Chapter 6

Ecosystems and Ecosystem Services

M.L. Brusseau

Chapter cover art: Ecosystem services—sources, benefits, and drivers of change From: https://www.epa.gov/enviroatlas/ecosystem-services-enviroatlas

The environment is much more than its physical compo- region at the Earth’s surface (called the critical zone),
nents such as soil, rocks, water, and air. Lifeforms are and particularly soil, is a key component of ecosystems that
present in the environment, and these lifeforms are sup- support all terrestrial life. These concepts are presented in
ported by the environment and have myriad impacts on this chapter.
the environment. These impacts can in turn affect the ability
of the environment to support life. This integrated system of
organisms and the physical components of the environment
6.1 ECOSYSTEMS
is called an ecosystem. Organisms such as humans receive
many benefits from ecosystems and can also be negatively Chapters 2–4 covered the physical and chemical properties
impacted by environmental components or processes. The of the environment. The biological component of the

Environmental and Pollution Science. https://doi.org/10.1016/B978-0-12-814719-1.00006-9

Copyright © 2019 Elsevier Inc. All rights reserved. 89
90 PART I Fundamental Concepts

environment—life, is covered in Chapter 5 specifically for l Organism: An individual organism (single cell or
microorganisms and more generally in this chapter. animal). Organismal ecologists study adaptations, ben-
The fundamental operational unit for characterizing the eficial features arising by natural selection, that allow
presence, activities, and impacts of lifeforms in the envi- organisms to live in specific habitats. These adaptations
ronment is an ecosystem. An ecosystem consists of a com- can be morphological, physiological, or behavioral.
munity of organisms living in a discrete designated area, l Population: A population is a group of organisms of the
together with the nonliving components of that area (the local same species that live in the same area at the same time.
physical environment). It is important to understand the dif- Population ecologists study the size, density, and
ference between “community” and “ecosystem.” A com- structure of populations and how they change over time.
munity is the aggregate of all of the populations of the l Community: A biological community consists of all the
different species that live together in the designated area. In populations of different species that live in a given area.
other words, a community is the biotic, or living, component Community ecologists focus on interactions between
of an ecosystem. Conversely, an ecosystem includes the populations and how these interactions shape the
physical environment in addition to the biotic component. community.
The sizes of ecosystems vary across a wide range. They l Ecosystem: An ecosystem consists of all the organisms
can be small, such as a tidal pool, a decaying tree trunk, or in an area, the community, and the abiotic factors that
the earth beneath a boulder. They can also be very large, influence that community. Ecosystem ecologists often
such as the Sahara desert or the Amazon rainforest. Very focus on flow of energy and recycling of nutrients.
large ecosystems are sometimes referred to as “Ecore- l Biome: A biome is a set of ecosystems sharing similar
gions.” In addition, ecosystems can often be subdivided into characteristics (Information Box 6.1).
smaller individual subecosystems. For example, the l Biosphere: The biosphere is planet Earth, viewed as the
Amazon rainforest basin has several subecosystems, at dif- aggregation of all of the ecosystems and biomes into one
ferent scales. The largest set of these subecosystems include integrated and interdependent system. Ecologists
(i) floodplain forests or varzea (the areas close to rivers that working at the biosphere level may study global pat-
are flooded during the rainy season); (ii) terra firma rain- terns—for example, climate or species distribution—
forest (forest that is off the floodplain); (iii) igapo (forests interactions among ecosystems, and phenomena that
around freshwater lakes and lower reaches of rivers); (iv) affect the entire globe, such as climate change.
the Cerrado savanna, significantly drier than the interior
The different levels of ecological investigation presented
Amazon forest; and (v) the Amazon River aquatic eco-
earlier are listed from small to large. They build progres-
system. Each one of these major subecosystems can be
sively—populations are comprised of individual organisms,
divided into smaller individual ecosystems, such as the tree
while communities are comprised of populations, and sim-
canopy, the forest land surface, and streams and ponds
ilarly a biome consists of a set of ecosystems. Each level has
within a single forest area. Thus we recognize that eco-
emergent properties, meaning new properties that are not a
systems can be defined at different telescoping scales. Arti-
part of any of the individual components comprising that
ficial boundaries are imposed to define a specific ecosystem
level, but instead emerge from the various synergistic and
when scientists wish to examine that particular ecosystem.
antagonistic interactions and relationships among the com-
Ecosystems also vary greatly in their properties, both in
ponents. Ecological studies will often incorporate more
the physical nature of the environment and in the types of
than one level to examine interactions among the levels.
lifeforms present. The major types of large-scale eco-
There are several different types of interactions among
systems are categorized into biomes. These are very large
individuals and populations within a community. Compe-
regions of the Earth’s surface that have unique fauna and
