires significant financial
investment. In 1999, the total funds allocated towards nature
reserve protection was an estimated USD 6 billion; USD 300
billion was estimated as necessary for a comprehensive global
conservation programme (James et al. 1999). This high level
of funds required for effective forest protection, plus the need

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Wetlands (2013) 33:257267

for transparency in global carbon and biodiversity offset markets, has led to increased recognition of the importance of
long-term post-implementation monitoring to assess the effectiveness of conservation interventions (i.e., management
actions/initiatives) for achieving their stated aims and enabling
intervention adaptation to maximise conservation success
(e.g., Ferraro and Pattanayak 2006; Lovett et al. 2007;
CCBA 2008; Gardner 2010; Lindenmayer and Likens
2010). Development of robust conceptual models to help
understand ecosystem function, and how human activities
influence ecosystem processes, functioning and service provision, is considered critical for developing appropriate questions for establishing the impacts of human activities on an
ecosystem (Lindenmayer and Likens 2009; Gardner 2010).
These models illustrate linkages between ecosystem components, thereby assisting us in predicting and interpreting the
impacts of human activities on these, particularly regarding
indirect impacts that may not be immediately obvious.
Tropical peat-swamp forests (PSF, Table 1) are a key
habitat for both reducing carbon emissions and biodiversity
conservation. PSFs are excellent carbon stores, with an estimated tropical peatland carbon pool of 88.6 Gt, or 1519 % of
the global peat carbon pool (Page et al. 2010). Consequently,
massive carbon emissions accompany PSF degradation and
burning (van der Werf et al. 2008; Hooijer et al. 2009): an

Table 1 Characteristics of tropical peat-swamp forests

estimated 0.812.57 Gt of carbon were released from peat

fires in Indonesia during the 1997 El Nio dry-season fires,
which was equivalent to 1340 % of mean annual global
emissions from fossil fuel combustion (Page et al. 2002).
PSFs also harbour diverse biological communities, including
many PSF-endemic species (Ng et al. 1994; Page et al. 1999;
Morrogh-Bernard 2009; Yule 2010; Posa et al. 2011). This
includes the worlds largest populations of the endangered
Bornean orang-utan (Pongo pygmaeus, Morrogh-Bernard et
al. 2003; Wich et al. 2008) and southern gibbon (Hylobates
albibarbis, Cheyne et al. 2008) in the Sabangau PSF,
Indonesian Borneo. Consequently, not only are payments for
REDD+ credits (Ebeling and Yasue 2008) on PSF likely to
offset the high opportunity costs of forest protection, but these
forests also contain higher-than-average numbers of threatened mammals, indicating great potential of REDD+ in PSF
for simultaneously achieving climate change and biodiversity
conservation objectives (Venter et al. 2009b). South-east
Asian PSF is also being rapidly converted and degraded
with 5.1 Mha lost between 1990 and 2008 (Miettinen and
Liew 2010)supporting arguments for the additionality of
PSF conservation schemes, such as REDD+.
This suite of anticipated benefits has led to a recent proliferation in forest protection and REDD+ pilot projects in PSF,
particularly in Indonesia (Paoli et al. 2010). Accompanying

Characteristic

Description

References

Geographical
distribution
Est. total area
covered
Peat type

South-east and East Asia, Tropical Central

and South America, and southern Africa
441,025 km2; highest country coverage in
Indonesia, with 206,950 km2
Fibric at surface to sapric near bottom
deposits
0.330+ m
65 %

Page et al. (2010)

Peat depth range

Organic content
peat
Altitude

Peat carbon
density
Above-ground
biomass
Canopy height

Dominant trees

Major threats

Page et al. (2010)

Wsten et al. (2008)
Page et al. (2010)
Page et al. (2010)

Mostly very low altitude, but some

high-altitude mountainous deposits
also exist
2,772 t C/ha (Indonesia)

Page et al. (2010)

100150 t C/ha (Indonesia)

Page et al. (2006)

