BioMERGE
ASWIII
Malaysian Borneo, 2005
ASWIII. BIODIVERSITY AND ECOSYSTEM FUNCTIONING:
MULTI-TROPHIC AND CROSS-BIOME SYNTHESES
WELCOME! To previous attendees of BioMERGE workshops and to new participants, welcome to Adaptive
Synthesis Workshop III. Our time here is short and the scientific and environmental goals of the workshop are
enormous. Our purpose is to structure 5 major activities of BioMERGE for the next year, all leading up to the final
workshop to be held in 2006. Additionally, many of us will be working with DIVERSITAS where our interests
overlap.
All of you are here because we are interested in understanding the ecological significance of biodiversity. Some of
us are motivated by the implications and applications that arise from biodiversity and ecosystem functioning (BEF )
research because of the dramatic declines we are witnessing in biodiversity all across the world. Others are
motivated by the intellectual challenge of understanding how nature functions when it is made up of so many
individuals from so many species all enmeshed in a complex, dynamic web that varies enormously over space and
time. Either way, the first generation of BEF research opened up a new terrain to be explored and we are here to
foster a second generation of BEF research – one that is much larger in scope and more true to the complexities of
the real world.
Can we accomplish these goals in so short a time? You bet! In reality, as a group, we have all contributed to the
transformation of the field of biodiversity and ecosystem functioning (BEF) research from a field dominated by
small-scale, experimental and theoretical research (and a lot of arguments), into a larger scale science. In this short
workshop, our goals are to consider the future of biodiversity (i.e., extinction scenarios), the significance of trophic
levels, how to link ecosystems, what the applications and implications of this 2 nd generation BEF research are, and
how to put together a volume on the topic.
This document provides an overview of BioMERGE and the 5 major goals for this workshop. The workshop is
meant to be a hands-on exercise in which explicit activities initiated here are carried through to completion as a
network over this and the following year. Note that all related documents and references will be posted on our
website at http://www.columbia.edu/cu/biomerge/workshop3.html.
SCHEDULE OVERVIEW
DAY
TIME
Thursday (1 Sept) 6pm
Friday (2 Sept)
8:00 am
9:00 – 10:00
10:00-10:30
10:30-12:00
12:00-1:30
1:30-3:00
3:00-3:30
3:30-5:30
Saturday (3 Sept)
8:00 am
10:00-10:30
12:00-1:30
3:00-3:30
Sunday (4 Sept)
8:00 am
10:00-10:30
12:00-1:30
3:00-3:30
3:30-5:30
TBA
TASK
Dinner
Introduction, Logistics
Update presentations
Break
Workshop Assignments
Lunch
Working groups
Break
Working groups
Plenary
Break
Lunch
Break
Implications and
Applications
Break
Lunch
Break
Final report, planning
for next meeting
Dinner w/Diversitas
1
DESCRIPTION
Naeem, Bunker
Current status of working groups.
Outline of workshop goals.
Assignments of ASWIII workshop groups.
(1) Cross-System (Naeem)
(2) Multi-trophic (Cardinale)
(3) Extinction Scenarios (Bunker)
Workshop representatives
short discussion of progress
problems arising
(4) Whole-Group Brainstorming (Symstad)
Assignment of Policy/Management Group
(5) Outline of BioMERGE book.
Next meeting
BioMERGE
ASWIII
Malaysian Borneo, 2005
Overview of BioMERGE. The NSF Research Coordinating Network (RCN) entitled BioMERGE (Biotic
Mechanisms of Ecosystem Regulation in the Global Environment) was established in 2002 to usher Biodiversity and
Ecosystem Functioning (BEF) research into a new era – or what we might consider as 2nd generation BEF. BEF is a
body of ecological research that explicitly examines the consequences of changes in biodiversity on ecosystem
functioning. Prior to 2002, BEF consisted almost entirely of small-scale experimental and theoretical work whose
findings did not always agree with observational studies, a disagreement that hindered progress in BEF research.
