The relative cost of saving species

Abstract

Although biodiversity conservation underpins socioeco-nomic welfare and human well-being, the links between biodiversity and human well-being are often overlooked. In many cases, conservation goals are seen as being different from human well-being goals or even opposed to develop-ment goals, promoting a trade-off between investing in nature and investing in economic development or social well-being. Nevertheless, in spite of its invisibility in markets, the benefits obtained from conserving biodiversity usually exceed the opportunity costs of its conservation. Moving from biodiversity intrinsic values to instrumental values People's attitudes and preferences are usually the driving force for promoting biodiversity conservation policies. These attitudes and preferences could be related to spiritual, moral, ethical, utilitarian, ecological, or scientific motivations. Therefore, understanding which factors cause changes in people's attitudes and preferences toward biodiversity is essential for designing successful conservation policies. Moreover, understanding people's preferences toward biodi-versity, as well as the underlying factors promoting them, helps identify which type of relationship is established between nature and society (e.g., utilitarian, affective). How people perceive biodiversity determines the choices they make regarding how to conserve it (Gatzweiler 2008). The broad spectrum of attitudes toward species may be grouped under two main conflicting and antagonistic head-ings: (1) affection, which represents people's affective and emotional responses to species, versus (2) utility, which represents people's self-interest related to the instrumental value of species. This model of two primary motivations for biodiversity conservation has been identified in several scientific investigations during the twentieth century; these studies have used various terms in describing this model, including anthropomorphism versus anthropocentrism, affec-tion versus economic self-interest, empathy/identification versus instrumental self-interest, and affection versus utility (see Serpell 2004 for a review).

Full text

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The relative cost of saving species
Although biodiversity conservation underpins socioeco-
nomic welfare and human well-being, the links between
biodiversity and human well-being are often overlooked. In
many cases, conservation goals are seen as being different
from human well-being goals or even opposed to develop-
ment goals, promoting a trade-off between investing in nature
and investing in economic development or social well-being.
Nevertheless, in spite of its invisibility in markets, the benefits
obtained from conserving biodiversity usually exceed the
opportunity costs of its conservation.
Moving from biodiversity intrinsic values to
instrumental values
People’s attitudes and preferences are usually the driving
force for promoting biodiversity conservation policies. These
attitudes and preferences could be related to spiritual, moral,
ethical, utilitarian, ecological, or scientific motivations.
Therefore, understanding which factors cause changes in
people’s attitudes and preferences toward biodiversity is
essential for designing successful conservation policies.
Moreover, understanding people’s preferences toward biodi-
versity, as well as the underlying factors promoting them,
helps identify which type of relationship is established
between nature and society (e.g., utilitarian, affective). How
people perceive biodiversity determines the choices they make
regarding how to conserve it (Gatzweiler 2008).
The broad spectrum of attitudes toward species may be
grouped under two main conflicting and antagonistic head-
ings: (1) affection, which represents people’s affective and
emotional responses to species, versus (2) utility, which
represents people’s self-interest related to the instrumental
value of species. This model of two primary motivations for
biodiversity conservation has been identified in several
scientific investigations during the twentieth century; these
studies have used various terms in describing this model,
including anthropomorphism versus anthropocentrism, affec-
tion versus economic self-interest, empathy/identification
versus instrumental self-interest, and affection versus utility
(see Serpell 2004 for a review).
This attitudinal structure toward species based on two
antagonistic dimensions is rooted in two kinds of human
values—intrinsic value and instrumental value. Intrinsic value
has been defined as the nonhuman species’rights to exist that
cannot be measured in terms of markets; it is strongly related
to the human moral obligation to share the planet (Callicott
1986). In the words of Paul R. Ehrlich and Anne Ehrlich
(1981, 48), “Our fellow passengers on Spaceship Earth, who
are quite possibly our only living companions in the entire
universe, have a right to exist.”Thus, intrinsic value proposes
that biodiversity has a value in itself and is valued as an end in
itself, independent of its usefulness (Brondízio et al. 2010). On
the contrary, instrumental value is a function of utility and
assumes that biodiversity is valuable only as a means or tool
for people to obtain human welfare, satisfaction, or happiness.
Instrumental value lies not in the object itself but in the uses
or potential uses it gives or could give.
This dichotomy of values constitutes the core of modern
environmentalism in Western culture and has guided the
conservation debate in the last decades (Pascual et al. 2010). In
fact, the conservation debate in modern Western societies has
moved away from biocentric ethics, which refers to promoting
conservation based on the idea of any theory that considers all
live beings as possessing intrinsic value, to anthropocentric
arguments, which consider that biodiversity conservation
should be supported because it is a tool to enhance human
quality of life, satisfying as many individual preferences,
desires, and needs as possible (such as the importance of many
plants for medicinal compounds, a plant’s ornamental value,
or the satisfaction that people feel just knowing that
biodiversity will exist in the future). Because the way people
relate to nature is reflected not only by their behavior, but also
by the rules they apply to articulate values for nature
(Brondízio et al. 2010), this conservation debate actually
represents how people perceive human–nature relationships
and support different biodiversity conservation policies.
As J. Baird Callicott claimed in a 1986 article, a biodiversity
conservation movement based on intrinsic values (or on
biocentric ethics) is ultimately emotional and derived from
feelings and affects toward nonhuman living species and is one
in which all of humanity feels responsible. Consequently, the
intrinsic value of species could be determined by how they
make humans feel, which depends ultimately on certain
factors that promote human affections toward species. Several
studies have demonstrated that humans devote attention to,
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and feel affection for, those species that are phylogenetically
close to them and to those that are physically or behaviorally
similar to them (i.e., vertebrates, mostly mammals). In
addition to those factors, Konrad Lorenz showed in a 1971
study that humans have an innate tendency to feel affection
for nonhuman species with human neonates’physical features,
such as large eyes, large rounded forehead, or short nose.