Variable, but typically low; e.g. 1245 m

in Sabangau, with small open interior areas,
in which tree height does not exceed 1.5 m
Variable, but with many swamp specialists. In
Indonesia, important species include members
of the Anacardiaceae, Annonaceae,
Apocynaceae, Clusiaceae, Dipterocarpaceae,
Ebenaceae, Euphorbiaceae, Fagaceae,
Hypericaceae, Lauraceae, Myristicaceae,
Myrtaceae, and Sapotaceae families
Drainage, fire, conversion, logging

Page et al. (1999)

Page et al. (2010)

Page et al. (1999);

Morrogh-Bernard
(2009)

See citations and

discussion in text

Wetlands (2013) 33:257267

this is an increase in the need to establish reliable ecological

monitoring programmes for assessing the effectiveness of
existing PSF management interventions, adapting interventions to increase their success and considering the potential
utility of novel interventions. Conceptual models of ecosystem function have proven useful for investigating these issues
in other habitats (Lindenmayer and Likens 2009; Gardner
2010). Although various models and allometric equations do
exist for certain PSF ecosystem processes, such as drainage,
plus carbon fluxes and emissions (e.g., Wsten et al. 2008;
Henson 2009), these are rather specific, with the result that
indirect or knock-on effects of the events depicted may not
become immediately obvious to conservation managers.
Thus, in light of their conservation importance, development
of a general conceptual model of ecosystem function for PSF
systems is timely and necessary.

Basis for a Conceptual Model of Ecosystem Function

Specific to Tropical Peat-swamp Forest
Possibly the most successful and widely-cited example of a
conceptual model of ecosystem function guiding ecological
monitoring research is that developed by Bormann and
Likens (1967) for use in the Hubbard Brook Experimental
Forest, New Hampshire, USA. Here, nutrient cycling between the nutrient pools within a terrestrial ecosystem is
depicted, alongside nutrient inputs and outputs from the
external biosphere. Subsequent model expansions illustrate
more explicitly the paths between different nutrient pools
(Likens and Bormann 1995), plus the influence of controllers driving ecosystem patterns and processes, and their
impacts on ecosystem services (Groffman et al. 2004).
These terrestrial forest conceptual models lack an essential
component present in wetlands, howeversurface/sub-surface water as a major within-system nutrient poolthe importance of which has been recognised in conceptual
frameworks and models created by scientists working in various wetland habitats (Canadian and Scottish temperate peat
bogs: Frolking et al. 2010 and Smith et al. 2010, respectively;
North American depressional wetlands: Euliss et al. 2004;
Texan riparian habitats: Arnold et al. 2001; Mississippi Delta
wetlands: Wicker et al. 1982; freshwater wetlands and mangroves: Mazzotti et al. 2005; Berger et al. 2008; and particularly Floridian cypress domes and other wetlands: Odum
1983; Brown 1988; Barnes 2005; Browder et al. 2005;
Davis et al. 2005; Havens and Gawlik 2005; Ogden 2005;
Ogden et al. 2005b; Rudnick et al. 2005). Conceptual models
incorporating water as a key component have been also been
useful in analyses concerning the use of wetlands in wastewater treatment (Heliotis and DeWitt 1983; Ko et al. 2004).
As in other wetland habitats, surface/sub-surface water is a
critical component of the PSF ecosystem and plays a major

259

role in nutrient cycling (Wsten et al. 2006; Couwenberg et al.

2010; Dommain et al. 2010; Hooijer et al. 2010; Moore et al.
2010).
Tropical PSFs form over millennia in poorly-drained areas
with high rainfall, where near-permanent water-logging creates anaerobic conditions and substrate acidification, leading
to peat formation, as the organic matter decomposition rate is
exceeded by the addition of dead forest materials (Rieley et al.
1992; Page et al. 2004; Wsten et al. 2008). Given this slow
rate of decomposition, PSF food webs are thought to be
supported mainly by leaching of dissolved organic carbon
and other nutrients from freshly-fallen leaves (Weiss et al.
2002; Yule and Gomez 2009; Yule 2010), with surface/subsurface water-borne nutrients present in forms both available
and unavailable for uptake by forest biota. Recognising these
key ecosystem characteristics is important for generating a
useful general conceptual model for the ecosystem.

