2 Ecosystems and Their Services

EXECUTIVE SUMMARY
 An ecosystem is a dynamic
dy namic complex of plant, animal, and microorganism com-
munities and the nonliving environment, interacting as a functional unit. Hu-
mans are an integral part of ecosystems.

 A well-defined ecosystem has strong interactions among its components and

weak   interactions across its boundaries. A useful ecosystem boundary is the
place where a number of discontinuities coincide, for instance in the distribu-
tion of organisms, soil types, drainage basins, or depth in a water body. At a
larger scale, regional and even globally distributed ecosystems can be evalu-
ated based on a commonality of basic structural units.

 Ecosystem services are the benefits people obtain from ecosystems. These
include provisioning services such as food and water; regulating
regulating services such
as flood and disease control; cultural services such as spiritual, recreational,
and cultural benefits; and supporting services, such as nutrient cycling, that
maintain the conditions for life on Earth.

 Biodiversity is the variability among living organisms. It includes diversity within

and among species and a nd diversity within and among ecosystems. Biodiversity is
the source of many ecosystem goods, such as food and genetic resources, and
changes in biodiversity can influence the supply of ecosystem services.

 People seek many services from ecosystems and thus perceive the condition
of an ecosystem in relation to its ability to provide desired services. The
The ability
of ecosystems to deliver services can be assessed by a variety of qualitative
and quantitative methods.

 An assessment of the condition of ecosystems, the provision of services, and

their relation to human well-being requires an integrated approach. This en-
ables a decision process to determine which service or set of services is val-
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ued most highly and how to develop approaches to maintain services by
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managing the system sustainably.
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Introduction
Privacy Policy Millionsof species populate Earth. The vast majority gain energy to sup-
port their metabolism either directly from the sun, in the case of plants,
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or, in the case of animals and microbes, from other organisms through
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feeding on plants, predation, parasitism, or decomposition. In the pursuit
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of life and through their capacity to reproduce, organisms use energy, wa-
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50   Ecosystems and Human Well-being: A Framework for Assessment 

ter, and nutrients. Terrestrial plants obtain water principally from soil,
while animals get it mainly from free-standing water in the environment
or from their food. Plants obtain most of their nutrients from the soil or
water, while animals tend to derive their nutrients from other organisms.
Microorganisms are the most versatile, obtaining nutrients from soil, wa-
ter, their food, or other organisms. Organisms interact with one another
in many ways, including competitive, predatory,
predatory, parasitic, and facilitative
ways, such as pollination, seed dispersal, and the provision of habitat.
These fundamental linkages among organisms and their physical and
biological environment constitute an interacting and ever-changing sys-
tem that is known as an ecosystem. Humans are a component of these
ecosystems. Indeed, in many regions they are the dominant organism.
Whether dominant or not, however, humans depend on ecosystem prop-
erties and on the network of interactions among organisms and within
and among ecosystems for sustenance, just like all other species.
As organisms interact with each other and their physical environment,
they produce, acquire, or decompose biomass and the carbon-based or or-
ganic compounds associated with it. They also move minerals from the wa-
ter, sediment, and soil into and among organisms, and back again into the
physical environment. Terrestrial plants also transport water from the soil
into the atmosphere. In performing these functions, they provide materials
to humans in the form of food, fiber, and building materials and they con-
tribute to the regulation of soil, air, and water quality.
These relationships sound simple in general outline, but they are in
fact enormously complex, since each species has unique requirements for
life and each species interacts with both the physical and the biological
environment. Recent perturbations, driven principally by human activi-
ties, have added even greater complexity by changing, to a large degree,
the nature of those environments.

This website stores data such as

Ecosystem Boundaries and Categories
cookies to enable essential site
functionality, as well as marketing,
Although
personalization, the notion
and analytics. You of an ecosystem is ancient, ecosystems first became a
may change unityourof
settings at any
study time
less than a century ago, when Arthur Tansley provided an
or accept the default settings.
initial scientific conceptualization in 1935 (T
(Tansley
ansley 1935) and Raymond
Lindeman did the first quantitative study in an ecosystem context in the
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early 1940s (Lindeman 1942). The first textbook built on the ecosystem
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concept, written by Eugene Odum, was published in 1953 (Odum 1953).
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Thus the ecosystem concept, so central to understanding the nature of life
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on Earth, is actually a relatively new research and management approach.
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Ecosystems and Their Services 51