Given that changes in biodiversity are among the most prevalent features of a human-dominated Earth, and given
that stemming biodiversity loss, conserving what species are present, and restoring ecosystems that have
experienced enormous changes in biodiversity due to habitat fragmentation, habitat destruction, invasion, pollution,
or over exploitation, are among the most intractable of contemporary environmental issues, understanding the
environmental consequences of biodiversity loss represents one of the most important areas of ecological research.
The disagreement between observation and both theory and experiment needed to be addressed if BEF research was
to advance.
BioMERGE has many functions, but its primary task is to incorporate the findings of predominately small-scale
experimental BEF research (and theory associated with these experiments) into larger scale studies. The goal is to
determine large-scale ecosystem consequences of biodiversity loss without having to resort to large-scale
experiments. BEF experiments manipulate biodiversity by a variety of methods, but even for small numbers of
species (e.g., > 10), experiments get quickly out of hand as the number of combinations of species becomes
formidable. It is generally agreed that comprehensive, large scale BEF experiments are intractable. Thus, either
BEF figures out how to relate its results to the larger scales of the real world, or its utility remains constrained to
theory and small scale phenomena.
ADAPTIVE SYNTHESIS WORKSHOPS BioMERGE accomplishes its task by a series of 4 workshops over 5 years.
These workshops focus on synthesizing studies of biodiversity independent of function (e.g., taxonomy,
biogeography, macroecology) and studies of the functional significance of biodiversity (e.g., BEF, and to a certain
extent, community and ecosystem ecology). They are referred to as Adaptive Synthesis Workshops (ASW) because
they do not have a fixed agenda, but rather the agenda adapts to the changing face of BEF research.
ASWI met in Seattle, WA, USA, in 2002 where it established a conceptual framework (Naeem and Wright 2003).
ASWII met at the Missouri Botanical Garden in St. Louis, MO, USA to put the BioMERGE framework to test. The
workshop explored the theoretical implications of the framework and applied the framework to data from
grasslands, a marine system, and a tropical system. Results from the theory group, more a general consideration of
the many issues than any formal models, are to appear in a symposium volume (Naeem et al. in press), the marine
group’s findings were published in Science (Solan et al. 2004), and those of the tropical group have been submitted
to Science. The grassland group’s work will be presented here at ASWIII in Malaysia. The goals of ASWIII are
discussed below.
ASWIV, Switzerland, 2006, is the final workshop in which BEF is brought into the realm of large scale
environmental research. This workshop will launch generation 2 of BEF – a new frontier in which biodiversity is
integrated into mainstream research rather than being an independent, primarily academic field that does not impact
either environmental policy, conservation biology, global change biology, or other major areas of ecological and
environmental research. Because we have the methods in hand and three worked examples, by the time of the
workshop there will be further developments. The volume is not meant to be the typical symposium volume, but
more of a manual and guide to understanding why biodiversity matters in ways we seldom realize and how best to
go about managing biodiversity to the betterment of human well-being.
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BioMERGE
ASWIII
Malaysian Borneo, 2005
ASWIII, KOTA KINABALU, MALAYSIAN BORNEO, 2005. With a partially proven conceptual framework in hand,
BioMERGE moves into its final and major stage. This stage consists of coordinating efforts to take current BEF
research from its predominantly single-system/single-trophic-level status to a multi-system/multi-trophic discipline.
The setting of Borneo was chosen to focus our attention on tropical biodiversity, the biome holding the vast majority
of known biodiversity. Up till now, BEF has focused on grass or meadow lands, microbial microcosms, and a
handful of other systems. Nowhere else are the challenges of BEF more clear than in tropical systems: many species
with little more than their taxonomy known, experiments are not practical, logistics may be difficult. Borneo is
particularly appropriate as it represents Asian tropical diversity, which is substantially understudied in comparison to
the New World tropics, and it represents a region in which environmental challenges (Costa and Magnusson 2002,
Page et al. 2002, Aldhous 2004, Curran et al. 2004, Van Nieuwstadt and Sheil 2005) do not deter commitment to
sustainable development (http://www.undp.org.my/cms_undp/events/e119.asp). BioMERGE is a natural science
RCN, not a social science network, but its mission is clearly motivated by providing the scientific foundation for
understanding how biodiversity conservation can enhance ecosystem function and thereby improve ecosystem
services – the fundamental means by which human well-being is insured.
