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Provided for non-commercial research and educational use only. Not for reproduction, distribution or commercial use. This article was originally published in the Encyclopedia of Biodiversity, second edition, the copy attached is provided by Elsevier for the author’s benefit and for the benefit of the author’s institution, for non-commercial research and educational use. This includes without limitation use in instruction at your institution, distribution to specific colleagues, and providing a copy to your institution’s administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier’s permissions site at: http://www.elsevier.com/locate/permissionusematerial Guerry Anne D., Ruckelshaus Mary H., Plummer Mark L., and Holland Dan (2013) Modeling Marine Ecosystem Services. In: Levin S.A. (ed.) Encyclopedia of Biodiversity, second edition, Volume 5, pp. 329-346. Waltham, MA: Academic Press. © 2013 Elsevier Inc. All rights reserved.

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Page 1: Chapter 333 - Modeling Marine Ecosystem Services

Provided for non-commercial research and educational use only. Not for reproduction, distribution or commercial use.

This article was originally published in the Encyclopedia of Biodiversity, second edition, the copy attached is provided by Elsevier for the author’s benefit and for the benefit of the author’s institution, for non-commercial research

and educational use. This includes without limitation use in instruction at your institution, distribution to specific colleagues, and providing a copy to your institution’s administrator.

All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s website or repository, are prohibited.

For exceptions, permission may be sought for such use through Elsevier’s permissions site at:

http://www.elsevier.com/locate/permissionusematerial

Guerry Anne D., Ruckelshaus Mary H., Plummer Mark L., and Holland Dan (2013) Modeling Marine Ecosystem Services. In: Levin S.A. (ed.) Encyclopedia of Biodiversity, second edition, Volume 5, pp. 329-346. Waltham, MA:

Academic Press.

© 2013 Elsevier Inc. All rights reserved.

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En

Modeling Marine Ecosystem ServicesAnne D Guerry and Mary H Ruckelshaus, Stanford University, Stanford, CA, USAMark L Plummer and Dan Holland, National Oceanic and Atmospheric Administration, Seattle, WA, USA

r 2013 Elsevier Inc. All rights reserved.

GlossaryEcosystem services Wide array of benefits that ecosystems

and their biodiversity confer on humanity.

Marine Broadly defined to include coastal (on land,

within a narrow fringe adjacent to saltwater), intertidal,

nearshore, and open ocean.

Production function approach An approach that models

ecosystem services as the relationship between ecological

cyclopedia of Biodiversity, Volume 5 http://dx.doi.org/10.1016/B978-0-12-3847

and human inputs (e.g., the structure and functions of an

ecological system, human labor and capital) and outputs

valued by humans.

Valuation Act of estimating or setting the value of

something,

Value Relative worth, merit, or importance. Can be

measured in various ways, including but not limited to

monetary metrics.

Introduction to Marine Ecosystem Services two that inhibit the division of cancer cells grown in the la-

Humans have always benefited from marine ecosystems,

enjoying resources such as seafood and opportunities for ac-

tivities like recreation and marine transportation. These eco-

systems also provide indirect benefits by sequestering carbon

and playing key roles in the regulatory processes of other

global cycles. Benefits derived from these systems are broadly

characterized as the ecosystem services provided by marine

ecosystems. As identified by the Millennium Ecosystem As-

sessment (MA) (2005b), marine ecosystem services span four

major categories: provisioning, regulating, cultural, and sup-

porting services (Table 1). Broad assessments of marine and

coastal ecosystems services based on the MA can be found in

Agardy et al. (2005) and the United Nations Environment

Program (2006), as well as other synthesis documents (e.g.,

Peterson and Lubchenco, 1997). Costanza (2000), Patterson

and Glavovic (2008), and Wilson and Liu (2008) also provide

useful overviews of these services, as do descriptions of the

particular services provided by fish populations (e.g., Holm-

lund and Hammer, 1999), coral reef ecosystems (e.g., Moberg

and Folke, 1999) and mangroves (e.g., Ronnback, 1999).