Tansley’s formulation of an ecosystem included “not only the organ-

ism-complex, but also the whole complex of physical factors forming what
we call the environment” (Tansley (Tansley 1935:299). He noted that ecosystems
“are of the most varied kinds and sizes.” The main identifying feature of an
ecosystem is that it is indeed a system; its location or size is important, but
secondary.
Following Tansley Tansley and subsequent developments, we chose to use the
definition of an ecosystem adopted by the Convention on Biological Di-
versity (CBD): “a dynamic complex of plant, animal and micro-organism
communities and their nonliving environment interacting as a functional
unit” (United Nations 1992:Article 2).
Biodiversity and ecosystems are closely related concepts. Biodiversity
is defined by the CBD as “the variability among living organisms from all
sources including, inter alia, alia, terrestrial, marine and other aquatic ecosys-
tems and the ecological complexes of which they are part; this includes
diversity within species, between species and of ecosystems” (United Na-
tions 1992:Article 2). Diversity thus is a structural feature of ecosystems,
and the variability among ecosystems is an element of biodiversity. The
parties to the convention have endorsed the “ecosystem approach” as their
primary framework for action. (See Box 2.1.)
For analysis and assessment, it is important to adopt a pragmatic view of 
ecosystem boundaries,
boundaries , depending on the questions being asked. In one sense,
the entire biosphere of Earth is an ecosystem since the elements interact. At
a smaller scale, the guiding principle is that a well-defined ecosystem has
strong interactions among its components and weak  weak interactions across its
boundaries. (See also Chapter 5.) A practical approach to the spatial de-
limitation of an ecosystem is to build up a series of overlays of significant
factors, mapping the location of discontinuities, such as in the distribution
of organisms, the biophysical environment (soil types, drainage basins, depth
in a water body), and spatial interactions (home ranges, migration patterns,
This website stores
fluxes data
ofsuch as
matter). A useful ecosystem boundary is the place where a number
cookies to enable essential site
ofwellthese
functionality, as relative discontinuities coincide. At a larger scale, regional and
as marketing,
personalization, and analytics.
even globally Youdistributed ecosystems can be evaluated based on the com-
may change your settings at any time
monality
or accept the default of basic structural units. We use such a framework in the MA for
settings.
the global analysis of ecosystem properties and changes.
Privacy Policy The global assessment being undertaken by the MA is based on 10
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categories: marine, coastal, inland water, forest, dryland, island, moun-
tain, polar, cultivated, and urban. (See Box 2.2.) These categories are not
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ecosystems
Analytics themselves, but each contains a number of ecosystems. The
MA reporting categories are not mutually exclusive: their boundaries can
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52   Ecosystems and Human Well-being: A Framework for Assessment 

BOX 2.1 The Ecosy

Ecosyste
stem
m Approac
Approach:
h: A Bridge
Bridge Betw
Between
een the
the Environ
Environmen
mentt and
Human Well-being

The concept of an ecosystem provides a valuable framework for analyzing and act-

ing on theapproach
ecosystem linkages has
between
been people
endorsedand
bytheir environment.
the Convention on For that reason,
Biological the
Diversity
(CBD) and the Millennium Ecosystem Assessment (MA) conceptual framework is
entirely consistent with this approach. The CBD defines the ecosystem approach as
follows:
The Ecosystem Approach is a strategy for the integrated management of 
land, water and living resources that promotes conservation and sustain-
able use in an equitable way. Thus, the application of the ecosystem
approach will help to reach a balance of the three objectives of the Con-
vention: conservation; sustainable use; and the fair and equitable sharing
of the benefits arising out of the utilization of genetic resources. An eco-
system approach is based on the application of appropriate scientific meth-
odologies focused on levels of biological organization, which encompass
the essential structure, processes, functions and interactions among or-
ganisms and their environment. It recognizes that humans, with their
cultural diversity, are an integral component of many ecosystems.

According to the CBD, the term ecosystem can refer to any functioning unit at
any scale. This approach requires adaptive management to deal with the complex
and dynamic nature of ecosystems and the absence of complete knowledge or un-
derstanding of their functioning. It does not preclude other management and con-
servation approaches, such as biosphere reserves, protected areas, and single-species
conservation programs, or other approaches carried out under existing national policy
and legislative frameworks; rather, it could integrate all these approaches and other
methodologies to deal with complex situations. As described in the CBD, there is

no
cial,single way to
national, implement
regional, and the ecosystem
global approach, as it depends on local, provin-
conditions.
The conceptual framework of the MA provides a useful assessment structure that
can contribute to the implementation
i mplementation of the CBD’s ecosystem approach. By way of 
analogy,
This website stores decision-makers
data such as would not make a decision about financial policy in a
cookies to enable essential
country site examining the condition of the economic system, since informa-
without
functionality, as well as marketing,
tion on the economy of a single
personalization, and analytics. You
si ngle sector such as manufacturing would be insufficient.
insuf ficient.
may change your Thesettings
same applies to ecological systems or ecosystems. Decisions can be improved by
at any time
considering
or accept the default settings.the interactions among the parts of the system. For instance, the drain-
ing of wetlands may increase food production, but sound decisions also require in-
Privacy Policy formation on whether the potential added costs associated with the increased risk
of downstream flooding or other changes in ecosystem services might outweigh those
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Ecosystems and Their Services 53

and do overlap. Ecosystems within each category share a suite of biologi-

cal, climatic, and social factors that tend to differ across categories. More
specifically,, there generally is greater
specifically great er similarity within than between each
category in:


climatic conditions;
 geophysical conditions;
 dominant use by humans;
 surface cover (based on type of vegetative cover in terrestrial ecosystems
or on fresh water
water,, brackish water, or salt water in aquatic ecosystems);
 species composition; and
 resource management systems and institutions.
The factors characterizing ecosystems in each category are highly in-
terrelated. Thus, for example, grasslands are found in many areas where
potential evaporation exceeds precipitation. Grasslands, in turn, tend to
be used by humans either as rangeland or for agricultural purposes. The
areas used for rangeland tend to have pastoral, sometimes nomadic, re-
source management systems. Thus these factors—high potential evapora-
tion relative to precipitation, grassland cover
cover,, use for livestock, and pasto-
ral or nomadic management systems—tend to be found together. together. (This is
typical of the dryland system category in Box 2.2.)
We use overlapping categories in the global MA analysis because this
better reflects real-world biological, geophysical, social, and economic in-
teractions, particularly at these relatively large scales. For example, an im-
portant issue for ecosystems and human well-being in forested regions re-
lates to the impact of forest harvest or conversion on the timing, quantity,
and quality of water runoff. Given the importance of this interaction, it is
helpful to analyze an area dominated by forest land cover as a single ecosys-
tem even if it contains some freshwater and agricultural areas within it,
This website stores data such as
rather than analyzing the forest, agriculture, and freshwater ecosystems sepa-
cookies to enable essential site
rately,
functionality, as since this allows for a more holistic analysis of these interactions.
well as marketing,
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Ecosystem Services
or accept the default settings.