A Conceptual Model for Ecosystem Function

in Peat-swamp Forest
A conceptual model for PSF ecosystem functioning is
depicted in Fig. 1. This builds upon previous models for
ecosystem function in wetlands (Wicker et al. 1982; Heliotis
and DeWitt 1983; Odum 1983; Brown 1988; Arnold et al.
2001; Euliss et al. 2004; Ko et al. 2004; Barnes 2005;
Browder et al. 2005; Davis et al. 2005; Havens and
Gawlik 2005; Mazzotti et al. 2005; Ogden 2005; Ogden et
al. 2005b; Rudnick et al. 2005; Frolking et al. 2010) through
adopting and adapting the framework established previously
for terrestrial forests in the Hubbard Brook Experimental
Forest (Groffman et al. 2004), to link drivers of PSF ecosystem patterns and processes to ecosystem function and
services. Meteorological, geological and biological inputs
and outputs from the system (cf. Likens and Bormann 1995)
are not shown. This model fits within the DPSIR (Driving
forcePressureStateImpactResponse) framework for
reporting on environmental issues (Gabrielsen and Bosch
2003), thereby facilitating the development of policy
actions. Terminology follows Harrington et al. (2010) with
respect to ecosystem processes, ecosystem function and
ecosystem services. Because our knowledge of basic PSF
ecology is incomplete, many of the linkages illustrated
should be treated as hypotheses for future testing. I hope
that this paper stimulates research to test these hypotheses,
thereby facilitating the development of more accurate future
models.
Following Groffman et al. (2004), the model depicts the
two types of controllers that drive ecosystem patterns and
processes: state and variable-stochastic factors. State
factors represent the five controllers of soil formation that
explain and predict spatial patterns in soil properties (Jenny

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Wetlands (2013) 33:257267

Understanding these enables management interventions to be

targeted towards mitigating negative anthropogenic stochastic
factors, to maintain ecosystem function and services.
As in conceptual models of ecosystem function developed
for other wetland habitats, the surface/sub-surface water nutrient pool in this PSF model is considered an intra-system
component, with nutrients cycling through it in forms both
available and unavailable for uptake by biota. Water cycling
from the atmosphere to the surface/sub-surface water pool
includes precipitation, which is the main form of water and
nutrient input into ombrotrophic PSFs. In contrast to dryland
forests, where microbial decomposition of dead biota is an
essential pathway of nutrients back into living biota, PSF food
webs are believed to be based primarily upon leaching of
nutrients from freshly-fallen tree leaves (i.e., part of the arrow
connecting the biota and available nutrients/water boxes),
with water playing a key part in this process (Weiss et al.
2002; Yule and Gomez 2009; Yule 2010). Consequently,
altering the volume or flow speed of surface/sub-surface water
may influence nutrient availability for living biota, and subsequent nutrient cycling into and out of the biota and other
nutrient pools, thereby influencing ecosystem function and
service provision.

1961; Amundson and Jenny 1997). These are most useful as

a tool for understanding broad natural variation among soils
and ecosystems (Groffman et al. 2004). For example, in
PSF, peat depth is a state factor known to have an important
influence on hydrology and vegetation (Page et al. 1999),
plus fauna (Morrogh-Bernard et al. 2003; Cheyne et al. 2008).
The need for variable-stochastic factors in the model arises
because ecosystems are frequently not observed in their
expected state, implying that they are being influenced by
shorter-term factors (Groffman et al. 2004). These may have
either natural (e.g., extreme climatic events) or anthropogenic
(e.g., manipulation of species) causes that may influence ecosystem processes and, ultimately, service provision. For example, removal of certain tree species by loggers in PSF
(anthropogenic variable-stochastic factor) may result in lower
densities of these species than expected based on peat depth
(state factor). Two-way relationships between stochastic and
state factors can exist, as indicated by the dashed arrow between these two boxes in Fig. 1. Humans may influence
topography, for example; and the presence of underground
coal may influence human disturbance regimes. It is, of course,
the effects of anthropogenic variable-stochastic factors (human
activities) that are of most interest to conservation managers.