Among the four types of marine ecosystem services, pro-

visioning services such as food from capture fisheries, aqua-

culture, and wild foods are the most obvious and easily

valued. About 80 million tons of fish were landed in marine

capture fisheries worldwide in 2009, and fish account for ap-

proximately 16% of the annual animal protein consumption

by humans (FAO Fisheries Department, 2010). Globally, more

than 1.5 billion people rely on fish for almost 20% of their

animal protein. On average, each person living in 2009 ate

17.2 kg of fish (including fish from aquaculture; the pro-

portion from capture fisheries alone is difficult to calculate

given nonfood uses of wild fish) (FAO Fisheries Department,

2010). Other provisioning services include timber and fiber

from mangroves and seagrass beds, and biochemicals for

cosmetics and food additives. The potential also exists for

developing novel natural products from marine species with

medical applications (Carte, 1996). For example, researchers

recently found that three marine species collected from off-

shore oil and gas platforms in California’s Santa Barbara

Channel had potential for biomedical applications, including

boratory (Schmitt et al., 2006). In addition, the ocean may

become an important energy source: biofuels from algae and

power generation from wave and tidal energy have potential

for more widespread use. And finally, the world’s oceans

provide the highways for the global shipping trade.

Marine systems also provide a wide range of regulating

services. As vividly highlighted by the human losses wrought

by the 2005 hurricanes on the US Gulf Coast, coastal and

estuarine wetlands have value for their ability to reduce storm

surge elevations and wave heights (Danielsen et al., 2005;

Travis, 2005). Other regulating services provided by marine

systems include the transformation, detoxification, and se-

questration of wastes (Peterson and Lubchenco, 1997). And

‘‘blue carbon’’ – the role oceans can play as carbon sinks – is a

service that is gaining interest (Nellemann et al., 2009).

Human societies have always been drawn to the oceans,

which have proven to be a rich source of cultural services. In

1995, an estimated 39% of the world’s population lived with

100 km of a coast (Burke et al., 2001). In the US, people love

to live near the ocean; one study predicts average increases of

3600 people a day moving to coastal counties through 2015

(Culliton, 1998). Coastal tourism is a key component of many

economies around the world and is one of the fastest growing

and most profitable sectors of tourism (United Nations

Environment Program, 2006). Countless intangible benefits

are drawn from the oceans, too. For example, many com-

munities define their identities in relation to the coast, people

draw inspiration from oceans in numerous ways, and rich

ceremonial traditions are intricately linked to the sea.

Finally, the oceans provide essential supporting services

that underpin many of the globe’s ecological functions. The

oceans hold 96.5% of Earth’s water (Gleick, 1996) and are a

primary driver of the atmosphere’s temperature, moisture

content, and stability (Colling, 2001). The global cycles of

carbon, nitrogen, oxygen, phosphorus, sulfur, and other key

elements flow through the oceans (Peterson and Lubchenco,

1997), and marine ecosystems are responsible for approxi-

mately 40% of global net primary productivity (Schlesinger,

1991; Melillo et al., 1993). The oceans are home to vast res-

ervoirs of genetic and ecological diversity, arguably the most

fundamental source of supporting services as they are directly

19-5.00333-6 329

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Table 1 Ecosystem services provided by marine systems

Types of service Examples

Provisioning servicesFood production (capture fisheries; aquaculture; and wild

foods)Tuna, crab, lobster; salmon, oysters, shrimp, seaweed; mussels, clams

Fiber production Mangrove wood, seagrass fiberBiomass fuel production Mangrove wood, biofuel from algaeMaintenance of aquatic systems Shipping, tidal turbinesGeneration of genetic resources Individual salmon stocks, marine diversity for bioprospectingProduction of biochemicals, natural medicines, and

pharmaceuticalsAntiviral and anticancer drugs from sponges, carrageenans from seaweed

Regulating servicesClimate regulation Major role in global CO2 cycleWater regulation Natural stormwater management by coastal wetlands and floodplainsErosion regulation Nearshore vegetation stabilizes shorelinesWater purification and waste treatment Uptake of nutrients from sewage wastewater, detoxification of PAHs by marine

microbes, sequestration of heavy metalsDisease regulation Natural processes may keep harmful algal blooms and waterborne pathogens in

checkPest regulation Grazing fish help keep algae from overgrowing coral reefsPollination/Assistance of external fertilization Innumerable marine species require seawater to deliver sperm to eggNatural hazard regulation Coastal and estuarine wetlands and coral reefs protect coastlines from storms

Cultural servicesProvision of conditions that support or enhance ethical

values (nonuse)Spiritual fulfillment derived from estuaries, coastlines, and marine waters

Provision of conditions that support or enhanceexistence values (nonuse)

Belief that all species are worth protecting, no matter their direct value tohumans

Provision of recreation and ecotourism opportunities(consumptive and nonconsumptive uses)

Scuba diving, beachcombing, whale watching, boating, snorkeling; fishing,clamming

Supporting servicesNutrient cycling Major role in carbon, nitrogen, oxygen, phosphorus, and sulfur cyclesSoil formation Many salt-marsh surfaces vertically accrete; eelgrass slows water and traps

sedimentPrimary production Significant portion of global net primary productivityWater cycling Most of Earth’s water is in oceans; they are central to the global water cycle

Source: Adapted from Guerry A, Plummer M, Ruckelshaus M, and Harvey C (2011) Ecosystem service assessments for marine conservation. In: Kareiva P, Tallis H, Ricketts T, Daily

G, and Polasky S (eds.) Natural Capital: Theory and Practice of Mapping Ecosystem Services. Oxford: Oxford University Press.