Privacy Policy Ecosystem services are the benefits people obtain from ecosystems. This
definition is derived from two other commonly referenced and representa-
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tive definitions:
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Ecosystem services are the conditions and processes through which natural
ecosystems, and the species that make them up, sustain and fulfill human
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54   Ecosystems and Human Well-being: A Framework for Assessment 

BOX 2.2 Report
Reporting
ing Categ
Categori
ories
es Used
Used in the Millenn
Millennium
ium Ecosys
Ecosystem
tem Asses
Assessme
sment
nt

Social and ecological systems can be categorized in an infinite number of ways. For
the purposes of reporting the global Millennium Ecosystem Assessment (MA) find-

ings,
on 10we have developed
systems. a practical,
Thus, for example, thetractable, sufficiently
MA will report rich systems,”
on “forest classification based
defined to
be areas with at least
l east 40 percent canopy (tree) cover. Using this approach, a forest
system will contain a variety of different types of ecosystems, such as freshwater
ecosystems, agroecosystems, and so forth. But all areas within the boundaries of the
forest system as defined here will tend to share a suite of biological, climatic, and
social factors, so the system categories provide a useful framework
f ramework for analyzing the
consequences of ecosystem change for human well-being. Because the boundaries
of these reporting categories overlap, any place on Earth may fall into more than
one category. Thus a wetland ecosystem in a coastal region, for instance, may be
examined both in the MA analysis of “coastal systems” as well as in the analysis of 
“inland water systems.”
The following table lists the basic boundary definitions that will be used in the
global MA analysis. In a number of cases the MA will also examine conditions and
changes in ecosystems with reference to more than one boundary definition. For
example, although we use a boundary of 40 percent tree (canopy) cover as our basic
definition of the forest category, another widely accepted definition of “forests” is at
least 10 percent canopy cover.

Millennium Ecosystem Assessment Reporting Categories

Category Central Concept Boundary Limits for Mapping

Marine Ocean, with fishing typically a major
m ajor Marine areas where the sea is deeper than 50 meters.
driver of change
Coastal Interface between ocean and land, Area between 50 meters below mean sea level and 50
extending seawards to about the middle meters above the high tide level or extending landward to
of the continental shelf and inland to a distance 100 kilometers from shore. Includes coral reefs,
include all areas strongly influenced by intertidal zones, estuaries, coastal aquaculture, and
the proximity to the ocean seagrass communities.
Inland Permanent water bodies inland from the Rivers, lakes, floodplains, reservoirs, and wetlands;
water coastal zone, and areas whose ecology includes inland saline systems. Note that the Ramsar
R amsar
This website stores data suchand as
use are dominated by the Convention considers “wetlands” to include both inland
cookies to enable essential permanent,
site seasonal, or intermittent water and coastal categories.
functionality, as well as marketing,
occurrence of flooded conditions
personalization, and analytics. You
may change yourForest
settings atLands
anydominated
time by trees; often used for A canopy cover of at least 40 percent by woody plants
timber, fuelwood, and non-timber forest
or accept the default settings. taller than 5 meters. The existence of many other
products definitions is acknowledged, and other limits (such as
crown cover greater than 10 percent, as used by the Food
Privacy Policy and Agriculture Organization of the United Nations) will
also be reported. Includes temporarily cut-over forests and
Marketing plantations; excludes orchards and agroforests where the
main products are food crops.
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Ecosystems and Their Services 55

BOX 2
 2..2 continued

Millennium Ecosystem Assessment Reporting Categories

Category Central Concept Boundary Limits for Mapping

Dryland Lands where plant production is limited Drylands as defined by the Convention to Combat
C ombat
by water availability; the dominant uses Desertification, namely lands where annual precipitation is
are large mammal herbivory, including less than two thirds of potential
potential evaporation, from dry
livestock grazing, and cultivation subhumid areas (ratio ranges 0.50–0.65), through
semiarid, arid, and hyper-arid (ratio <0.05), but excluding
polar areas; drylands include cultivated lands, scrublands,
shrublands, grasslands, semi-deserts, and true deserts.
Island Lands isolated by surrounding water, As defined by the Alliance of Small Island States
with a high proportion of coast to
hinterland
Mountain Steep and high lands As defined by Mountain Watch using criteria based on
elevation alone, and at lower elevation, on a combination
of elevation, slope, and local elevation range. Specifically,
elevation >2,500 meters, elevation 1,500–2,500 meters

and
slopeslope >2 degrees,
>5 degrees elevation
or local 1,000–1,500
elevation m eters and
meters
range (7 kilometers
radius) >300 meters, elevation 300–1,000 meters and
local elevation range (7 kilometers
k ilometers radius) >300 meters,
isolated inner basins and plateaus less than 25 square
kilometers extent that are surrounded by mountains.
Polar High-latitude systems frozen for most of Includes ice caps, areas underlain by permafrost, tundra,
the year polar deserts, and polar coastal areas. Excludes high-
altitude cold systems in low
lo w latitudes.
Cultivated Lands dominated by domesticated plant Areas in which at least 30 percent of the landscape comes
species, used for and substantially under cultivation in any particular year. Includes orchards,
changed by crop, agroforestry, or agroforestry, and integrated agriculture-aquaculture
aquaculture production systems.
Urban Built environments with a high human Known human settlements with a population of 5,000 or
density more, with boundaries delineated by observing persistent
night-time lights or by inferring areal extent in the cases
where such observations are absent.