tition occurs when more than one organism or population
flora that have developed because of the specific geologic
uses a resource at the same time. Predation is one organism
factors present, such as climate (temperature and precipi-
or population feeding on another. Many species obtain their
tation). The major biomes are typically designated as
food by eating other organisms. Symbiotic Relationships are
Desert, Aquatic, Grasslands, Forests, and Tundra. These
the close relationships that exist when two or more species
are briefly described in Information Box 6.1. A map
live together and each provides benefits to the other. A
showing the distribution of major biomes on Earth is pre-
habitat is an area where an organism lives. A niche is the
sented in Fig. 6.1.
role or position that an organism has in its ecosystem.
Ecology is the science discipline that studies how
The feeding relationship in an ecosystem is called a food
organisms interact with each other and with their physical
chain. Food chains are usually in a sequence, with an arrow
environment. Ecological studies can be conducted at six
used to show the flow of food. A food web is a network of
general levels: organism, population, community, eco-
many food chains and is more complex, detailing the
system, biome, and biosphere (Fig. 6.2). These levels are
feeding interactions within the whole ecosystem
defined as follows:
 INFORMATION BOX 6.1 Major Biomes
Aquatic: Forests:
Aquatic biomes contain numerous species of plants and Today, forests occupy approximately one-third of the Earth’s
animals, both large and small. Although water temperatures land area, account for over two-thirds of the leaf area of land plants,
can vary widely, aquatic areas tend to be more humid and and contain about 70% of carbon present in living things. Forest
the air temperature on the cooler side. The aquatic biome can biomes have relatively high rates of precipitation and are domi-
be divided into two basic types, freshwater (i.e., lakes, ponds, nated by trees and other woody vegetation. Soils are generally rich,
streams, and wetlands) and marine (i.e., oceans and estuaries). with high acidity and decaying organic matter. Forests typically
The major property differentiating these two types of systems is have very large and diverse communities (large biodiversity).
the salinity of the water. See Chapter 3 for discussion of these Forest biomes are classified according to numerous charac-
two systems. teristics, with seasonality being the most widely used. Distinct
The plants and animals that live in each type of system have forest types also occur within each of these broad groups. There
evolved the ability to thrive under the respective conditions. are three major types of forests, classified according to latitude:
The physical conditions of aquatic systems have led to great tropical (e.g., the Amazon rain forest in Brazil), temperate (e.g.,
differences in the morphology of lifeforms living within these the forests of eastern United States), and boreal or taiga (e.g., the
systems versus those of terrestrial lifeforms. Examples include Scandinavian forest). The types of trees and other vegetation
massively large whales and the general streamlined shapes present differ greatly among these three forests.
of fish. Grasslands:
Deserts: Grasslands are characterized as lands dominated by grasses
Deserts cover about one-fifth of the Earth’s surface and occur rather than large shrubs or trees. Grasslands occur between the
where precipitation is less than 50 cm/year. Low magnitude of desert and mountain biomes, and are considered a transitional
rainfall is the defining characteristic of deserts, and it has a sig- biome. They typically receive moderate amounts of
nificant impact on the vegetation and animals present. Vege- precipitation.
tation is usually sparse, with a preponderance of smaller, There are two main types of grasslands, tropical grasslands
hardy plants. There are relatively few large animals in deserts (called savannas) and temperate grasslands. Savannas cover
because most are not capable of storing sufficient water and almost half of the African continent (about 5 million square
withstanding the heat (or cold). The dominant animals of warm miles, generally in central Africa) and large areas of Australia,
deserts are nonmammalian vertebrates, such as reptiles. South America, and India. The major examples of temperate
Mammals are usually small, like the kangaroo mice of North grasslands are the veldts of South Africa, the puszta of Hungary,
American deserts. the pampas of Argentina and Uruguay, the Eurasian steppes, and
Although most deserts, such as the Sahara of North Africa the plains and prairies of central North America.
and the deserts of the southwestern United States, Mexico, Tundra:
and Australia, occur at low latitudes, another kind of desert, Tundra is the coldest of all of the terrestrial biomes. Tundra
cold deserts, occur in the basin and range area of Utah and originates from the Finnish word tunturia, meaning treeless
Nevada and in parts of western Asia. More generally, desert plain. This biome has very little precipitation, freezing temper-
biomes can be classified according to several characteristics. atures, and covers about a fifth of the Earth’s land surface. It has
There are four major types of deserts: Hot and Dry (e.g., Chi- the lowest biodiversity of all of the biomes. A key property of this
huahuan, Sonoran, and Mojave in North America), Semiarid biome is permafrost, which is the frozen ground of the tundra.
(e.g., the Great Basin in North America), Coastal (e.g., the Tundra is separated into two types, arctic tundra (the Arctic
Atacama of Chile), and Cold (e.g., regions in Antarctic and region of the northern hemisphere) and alpine tundra (the tops
Greenland). of mountainous areas).