There are 5 major goals for ASWIII:
I. Linking ecosystems: Terrestrial and Aquatic
II. Including trophic levels: Introduce herbivory, microbial, and higher trophic levels
III. Developing extinction (biodiversity) scenarios
IV. Implications and Applications: Developing concrete, real-world implications and applications.
V. Draft proposal for BioMERGE/DIVERSITAS volume to be finalized at ASWIV, Switzerland, 2006.
Each of these are described in detail below. The goal is described with some possible products as models, an
overview provided, and some preliminary thoughts are offered to get the discussion going.
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BioMERGE
ASWIII
Malaysian Borneo, 2005
I. LINKING ECOSYSTEMS: TERRESTRIAL AND AQUATIC
GOAL. Initiate a study that builds links between aquatic, grassland, and forest groups.
OVERVIEW. No ecosystem is an island unto itself. All solar-based ecosystems are linked to one another by
evolutionary history, geography, and biogeochemistry. Field-based BEF, however, has focused almost exclusively
on grassland or non-woody ecosystems. Because change in biodiversity is an ubiquitous feature of modern-day
Earth, it stands to reason that if the loss of diversity impacts the functioning of the ecosystem within which it
resides, then neighboring ecosystems are likely to be affected as well.
There are many papers on the topic of cross-ecosystem linkages, especially aquatic-terrestrial linkages, perhaps the
Hubbard Brook experiments starting in the 1960s being the most well known. More recent studies include Beman et
al. (2005) who argued that agricultural runoff can fuel harmful algal blooms in sensitive marine areas. A standard
example is that of the marine-terrestrial link created by anadromous fish, such as salmon, that bring nutrients from
the sea up stream and eventually into the surrounding forests (Willson et al. 1998). In freshwater systems, Pace et
al. (2004) argued that internal production of organic carbon may be insufficient to support resident foodwebs and
that terrestrial ecosystems may supply important sources of carbon. In an old growth stream in Oregon, Ashkenas et
al. (2004) showed that terrestrial and aquatic organisms were tightly linked by their common use of added N.
Finally, the well known NYC watershed example, where managing the watershed appears to be cheaper than
building water treatment facilities (Chichilisky and Heal 1998), also speaks to the linkages between terrestrial and
aquatic systems.
PRELIMINARIES.
1. What are the key linkages? These linkages can be classified as either:
Additive – for example, the loss of biodiversity in several ecosystems leads to a loss of carbon storage, meaning that
the combined effect is a global increase in atmospheric CO2 without any complex interactions between the systems.
That is, global Cstored = sum of all individual ecosystem C storages.
Interactive – for example, loss of plant diversity of a watershed alters carbon and nutrient inputs into an estuary
which, in turn, leads to biodiversity loss and altered estuarine function. Note that the trick here is to disentangle
changes in function due to changes in inputs versus changes due to losses in biodiversity.
Multiplicative – increasing biodiversity across all trophic levels increases rates of nutrient cycling and lowers system
variability.
2. Develop a worked example from the marine/terrestrial studies.
Consider the following diagram. If we already can state how
ecosystem function is affected by change in biodiversity, then we
should be able to tackle such a system.
a. What would the system-level ecosystem function be?
b. What are the scenarios and how does biodiversity respond for each
of these? For example, elevated N deposition – plant biodiversity loss,
biomass increase, infaunal decline due to pollution?
4
Grassland
Forest
Estuary
BioMERGE
ASWIII
Malaysian Borneo, 2005
II. INCLUDING TROPHIC LEVELS: INTRODUCE HERBIVORY, MICROBIAL PROCESSES, AND
HIGHER TROPHIC LEVELS
GOAL. A BioScience like paper that describes how biodiversity loss across trophic levels compares to biodiversity
loss within trophic levels.