life. They maintain biodiversity and the production of ecosystem goods, such
This website stores data such as
as seafood,
cookies to enable essential site forage timber, biomass fuels, natural fiber, and many pharmaceu-
functionality, as well ticals,
as marketing,
industrial products, and their precursors (Daily 1997b:3).
personalization, and analytics. You
may change your settings at any time
or accept the defaultEcosystem
settings. goods (such as food) and
an d services (such as waste assimilation) rep-
resent the benefits human
huma n populations derive, directly or indirectly, from eco-
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The MA definition follows Costanza and his colleagues in including
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both
Analytics natural and human-modified ecosystems as sources of ecosystem ser-
vices, and it follows Daily in using the term “services” to encompass both
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56   Ecosystems and Human Well-being: A Framework for Assessment 

the tangible and the intangible benefits humans obtain from ecosystems,
which are sometimes separated into “goods” and “services” respectively.
respectively.
Like the term ecosystem itself, the concept of ecosystem services is
relatively recent—it was first used in the late 1960s (e.g., King 1966;
Helliwell 1969). Research on ecosystem services has grown dramatically
within the last decade (e.g., Costanza et al. 1997; Daily 1997a; Daily et al.
2000; de Groot et al. 2002).
It is common practice in economics both to refer to goods and services
separately and to include the two concepts under the term services. Al-
though “goods,” “services,” and “cultural services” are often treated sepa-
rately for ease of understanding, for the MA we consider all these benefits
together as “ecosystem services” because it is sometimes difficult to deter-
mine whether a benefit provided by an ecosystem is a “good” or a “ser-
vice.” Also, when people refer to “ecosystem goods and services,” cultural
values and other intangible benefits are sometimes forgotten.
Ecosystem services have been categorized in a number of different ways,
including by:
 functional groupings, such as regulation, carrier, habitat, production,
and information services (Lobo 2001; de Groot et al. 2002);
 organizational groupings, such as services that are associated with cer-
tain species, that regulate some exogenous input, or that are related to
the organization of biotic entities (Norberg 1999); and
 descriptive groupings, such as renewable resource goods, nonrenewable
resource goods, physical structure services, biotic services, biogeochemi-
cal services, information services, and social and cultural services
(Moberg and Folke 1999).
For operational purposes, we will classify ecosystem services along func-
tional lines within the MA, using categories of provisioning, regulating,
cultural, and supporting services. (See Figure 2.1.) We recognize that some
This website stores data such as
of the categories overlap.
cookies to enable essential site
functionality, as well as marketing,
Provisioning
personalization, and analytics.Services
You
may change your settings at any time
These
or accept the defaultare the products
settings. obtained from ecosystems, including:
 Food and fiber. This includes the vast range of food products derived
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from plants, animals, and microbes, as well as materials such as wood,
jute, hemp, silk, and many other products derived from ecosystems.
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
Fuel. Wood,
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energy.
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Ecosystems and Their Services 57

 FIGURE 2.
2.1
1 Ecos
Ecosys
yste
tem
m Se
Serv
rvic
ices
es

Ecosystem services are the benefits people obtain from ecosystems. These include provi-
sioning, regulating, and cultural services that directly affect people and supporting
s upporting services
needed to maintain the other services.

 Genetic resources. This
resources. This includes the genes and genetic information used
for animal and plant breeding and biotechnology
biotechnology..
 Biochemicals, natural medicines, and pharmaceuticals. 
pharmaceuticals.  Many medicines,
biocides, food additives such as alginates, and biological materials are
derived from ecosystems.


Ornamental resources.
resources. Animal
flowers are used  Animal although
as ornaments, products, the
suchvalue
as skins and resources
of these shells, and
is
often culturally determined. This is an example of linkages between
the categories of ecosystem services.
This website stores data such as
cookies to enable

essentialwater. Fresh
Fresh site
water.  Fresh water is another example of linkages between catego-
functionality, as well as marketing,
ries—in
personalization, and analytics. this
You case, between provisioning and regulating services.
may change your settings at any time
or accept the default settings.
Regulating Services
These are the benefits obtained from the regulation of ecosystem processes,
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including:
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  Air quality maintenance. Ecosystems
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maintenance. Ecosystems both contribute chemicals to and
Analytics extract chemicals from the atmosphere, influencing many aspects of air
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58   Ecosystems and Human Well-being: A Framework for Assessment 

 Climate regulation. Ecosystems
regulation. Ecosystems influence climate both locally and glo-
bally. For example, at a local scale, changes in land cover can affect
both temperature and precipitation. At the global scale, ecosystems play
an important role in climate by either sequestering or emitting green-
house gases.
 Water regulation. 
regulation.  The timing and magnitude of runoff, flooding, and
aquifer recharge can be strongly influenced by changes in land cover,
including, in particular, alterations that change the water storage po-
tential of the system, such as the conversion of wetlands or the replace-
ment of forests with croplands or croplands with urban areas.
 Erosion control. Vegetative
control. Vegetative cover plays an important role in soil reten-
tion and the prevention of landslides.
 Water purification and waste treatment. 
treatment.  Ecosystems can be a source of 
impurities in fresh water but also can help to filter out and decompose
organic wastes introduced into inland waters and coastal and marine

ecosystems.
 Regulation of human diseases. Changes
diseases. Changes in ecosystems can directly change
the abundance of human pathogens, such as cholera, and can alter the
abundance of disease vectors, such as mosquitoes.
 Biological control. Ecosystem changes affect the prevalence of crop and
livestock pests and diseases.
 Pollination. Ecosystem changes affect the distribution, abundance, and
Pollination. Ecosystem
effectiveness of pollinators.
 Storm protection. The
protection. The presence of coastal ecosystems such as mangroves
and coral reefs can dramatically reduce the damage caused by hurri-
canes or large waves.