FIG. 6.1 Major biomes on Earth. Reproduced from OpenStax College, Terrestrial Biomes. OpenStax CNX. 21 Jun 2013. http://cnx.org/contents/
cf87c2b7-d8ec-4cba-adba-6c30f5736f63@4.
92 PART I Fundamental Concepts

(Fig. 6.3). The levels of a food chain are called Trophic

levels. The trophic level of an organism is the level it holds
in the overall food pyramid (Fig. 6.4). The sun is the source
of energy in the vast majority of food chains. Green plants,
usually the first level of any food chain, absorb some of the
Sun’s light energy to make their own food via photosyn-
thesis. Green plants are therefore known as “Primary Pro-
ducers” in a food chain. The second level of the food
chain is called the Primary Consumer. These organisms
consume the green plants. Examples include insects, sheep,
caterpillars, and cows. The third level in the chain is Sec-
ondary Consumers, which eat the primary consumers and
other animal matter. They are commonly called carnivores
and examples include lions, snakes, and cats. The fourth
level is called Tertiary Consumers. These are animals that
eat secondary consumers. Quaternary Consumers eat ter-
tiary consumers. At the top of the levels are the apex
Predators.
Food chains and webs are an example of the flow of
matter and energy within an ecosystem. Other examples
of matter and energy flow are the various biogeochemical
cycles that occur within ecosystems. Important examples
are the carbon and nitrogen cycles, covered in
Section 6.4. Finally, physical cycles of geological and
hydrological processes are also important. These physical
and biogeochemical cycles are critical to the existence of
life, transforming energy and matter into usable forms to
support the functioning of ecosystems. A few general prin-
ciples govern energy and matter flow in ecosystems. Energy
flows through the ecosystem, usually entering as sunlight
and exiting as heat, and is not recycled. Conversely, matter
is constantly recycled within an ecosystem.

6.2 ECOLOGICAL PRINCIPLES

Ecosystems are dynamic systems—energy is constantly
flowing through an ecosystem, and materials are contin-
ually being recycled; individual organisms die while others
are being born; populations of different species are fluctu-
ating in their numbers; and climate is varying seasonally. In
essence, a static ecosystem is normally a dead ecosystem—
just as a static cell would normally be a dead cell or at least
dormant. An ecosystem whose composition and identity
remain generally constant despite fluctuations in physical
conditions and community composition is at steady state
and is said to be in equilibrium. A system in equilibrium
is still a dynamic system; it is not static.
Ecosystems may be knocked out of equilibrium by dis-
turbances, disruptive events that affect their composition or
functioning. These disturbances may be of natural (earth-
quake, volcanic eruption) or anthropogenic (pollution,
excessive resource use) origin. If a disturbance is severe
enough, it may change an ecosystem beyond the point of
FIG. 6.2 Levels of ecology. From https://en.wikipedia.org/wiki/Bio recovery, to where it is no longer resilient. This could lead
logical_organisation
to permanent impairment or loss of the ecosystem and the
 Ecosystems and Ecosystem Services Chapter 6 93

FIG. 6.3 Example of a food web. From https://pubs.usgs.gov/circ/circ1316/html/circ1316chap14.html

FIG. 6.4 Trophic levels.