OVERVIEW. Although BEF has been dominated by studies of autotrophs, there have been a number of studies that
have examined heterotrophic groups, multiple trophic levels (e.g., Mulder et al. 1999, e.g., Sankaran and
McNaughton 1999, Symstad et al. 2000), though largely in microcosms (e.g., Naeem et al. 1995, McGrady-Steed et
al. 1997, Naeem and Li 1997, Mikola and Setälä 1998, McGrady-Steed and Morin 2000, Fukami and Morin 2003,
Fox 2004, Heemsbergen et al. 2004, Wohl et al. 2004, e.g., Bezemer et al. 2005) or mesocosms (e.g., Duffy et al.
2001, Emmerson et al. 2001, Engelhardt and Ritchie 2001, e.g., Downing and Liebold 2002, Duffy et al. 2005,
Schlapfer et al. 2005). There is too much literature to list, but even though the literature is extensive, ranging from
marine, to freshwater, to terrestrial systems, how to apply these findings to large-scale, non experimental studies
remains unclear. Can we develop a plan of attack and a worked example?
PRELIMINARIES.
1. Trans-trophic Biodiversity Effects. If changes in biodiversity alter ecosystem functioning within a trophic
level, then it stands to reason that other trophic levels may respond to this effect. Consider the following. Augustine
et al. (2003) showed that ANPP was linearly correlated with consumption rates of ungulates. If loss of plant
biodiversity were to lead to a reduction in ANPP, would this mean a linear decline in consumption? In the review
by Bardgett et al. (2003), the authors suggest that herbivory can have positive (in nutrient rich systems), neutral, and
negative (in nutrient poor systems) impacts on soil biology and biochemistry. How would this affect rangelands or
habitats like the Serengeti where top predators are declining or the vast steppes of Inner Mongolia now experiencing
enormous increases in grazing? In another example, Duffy et al. (2005) demonstrated that the diversity of grazers
had trans-trophic impacts in an eel-grass community. Given the remarkable prevalence of intra-guild predation
(Arim and Marquet 2004), how does this relate to BEF? Schmitz (Schmitz 2003) demonstrated that spiders could
affect herbivore impacts on the distribution and abundance of plants which, in turn, would alter ecosystem function.
Bezemer et al. (2005) showed that manipulation of belowground fauna (nematodes and microbes) could have affects
all the way up the trophic ladder to parasitoid-aphid interactions. Finally, Tiunov and Scheu (2005) showed that
microbial diversity can affect decomposition, so do our deliberations about macrobes carry through to microbes? A
couple of important papers are Ives et al. (2005) and Statzner and Moss (2004).
2. BEF Trophic Traits. What are the relevant response and effect traits for heterotrophic organisms?
What is the role of phylogeny? Cattin et al. (2004) argue that food web structure only makes sense when one
includes phylogenetic constraints. There is a tendency to simply use traits indiscriminately, but if one is a
hummingbird, one does not eat seeds. If one is a grasshopper, one does not suck phloem. Is there a reasonable way
to bring this into our deliberations? Is phylogeny a surrogate for the degree of specialization and redundancy?
Are there scale-invariant food web properties that are relevant? Donald and Stewart (2003) argue that
predator:prey ratios are predictable – can these be used when a food web is not completely known? More generally
(perhaps pt 1?), how much can we rely on sampling the food web to characterize the system, rather than a complete
census? Are there food web properties that are predictable from environmental data?
Will the degree of dietary specialization be an important predictor of BEF? In a system with generalist consumers,
such as ungulate grazers, loss of consumers may have little effect on primary functioning, as remaining grazers may
compensate fully. In contrast, systems characterized by specialist consumers, such as tropical forests, may be more
sensitive to consumer losses if consumer extinctions confer a competitive advantage on their surviving hosts. Can
we quantify the degree of specialization of communities?
Is the BioMERGE system a logical division for constructing a table of relevant traits, or does this need to be
revisited? Currently, the BioMERGE system in the conceptual framework is as follows:
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BioMERGE
a.
b.
c.
d.