Cultural Services
These are the nonmaterial benefits people obtain from ecosystems through
This website stores data such as
spiritual enrichment, cognitive development, reflection, recreation, and
cookies to enable essential site
aesthetic
functionality, experiences, including:
as well as marketing,
personalization, and analytics. You
may change Cultural
 your diversity.
diversity. The
settings at any time  The
diversity of ecosystems is one factor influencing
or accept the default settings.
the diversity of cultures.

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Spiritual and religious values. 
values.  Many religions attach spiritual and reli-
gious values to ecosystems or their components.
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 Knowledge systems (traditional
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systems (traditional and formal). Ecosystems influence the
types of knowledge systems developed by different cultures.
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Ecosystems and Their Services 59

 Educational values. Ecosystems
values. Ecosystems and their components and processes pro-
vide the basis for both formal and informal education in many societies.
 Inspiration. Ecosystems provide a rich source of inspiration for art, folk-
Inspiration. Ecosystems
lore, national symbols, architecture, and advertising.


 Aesthetic
aspects ofvalues. Many
values. Manyaspeople
ecosystems, find
reflected in beauty or aesthetic
the support for parks,value in drives,”
“scenic various
and the selection of housing locations.
 Social relations. Ecosystems
relations. Ecosystems influence the types of social relations that
are established in particular cultures. Fishing societies, for example, dif-
fer in many respects in their social relations from nomadic herding or
agricultural societies.
 Sense of place. Many
place. Many people value the “sense of place” that is associated
with recognized features of their environment, including aspects of the
ecosystem.
 Cultural heritage values. Many
values. Many societies place high value on the mainte-
nance of either historically important landscapes (“cultural landscapes”)
or culturally significant species.
 Recreation and ecotourism. 
ecotourism.  People often choose where to spend their
leisure time based in part on the characteristics of the natural or culti-
vated landscapes in a particular area.
Cultural services are tightly bound to human values and behavior, as
well as to human institutions and patterns of social, economic, and politi-
cal organization. Thus perceptions of cultural services are more likely to
differ among individuals and communities than, say, perceptions of the
importance of food production. The issue of valuing ecosystem services is

addressed in Chapter 6.
Supporting Services
Supporting services are those that are necessary for the production of all
This website stores data such as
other
cookies to enable ecosystem
essential site services. They differ from provisioning, regulating, and
functionality, as well as marketing,
cultural services in that their impacts on people are either indirect or oc-
personalization, and analytics. You
may change your cursettings
overataany very
timelong time, whereas changes in the other categories have
or accept the default settings.direct and short-term impacts on people. (Some services, like
relatively
erosion control, can be categorized as both a supporting and a regulating
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Marketingpeople.) For example, humans do not directly use soil formation services,
although changes in this would indirectly affect people through the im-
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60   Ecosystems and Human Well-being: A Framework for Assessment 

regulation is categorized as a regulating service since ecosystem changes

can have an impact on local or global climate over time scales
s cales relevant to
human decision-making (decades or centuries), whereas the production
of oxygen gas (through photosynthesis) is categorized as a supporting ser-
vice since any impacts on the concentration of oxygen in the atmosphere
would only occur over an extremely long time. Some other examples of 
supporting services are primary production, production of atmospheric
oxygen, soil formation and retention, nutrient cycling, water cycling, and
provisioning of habitat.

 A Multisectoral Approach
Every part of Earth produces a bundle of ecosystem services. (See Box
2.3.) Human interventions can increase some services, though often at
the expense of other ones. Thus human interventions have dramatically
increased food provisioning services through the spread of agricultural tech-
nologies, although this has resulted in changes to other services such as
water regulation. For this reason, a multisectoral approach is essential to
fully evaluate changes in ecosystem services and their
t heir impacts on people.
The multisectoral approach examines the supply and condition of each
ecosystem service as well as the interactions among them. The MA has
adopted just such an approach.
When assessing ecosystem services, it is often convenient to bound the
analysis spatially and temporally with reference to the ecosystem service
or services being examined. Thus a river basin is often the most valuable
ecosystem scale for examining changes in water services, while a particu-
lar agroecological zone may be more appropriate for assessing changes in
crop production. When looking at interactions among services, the com-
bination of services provided by an ecosystem, or the variety of services
drawn on by a societysociety,, the question of boundaries becomes more complex.
Issues of boundaries, scale, and habitat heterogeneity are important and
arestores
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Biodiversity and Ecosystem Services
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Habitat modification, invasion, and many other factors are leading to
changes in biodiversity across
acros s many taxa within most ecosystems. Recently,
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theoretical and empirical work has identified linkages between changes in
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Ecosystems and Their Services 61

BOX 2.3 Analy

Analysi
siss of Ec
Ecos
osys
yste
tem
m Serv
Servic
ices
es

Any region of Earth produces a set of services that in turn influences human well-
being. It also receives
r eceives flows of energy, water,
water, organisms, pollutants, and other mate-

rials from adjacent

strategies regions and
and interventions releasesthe
influence similar materials
quantity into those
and quality regions.
of the Various
Various
services pro-
vided.
An ecosystem is typically composed of a number of different regions, such as
forest, agriculture, and urban areas, each of which produces a different bundle of 
services. In an ecosystem assessment, both the production of services
s ervices from each area
and the flows of materials between areas must be assessed.