94 PART I Fundamental Concepts

services they provide. Ecosystems may have more than one The degree to which individual components and pro-
state of equilibrium. Hence a perturbed ecosystem may be cesses are connected and interact among the different scales
able to transition to another state of equilibrium that is and levels of an ecosystem is another major factor
optimal for the current, postdisturbance, conditions. influencing resilience. This has been referred to as
The state of an ecosystem with respect to its stability or panarchy. Systems with higher levels of interconnectedness
instability is measured by two parameters—latitude and are generally considered to be more resilient.
precariousness. Latitude is the maximum extent to which Abiotic factors may also influence the resilience of an
a system can be altered before losing its ability to recover. ecosystem. Climate is one primary abiotic factor, acting
This point is designated as the critical threshold, or tipping through, for example, its control of water availability.
point, beyond which recovery is difficult or impossible. The properties and conditions of soil are another example.
Precariousness describes how close the ecosystem cur- Third, an ecosystem that comprises a higher diversity of
rently is to the critical threshold. habitats will generally be more resilient. Finally, the type
The response of ecosystems to disturbances, and their of disturbance itself can influence resilience, as a particular
ability to withstand or adapt to said disturbances, is deter- ecosystem may respond differently to different types of
mined by their resilience. Ecological resilience, then, rep- disturbances.
resents how readily and quickly an ecosystem returns to If an ecosystem’s resilience is overcome, it will then
equilibrium after being disturbed. This is a critical concept undergo a longer term series of changes through the process
in ecology for understanding the impacts of human activ- of ecological succession. Think of a lava flow, such as those
ities on ecosystems. In addition, ecological resilience has that still occur on the island of Hawaii on the slopes of
become a central concept for conservation practices and Mauna Loa and Kilauea, burning through portions of the
ecosystem restoration and management efforts. native rain forest. The lava cools quickly, but lays barren
A resilient ecosystem is one that has the capability to for many years, too hostile to support life. Eventually rain
maintain key functions and processes in the face of stresses, and wind erosion create tiny fissures in the lava, where life
either by resisting or adapting to change. Possessing a high can establish a foothold. Microorganisms, usually bacteria,
degree of resilience is a positive attribute in the vast majority are often the first forms of life to become established. In
of cases. However, resilience is not always a positive feature fact, very few disturbed sites are microbiologically sterile.
of an ecosystem. An ecosystem may be locked into an unde- Microorganisms are a prerequisite for plant growth.
sirable state, thus operating at suboptimal levels. An example Lichens and cyanobacteria are the next colonists to arrive
of such a condition is a lake under permanent eutrophic on the flow. Continual breakdown of the parent lava by
conditions, where an overabundance of nutrients results in acids secreted by the lichens produces a thin layer of soil
depleted oxygen levels that in turn lead to loss of desirable in which the first higher order plants can root. Generally,
aquatic species and the proliferation of undesirable ones. small ferns appear first, followed by grasses and shrubs
Two critical characteristics determine the resilience of a as the fissures widen due to the action of the plant roots.
particular ecosystem, resistance and adaptive capacity. Cyanobacteria fix nitrogen, which also supports the plant
Resistance is the ability of an ecosystem to remain at equi- life. Each stage of succession conditions the lava substrate
librium in spite of a disturbance. It represents the ease or to favor the next stage, and finally the rain forest is restored.
difficulty of changing the system. An ecosystem that is This process may take many tens to hundreds or even thou-
more resistant will likely exhibit fewer deleterious effects sands of years. These long time scales for recovery make it
of a perturbation. Adaptive capacity is the ability of an eco- imperative that anthropogenic disturbances are minimized
system to adapt to the change in conditions brought about and that ecosystems are protected and managed to maintain
by the disturbance. their resilience.
Many factors influence the resistance and adaptive
capacity of an ecosystem and thus its resilience. One major
6.3 THE CRITICAL ZONE
factor that is generally considered essential for resilience is
biodiversity. There are several types or levels of biodi- The Critical Zone is the upper surface layer of Earth—from
versity. Genetic diversity represents biological diversity the tops of the trees to the bottom of the groundwater
within a single population of a species. Community (Fig. 6.5). The term was coined for this purpose in 1998
diversity refers to the numbers and types of populations by Gail Ashley and represents the concept of ecosystems
or species present in the ecosystem. Functional diversity applied to that region of Earth that supports all terrestrial
refers to the differentiation of populations based on their life, including humans. It is a living, breathing, constantly
function or role in the ecosystem. Redundancy refers to evolving boundary layer where rock, soil, water, air, and
overlap in populations for a particular role or niche. Pos- living organisms interact. It also borders the most critical
sessing greater levels of biodiversity provides the eco- zone of the atmosphere that affects human life—the atmo-
system with more opportunities and abilities to resist and spheric boundary layer (see Chapter 4). These complex
adapt to changes. interactions regulate the natural habitat and determine the
 Ecosystems and Ecosystem Services Chapter 6 95