If intrinsic values (and the underlying factors determining
them) have mostly been responsible for the biodiversity
conservation policies of the late twentieth and early twenty-
first centuries, does this mean that conservation policies have
been biased toward those species that evoke positive affections
and emotions in humans? Scientific research has demonstrat-
ed that, in fact, likability factors, such as different physical
traits (e.g., body size, body shape, eye size), as well as whether
the species is phylogenetically close to humans, play an
important role in defining the conservation policies adopted
for a species. As a consequence, the majority of species (i.e.,
invertebrates, plants, fungi, and microorganisms), which are
key to maintaining ecological functions as well as sustaining
life, are underrepresented in conservation priority-setting
schemes. Biodiversity conservation policies based exclusively
on the intrinsic values of nonhuman species trigger people to
desire to protect those higher life-forms (i.e., mammals and
birds) that produce positive emotions and feelings.
In order to counteract this “vertebrate chauvinism”in
biodiversity conservation strategies (so named by Thomas E.
Lovejoy; see Kellert 1986), a pragmatic movement has
appeared under an anthropocentric utilitarianism lens. Conser-
vation arguments based on the benefits society obtains from
nature have exponentially increased in scientific literature
under the rubric of the term ecosystem services (see Figure 1).
In fact, this notion has involved the development of new
conceptual and analytical frameworks for understanding
human-nature relationships, such as the Millennium
Ecosystem Assessment, The Economics of Ecosystems and
Biodiversity, and the Intergovernmental Platform on Biodi-
versity and Ecosystem Services.
In this debate, the main issue is whether these two
approaches to species conservation are complementary or
opposed. The question here focuses again on the society–
nature relations model. If one considers humanity’s relation
with nature to be innate to human evolutionary history—the
biophilia hypothesis (Kellert and Wilson 1993)—then one
should conclude that all humans have an innate need to
connect with nature from both dimensions: ethical or
emotional and utilitarian (Brondízio et al. 2010). Under this
relations model, biocentric and anthropocentric arguments
should coexist in biodiversity conservation strategies. From
this point of view, justifying conservation exclusively on the
basis of ethical considerations about the right of species to
exist (i.e., intrinsic value) ignores an important motivation that
people have for preserving species: the importance of
biodiversity as a source of human well-being through the
delivery of ecosystem services (i.e., instrumental value). In
fact, the Millennium Ecosystem Assessment (2005) acknowl-
edges that people can make decisions concerning biodiversity
based on their own and others’well-being as well as based on
ethical concerns about species, thus recognizing instrumental
and intrinsic values.
Conceptualizing biodiversity as a source of
ecosystem services
There is a growing awareness of the importance of
biodiversity for human well-being through the delivery of
ecosystem services to society (see Figure 2). Ecosystem services
have been defined as the direct and indirect contributions of
ecosystems to human well-being (Wittmer and Gundimeda
2010). They have been classified into three main categories:
(1) provisioning services, such as food, fiber, raw materials,
resources for medicine, and fuel; (2) regulating services, such
as climate regulation, air purification, flood control, water
purification, erosion control, renewal of soil fertility, pollina-
tion, and pest control; and (3) cultural services, such as
recreational activities and ecotourism, aesthetic values, spiri-
tual values, and scientific and local ecological knowledge (see
Table 1). In addition to the inherent importance of
biodiversity for most of the provisioning services, such as
food, medicines, and fibers, biodiversity influences recrea-
tional activities (e.g., recreational fishing) and the aesthetic
experiences of people, promoting psychological well-being;
moreover, biodiversity plays a key role in sustaining a number
of regulating services, including water purification, flood
buffering, global climate and microclimate regulation, air
purification, soil formation, and pollination.
Although all components of biodiversity, from genotypes
to ecological communities, are essential to the delivery of
ecosystem services, there is ample evidence supporting the
positive and key effect of functional diversity on delivering
many ecosystem services (Díaz et al. 2006), particularly
regulating services (see Figure 2). Here, functional groups
are understood as those that perform particular ecosystem
functions (e.g., nitrogen fixation, seed dispersal, biomass
0
Publication year
Number of references to ecosystem
services per year
Number of total references to ecosystem
services
1992
2011
2010
2009
2007
2008
2006
2005
2004
2003
2002
2001
2000
1999
1998
1997
1996
1995
1994
100
200
300
400
500
600
700
800
900
0
3500
3000
2500
2000
1500
1000
500
TEEB
IPBES
MA
References to ecosystem
services per year Total references to ecosystem
services
Figure 1. Number of publications using the term of “ecosystem
services”between the period 1990–2011 years (search carried out in
ISI Web of Science in March 2012). Reproduced by permission of Gale, a
part of Cengage Learning.
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decomposition, biomass production) through different physi-
ological, structural, behavioral, or phenological characteristics
or traits (e.g., root type, leaf texture, plant height). In addition,
the diversity of species and populations within a functional
group—that is, its amount of functional redundancy—is the
main biological component that helps to preserve ecosystem
services (Elmqvist et al. 2003). In fact, the scientific evidence
suggests that the relationship between biodiversity and
ecosystem services supply is mostly based on functional
diversity and functional redundancy, rather than on species
richness.