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Among the most important factors identified is the degree of func-
tional redundancy found within an ecosystem. This indicates the substi-
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the impact created by the loss of one or more species is compensated for by
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62   Ecosystems and Human Well-being: A Framework for Assessment 

others (Naeem 1998). For example, in many ecosystems there are several
species that fix nitrogen (known as a functional group of species). If the
loss of any one of them is compensated for by the growth of others and
there is no overall loss in nitrogen fixation, then there is functional re-
dundancy in that ecosystem.
Some species make unique or singular contributions to ecosystem func-
tioning, however, and therefore their loss is of greater concern (Walker
1992). Small changes in the biodiversity of diverse systems may lead to
only small changes in the functioning of an ecosystem, including its pro-
duction of services, providing no species with unique roles are lost (Jones
et al. 1994; Power et al. 1996). But the possibility of significant losses of 
function increases as more species are lost and as redundancy is reduced—
that is, there is an asymptotic relationship between biodiversity and eco-
system functioning. For example, the high diversity of South African fynbos
ecosystems ensures steady rates of production because many plant species
can compensate for losses by growing when others cannot (Cowling et al.
1994). Greater redundancy represents greater insurance that an ecosys-
tem will continue to provide both higher and more predictable levels of 
services (Yachi and Loreau 1999).
The MA will seek to evaluate biodiversity and potential declines in
biodiversity for different ecosystems under a set of different scenarios for
plausible changes in driving forces. This work will extend previous studies
that developed scenarios for biodiversity change (Sala et al. 2000). For
provisioning and supporting services, the MA will identify which ecosys-
tem functions are associated with these services and link their response to
declining biodiversity,
biodiversity, using the
t he fundamental asymptotic relationship be-
tween biodiversity and ecosystem functioning. Both magnitudes and sta-
bility responses to biodiversity loss can be considered using this funda-
mental relationship.

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Ecosystem Condition and Sustainable Use
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People
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thus
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condition of an ecosystem in relation to its ability to
pro-
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vide the services desired. The ability of ecosystems to deliver particular
services can be assessed separately with various methods and measures.
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An adequate assessment of the condition of ecosystems, the provision of 
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decision process (see Chapter 8) can then determine which set of ser-
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Ecosystems and Their Services 63

vices are valued most highly (see Chapter 6) and can manage the system
in a sustainable way.
In a narrow sense, the sustainability of the production of a particular
ecosystem service can refer simply to whether the biological potential of 
the ecosystem to sustain the yield of that service (such as food produc-
tion) is being maintained. Thus a fish provision service is sustainable if 
the surplus but not the resource base is harvested, and if the fish’s habitat
is not degraded by human activities. In the MA, we use the term “sus-
tained yield management” to refer to the management and yield of an
individual resource or ecosystem service.
More generally, however, sustainability is used in the context of “sus-
tainable development” to refer to a pattern of development that meets
current needs without diminishing prospects for future generations. We
use sustainability, and sustainable management, to refer to this goal of en-
suring that a wide range of services from a particular ecosystem is sustained.
The MA will consider criteria and methods to provide an integrated
approach to ecosystem assessment. The condition and sustainability of each
category of ecosystem services is evaluated in somewhat different ways, al-
though in general a full assessment of any service requires considerations of 
stocks, flows, and resilience.

Condition of Provisioning Services

The flows of provisioning services do not accurately reflect their condi-
tion, since a given flow may or may not be sustainable over the long term.
The flow is typically measured in terms of biophysical production, such as
kilograms of maize per hectare or tons of tuna landings. The provisioning
of ecological goods such as food, fuelwood, or fiber, depends both on the
flow and the “stock” of the good, just as is the case with manufactured
goods. (In economics, “stock” refers to the total merchandise kept on hand
by a merchant; in this section, we use “stock” in its economic sense to
showdata
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as of ecosystem goods can be incorporated into the
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economic framework of stocks and flows.) The quantity of goods sold by a
functionality, as well as marketing,
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pro duction of new goods
or the depletion of built-up stocks. Indeed, production of biological re-
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sources has often been maintained in the short term at a higher rate than
its sustainable yield. In the long term, the production of overharvested
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graded even while output has been temporarily maintained or increased
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64   Ecosystems and Human Well-being: A Framework for Assessment 

by more intensive harvesting. Numerous fisheries around the world have

been overharvested, exhibiting a general pattern of rapid growth in land-
ings (production) followed by the eventual collapse of the fishery. (See
Box 2.4.) Similar patterns can be found with virtually all other provision-
ing services.
Agricultural production, for example, can be maintained through the
addition of fertilizers and through new crop varieties even while the pro-
ductive potential of the ecosystem is degraded through soil erosion. Some
40 percent of agricultural land has been strongly or very strongly degraded
in the past 50 years by erosion, salinization, compaction, nutrient deple-
tion, biological degradation, or pollution even while overall global food
production has increased (WRI et al. 2000). So long as manufactured capital
can compensate for losses of the natural capital of the ecosystem, agricul-
tural production can be maintained. In this case, however
however,, manufactured
and natural capital are not perfectly substitutable, and once a critical level
of soil degradation is reached, agricultural output will decline. A complete
accounting of the condition of food production would reveal that it has
been degraded because the underlying capability of the ecosystem to main-
tain production has been degraded.
Historically, it has not been common for environmental or resource
assessments to include measures of the productive potential of biological
resources when monitoring the condition of the resource. Thus although
all countries have considerable information on the production of grain,
fisheries, and timber, relatively little is known about the actual condition
of these services since the productive potential of the resource has rarely
been evaluated. The Pilot Analysis of Global Ecosystems, which was pre-
pared by the World Resources Institute and the International Food Policy
Research Institute to assist in the MA design, attempted to provide a more
complete assessment of the condition of ecosystem services along these
lines (Matthews et al. 2000; Revenga et al. 2000; White et al. 2000; Wood
Wood
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Condition
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and analytics. You Cultural, and Supporting Services
may change your settings at any time
In the case of regulating services, as opposed to provisioning services,
or accept the default settings.
the
level of “production” is generally not relevant. Instead the condition of 
the service depends more on whether the ecosystem’s capability to regu-
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late a particular service has been enhanced or diminished. Thus if forest
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Ecosystems and Their Services 65