ASWIII
Malaysian Borneo, 2005
Autotrophs. Organisms that rely heavily on
inorganic carbon and nutrients. Relevant traits are
well developed among plant biologists. For
example, mass-specific photosynthetic traits,
symbiotic association with N-fixers, C3 or C4
pathways.
Decomposers. Decomposers are defined here as
microbes (fungi and prokaryotes) that mineralize
dead organic matter. In contrast, soil arthropods
and invertebrates, detritivores, and so forth, are
heterotrophs that derive their resources from the
DOM chain. Would the relevant traits be
exclusively exoenzymes (e.g., cellulose or lignin
degraders, protein degraders, polycyclic aromatic
hydrocarbon degraders, etc.)? Would anaerobic,
aerobic life styles be important? Would taxonomy
be important (e.g., bacteria, archaea, fungi)?
Decomposer-derived heterotrophs. These are
generally small organisms, protists and
invertebrates, that feed on decomposer microbes,
either directly (e.g., protists) or indirectly (e.g.,
detritivores) in the sense that their carbon and
nutrients are acquired through the decomposer
chain. Would the relevant traits be body size,
metabolic rates, etc?
Autotroph-derived heterotrophs. These are the more commonly studied macrobes usually referred to
simply as “animals” consisting of herbivores, omnivores, carnivores, and so forth. What traits would be
important? Body size, metabolic rates, reproductive rates, symbiotic associations (e.g., termites,
ungulates). If they are parasites or facilitators (e.g., pollinators), should they be classified differently? For
example, pollinators are, effectively, herbivores, but their loss is quite different from the loss of leaf-eating
herbivores.
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BioMERGE
ASWIII
Malaysian Borneo, 2005
III. BIODIVERSITY (EXTINCTION) SCENARIOS
GOAL. Develop, with worked examples, a foundation for
the construction and implementation of extinction
scenarios, which we propose are more aptly named
biodiversity scenarios. The model should be a review
paper for Ecology Letters.
OVERVIEW. A critical part of non-experimental BEF is
the use of extinction scenarios to describe what we will
call biodiversity futures. In reality, the purpose of
extinction scenarios is to generate plausible states of
biodiversity in the near or distant future, thus extinction,
neither global nor local, has to be part of the exercise. In
fact, biodiversity scenarios is a more appropriate term than
extinction scenarios. In many cases, relative abundance is
what changes and, in the case of invasions, new species
may be added. For example, woody encroachment in
grasslands due to climate change does not necessarily
involve the loss of grasses or forbs.
PRELIMINARIES. Most experimental and modeling
approaches employ a two-step random extinction approach to biodiversity scenarios. The first step is to use a
combinatorial approach to consider all possible combinations. The second step is to explore a subset of these
combinations as the total number of combinations is generally an intractable number, even for simulations.
An alternative approach is to explore a specific subset of possible biodiversity futures. For example, examine the
community with a top predator removed, or in the presence of an invader, or with an environmentally sensitive or
over-exploited species removed. Exploring the absence or addition of even just one species reflects a biodiversity
scenario.
These two, the stratified combinatorial random extinction approach and the single-species exclusion or addition,
represent the extreme conditions.
The goal of this working group is to review and develop biodiversity scenarios for non-experimental BEF research
in which the following are considered:
1. Classes of drivers of biodiversity change or simply, biodiversity drivers
Biotic
Abiotic
Anthropogenic
Others?
2. Classes of scenarios (pros, cons, situations for which they are relevant)
Stratified combinatorial random extinctions
Nested
↑
Others…
↓
Single species exclusions or additions
3. Statistical issues
What are the null expectations for both mean levels and variance of functioning
What issues are relevant for the use of parametric statistical methods?
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BioMERGE
ASWIII
Malaysian Borneo, 2005
IV. IMPLICATIONS AND APPLICATIONS
Concrete, real-world implications of 2nd generation BEF.
GOAL Develop concrete, real-world implications and applications of BEF for a policy piece in Science or Nature.
Use the BioMERGE examples as a basis.