reservoirs of Earth—the atmosphere, the terrestrial bio-

sphere, the oceans, and the geosphere (soil, sediments,
and rocks). The six most common elements comprising
organic molecules—the backbone of lifeforms—are
carbon, nitrogen, hydrogen, oxygen, phosphorus, and
sulfur. These elements reside in the different reservoirs to
different extents and consist of a variety of chemical forms
both organic and inorganic.
Carbon is an essential element in the bodies of all living
organisms and an essential source of energy for many
organisms. As discussed in Chapter 4, carbon in the form
of CO2 in the atmosphere is a greenhouse gas that helps reg-
ulate Earth’s temperature to make it hospitable for life.
Carbon, in the form of oil, natural gas, and coal, is a critical
source of energy for human activities. Carbon is also a key
element for the manufacture of everyday products such as
plastics. Carbon is central to current issues of global climate
change (Chapter 25). The carbon cycle describes the
FIG. 6.5 The critical zone. Reproduced from http://criticalzone.org/ movement of carbon among the various reservoirs
national/research/the-critical-zone-1national/ based on a figure from (Fig. 6.6).
Chorover, J., Kretzschmar, R., Garcia-Pichel, F., Sparks, D.L., 2007. Soil Carbon cycles through the environment at different
biogeochemical processes in the critical zone. Elements 3, 321–326.
spatial and temporal scales. The carbon cycle can be char-
acterized as two interconnected subcycles. One subcycle
availability of life-sustaining resources, including food pro- consists of carbon exchange among living organisms,
duction and water quality. which occurs over short time scales of days–weeks–
The critical zone is imprinted by different types of months. The second subcycle comprises long-term cycling
events over an extremely wide range of time scales, from of carbon through geologic processes. It is important to note
seconds, hours, years, millennia, and geologic time. The that these two subcycles are linked.
present structure and functioning of the critical zone reflects The movement of carbon through the food chain repre-
the aggregate impacts of these various events. The critical sents the rapid carbon subcycle. Plants move carbon from
zone and human society are closely intertwined, impacting the atmosphere into the terrestrial biosphere through photo-
each other in many ways. Human activities can have a synthesis. These so-called autotrophs use energy from the
major impact on the conditions and functioning of the sun to convert CO2 into complex organic molecules such
critical zone. Various examples of these impacts are as glucose (sugar). Animals that eat these plants digest
covered in many later chapters of this book. Understanding the sugar molecules to obtain energy for their bodies in
the complex web of physical, chemical, and biological pro- the process termed respiration. These organisms are called
cesses of the critical zone requires a systems approach heterotrophs. Respiration, excretion, and decomposition
across a broad array of sciences: hydrology, geology, soil release the carbon back into the atmosphere or soil, con-
science, biology, ecology, geochemistry, geomorphology, tinuing the carbon cycle. A similar cycle occurs in the
and more. ocean, except that the marine autotrophs obtain their carbon
The critical zone provides many benefits to humans. in the dissolved form (HCO–3).
These have been termed “ecosystem services” and will be The movement of carbon between the primary reser-
covered in a following section. Certain interactions between voirs of Earth, the atmosphere, ocean, terrestrial biosphere
humans and the critical zone can lead to negative health out- (land), and the geosphere is generally very slow, occurring
comes. This is covered in Chapter 27. An essential com- over scales of hundreds to thousands of years or more.
ponent of the critical zone is soil. Soil health and the Carbon dioxide is the major form of carbon in the atmo-
interaction between soil and humans are also covered in sphere. The level of carbon dioxide in the atmosphere is
Chapter 27. greatly influenced by exchange between it and the ocean
and land reservoirs.
Carbon dioxide in the atmosphere dissolves in the
6.4 BIOGEOCHEMICAL CYCLES
ocean, after which it is transformed into different forms
Biogeochemical cycles are critical to the existence of life, (Fig. 6.7). Phytoplankton convert the carbon through pho-
transforming energy and matter into usable forms to support tosynthesis and become food sources for higher level
the functioning of ecosystems, as noted previously. These organisms. Some organisms combine carbon with calcium
cycles describe the movement of matter between the major ions in the seawater to form calcium carbonate (CaCO3), a
96 PART I Fundamental Concepts

CO2

Atmosphere Carbon cycle

Valcanoes

Fossil fuel
emissions

Vegetation Rivers
Soil
organic matter

Surface ocean

Marine biota

Dissolved organic
carbon
Deep ocean

Sediments

FIG. 6.6 The carbon cycle. From http://www.noaa.gov/resource-collections/carbon-cycle

major component of the shells of marine organisms. Marine carbon in the form of CO2 upon eruption. Clearly, this
organisms die and eventually form sediments on the ocean process occurs over many hundreds of thousands or mil-
floor. Over many hundreds of thousands of years, the sed- lions of years.
iment is subjected to geological processes to form limestone Carbon is stored in soil as a result of the decomposition
and other marine sedimentary rock. These solid forms of of living organisms or from weathering of terrestrial rock
carbon at the ocean bottom represent the largest carbon res- and minerals. Carbon cycling can be quite complex in the
ervoir on Earth. shallow subsurface as shown in Fig. 6.9. Carbon is stored
On land, carbon exists in living plants and animals gen- deeper underground in the form of decomposed remains
erated through the food web as discussed previously. of plants and animals that have been converted into
Transfer of carbon between land and the atmosphere occurs petroleum, natural gas, coal, and oil shale.
through photosynthesis–respiration processes associated The nitrogen cycle is another biogeochemical cycle
with these living organisms. In addition, human activities critical to life (Fig. 6.10). Nitrogen is especially important
such as fossil-fuel consumption are another source of to ecosystem dynamics because many ecosystem processes,
carbon to the atmosphere. Both of these sources are such as primary production and decomposition, are limited
depicted in Fig. 6.8. These transfer processes occur over rel- by the available supply of nitrogen. While it is an essential
atively short time scales. Another way for carbon to transfer element, the vast majority of organisms cannot make direct
from land to the atmosphere is through volcanoes and other use of the primary source of nitrogen, nitrogen gas (N2) in
geothermal systems. Carbon-containing sediments and the atmosphere. The conversion of N2 into a more useable
rocks on the ocean floor move deep within the Earth by form of nitrogen, ammonia, is carried out by nitrogen-fixing
the process of subduction, which is the movement of one bacteria. Once this occurs, various processes transfer the
tectonic plate beneath another. These materials can be nitrogen within the ecosystem, and ultimately back to the
transformed into molten rock (magma) under the high tem- atmosphere, as depicted in Fig. 6.10.
peratures and pressures present at great depth. This process Human activity can release nitrogen into the envi-
can lead to the formation of volcanoes, which releases ronment by two primary means. One is through the
 Ecosystems and Ecosystem Services Chapter 6 97