330 Modeling Marine Ecosystem Services

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linked to the rate of evolution and therefore the ability to

adapt to a changing climate (Pergams and Kareiva, 2009).

Since the MA, much work has been done in developing

and applying new methodologies to model, map, and value

ecosystem services (e.g., UNEP and IOC-UNESCO, 2009;

Kareiva et al., 2011; UK National Ecosystem Assessment, 2011).

A great deal of the original MA was focused on terrestrial

systems, but new workFand new political contextsFhave

sparked the expansion of an ecosystem services framework to

marine systems. For example, the UK recently completed a

groundbreaking, countrywide assessment of both terrestrial

and marine ecosystem services and their value (Stokstad, 2011;

UK National Ecosystem Assessment, 2011). Although agri-

cultural production has increased between 1950 and the pre-

sent, landings of fish and shellfish from UK waters have

declined since the 1960s from almost 900,000 t to slightly

more than 500,000 t in 2008 (UK National Ecosystem As-

sessment, 2011). Protection of UK coastlines from storm-

induced flooding and erosion has declined in response to a

10% loss of natural habitats such as dunes, kelps, seagrasses

and marshes over the past 60 years (UK National Ecosystem

Assessment, 2011).

The MA and subsequent assessments have raised awareness

of ecosystem services, the explicit human dependence on

them, and the threatened status of many of them. Bringing

ecosystem services into the active management of terrestrial

and marine ecosystems, however, requires more than just a

catalog of services and their total values. More pragmatic is the

assessment of the ecological and economic consequences of

management activities in particular places. An understanding

of how changes in ecosystems are likely to lead to changes

in ecosystem services can most clearly provide information to

decision makers. Modeling marine ecosystem services can play

an important role in providing such insights.

Foundations

Marine ecosystems provide both great opportunities and

challenges for the application of the framework of ecosystem

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services. The concept of ecosystem services has a long history

but has seen a relatively recent resurgence of interest from

ecologists, economists, and conservation practitioners. Be-

cause ecosystem services are reviewed elsewhere in this vol-

ume, we focus here on the conceptual frameworks and

methodologies that pertain to marine applications, as well as

the challenges facing these applications.

Why Model Marine Ecosystem Services?

Models are important tools for examining the dynamic na-

tures of and interconnections among the biophysical and

human elements of marine ecosystems. They provide a way of

exploring future scenarios that lie outside the range of past

experiences, as well as possible unexpected consequences of

policy actions. An ecosystem services framework provides

important insights into the challenge of pursuing ecosystem-

based management. Ecosystem services are the currencies

through which the consequences of ecosystem change flow to

people. Using such a framework, ecosystem services and their

values can be used as a set of metrics for assessing alternative

management interventions and their potential impacts on

economic or social well-being.

Models of ecosystems that incorporate both biophysical

and human components can present policy makers with an

enormous set of potential indicators for gauging the changes

brought about by policies. Ecosystem services – the ‘‘ecological

endpoints’’ of the system that are directly connected

to human well-being (Boyd, 2007) – can provide a guide for

winnowing this set down. In the context of ecosystem mod-

eling, such a framework avoids the potential problem of

double counting the value of ‘‘intermediate’’ ecological elem-

ents (e.g., forage fish) that support other ‘‘endpoint’’ elements

with direct value (e.g., fishery harvests) because the value of

the intermediate elements are embedded in the value of the

endpoints (Boyd and Banzhaf, 2007).

Ecosystem models that focus on ecosystem services also

provide the opportunity to overcome problems arising from

traditional management in the marine realm, which generally

proceeds sector by sector, with distinct bodies making isolated

decisions. These single-sector decisions often affect a broad set

of ecosystem services, many of which are outside the scope of

their authority. As a result, there has been little effort to con-

sider the ways in which single decisions impact the full suite of

things people care about and need.