ECOSYSTEM FUNCTIONS
AND SERVICES

ECOSYSTEM
ATMOSPHERE

STATE FACTORS
Time
Topography
Parent material (peat;
water table)
Climate
Biota*

Flora
Fauna
Microbes
Dead organisms

BIOTA

GROUND
WATER +
SURFACE
FLOW

SURFACE / SUBSURFACEWATER

AVAILABLE
NUTRIENTS
PEAT + MINERAL SOIL
Water

Peat

INTRA-SYSTEM CYCLING

Wood production (including for local

needs; e.g., housing)
Water quality
Water quantity (slowing wet-season
floodwaters and maintaining dry-season
base flows)
Air quality (gas exchange)
Storage of elements (e.g., C, N, metals)
Maintenance of biodiversity, including
fish and NTFPs (e.g., food and medicine)
Fire prevention
Maintenance of local climate (e.g., rain)
Crop pollination and seed dispersal
Pest control
Genetic resources
Cultural, spiritual and aesthetic values

STOCHASTIC FACTORS
NATURAL
Extreme climatic events
Eruptions of native biota (e.g., population booms, pathogen outbreaks)
Lightning fires
Natural colonisation by new species
Geological events

ECOSYSTEM
MANAGEMENT

ANTHROPOGENIC
Human-accelerated climate change
Manipulation of species (e.g., logging, hunting, invasive alien species)
Alteration of nutrient cycles (e.g., timber removal, agriculture)
Ecosystem restoration
Human-induced disturbance to hydrological cycles (e.g., drainage)
Fire subsequent to lowered water tables
Pollution and addition of novel chemicals (e.g., fertilisers, lime)

Fig. 1 A conceptual model for ecosystem processes and function in

tropical peat-swamp forest. NTFP non-timber forest product. The
arrow illustrating cycling of nutrients from the biota to the peat +
mineral soil nutrient pools is one-way, because the return cycle requires
cycling into available nutrients prior to uptake by biota. Asterisk This

set of organisms (the comprehensive set of organisms that could occur

in the ecosystem) is distinguished from those that are actually observed
in the ecosystem, which are state variables within the ecosystem box,
not state factors (controllers, Groffman et al. 2004)

Wetlands (2013) 33:257267

261

Impacts of Human Activities

Two of the arrows in Fig. 1 are particularly important for
understanding the impacts of human activities on PSFs: that
joining variable-stochastic factors to ecosystem processes
(circled in Fig. 1), and that joining ecosystem processes to
its functions and services (with the cross in Fig. 1 indicating
potential disruption to these). In light of their greater relevance for conservation management, I concentrate here on
the impacts of (selected) anthropogenic variable-stochastic
events.
Building upon an earlier framework devised by Gardner
et al. (2009), Fig. 2 expands upon the circled arrow in Fig. 1
to illustrate how natural and anthropogenic events might
influence PSF biodiversity and, through this, ecosystem
processes, function and services. The natural drivers of
change and anthropogenic drivers of change boxes here
are equivalent to the drivers in Ogden et al.s (2005b)
model for Floridian wetlands, with their stressors equating
to my landscape modification box, their ecological
effects equating to my proximate mechanisms box, and
their attributes equating to my three shaded boxes. The
dashed line between Natural drivers of change and
Invasion of new species is in recognition of the belief