FIG. 6.7 The oceanic carbon cycle. From Office of Biological and Environmental Research of the U.S. Department of Energy Office of Science. https://
science.energy.gov/ber/

combustion of fossil fuels, which releases nitrogen oxides by most people. This situation has resulted in the fact that
into the atmosphere (Chapter 17). The second is through the full value and importance of the benefits are not con-
the use of fertilizers that contain nitrogen and phosphorus sidered in decision-making for most human activities. Con-
compounds in agriculture. Nitrogen can be taken up by comitantly, the full costs of negative impacts to the
surface runoff during precipitation events and eventually environment, and the subsequent loss of benefits, are not
deposit into lakes, streams, and rivers. A major effect from fully considered. The concept of Ecosystem Services was
fertilizer runoff is eutrophication, a process whereby developed to improve our ability to recognize and assign
nutrients from the runoff cause the overgrowth of algae value to the benefits provided by the environment and to
and a number of resulting problems (Chapter 16). The phos- better assess the impacts of human activities on the systems
phorus and sulfur cycles are other critical biogeochemical that provide these benefits.
cycles. More discussion of the nitrogen and phosphorous A landmark event in the development of this concept
cycles is presented in Chapter 16. was the Millennium Ecosystem Assessment (MEA), con-
ducted from 2001 to 2005 under the auspices of the United
Nations (WRI, 2005). It was governed by a multistake-
6.5 ECOSYSTEM SERVICES
holder board that included representatives of international
It has long been understood that human life and well-being institutions, governments, business, NGOs, and indigenous
depend upon resources and other benefits provided by the peoples. Its purpose was to assess the consequences of eco-
environment. However, to this day, the value of these ben- system change for human well-being and provided an
efits, as well as the reduction or loss of benefits due to appraisal of the condition and trends in the world’s eco-
human impacts on the environment, is not fully recognized systems and the services they provide, as well as the
98 PART I Fundamental Concepts

comparable period of time in human history, largely

to meet rapidly growing demands for food, fresh water,
timber, fiber, and fuel due to population growth and
economic development.
2. The changes that have been made to ecosystems have
contributed to substantial net gains in human well-being
and economic development. However, these gains have
been achieved at growing costs in the form of the deg-
radation of many ecosystem services, increased risks of
nonlinear changes, and the exacerbation of poverty for
some groups of people. These problems, unless
addressed, will substantially diminish the benefits that
future generations obtain from ecosystems.
3. The degradation of ecosystem services could grow sig-
nificantly worse during the first half of this century and
is a barrier to achieving the Millennium Development
Goals (see Chapter 28).
4. The challenge of reversing ecosystem degradation while
meeting increasing demands for services can be partially
met under some scenarios considered by the MEA but will
involve significant changes in policies, institutions, and
practices that are not currently under way.
Ecosystem services are classified into four groups—
Provisioning, Regulating, Cultural, and Supporting
(Fig. 6.11). Humans receive both direct and indirect ben-
FIG. 6.8 The terrestrial carbon cycle. From Office of Biological and efits from these services. Provisioning services are material
Environmental Research of the U.S. Department of Energy Office of
and energy resources obtained from ecosystems: food (such
Science. https://science.energy.gov/ber/
as crops, fruit, fish), fuel, building materials, fiber (such as
timber, cotton, and wool), biochemicals (such as natural
scientific basis for action to conserve and use them sus-
medicines, pharmaceuticals), and ornamental resources
tainably. The main findings of the assessment are:
(such as flowers, shells, stone). Regulating services are ben-
1. Over the past 50 years, humans have changed eco- efits obtained from processes carried out by ecosystems:
systems more rapidly and extensively than in any maintenance of air quality, climate and water regulation,

FIG. 6.9 Carbon cycling in the shallow subsurface envi-

ronment. From Office of Biological and Environmental
Research of the U.S. Department of Energy Office of
Science. https://science.energy.gov/ber/
 Ecosystems and Ecosystem Services Chapter 6 99