Reviews probing for evidence of links between biodiversity
and the delivery of ecosystem services have shown that the
functional role of microorganisms, vegetation, and inverte-
brates is the component of biodiversity that most influences
ecosystem service supply. Table 1 shows, in a simplified way,
the links between biodiversity components and the delivery of
provisioning and regulating services. Cultural services are not
considered in Table 1 because they are generally coproduced
by a deeper relationship between people and ecosystems. In
fact, most cultural services (e.g., recreational activities,
aesthetic enjoyment, spiritual values) require people’s experi-
ences in nature. That is, only people who visit nature will
benefit from cultural services through experiential enjoyment.
A number of scientific studies have reviewed the role of
functional diversity and species in the delivery of ecosystem
services (Kremen 2005; Balvanera et al. 2006; Díaz et al. 2006;
Luck et al. 2009; Cardinale et al. 2012). As previously
mentioned, however, little information exists about the state
and tendencies of the key functional groups that provide
services to society, despite the essential role they play either in
ecological functioning or in the delivery of ecosystem services.
Focusing more scientific and political attention on these
overlooked functional groups is an undertaking of great
importance for the implementation of future biodiversity
conservation strategies.
The social and economic importance
of biodiversity
Regarding the social dimension, biodiversity conservation
has been considered an important tool for achieving poverty
reduction and social justice because priority areas of
conservation seem to be efficient targets for maintaining
human well-being through the services they deliver (Turner
et al. 2012). Emerging evidence for this has been found by the
World Bank, which estimated that ecosystem services directly
support more than one billion people living in poverty (World
Bank 2011). Similarly, a 2010 study carried out in Costa Rica
and Thailand by Kwaw S. Andam and colleagues has shown
that those districts with protected areas experienced between
10 percent and 30 percent less poverty. In addition, WWF
International and the International Union for Conservation of
Nature offered some different examples in two reports—
Safety Net: Protected Areas and Poverty Reduction (Dudley et al.
2008) and Can Protected Areas Contribute to Poverty Reduction?
Opportunities and Limitations (Scherl et al. 2004), respective-
ly—in which they illustrated how protected areas mostly
contribute to the wider aspects of human well-being (i.e.,
promoting basic material resources for living, increasing
education rates, enhancing good social relationships in those
cases of comanagement) rather than to poverty reduction per
se (i.e., increasing per capita income). In fact, biodiversity
conservation directly contributes to poverty reduction in at
Ecosystem services
Ecosystems
Indirect drivers of change Human well-being of the
ecosystem service beneciaries
Provisioning
Cultural
Regulating
• Genotypes
• Species
• Communities
• Functional diversity
Direct drivers of change
• Land-use changes
• Climate change
• Over exploitation
• Pollution
• Invasive alien species
• Basic materials for a good life
• Health
• Good social relationships
• Security
• Freedom of choice and action
Biodiversity
Figure 2. Complex links between biodiversity and human wellbeing.On one
hand, biodiversity is affected by indirect and direct drivers of change but,
on the other hand, biodiversity influence on human wellbeing through the
delivery of ecosystem services. Among all the ecosystem services,
regulating services are the base for the maintenance of others and
provisioning services depend on both regulating, which are strongly related
to ecosystem’s integrity and those cultural services related to knowledge
(local ecological knowledge and scientific knowledge). Reproduced by
permission of Gale, a part of Cengage Learning.
Ecosystem services
Organizational level
at which biodiversity
is involved
Main taxonomic groups
involved
Provisioning
Food Genes, species
populations,
communities
Mainly vegetation (plant crops,
wild fruits, etc.), fish, birds, and
mammals. In specific cases,
fungi, invertebrates, and
other vertebrates
Fibers Species populations,
communities,
functional groups
Vegetation and vertebrates
Medicine Genes, species
populations
Microorganisms, fungi,
vegetation, and animals
Regulating
Air purification Species populations,
functional groups
Microorganisms and vegetation
Water purification Communities,
functional groups
Microorganisms, vegetation,
and aquatic invertebrates
Hydrological regulation,
erosion control and
flood mitigation
Species populations,
communities,
functional groups
Vegetation
Renewal of soil fertility Communities,
functional groups
Soil microorganisms, nitrogen-
fixing plants, soil invertebrates,
and waste products of animals
Pollination Species populations,
functional groups
Insects, birds and mammals
Table 1. Links between ecosystem services delivery and biodiversity
(organizational level of biodiversity and main taxonomic groups
involved). Reproduced by permission of Gale, a part of Cengage Learning.
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least five ways: food security, health improvements, income
generation, reduced vulnerability for human populations, and
ecosystem services delivery. Consequently, protecting biodi-
versity conservation priority areas delivers direct benefits to
society and alleviates poverty (Turner et al. 2012).
Regarding the economic dimension, the contributions of
biodiversity to human well-being have major economic
significance, even though their value is not recognized by
conventional markets. The economic value of biodiversity can
be divided into insurance value and output value (Pascual et al.
2010; see Figure 3). Insurance value refers to the ecosystem’s
capacity to maintain the delivery of ecosystem services to
society in the face of disturbance. Output value is obtained
from the aggregation of the ecosystem service benefits and
usually is recognized as total economic value (TEV).