Fig. 2 Potential mechanisms

through which natural and
anthropogenic stochastic events
might influence tropical peatswamp forest biodiversity.
Mobile link species are those
that connect habitats through
space and time through their
functional processes (Gilbert
1980). Modified from Gardner
et al. (2009)

that pristine PSF is highly resistant to invasion by new

species, owing to its specialised conditions (Yule 2010).
Landscape modification is therefore likely to be a more
important influence in this regard.
The depicted chain of events is equally as applicable to
both flora and fauna, but proximate events are not always
required for influences on flora to be felt; e.g., logging leads
to direct population impacts on tree species. Changes in
species interactions and populations, plus community structure can be considered as representing changes to the biota
nutrient pool in Fig. 1, with either subsequent direct (e.g.,
maintenance of biodiversity) and/or indirect (e.g., element
storage) impacts on ecosystem services. Similar models
could be developed to illustrate impacts on other nutrient
pools. Where useful, more complex models depicting multiple steps within the boxes in Figs. 1 and 2 (cf. Odum 1983;
Ogden et al. 2005a) can also be constructed from this
foundation.
The connections between species (where the flow chart in
Fig. 2 ends), the resultant processes within an ecosystem,
and its functions and resultant services, are complex
(Cardinale et al. 2000; Daz and Cabido 2001; Lerdau and
Slobodkin 2002; Luck et al. 2003; Kremen 2005; Daz et al.
2007; Luck et al. 2009) and may involve activities of

Natural drivers of
change

Anthropogenic drivers
of change

Altered disturbance
regime

Landscape modification
Invasion of
new species

Changes in hydrology
Peat loss and degradation
Forest loss and degradation
Non-forest/mosaic land-use practice
Landscape mosaic design
Forest regeneration

Proximate mechanisms
Changes to food resources
Changes to breeding resources
Dispersal constraints
Physiological tolerance limits
Changes in species behaviour

Altered species
interactions
Trophic cascades,
mobile link species

Population change

Community change

Births, deaths,
immigration, emigration

Composition, structure,
function

Overharvesting

262

organisms at several trophic levels (Bardgett and Wardle

2003; Kremen et al. 2007). It can therefore be difficult to
pinpoint the relative importance of each species for the
provision of these services (Ehrlich and Mooney 1983;
Kremen et al. 2002). For example, PSF carbon sequestration
and storage is dependent upon (i) its uptake from the atmosphere by flora (which varies both between different species
and with respect to flooding/drainage, plus numerous other
factors) and, subsequently, fauna; (ii) cycling from living to
dead biota; and (iii) subsequent deposition and storage in
peat, rather than cycling back into other nutrient pools. This
process is dependent upon a large number of species and
variables, which interact to sequester and store carbon in
intact PSF, but evidently do not in degraded/drained PSF.
Considerations of this nature often lead to desire to conserve
ecological integrity to ensure continued service provision
(Karr and Dudley 1981; Angermeier and Karr 1994;
Grumbine 1994).
Despite these difficulties, insight into the role of different
organisms in ecosystem service provision and, by inference,
the likely impacts of changes in their populations can be
gained from (i) allocating relevant ecosystem functions (referred to as ecosystem properties by de Bello et al. 2010)
to each service and identifying which organisms or groups
of organisms control these; (ii) identifying the key characteristics and mechanisms through which these organisms influence ecosystem functions and, hence, services; and (iii)
using this basis to determine how changes in these organisms populations might influence ecosystem service provision (Luck et al. 2009; de Bello et al. 2010). Following this
process will aid in understanding the links between the
Ecosystem, and Ecosystem Functions and Services
boxes in Fig. 1 and, thus, in managing PSFs and other
wetlands to maintain these services.
The different nutrient pools in Fig. 1 are inter-connected.
Thus, it may be expected that any variable-stochastic event
that impacts one nutrient pool will impact all ecosystem
processes and, thus, influence all ecosystem functions and
services. While this will be true to an extent, certain stochastic events will clearly have greater impacts on some
ecosystem processes than on others, with subsequently different impacts on overall ecosystem function and service
provision (see below for practical illustrations relevant for
PSF conservation management). For conservation monitoring purposes, it is therefore necessary to identify measurable
variables for each potential threat faced relating to (i) the
landscape changes and proximate mechanisms likely to
result from the threat; (ii) which organisms or groups of
organisms are most likely to be influenced by these landscape changes and proximate mechanisms; (iii) the ecosystem processes and functions upon which the affected
organisms or groups of organisms exert influence; and (iv)
which ecosystem services are most likely to be influenced