OVERVIEW The majority of policy/management BEF has focused on experimental findings, their interpretations,
future research, and many other issues, but it is fair to say that the connections between biodiversity – ecosystem
function – and human well being are tentative at best. The Ecological Society of America commissioned a white
paper on BEF, but this appeared in Ecological Monographs (Hooper et al. 2005). SciDev.Net
(http://www.scidev.net/index.cfm) has a policy dossier on biodiversity in which BEF figures, but only in a small
way. The Millennium Assessment explicitly issued its Biodiversity Synthesis Report (Duraiappah and Naeem
2005), but the examples provided were few. Symstad et al. (2003) reviewed the possible implications for BEF when
juxtaposed against real-world issues (e.g., large-scale, long-term), but have we progressed? For example, (Roscher
et al. 2005) argued that the positive species-richness/productivity relationship holds up over larger scales – does this
study alter our convictions of the policy implications? Two pieces on implications and applications have questioned
the utility of BEF (Schwartz et al. 2000, Cameron 2002) in part because many species are rare – what is the
relevance of such concerns?
PRELIMINARIES
1. Clarifiying the link between ecosystem function and
ecosystem services. The Millennium Assessment explicitly
links biodiversity with human well-being via its impacts on
ecosystem services in its Biodiversity Synthesis Report
(Duraiappah and Naeem 2005). Unfortunately, examples
were very limited, but it does provide the skeletal framework
for policy and management. The critical link for policy and
management is clearly the link between ecosystem function
and ecosystem services, yet this linkage is very poorly
developed.
Kremen (2005) argues that [BEF] research should:
(1) identify important 'ecosystem service providers'
(2) determine the aspects of community structure that influence function in real landscapes, especially compensatory
community responses that stabilize function, or non-random extinction sequences that rapidly erode it
(3) assess key environmental factors influencing provision of services
(4) measure the spatio-temporal scale over which providers and services operate
2. Are there appropriate BEF indices? Indices are absolutely critical to policy, like it or not. Is there an index of
BEF for policy? For example, The Convention on Biological Diversity has established 2010 targets for all nation
members to achieve in an effort to stop the loss of biodiversity. Mace argues that these targets, based on species
richness, are inappropriate (2005) in her perspective piece on Scholes and Biggs (2005) paper that argues that what
needs to be preserved is biodiversity intactness.
3. Provide a concrete example using the three case studies (grasslands, marine, and forest) in BioMERGE.
What are the specific policy/management implications of the three BioMERGE studies thus far? In the Solan et al.
study, there were clear implications for pollution, indicator species, and other environmental issues. Are there
indicator ecosystem functions as well as indicator species? Can the differences in C storage in the tropical system
be translated into global warming and forest management policy? Can the grassland studies be applied to
rangelands or hay meadows?
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BioMERGE
ASWIII
Malaysian Borneo, 2005
V. DRAFT PROPOSAL FOR BIOMERGE/DIVERSITAS VOLUME.
GOAL Develop a draft book proposal.
Overview. BioMERGE ends in its fifth year. It is explicitly a non renewable program of NSF. The grant includes
the preparation of a final volume. It would be valuable at this meeting to consider the scope of the book and to
invite authors where necessary.
PRELIMINARIES
Title. The Ecological and Environmental Consequences of Biodiversity Loss
Editors: Shahid Naeem, Andy Hector, Daniel Bunker
Chapters
1) Introduction
2) Review of BEF
i) Theoretical
ii) Empirical
iii) Challenges
3) BioMERGE
i) Framework
ii) Examples
(a) Aquatic
(b) Terrestrial
1. Grassland
2. Forest
(c) Linked
(d) Multi-trophic
4) Implications and Applications
i) Research
(a) Data needs
(b) Models
(c) Future
ii) Ecosystem services
iii) Economic
iv) Management/ Policy
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BioMERGE
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Malaysian Borneo, 2005
LIST OF PARTICIPANTS. Tentative assignments to working groups. Please let us know if you have particular
interests in alternative groups. Note that many of us will move between groups as the workshop progresses. The
participant list will be posted on the website and updated as necessary. x: core participant; *: group leaders.