The components of TEV usually are represented using a
value taxonomy (see Figure 4). The main distinction made
is between use and nonuse values (Turner et al. 2008). Use
values include direct use,indirect use,andoption values.Direct
use values are derived from the conscious use and
enjoyment of ecosystem services by individuals. They may
be extractive, such as agriculture or fishing, or they may be
nonextractive, such as recreational activities, nature tourism,
and aesthetical enjoyment of landscapes. Hence, extractive
direct use values are strongly related to provisioning
services, whereas nonextractive direct use values are
strongly related to cultural services. Indirect use values are
associated with the regulating services that are provided
by biodiversity and ecosystems, without entailing conscious-
ness in their use. Finally, option values are related to the
future direct and indirect uses of biodiversity by individuals.
Thesevaluesarerelatedtotheindividualsatisfaction
derived from ensuring that an ecosystem service will be
available for the future.
Nonuse values reflect the value that arises from people’s
feelings attributable to the existence of biodiversity and
ecosystem services, either related to the satisfaction that
individuals derive from the knowledge that an ecosystem
service will be available to future generations (bequest value)
or related to the individual satisfaction obtained from the
knowledge that a species, species population, community,
landscape, or ecosystem service continues to exist (existence
value). Among nonuse values, some authors also consider
altruist values, which are related to the satisfaction derived
from ensuring that biodiversity and ecosystem services are
available to other people in the current generation. Hence,
whereas bequest values are related to intergenerational equity
concerns, altruist values are related to intragenerational
concerns (Pascual et al. 2010). In addition, because nonuse
values are derived from human satisfaction related to moral,
ethical, or religious issues, they are connected to particular
cultural services, such as spiritual values. Thus, the boundaries
of nonuse values are not clear-cut because of their strong
relationship with different moral motivations—that is, a
person’s motivation to support biodiversity conservation is
composed of intertwined value types and not just one value
type. In fact, the existence and bequest values are sometimes
considered a way to capture the intrinsic values that exist
under the ecosystem service framework.
In practical terms, the valuation of TEV is estimated by the
aggregation of the different ecosystem service values provided
by a particular ecosystem that are feasible to quantify. The
estimation of the TEV of biodiversity and ecosystem services
in a particular area or ecosystem should help to assess the
contributions of these services to the economy of the region.
The most interesting example illustrating the contributions of
biodiversity to economic or financial value is the case of the
Catskill watershed, the main source of water for New York
City. The citizens of New York avoided spending $6 billion to
$8 billion on the construction of a water filtration plant by
instead investing $1.5 billion to buy lands in the upper
watershed to restore and conserve them (Chichilnisky and
Heal 1998). From a scientific point of view, one of the first
attempts to estimate the value of ecosystem services on a
global basis was a 1997 study by Robert Costanza and
colleagues encompassing seventeen ecosystem services deliv-
ered by sixteen biomes around the world. These researchers
estimated that the value of ecosystem services ranged from
$16 trillion to $54 trillion per year, with an average annual
value of over $33 trillion, a figure higher than the annual
global gross domestic product. In a 2007 study, Will R.
Turner and colleagues demonstrated that the areas identified
a decade earlier by Costanza and colleagues as priority areas
Insurance value
Biodiversity
Functions
The capacity of
ecosystems to
supply services
(e.g., CO2
sequestration)
Resilience
Ecosystem services
Provisioning
(e.g., food)
Regulating
(e.g., favorable clima)
Cultural
(e.g., recreational
activities)
Benets
Basic material for
a good life
(e.g., food)
Health
(e.g., favorable clima
and psychological
well-being)
Output value
Structure
Functioning
Figure 3. The “ecosystem service cascade”framework, which assess
links between the insurance value and the output value or Total
Economic Value (TEV). Reproduced by permission of Gale, a part of
Cengage Learning.
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because of the high economic value of their ecosystem
services spatially overlap with high (terrestrial) biodiversity
priority areas. This means that a general positive concor-
dance exists between biodiversity conservation priority areas
(i.e., areas with a high diversity of species) and areas with
high ecosystem services value. Among all terrestrial areas,
Turner and colleagues found the following areas of high
concordance: Congo, the Amazon, central Chile, the
Western Ghats region of India, and parts of Southeast Asia.
Consequently, this study shows that tropical forests offer the
best opportunities for synergies between conserving biodi-
versity because of their intrinsic value (high biodiversity) and
their instrumental value (high ecosystem service value). In
fact, sustainable strategies of tropical forest management
show a positive benefit-cost ratio—that is, net marginal
benefits—derived from not only timber but also non-timber
forest products, water supply, hydrological regulation, flood
prevention, carbon sequestration, and the preservation of
habitat for endangered species. Unsustainable management,
by contrast, was associated with higher private benefits
through timber harvesting or conversion to agriculture lands
but reduced social benefits because of the degradation of
regulating and cultural services. Unfortunately, estimates of
land-use changes in forests reveal the highest losses in
tropical forests, specifically high economic losses. For
example, the economic losses expected in Brazil between
the years 2000 and 2050 as a result of the loss of tropical
forests range from 2 percent to 6 percent of the country’s
2050 gross domestic product ($3.6 billion to $11 billion in
2050 dollars) (Chiabai et al. 2011). These economic losses
are underestimated because they include only the degrada-
tion of wood and non-wood forest products (provisioning
services), carbon sequestration as regulating service, and the
cultural services of ecotourism and existence value. For a
review of the economic losses derived from unsustainable
management of forests, see the 2011 study by Aline Chiabai
and colleagues.