BOX 2.4 Collapse of the Atlantic Cod Fishery

The Atlantic cod stocks off the east coast of Newfoundland collapsed in 1992,
forcing the closure of the fishery after hundreds of years of exploitation. Until the

late
shore1950s, the fishery
small-scale was
fishers. exploited
From by1950s,
the late migratory seasonal
offshore fleets
bottom and resident
trawlers in-
began ex-
ploiting the deeper part of the stock, leading to a large catch increase and a strong
decline in the underlying biomass. Internationally agreed quotas in the early 1970s
and, following the declaration by Canada of an Exclusive Fishing Zone in 1977,
national quota systems ultimately failed to arrest and reverse the decline.
Two factors that contributed to the collapse of the cod stock were the shift to
heavy fishing offshore and the use of fishery assessment methods that relied too
much on scientific sampling and models based on the relatively limited time series
and geographical coverage of the offshore part of the
t he fish stocks. Traditional
Traditional inshore
fishers, whose landings account for one third to one half of the total, had noticed
the decline in landings even before the mid-1980s, ahead of the scientists involved
in fisheries assessment work but these observations could not be used in stock as-
sessments because of technical difficulties in converting the catches into a suitable
form. Finlayson (1994) noted that “science will confer the status of ‘valid’ only on
very specific forms of data presented in a very specialized format.”

Northern Cod Off Newfoundland, Canada (NAFO area 2J3KL)

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 Source: Myers et al. 1995.
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Marketing The evaluation of the condition of cultural services is more difficult.

Some cultural services are linked to a provisioning service (such as recre-
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66   Ecosystems and Human Well-being: A Framework for Assessment 

provisioning or regulating services, assessing the

t he condition of cultural ser-
vices depends heavily on either direct or indirect human use of the
service. For example, the condition of a regulating service such as water
quality might be high even if humans are not using the clean water pro-
duced, but an ecosystem provides cultural services only if there are people
who value the cultural heritage associated with it.
Information about the condition of cultural services can be obtained
by identifying the specific features of the ecosystem that are of cultural,
spiritual, or aesthetic significance and then examining trends in those fea-
tures. For example, salmon are a totemic or revered species in almost all
parts of the world where they are found, and thus the degradation of wild
salmon stocks represents degradation of a cultural service provided by the
ecosystem. But cultural service information such as this would be difficult
to obtain and to quantify: tigers, for instance, remain totemic species even
in areas where they have been extinct for decades. Recognizing that the
concept of cultural services is relatively new, the MA will explore meth-
ods for evaluating the condition and value of these services.
Supporting services maintain the conditions for life on Earth but may
affect people only indirectly (by supporting the production of another ser-
vice, as soil formation supports food production) or over very long time
periods (such as the role of ecosystems in producing oxygen). Because the
link to human benefits is indirect, as opposed to the other ecosystem ser-
vices just discussed, a normative scale for assessing the condition of a ser-
vice is not always practical. For example, primary production is a funda-
mental supporting service, since life requires the production of organic
compounds. But if global primary production were to increase by 5 per-
cent over the next century, it would be difficult to categorize the change
as an enhancement or degradation of the service, though it certainly would
be a significant change. In such cases the MA will report on the current
biophysical state (production, flux, and stocks) of the supporting service.
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 Variability
ariability, , site
Resilience, and Thresholds in Services
functionality, as well as marketing,
Whenever
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and analytics. You individuals and governments generally invest in
vari-
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ous types of insurance that can buffer human welfare against natural
or accept the default settings.
vari-
ability. Such investments may be as basic as establishing limited stores of 
food, medicine, and potable water for disaster relief to more elaborate in-
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vestments such as building dams, levies, and canals to guard against 100-
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assessing
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also their dynamics or, more specifically,
specifically, their variability and stability
st ability..
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Ecosystems and Their Services 67

Three characteristics of ecosystem services are important in such an

assessment:
assessmen t: ecosystem variability
variabilit y, resilience, and thresholds.
thresholds . (See Box 2.5.)
There are many other properties of stability in dynamic systems (such as
resistance, sensitivity
s ensitivity,, persistence, reliability
reliability,, predictability,
predictability, and so forth),
but the MA will limit its focus to these three important and well-studied
stability properties.
Variability  in ecosystem services consists of changes in stocks or flows
Variability
over time due to stochastic, intrinsic, and extrinsic factors, all of which
must be disentangled to understand system behavior properly. Stochastic
variability is due to random or uncontrolled factors creating variability
that is often considered background or “white” noise in system behavior.

BOX 2.5 Dynami

Dynamics
cs and Sta
Stabili
bility
ty in
in Ecosy
Ecosyste
stem
m Servi
Services
ces

This figure illustrates the level of provisioning

provisioni ng of an ecosystem service that has been
perturbed twice. Hypothetically, such a service exhibits stochastic (random or un-
controlled) and inherent variability (fluctuations above and below the two hori-
zontal lines, which represent different system states). The system recovers after the
first perturbation, with its resilience being measured by the
t he duration of the recovery
phase or return time to its first state. Note that crossing the threshold of the second
state does not cause a shift when in the first state. The second perturbation causes
the service to cross the second threshold, which leads to a regime shift or cata-
strophic change to an alternative stable state. The long dashed lines illustrate two
thresholds. Only when a system crosses a threshold does it switch to an alternate
state.