Wetlands (2013) 33:257267

by changes in the expression of these service-providing

organisms (or service providers, Luck et al. 2009). Two
examples of such a process, based largely on observations
from the PSF of Sabangau District, Central Kalimantan
(Page et al. 1999; Morrogh-Bernard et al. 2003) are provided in Fig. 3. The forests in Sabangau are being managed in a
very similar manner to that planned for REDD+ forests in the
region and, hence, provide a particularly useful case study.
Note that the chains of cause and effect depicted in Fig. 3
represent examples and are not the only plausible scenarios
resulting from either the depicted threats, landscape change,
proximate mechanisms, population changes or influences on
the system. Links from the two illustrated population
changes to decreased biodiversity maintenance are intended
to illustrate this fact. Relevant references on the effects of
drainage include Page et al. (2002, 2009a, b) and Wsten et
al. (2006, 2008); and for flying-fox (Pteropus vampyrus)
hunting include Struebig et al. (2007), Harrison et al. (2011)
and McConkey and Drake (2006).
The two examples highlighted in Fig. 3 provide a habitatspecific illustration of how threats may differ in their complexity and subsequent influences on ecosystem function
and service provision. Even considering the probable
changes in tree species composition arising from prolonged

Flying fox
hunting

Peat drainage

Threat

Forest/peat loss
& degradation
(C storage)

Landscape
change

Decreased trees/
food resources
(C sequestration)

Proximate
mechanism

Decreased flora &

fauna populations

Decreased flying
fox population
(pollination; seed
dispersal)

Decreased biota
nutrient pool
(C storage +
sequestration)

Decreased carbon
sequestration and
inc. CO2 emissions

Population
change

Influence on
system

Decreased
biodiversity
maintenance

Decreased
pollination and
seed dispersal

Impact on
ecosystem
service

Fig. 3 Chains of cause and effect providing examples of specific

impacts of anthropogenic threats on populations and resultant impacts
on tropical peat-swamp forest ecosystem services. Dashed lines indicate leaps in the chain, where effects at each step are not needed for
an impact on ecosystem services to be felt. Ecosystem functions
exhibited by affected organisms or groups of organisms are enclosed
in parentheses

Wetlands (2013) 33:257267

decreased pollination and seed dispersal by flying foxes

(Pterpous vampyrus) and subsequent knock-on effects of
this, peat drainage would still be expected to impact a
greater number of non-illustrated ecosystem processes and
services than would flying-fox hunting. Further, the example of peat drainage illustrates not only (selected) impacts of
changes in surface/sub-surface water on the ecosystem, but
also how a state factor (peat) can be influenced by an
anthropogenic variable-stochastic factor (peat drainage) that
is mediated by a natural stochastic factor (drought), and how
this can lead to changes within an ecosystem nutrient pool
(biota), which influences another nutrient pool (peat and
mineral soil), producing an impact on a desired ecosystem
function/service (carbon storage and sequestration). In this
way, this model provides a framework for understanding
and evaluating the causes and consequences of changes in
the ecosystem, in addition to providing a framework for
research hypothesis generation.
Finally, anthropogenic and natural stochastic factors may
have very similar, or even synergistic, impacts on the ecosystem. Drought is one natural variable-stochastic factor that can
lead to severe short-term negative impacts on PSF through
increasing fire vulnerability: charcoal recovered from peat
cores dating back to 830,000 years B.P. indicate that fire is
not exclusively associated with modern human activities
(Anshari et al. 2001; Hope et al. 2005). Despite this history,
plus the fact that fire has been viewed as beneficial in some
other wetland ecosystems (e.g., cypress domes, Ewel and
Mitsch 1978), fires resulting from the interaction between
extreme climatic events (i.e., El Nio-associated drought)
and human-induced peat drainage is having serious negative
impacts on contemporary PSFs, at least in Southeast Asia.
Peat decomposition and forest fires caused by low peat water
levels leads to peat/forest loss and degradation, severely
impacting ecosystem functioning and multiple ecosystem
services (Wsten et al. 1997; Morrogh-Bernard et al. 2003;
Wsten et al. 2006; Chokkalingam et al. 2007; Wsten et al.
2008; Harrison et al. 2009; Hirano et al. 2009; Page et al.
2009b; Moore et al. 2010). The severity of these impacts
increases along the disturbance (and drainage) gradient from
primary forest, to degraded, heavily degraded and non-forest
(Page et al. 2009a). Similarly, natural colonisation by new
species, in particular fauna, would also surely have happened
from time to time, despite the natural resistance to this
exhibited by intact PSF (Yule 2010). Thus, the impacts of
human activities are probably most likely to exacerbate, accelerate or mimic the impacts of natural stochastic events.