One of the main contributors to the study of the
economics of biodiversity in the early twenty-first century
has been the international project called the Economics of
Ecosystems and Biodiversity (Wittmer and Gundimeda
2010). This project, which quantifies the costs of biodiversity
loss and ecosystem services degradation, has analyzed seven
different scientific studies that estimate the TEV of
sustainable management practices in different ecosystems
andcomparedthefigurestothose arising from management
regimes involving land-use intensification or unsustainable
practices. Overall, this comparative analysis demonstrates
that the global benefits of biodiversity conservation are about
250 percent greater than the benefits from conversion. Thus,
the economic value of managing the ecosystem more
sustainably (in all of the cases) is higher than the economic
value of converting the ecosystem or managing it unsustain-
ably, even though the private benefits of the provisioning
services captured in current markets (e.g., timber, aquacul-
ture, agriculture) would favor conversion or unsustainable
practices.
The following three studies, which were discussed in a
2002 article by Andrew Balmford and colleagues, offer a
Use values
Option values
(future direct and
indirect uses)
• Extractive uses (e.g.,
fishing or agriculture)
• Non extractive uses
(e.g., recreation or
aesthetics)
Uses derived from
regulating services
(e.g., pollination or
flood protection)
Satisfaction that the
ecosystem services
are available to
future generations
Satisfaction of
continued existence
of biodiversity and
ecosystem services
Bequest
values
Direct use
values
Indirect use
values
Existence
values
Non use values
Total Economic Value (TEV)
Decreasing tangibility to individuals
Figure 4. Components of the Total Economic Value (TEV) of biodiversity and ecosystems (Turner et al. 2008). Reproduced by permission of Gale, a part
of Cengage Learning.
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comparison between the net benefits of conserving ecosystems
and the net benefits of converting them:
1. A study carried out in a tropical forest of Mount
Cameroon estimated that the social benefits obtained
from low-impact logging practices, such as non-
timber forest products, flood prevention, carbon
sequestration, and nonuse values, amount to $1,376
per acre. However, the conversion of forests to small-
scale agriculture for food yielded the greatest private
benefits, about $809 per acre.
2. Research in Thailand to study which management
practice is more profitable—conserving mangroves
or converting the land to aquaculture—demonstrates
that the net marginal benefits of conserving man-
groves exceed at least $324 per acre. The estimated
benefits of mangrove ecosystems, which include
timber, charcoal, non-wood forest products, offshore
fisheries, and storm protection, yield a minimum of
$405 and a maximum of $14,569 per acre. In
contrast, the conversion to shrimp farming in order
to provide food generates benefits of only about $81
per acre.
3. Draining freshwater marshes in agricultural areas in
Canada involves a total economic loss of $1,376 per
acre. The social benefits of marshes are related to
provisioning services, such as hunting; regulating
services, such as hydrological regulation; and
cultural services, such as recreational activities.
The TEV value estimated for these services was
on average $2,347 per acre, whereas the value of
land converted from wetlands to agriculture totaled
$971 per acre.
In these three cases, the average economic losses attribut-
able to conversion or to unsustainable practices amounted to
54.9 percent of the TEV of the sustainable ecosystems. These
three studies provide good examples of cases in which the
conversion of ecosystems for forestry, aquaculture, or
agriculture does not make sense from either an environmental
viewpoint or a socioeconomic perspective.
The value of species conservation
As species populations are becoming increasingly more
threatened worldwide because of the effects of drivers of
change (see Figure 2), it is important to acknowledge an
important consideration—the willingness of society to
conserve these species. One of the most commonly used
measures of people’s awareness of species conservation is
their willingness to pay (WTP) for the preservation of a
particular species. Several studies have found that indivi-
duals are willing to pay a small portion of their income
toward the aim of conserving endangered and rare species
(Richardson and Loomis 2009). The WTP for conserving
species could be related to a variety of reasons, such as
recreational use (direct use value), pollination or pest
control (indirect use values), bequest values, or mostly
existence values.
Although this indicator has been frequently used by
scientists since the early 1980s, its use is clearly on the rise
in the twenty-first century. In a 2008 article, Berta Martín-
López and colleagues reviewed 60 studies, published from
1980 to 2005, measuring the WTP for conserving species (see
Figure 5a). Overall, scientists have tended to focus on
estimating WTP for protecting vertebrates, mainly mammals
and birds (see Figure 5b). The economic valuation of species
has also focused on studying those species that live in marine
and forest ecosystems, whereas species that live in drylands
and human-made habitats have been understudied. In
addition, the economic studies have focused (among all the
components of TEV) on nonuse values, mostly existence
values (see Figure 5c). Whereas direct use values, mainly
recreational activities, generate the highest economic value for
species protection ($54.4 per person in 2005 dollars), indirect
use values generate the lowest ($33.9 per person). It seems,
however, that people refer to both use and nonuse values in
determining their WTP for biodiversity conservation. This
indicates that economic valuations of species should explore
all components of TEV rather than individual elements. In
general, people hold a higher WTP for protecting mammals
and birds, in comparison with reptiles and crustacean, which
are less valued (see Figure 5d).
It is interesting to note that people around the world are
highly willing to pay for big species (both in length and
weight) with relative big eyes (see Figure 6). Consequently,
the economic value of species conservation obtained through
this technique is biased not only toward mammals and birds
but also toward those species with neonates’physical
characteristics, as Konrad Lorenz showed in a 1971 study.
This indicates that people’s motivations that underlie
existence values (i.e., phylogenetic and physical similarity)
also explain society’s prioritization regarding species conser-
vation.