FIG. 6.10 The nitrogen cycle. From Office of Biological and Environmental Research of the U.S. Department of Energy Office of Science. https://
science.energy.gov/ber/

erosion control, water purification and detoxification, cycling, primary production, production of atmospheric
natural hazard protection, and bioremediation of waste. oxygen, and provision of habitat).
Cultural services are the nonmaterial benefits that people The current status of ecosystems and the services they
obtain from the environment that support physical and provide based on the MEA is presented in Information
mental well-being. These include spiritual enrichment, cog- Box 6.2. Inspection shows that many services are in decline,
nitive development, recreation, art, social relations, esthetic due mostly to human impacts. A primary concept of eco-
values, cultural heritage values, and ecotourism. Supporting system services is that human impacts on ecosystems can
services are the services that are necessary for the pro- impair the condition and functioning of ecosystems, and
duction of all other ecosystem services, in other words, that this impairment can lead to reduced or lost services,
the services that maintain ecosystem health and functioning which in turn can negatively impact human well-being.
(soil formation and retention, water cycling, nutrient In other words, it recognizes that humans are an integral
part of ecosystems and that a dynamic interaction exists
between them and other parts of ecosystems, with human
activities directly and indirectly driving changes in eco-
systems that in turn cause changes in human well-being.
The connection between human well-being and ecosystem
services is illustrated in Fig. 6.12. Illustrations of how
human activities create “drivers for change” of ecosystems,
and how these changes impact services and human well-
being are presented in Fig. 6.13.
Metro nature is a relatively new term that can be defined
as the integration of nature into urban areas or cities (Wolf
and Robbins, 2015). An example of metro nature would be
the iconic Central Park in New York, which is critically
FIG. 6.11 Ecosystem services. Reproduced from http://www.ceeweb.org/ important to the community for recreational and other
work-areas/priority-areas/ecosystem-services/what-are-ecosystem-services/.
100 PART I Fundamental Concepts

INFORMATION BOX 6.2 Status of Ecosystems and Ecosystem Services

From the Millennium Ecosystem Assessment (World Resources Institute (WRI), 2005. Millennium ecosystem assessment, eco-
systems and human well-being: synthesis. Island Press, Washington, DC.
Legend: ▲ Enhanced, . Degraded, +/ Both.
Status indicates whether the condition of the service globally has been enhanced (if the productive capacity of the service has
been increased, for example) or degraded in the recent past. Definitions of “enhanced” and “degraded” are provided in the following
note. A fourth category, supporting services, is not included here as they are not used directly by people.
Service Subcategory Status Notes

Provisioning Services
Food Crops ▲ Substantial production increase
Livestock ▲ Substantial production increase
Capture . Declining production due to overharvest
Fisheries
Aquaculture ▲ Substantial production increase
Wild foods . Declining production
Fiber Timber +/ Forest loss in some regions, growth in others
Cotton, +/ Declining production of some fibers, growth in others
hemp, silk
Wood fuel . Declining production
Genetic resources . Lost through extinction and crop genetic resource loss
Biochemicals, natural medicines, pharmaceuticals . Lost through extinction, overharvest
Fresh water . Unsustainable use for drinking, industry, and irrigation; amount
of hydro energy unchanged, but dams increase the ability to use
that energy
Regulating Services
Air quality regulation . Decline in ability of atmosphere to cleanse itself
Climate regulation Global ▲ Net source of carbon sequestration since mid-century
Regional and . Preponderance of negative impacts
local
Water regulation +/ Varies depending on ecosystem change and location
Erosion regulation . Increased soil degradation
Water purification and waste treatment . Declining water quality
Disease regulation +/ Varies depending on ecosystem change
Pest regulation . Natural control degraded through pesticide use
a
Pollination . Apparent global decline in abundance of pollinators
Natural hazard regulation . Loss of natural buffers (wetlands, mangroves)
Cultural Services
Spiritual and religious values . Rapid decline in sacred groves and species
Esthetic values . Decline in quantity and quality of natural lands
Recreation and ecotourism +/ More areas accessible but many degraded
a
Indicates low to medium certainty. All other trends are medium to high certainty.
Note: For provisioning services, we define enhancement to mean increased production of the service through changes in area over which the service is
provided (e.g., spread of agriculture) or increased production per unit area. We judge the production to be degraded if the current use exceeds sustainable
levels. For regulating and supporting services, enhancement refers to a change in the service that leads to greater benefits for people (e.g., the service of
disease regulation could be improved by eradication of a vector known to transmit a disease to people). Degradation of regulating and supporting services
means a reduction in the benefits obtained from the service, either through a change in the service (e.g., mangrove loss reducing the storm protection
benefits of an ecosystem) or through human pressures on the service exceeding its limits (e.g., excessive pollution exceeding the capability of ecosystems to
maintain water quality). For cultural services, enhancement refers to a change in the ecosystem features that increase the cultural (recreational, esthetic,
spiritual, etc.) benefits provided by the ecosystem.