Applications
This conceptual model of PSF ecosystem function is important for three main reasons. First, as resource investment in

263

PSF protection increases, so does the need to establish effective PSF ecological monitoring programmes. Considering the
value placed on good conceptual models as a tool for guiding
the development of these programmes (Ogden et al. 2005b;
Lindenmayer and Likens 2009; Gardner 2010), this model
therefore represents an important step towards satisfying this
need. Further, although my discussion is centred on biodiversity, the underlying rationale is equally relevant for monitoring
the effectiveness of management interventions towards
achieving emission reduction and other goals, as illustrated
in Fig. 3 and discussed above. This model therefore has
potential relevance for all PSF conservation projects, including REDD+ and biodiversity banking, which are subject to
rigorous monitoring requirements (CCBA 2008; VCS 2008;
Yaap et al. 2010). In particular, conceptual models of ecosystem function will be useful for understanding which of the
potentially large suite of emission reduction interventions
conducted simultaneously by a REDD+ project are actually
leading to emission reductions and positive co-benefits.
Second, habitat-specific conceptual models also aid in the
development of new lines of enquiry into the impacts of
anthropogenic and natural variable-stochastic events. One
such question might be: what is the impact of humaninduced hydrological changes on available nutrient supply
and, hence, on primary productivity, fauna and ecosystem
services? When drained, aerobic conditions lead to peat oxidisation, resulting in decomposition of peat and the leaf litter
that would contribute towards future peat formation under
natural wet conditions, thereby altering the entire ecosystem
(Wsten et al. 1997; Couwenberg et al. 2010). Construction of
drainage canals in PSF leads to increased water output rates
from the system (Wsten et al. 2006, 2008). Coupled with
increases in peat and leaf litter decomposition, this would be
expected to lead to increased dissolved carbon and other
nutrient discharge rates (McDowell and Likens 1988), resulting in loss of these nutrients from the ecosystem as the water
flows out to sea and contributing towards global sea level rises
(Sahagian et al. 1994; Alley et al. 2002). Any such effects
would be above and beyond the already-quantified effects of
peat drainage on fire susceptibility and peat degradation
(Furukawa et al. 2005; Jauhiainen et al. 2005; Wsten et al.
2006, 2008; Miettinen and Liew 2010).
Depending on the severity and time frame of drainage,
these changes may lead to increased or decreased short- or
long-term nutrient availability and habitat productivity. For
example, in the Sabangau Forest, the existence of the apparently unique tall-interior forest on the thickest part of
the peat dome is thought to be due to decomposition of the
uppermost peat layer in this driest part of the forest over the
past 2,500+ years, resulting in a more productive forest than
found elsewhere in Sabangau (Page et al. 1999). Increases in
diversity and abundance of forest fauna are thought to have
followed; for example, orang-utan (Pongo pygmaeus