In addition, WTP experiments for species conservation
show that visitors usually allocate higher values to species
protection than local stakeholders, because visitors are likely
to have a recreational component to their WTP and are likely
to be more knowledgeable about biodiversity. Reaching the
opposite conclusion, however, was a 2011 study by Adriana
Ressurreição and colleagues that centered on marine biodi-
versity in southern Europe. When comparing residents and
visitors with similar socioeconomic profiles, local residents
were found to be more likely to attach higher values to species
protection than visitors. This study also demonstrated that
people are able to perceive the importance of biodiversity
conservation in wider terms than the single-species approach
and that they therefore attach greater economic benefits to
the conservation of entire ecosystems rather than partial
conservation plans of particular species.
To give another illustration of the economic contributions
of species to human welfare, Justin G. Boyles and colleagues
in a 2011 study found that insectivorous bats provided pest
control services worth more than $3.7 billion annually to the
U.S. agricultural sector. This figure represents the cost of
pesticide applications that the agricultural industry does not
have to make because bats eat so many insects.
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Similarly, John E. Losey and Mace Vaughan, in a 2006
study, estimated the economic value of insects in the
United States through an analysis of dung burial, pest
control, and pollination regulating services, as well as the
role of insects in wildlife nutrition, in order to estimate
their contribution to nature tourism and recreational
activities. These authors estimated that the value of such
ecosystem services provided by insects totaled almost $60
billion per year. The annual contribution of dung beetles
with regard to decomposing cattle waste was estimated at
$380 million. Regarding pollination, one-third of human
food comes from plants pollinated by wild pollinators, and
native insect pollinators were found to be responsible for
almost $3.7 billion of fruits and vegetables produced in the
United States per year. The value of pest control by native
insects was estimated at approximately $4.5 billion per year.
Finally, the key role of insects as a food source for much of
the wildlife involved in hunting, fishing, and bird-watching
activities, was estimated to be valued at $49.9 billion
annually. Despite the clear benefits they supply, insects are
pitifully underrepresented in conservation policies and
conservation programs involving insects are severely under-
funded.
Economic benefits of biodiversity versus
conservation costs
Based on all the estimates of the economic value of
ecosystems, species, or the ecosystem services provided by
biodiversity, increased investment in biodiversity conservation
is justified. According to a 2002 study by Balmford and
colleagues, expanding the biodiversity conservation area
network to cover 15 percent of the landscape and 30 percent
of the seascape would cost approximately $45 billion per year.
In this case, the costs of biodiversity conservation include five
main components: acquisition, management, damage, trans-
action, and opportunity (see Table 2). The ecosystems within
that network would supply a diverse flow of ecosystem
services with net annual benefits greater than $4.4 trillion per
year. Consequently, investing in biodiversity conservation
would help to maintain global ecosystem services that are
(b)
Fish
11%
Reptile
2%
Crustacean
1%
Mammal
63%
Bird
23%
0
Mammal Bird Reptile Fish Crustacean
10
20
Mean WTP ($ 2005)
30
40
50
60
70
43.4 44.5
37.6
17.0
24.9
(d)
Use 13%
(c)
Direct use
84%
Option
11%
Indirect use
5%
Non use
87%
Year
00
10
20
30
40
50
60
70
Before
1980
1981–
1985
1986–
1990
1991–
1995
1996–
2000
2001–
2005
5
Number of economic valuation studies
Number of accumulative studies
10
15
20
(a)
Accumulative studies
Figure 5. (a) Historical trend of publications studying the individuals’willingness to pay (WTP) for species conservation. (b) Distribution of publications
for species conservation among taxonomic groups. (c) Distribution of publications for species conservation among components of the Total Economic
Value (TEV). (d) Average economic value (WTP per person) per taxonomic group. Reproduced by permission of Gale, a part of Cengage Learning.
Grzimek’s Animal Life Encyclopedia 863
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worth about 100 times more than the costs of designating and
managing the conservation network. Therefore, by spending
$1 on biodiversity conservation, humanity will reap $100 in
benefits because of all the ecosystem services provided by
biodiversity. Although this conclusion should be regarded as
indicative rather than precise because of the assumptions,
generalizations, and various findings of different valuation
studies, it seems clear that the benefits of biodiversity
conservation do outweigh its conservation costs.
Nevertheless, the economic benefits and costs of biodiver-
sity conservation through the use of protected areas vary
significantly depending on the spatial scale (Wittmer and
Gundimeda 2010). At the global scale, the economic benefits
of conservation are more important than its costs. At the local
scale, however, the situation is ambiguous because sometimes
benefits outweigh the costs and other times the opposite holds
true. Although the management costs of protected areas are
usually placed at the national or international levels, the costs
of lost access to natural resources, costs related to wildlife
conflicts, and opportunity costs stemming from forgoing
conversions to nonnatural systems (that produce high
commercial value) are often placed at the local scale.
Protected areas, however, provide several benefits at the local
level:
1. the supply of clean water, particularly from moun-
tains and well-managed natural forests
2. the maintenance of food security because protected
areas act as an habitat for insects and other species
that provide pollination and pest control, and the
maintenance of the subsistence economy through
game, fish, wild fruits, and natural medicines
3. a reduction in risks from unpredictable events and
natural hazards, such as landslides or floods being
mitigated by soil stabilization or hydrological regulation
4. the support of nature tourism, benefiting local people
5. assistance in protecting cultural heritage and spiritual
values such as traditional practices and sacred places
For all these services, there is evidence that local economic
benefits provided by ecosystems in protected areas exceed the
costs of conservation. In summary, the overall magnitude of
the global benefits of biodiversity conservation, which was
estimated at $33 trillion annually in 1997 by Robert Costanza
and colleagues, suggests that the economic benefits clearly
exceed the costs of conservation, which Alexander N. James
and colleagues estimated at only $6 billion per year in 1999.