activities (Fig. 6.14). Potential benefits of exposure to services they provide. This can happen via pollution events,
nature are presented in Table 6.1. The concept of metro impairment or destruction of habitat, or by overuse of
nature is an example of applying ecosystem services con- resources. Methods to restore ecosystems and thus eco-
cepts to a specific case. system services are discussed in Chapter 20.
Soil is a central component of the environment. It pro- The Tragedy of the Commons is a concept related to
vides many critical services to humans. However, it can also overuse of resources provided by the environment, that
have deleterious impacts on humans, affecting their health is, an ecosystem service. Central to the concept is the idea
and well-being. The complex interactions between soil that individuals will put their own personal interests above
health and human health are discussed in Chapter 27. those of the common good (the general public). Thus when
As will be discussed in several upcoming chapters in multiple individuals share a common resource, each will
this text, many human activities can disturb ecosystems, attempt to maximize their use of the resource and the ben-
disrupting their functioning and thereby reducing the efits they receive. However, the cumulative impact of these
 Constituents of well-being
Ecosystem services Security
Personal safety
Provisioning Secure resource access
Food Security from disasters
Freshwater
Wood and fiber
Fuel Basic material
... Freedoms
for good life
Adequate livelihoods of choices
Supporting Regulating Sufficient food and action
Climate regulation Shelter
Nutrient cycling Access to goods Opportunity to be
Soil formation Flood regulation able to achieve
Disease regulation
Primary production what an individual
... Water purification
... Health values doing
Strength and being
Feeling well
Cultural Access to clean air
Aesthetic and water
Spiritual
Educational
Recreational Social relations
... Social cohesion
Mutual respect
Ability to help others
Arrow’s color Arrow’s width
Potential for mediation by Intensity of linkages between ecosystem Source: Millennium Ecosystem Assessment
socio-economic factors services and human well-being
Low Weak
Medium Medium

High Strong

FIG. 6.12 The connections between ecosystem services and human well-being. From the Millennium Ecosystems Assessment report, World Resources
Institute (WRI), 2005. Millennium Ecosystem Assessment, Ecosystems and Human Well-Being: Synthesis. Island Press, Washington, DC.

Short-term
Global
Long-term

Regional

Local

Human well-being Indirect drivers of change

and poverty reduction Demographic
Basic material for a good life
Economic (e.g., globalization, trade,
market, and policy framework)
Health
Sociopolitical (e.g., govemance,
Good social relations
institutional and legal framework)
Security
Science and technology
Freedom of choice and action
Cultural and religious (e.g., beliefs,
consumption choices)

Ecosystem services Direct drivers of change

Provisioning Changes in local land use and cover
(e.g., food, water, fiber, and fuel)
Species introduction or removal
Regulating Technology adaptation and use
(e.g., climate regulation, water, and disease)
External inputs (e.g., fertilizer use,
Cultural pest control, and irrigation)
(e.g., spiritual, aesthetic, recreation, Harvest and resource consumption
and education)
Climate change
Supporting
Natural, physical, and biological
(e.g., primary production, and soil formation)
drivers (e.G., evolution, volcanoes)

Life on earth - biodiversity

Strategies and interventions Source:Millennium Ecosystem Assessment

FIG. 6.13 The connections between drivers for change in ecosystems and the impacts on services and human well-being. From the Millennium Eco-
systems Assessment report, World Resources Institute (WRI), 2005. Millennium Ecosystem Assessment, Ecosystems and Human Well-Being: Synthesis.
Island Press, Washington, DC.
102 PART I Fundamental Concepts

FIG. 6.14 An example of “metro nature,” the iconic Central Park in New York. Reproduced from Jean-Christophe BENOIST [CC BY 3.0 (https://
creativecommons.org/licenses/by/3.0)], from Wikimedia Commons.

sustainable management practices to regulate resource use

TABLE 6.1 Health Benefits From Contact With Nature to preserve the long-term benefits accruing from ecosystem
Health/Well-Being Benefits services. Sustainable management and development con-
cepts are discussed in Chapter 32.
Reduced stress
Better sleep
QUESTIONS AND PROBLEMS
Improved mental health
Reduced depression 1. Define ecosystems and biomes and discuss the
Reduced anxiety differences
Greater happiness, well-being 2. What biome do you live in?
3. Identify some examples of ecosystem services
Reduced aggression
important to you.
Lower blood pressure 4. What is the critical zone?
Improved congestive heart failure 5. Discuss the different types of biodiversity, using a spe-
cific example ecosystem.
Reduced obesity
6. Discuss a specific example of the tragedy of the
Reduced diabetes commons.
Better eyesight 7. Describe the critical components and transfer processes
for the sulfur or phosphorous cycle.
Reduced mortality

Modified from Table 1, Frumkin, H., Bratman, G.N., Breslow, S.J., et al.,
2017. Nature contact and human health: a research agenda. Environ.
REFERENCES
Health Perspect. 125 (7), 075001. doi:10.1289/EHP1663.
Wolf, K.L., Robbins, A.S.T., 2015. Metro nature, environmental health and
economic value. Environ. Health Perspect. 123 (5), 390–398.
World Resources Institute (WRI), 2005. Millennium Ecosystem
individual decisions results in overuse of the resource. Assessment, Ecosystems and Human Well-Being: Synthesis. Island
This in turn ultimately results in a reduction or loss of ben- Press, Washington, DC.
efits for all of the users. This concept illustrates the need for