264

wurmbii) population density is higher in the tall-interior

forest than in any other habitat-sub type in Sabangau, due
to higher fruit availability in this habitat sub-type (MorroghBernard et al. 2003). Presumably, over time and in the
absence of human disturbance, the tall-interior forest peat
would eventually decompose down to the new lower water
level, whereupon the forest would revert to a lowerproductivity forest; or eventually change into a non-swamp
habitat, if drainage continued to the point where peat formation was no longer possible. Subsequent impacts on
biodiversity and ecosystem services would likely be approximately neutral overall, as communities have time to adapt
to changes in the forest.
Anthropogenic drainage is unlikely to have such neutral impacts, however, owing to the pace of change and
because drainage is generally also accompanied by other
threats, such as logging and conversion to plantations.
Some short-term increases in forest productivity may be
expected as decomposition rates and the available nutrient pool increase. Any such benefits are likely to be
short lived, however, as increased nutrients are lost to
the atmosphere through fire and oxidation, and to external water bodies through increased nutrient dissolvability and discharge rates, reducing total nutrient
stocks; and fire and peat structural changes create conditions unsuitable for peat-swamp forest tree survival
and reproduction. This model provides a framework
for thinking about how these different scenarios might
play out, and how natural and anthropogenic variablestochastic events might interact to produce varying
impacts on ecosystem processes, function and services.
Third, through stimulating thinking about the broader
impacts of specific human activities on PSFs, this conceptual model can also aid in identifying new conservation
problems and solutions to known conservation problems.
For example, intensive flying fox hunting and subsequent
population declines will result in reduced pollination and
seed dispersal services by these animals (McConkey and
Drake 2006; Harrison et al. 2011; Fig. 3). Seed dispersal is a
known barrier to PSF regeneration (Page et al. 2009a) and
flying foxes are considered particularly important for
breaching this barrier, owing to their ability to fly to and
over regenerating areas (Corlett 2009). They are also known
to pollinate and/or disperse seeds of tree species frequently
used in PSF reforestation/regeneration activities, such as
Palaquium spp., Madhuca mottleyana and Calophyllum
spp. (Struebig et al. 2007; Harrison et al. 2011; see Graham
2009 for a full list of species used in PSF restoration projects).
Thus, flying fox hunting could be problematic for PSF projects conducting reforestation and/or forest regeneration activities, and controlling flying fox hunting to maintain healthy
populations could even help increase carbon sequestration for
REDD+ projects.

Wetlands (2013) 33:257267

In conclusion, I highlight in this paper the importance of

creating a habitat-specific conceptual model of ecosystem
function for PSF and selected potential applications of this.
This allows for consideration of habitat-specific issues that
may have particularly important influences on PSF, such as
the impacts of fire, which differ markedly in PSF compared
to some other wetlands, e.g., cypress swamps (Ewel and
Mitsch 1978). Such considerations also indicate an anticipated need for development of habitat-specific conceptual
models to ensure effective management in other wetland
types. At the very least, the applicability of any models
established for other areas/habitat types must be carefully
assessed before application in any particular wetland site.
Combining previous wetland models with those developed
for dryland forests, while incorporating habitat-specific considerations to link drivers of ecosystem patterns and processes with its function and services, offers particularly high
potential in forested wetland habitats.
Acknowledgments I am grateful to CIMTROP and RISTEK for
long-term support of my research in Indonesia and permissions to
work in the country. I thank all colleagues who have engaged in
discussion and debate with me over the years on PSF ecology and
conservation issues, in particular Simon Husson, Helen MorroghBernard, Susan Cheyne, Laura DArcy, Suwido Limin, Laura Graham,
Jack Rieley and Susan Page. This manuscript also benefited from discussions and information from members of the Ecosystem Modelling
breakout group at the 2011 CIFOR Workshop on Tropical Wetland
Ecosystems of Indonesia: Science Needs to Address Climate Change
Adaptation and Mitigation in Sanur, Bali, and comments from the editor
and four anonymous reviewers. The writing of this paper was supported
financially by the Orangutan Tropical Peatland Project (OuTrop) and was
conducted as part of the OuTrop multi-disciplinary research project,
conducted in collaboration with CIMTROP, University of Palangka Raya.

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