Name
Daniel Bunker
Brad Cardinale
Jerome Chave
Sandra Diaz
Amy Downing
Emmett Duffy
Katia Engelhardt
Andy Hector
Claire Jouseau
Jennie McLaren
Shahid Naeem
Natalia PerezHarguindeguy
Oliver Phillips
Chris Phillipson
Mahesh Sankaran
Martin Solan
Diane Srivastava
Amy Symstad
Justin Wright
Linkages
Trophic
Extinctions
*
Implications
x
x
x
x
x
x
x
x
x
x
x
Book
*
x
x
x
x
x
x
*
x
x
*
x
x
x
nperez@com.uncor.edu
x
x
x
x
x
x
x
*
x
x
x
x
x
x
x
x
o.phillips@geography.leeds.ac.uk
chrisp@uwinst.unizh.ch
mahesh@nrel.colostate.edu
m.solan@abdn.ac.uk
srivast@zoology.ubc.ca
asymstad@usgs.gov
justin.wright@duke.edu
*
x
x
x
x
x
x
x
x
*
x
x
x
x
x
10
e-mail
deb37@columbia.edu
bjcardinale@facstaff.wisc.edu
chave@cict.fr
sdiaz@com.uncor.edu
aldownin@cc.owu.edu
jeduffy@vims.edu
kengelhardt@al.umces.edu
ahector@uwinst.unizh.ch
cfj2101@columbia.edu
jmclaren@interchange.ubc.ca
sn2121@columbia.edu
BioMERGE
ASWIII
Malaysian Borneo, 2005
LITERATURE CITED
Aldhous, P. 2004. Land remediation: Borneo is burning. Nature 432:144-147.
Arim, M., and P. A. Marquet. 2004. Intraguild predation: a widespread interaction related to species biology.
Ecology Letters 7:557-564.
Ashkenas, L. R., S. L. Johnson, S. V. Gregory, J. L. Tank, and W. M. Wollheim. 2004. A stable isotope tracer study
of nitrogen uptake and transformation in an old-growth forest stream. Ecology 85:1725-1739.
Augustine, D. J., S. J. McNaughton, and D. A. Frank. 2003. Feedbacks between soil nutrients and large herbivores
in a managed savanna ecosystem. Ecological Applications 13:1325-1337.
Bardgett, R. D., and D. A. Wardle. 2003. Herbivore-mediated linkages between aboveground and belowground
communities. Ecology 84:2258-2268.
Beman, J. M., K. R. Arrigo, and P. A. Matson. 2005. Agricultural runoff fuels large phytoplankton blooms in
vulnerable areas of the ocean. Nature 434:211-214.
Bezemer, T. M., G. B. De Deyn, T. M. Bossinga, N. M. van Dam, J. A. Harvey, and W. H. Van der Putten. 2005.
Soil community composition drives aboveground plant-herbivore-parasitoid interactions. Ecology Letters
8:652-661.
Cameron, T. 2002. 2002: the year of the diversity ecosystem function debate. Trends in Ecology & Evolution
17:495-496.
Cattin, M.-C., L.-F. Bersier, C. B. Richter, R. Baltensperger, and J.-P. Gabriel. 2004. Phylogenetic constraints and
adaptation explain food-web structure. Nature 427:835-839.
Chichilisky, G., and G. Heal. 1998. Economic returns from the biosphere. Nature 391:629-630.
Costa, F., and W. Magnusson. 2002. Selective logging effects on abundance, diversity, and composition of tropical
understory herbs. Ecological Applications 12:807-819.
Curran, L. M., S. N. Trigg, A. K. McDonald, D. Astiani, Y. M. Hardiono, P. Siregar, I. Caniago, and E. Kasischke.
2004. Lowland forest Loss in protected areas of Indonesian Borneo. Science 303:1000-1003.
Donald, D. B., and R. S. Anderson. 2003. Resistance of the prey-to-predator ratio to environmental gradients and to
biomanipulations. Ecology 84:2387-2394.
Downing, A. L., and M. A. Liebold. 2002. Ecosystem consequences of species richness and composition in pond
food webs. Nature 416:837-841.
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