Beyond 2020 biodiversity targets
In spite of efforts to improve biodiversity conservation
based on higher investments, biodiversity continues to decline
and the main target for 2010—that is, to halt the decline of
biodiversity—were not reached (Butchart et al. 2010). The
economics of biodiversity has been given a key role to play in
the strategies outlined to achieve the 2020 targets: (1) to deal
with the underlying causes of biodiversity loss through
(b)
–1
0,0
Log mean WTP ($2005)
Log (species weight)
0,5
1,0
1,5
2,0
2,5
1357
(a)
1
0,0
Log mean WTP ($2005)
Log (species height)
0,5
1,0
1,5
2,0
2,5
234
(c)
0
0
Log mean WTP ($2005)
Log (eye size of species)
0,5
1
1,5
2,0
2,5
0,2 0,4 0,6
Figure 6. Correlations between average WTP ($2005) for conserving
species and species’physical traits: (a) length, (b) weight, and (c)
relative eye size. Reproduced by permission of Gale, a part of Cengage
Learning.
Table 2. Different types of conservation costs. Reproduced by
permission of Gale, a part of Cengage Learning.
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mainstreaming biodiversity across society; (2) to diminish the
pressures on biodiversity, (3) to improve the state of
biodiversity by protecting ecosystems, species, and genetic
diversity, (4) to enhance the benefits obtained from biodiver-
sity and ecosystem services, and (5) to implement conservation
strategies based on participatory planning, knowledge man-
agement, and capacity building. In this sense, it has been
recommended that the economic values of biodiversity should
be integrated into national accounts and environmental
decision-making. It should be noted, however, that capturing
the economic values of biodiversity in terms of markets—
sometimes by creating new markets, through such initiatives
as wetland banking, endangered species credits, and payments
for ecosystem services—could obscure other dimensions of
biodiversity value, including intrinsic, ecological, and socio-
cultural values. Consequently, while the economic criteria of
biodiversity conservation (i.e., that the economic benefits of
biodiversity exceed the costs of conservation) is essential to
designing conservation policies, it is also important to keep in
mind that (1) biodiversity conservation is a combination of
intrinsic and instrumental values; (2) within instrumental
values, biodiversity conservation should be viewed as a diverse
flow of ecosystem services in order to promote sustainable
land-use management and avoid social conflicts; and (3)
criteria of social equity, of either an intragenerational and
intergenerational nature, should be considered.
Resources
Books
Brondízio, Eduardo S., Franz W. Gatzweiler, Christos Zografos,
and Manasi Kumar. “The Socio-Cultural Context of Ecosys-
tem and Biodiversity Valuation.”In The Economics of Ecosystems
and Biodiversity: Ecological and Economic Foundations, edited by
Pushpam Kumar, 149–181. London: Earthscan, 2010.
Callicott, J. Baird. “On the Intrinsic Value of Nonhuman
Species.”In The Preservation of Species: The Value of Biological
Diversity, edited by Bryan G. Norton, 138–172. Princeton, NJ:
Princeton University Press, 1986.
Dudley, Nigel, Stephanie Mansourian, Sue Stolton, and Surin
Suksuwan. Safety Net: Protected Areas and Poverty Reduction.
Gland, Switzerland: WWF International, 2008. Accessed July
27, 2012. http://assets.panda.org/downloads/safety_net_final.
pdf
Ehrlich, Paul R., and Anne Ehrlich. Extinction: The Causes and
Consequences of the Disappearance of Species. New York: Random
House, 1981.
Kellert, Stephen R. “Social and Perceptual Factors in the
Preservation of Animal Species.”In The Preservation of Species:
The Value of Biological Diversity, edited by Bryan G. Norton,
50–73. Princeton, NJ: Princeton University Press, 1986.
Kellert, Stephen R., and Edward O. Wilson, eds. The Biophilia
Hypothesis. Washington, DC: Island Press, 1993.
Lorenz, Konrad. Studies in Animal and Human Behaviour. Vol. 2.
Translated by Robert Martin. Cambridge, MA: Harvard
University Press, 1971.
Millennium Ecosystem Assessment. Ecosystems and Human Well-
Being: Biodiversity Synthesis. Washington, DC: World
Resources Institute, 2005. Accessed July 27, 2012. http://www
.maweb.org/documents/document.354.aspx.pdf
Pascual, Unai, Roldan Muradian, Luke Brander, et al. “The
Economics of Valuing Ecosystem Services and Biodiversity.”
In The Economics of Ecosystems and Biodiversity: Ecological and
Economic Foundations, edited by Pushpam Kumar, 183–255.
London: Earthscan, 2010.
Scherl, Lea M., Alison Wilson, Robert Wild, et al. Can Protected
Areas Contribute to Poverty Reduction? Opportunities and
Limitations. Gland, Switzerland: International Union for
Conservation of Nature, 2004. Accessed July 27, 2012. http://
data.iucn.org/dbtw-wpd/edocs/2004-047.pdf
Turner, R. Kerry, Stavros Georgiou, and Brendan Fisher. Valuing
Ecosystem Services: The Case of Multi-functional Wetlands.
London: Earthscan, 2008.
Wittmer, Heidi, and Haripriya Gundimeda, eds. The Economics of
Ecosystems and Biodiversity for National and International Policy
Makers. London: The Economics of Ecosystems and Biodi-
versity, 2010.
World Bank. The Changing Wealth of Nations: Measuring
Sustainable Development in the New Millennium. Washington,
DC: World Bank Publications, 2011.
Periodicals
Andam, Kwaw S., Paul J. Ferraro, Katharine R.E. Sims, et al.