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68   Ecosystems and Human Well-being: A Framework for Assessment 

In contrast, intrinsic (inherent) variability is due to the structural

str uctural proper-
ties of an ecosystem, such as oscillations in systems where predation or
disease regulate the number of animals. Examples of extrinsic variability,
due to forces outside the system, include seasonality in temperate systems
and longer-term climate systems such as El Niño–La Niña cycles.
Resilience is most often considered a measure of the ability of a system
to return to its original state after a perturbation—a deviation in condi-
tions that is outside the range experienced over a decade or more, such as
a large-scale fire or an unusually severe drought. When the duration of the
recovery phase is short in comparison to other systems, the system is con-
sidered to be more resilient than the others.
Thresholds or breakpoints in ecosystems represent dramatic, usually
sudden (less than a decade) deviations from average system behavior. behavior. Such
dramatic shifts—also known as regime shifts, catastrophic change, or en-
tering alternative stable states—are often primed by a steady change in
internal or external conditions that increases a system’s susceptibility to
being triggered to enter an alternative state (Scheffer et al. 2001; Carpen-
ter 2003). For example, on a global scale, small, steady increases in global
warming may lead to a sudden reorganization of Earth’ Earth’ss ocean circulation
patterns (Broecker 1997). On a local scale, the increase in grazing animals
by ranchers or herders may be responsible for shifts in steppe (grass-
dominated) to tundra (moss-dominated) ecosystems (Zimov et al. 1995).
While management goals are often conceived in terms of stocks and
flows, reducing system variability and improving predictability are a re often
key parts of management strategies. Examples of such interventions
include irrigating crops during droughts, using biocides during pest out-
breaks, controlled burning to prevent catastrophic
catastr ophic fires, and culling herds
to prevent a population explosion. Maintaining forests to prevent ero-
sion or coral reefs to prevent wave impacts in the face of severe storms
are examples of managing ecosystems for their insurance value. Ecosys-
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thresholds You
is sometimes overlooked. In part this is because the mecha-
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additi
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probability of perturba-
perturba-
tions, and the time frame over which such events occur is too long.
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behavior,,
behavior
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however
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Ecosystems and Their Services 69

ecosystem stocks and flows as they are related to ecosystem goods and
services, but also their stability properties. Much of this will be done by
extrapolation from expert assessment of paleo records (for instance, cli-
mate records derived from ice cores) and historical records (such as long-
term fisheries, forestry, or agricultural records) to obtain guidelines on the
norms of system variability, resilience, known thresholds, and the envi-
ronmental stresses that cause ecosystems to be triggered by perturbations
to enter into alternative states.

 Ecosystem Health and Other Related Concepts

Ecosystem health is a concept that has often been applied to the evalua-
tion of ecosystems (Rapport et al. 1995). This has become a subdiscipline
in the life sciences, with its own journals and professional organizations,
such as the International Society for Ecosystem Health (ISEH) and the
Aquatic Ecosystem Health and Management Society. The term is used
sometimes to mean the links between ecosystems and human health. For
example, the mission of ISEH is to “encourage the understanding of the
critical linkages between human activity, ecological change and human
health” (Rapport et al. 1999:83). It is also used to refer to the health of the
ecosystem itself: “an ecological system is healthy…if it is stable and sus-
tainable—that is, if it is active and maintains its organization and au-
tonomy over time and is resilient to stress” (Costanza et al. 1992:9).
This concept has generated debate and alternative approaches within
the scientific literature (e.g., Reid 1996; de Leo and Levin 1997). One
method measures health as a departure from some preferred (often “natu-
ral”) state. Another, which is consistent with the approach used in the MA
to examine the condition of ecosystem services, relates health to the abil-
ity of an ecosystem within its surrounding landscape to continue to provide
a particular set of services. This considers whether the ecosystem and its
external inputs (such as energy or fertilizer) are sustainable in the long
termdata
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health as its primary organizational framework, the concept could be use-
fully applied within an assessment that used the MA framework.
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70   Ecosystems and Human Well-being: A Framework for Assessment 

characteristic of a particular locale or deemed satisfactory to society” (Cairns

1977:56) or “the capability of supporting and maintaining a balanced, in-
tegrated, adaptive community of organisms having species composition,
diversity,, and functional organization comparable to that of natural habi-
diversity
tats of the region” (Karr and Dudley 1981:171). Another example is the
“ecological footprint,” which expresses the impact of human activity on
ecosystems in terms of areas required to provide the services used by an
individual or community.

Substitution of Services
Substitutes are available for some ecosystem services, although often the
cost of a technological substitution will be high and it may not replace all
the services lost. For example, water treatment plants can now substitute for
ecosystems in providing clean drinking water,
water, although this may be expen-
sive and will not overcome the impacts of water pollution on other compo-
nents of the ecosystem and the services they provide. Another outcome of 
substitution is that often the individuals gaining the benefits are not those
who originally benefited from the ecosystem services. For example, local
coastal fish production can be replaced by shrimp aquaculture in tropical
regions, but the individuals making a living from capture fisheries are not
those who would profit from thet he new shrimp aquaculture facilities.
Therefore, a full assessment of ecosystems and their services must
consider:
 information on the cost of a substitute,
 the opportunity cost of maintaining the service,
 cross-service costs and impacts, and
 the distributional impacts of any substitution.

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