“Protected Areas Reduced Poverty in Costa Rica and
Thailand.”Proceedings of the National Academy of Sciences of the
United States of America 107, no. 22 (2010): 9996–10001.
Accessed July 27, 2012. doi:10.1073/pnas.0914177107.
Balmford, Andrew, Aaron Bruner, Philip Cooper, et al.
“Economic Reasons for Conserving Wild Nature.”Science
297, no. 5583 (2002): 950–953.
Balvanera, Patricia, Andrea B. Pfisterer, Nina Buchmann, et al.
“Quantifying the Evidence for Biodiversity Effects on
Ecosystem Functioning and Services.”Ecology Letters 9, no. 10
(2006): 1146–1156. Accessed July 27, 2012. doi:10.1111/
j.1461-0248.2006.00963.x.
Boyles, Justin G., Paul M. Cryan, Gary F. McCracken, and
Thomas H. Kunz. “Economic Importance of Bats in
Agriculture.”Science 332, no. 6025 (2011): 41–42.
Butchart, Stuart H.M., Matt Walpole, Ben Collen, et al. “Global
Biodiversity: Indicators of Recent Declines.”Science 328, no.
5982 (2010): 1164–1168.
Cardinale, Bradley J., J. Emmett Duffy, Andrew Gonzalez, et al.
“Biodiversity Loss and Its Impact on Humanity.”Nature 486,
no. 7401 (2012): 59–67.
Chiabai, Aline, Chiara M. Travisi, Anil Markandya, et al.
“Economic Assessment of Forest Ecosystem Services Losses:
Cost of Policy Inaction.”Environmental and Resource Economics
50, no. 3 (2011): 405–445.
Chichilnisky, Graciela, and Geoffrey Heal. “Economic Returns
from the Biosphere.”Nature 391 (1998): 629–630.
Grzimek’s Animal Life Encyclopedia 865
Extinction The relative cost of saving species
© Cengage Learning. All rights reserved. No distribution allowed without express authorization.

Costanza, Robert, Ralph d’Arge, Rudolf de Groot, et al. “The
Value of the World’s Ecosystem Services and Natural
Capital.”Nature 387, no. 6630 (1997): 253–260.
Díaz, Sandra, Josepth Fargione, F. Stuart Chapin III, and David
Tilman. “Biodiversity Loss Threatens Human Well-Being.”
PLoS Biology 4, no. 8 (2006): e277. Accessed July 27, 2012.
doi:10.1371/journal.pbio.0040277.
Elmqvist, Thomas, Carl Folke, Magnus Nyström, et al.
“Response Diversity, Ecosystem Change, and Resilience.”
Frontiers in Ecology and the Environment 1, no. 9 (2003):
488–494.
Gatzweiler, Franz W. “Beyond Economic Efficiency in Biodi-
versity Conservation.”Journal of Interdisciplinary Economics 19,
nos. 2–3 (2008): 215–238.
James, Alexander N., Kevin J. Gaston, and Andrew Balmford.
“Balancing the Earth’s Accounts.”Nature 401, no. 6751
(1999): 323–324.
Kremen, Claire. “Managing Ecosystem Services: What Do We
Need to Know about Their Ecology?”Ecology Letters 8, no. 5
(2005): 468–479.
Losey, John E., and Mace Vaughan. “The Economic Value of
Ecological Services Provided by Insects.”BioScience 56, no. 4
(2006): 311–323.
Luck, Gary W., Richard Harrington, Paula A. Harrison, et al.
“Quantifying the Contribution of Organisms to the Provision
of Ecosystem Services.”BioScience 59, no. 3 (2009): 223–235.
Martín-López, Berta, Carlos Montes, and Javier Benayas.
“Economic Valuation of Biodiversity Conservation: The
Meaning of Numbers.”Conservation Biology 22, no. 3 (2008):
624–635. Accessed July 27, 2012. doi:10.1111/j.1523-
1739.2008.00921.x.
Naidoo, Robin, Andrew Balmford, Paul J. Ferraro, et al.
“Integrating Economic Costs into Conservation Planning.”
Trends in Ecology and Evolution 21, no. 12 (2006): 681–687.
Ressurreição, Adriana, James Gibbons, Tomaz Ponce Dentinho,
et al. “Economic Valuation of Species Loss in the Open Sea.”
Ecological Economics 70, no. 4 (2011): 729–739.
Richardson, Leslie, and John Loomis. “The Total Economic
Value of Threatened, Endangered, and Rare Species: An
Updated Meta-analysis.”Ecological Economics 68, no. 5 (2009):
1535–1548.
Serpell, James A. “Factors Influencing Human Attitudes to
Animals and Their Welfare.”Animal Welfare 13, supp.
1 (2004): S145–S151.
Turner, Will R., Katrina Brandon, Thomas M. Brooks, et al.
“Global Conservation of Biodiversity and Ecosystem
Services.”BioScience 57, no. 10 (2007): 868–873.
Turner, Will R., Katrina Brandon, Thomas M. Brooks, et al.
“Global Biodiversity Conservation and the Alleviation of
Poverty.”BioScience 62, no. 1 (2012): 85–92.
Other
“The Economics of Ecosystems and Biodiversity.”Accessed July
27, 2012. http://www.teebweb.org
“Intergovernmental Platform on Biodiversity and Ecosystem
Services.”Accessed July 27, 2012. http://www.ipbes.net
“Millennium Ecosystem Assessment.”Accessed July 27, 2012.
http://www.maweb.org
Berta Martín-López
Marina García-Llorente
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