Using an ecosystems services framework can highlight

trade-offs among multiple objectives so that decisions to re-

solve those trade-offs can be made transparently. Increasingly,

scientists and managers are pointing to links between diver-

sity, productivity, and resilience attributes of marine systems

and their response to human interventions in conserving,

harvesting, and regulating marine ecosystem services (Liu

et al., 2007a; Levin and Lubchenco, 2008; Murawski et al.,

2009; Chan and Ruckelshaus, 2010). Our ability to model and

assess trade-offs among objectives of multiple management

sectors (e.g., fisheries, wave or wave energy, recreation) and

ecosystem services (e.g., value of fishery landings, kilowatt

hours of energy generated, revenue from recreational activ-

ities) is needed in order to inform more complex cases

of ecosystem-based management that can accommodate a

broader suite of actors (Foley et al., 2010).

Pioneering efforts to use the framework of ecosystem ser-

vices to understand the connections between activities in one

sector and their impacts on others are underway in the marine

environment. Decision makers at various levels of government

from around the globe, including the Interagency Ocean

Policy Task Force (IOPTF) (2010), the European Commission

(2010), and the recently approved UN Intergovernmental

Science-Policy Platform on Biodiversity and Ecosystem Ser-

vices (IPBES) have recognized that new approaches are needed

to ensure the sustainability of benefits people derive from

oceans and coasts (UN General Assembly (UNGA), 2010).

Among these parties, there is currently a great deal of em-

phasis on marine (or ‘‘maritime’’) spatial planning. This ap-

proach to comprehensive management of marine and coastal

systems analyzes current and anticipated uses, identifies areas

most suitable for particular activities, and provides a process

to better determine how oceans are used and protected now

and for future generations (Douvere and Ehler, 2009; Ehler

and Douvere, 2009). Ecosystem services provide a framework

and a common language for this type of planning process,

allowing agencies and private interests to articulate their goals

transparently using common terms and to better understand

how their decisions interact with those of other users and

activities.

Connecting Social and Ecological Systems

Several related conceptual frameworks provide the context

within which ecosystem service information is critical to

understanding feedbacks between humans and ecosystem

conditions in marine and other environments. One of the

most important is the notion that humans are an integral part

of ecosystems, and so ecosystem models should encompass

their behavior (Holland et al., 2010). Similarly, the theory of

complex adaptive systems encompasses humans as parts of

coupled systems and presents humans as participating in the

dynamics of the system (Levin, 1998; Levin and Lubchenco,

2008). The production of ecosystem services involves a com-

bination of ecological functions and human actions and val-

ues (Figure 1). In the marine environment, for example,

marine ecosystems have fish populations that offer oppor-

tunities for commercial and recreational harvests, two of the

most valuable services derived from these systems. The human

action of harvest, however, is what transforms the potential

ecological supply into the actual provision of the ecosystem

service (Tallis et al., 2011). Human values in the form of the

demand for seafood, the valuation of recreation, and the costs

of commercial harvest or recreational angling all determine

the value of these services.

Building models to account for the humans who interact

with or are affected by an ecological system can therefore

provide fundamental insights into the provision and value of

ecosystem services. Simple economic models can focus on

individual decisions to use more or less of a marine resource,

for example. An ecological change that makes it harder to

harvest fish is likely to produce a lower level of effort that can

be modeled in a simple framework of the demand for that

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Figure 1 The production of ecosystem services involves a combination of ecological functions and human actions and values. Recreationalfishing as an ecosystem service, for example, depends on the healthy functioning of natural systems that support fish populations. The humanaction of harvest transforms the potential ecological supply into the actual provision of the ecosystem service.

332 Modeling Marine Ecosystem Services

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activity. More complicated models involve complex decisions

about where and when to fish, which species to fish for, and so

forth (e.g., Sanchirico and Wilen,1999; Wilen et al., 2002).

Similarly, human behavior has more latitude to make adjust-

ments over longer periods of time, and so modeling short-run

versus long-run behavior of a system can account for such

differences (Holland and Brazee, 1996). As they mature,

models of marine ecosystem services can include how humans

interact with biophysical components of the system and can

incorporate realistic mechanisms for changing those inter-

actions based on economic and other incentives. We include

some examples of these kinds of truly linked human and

biophysical models in the ‘‘Production Function Modeling’’

section.

The coupled social–ecological systems model developed by

Ostrom (2009) is one key conceptual framework that includes

interactions between biophysical and social systems. Much of

the focus of research guided by this conceptual model is to

identify relevant attributes for understanding the dynamics

of a coupled social–ecological system (e.g., the lobster fishery

in Maine and its fishing community) and the effects of dif-

ferent management approaches (e.g., regulation, incentives)

on ecological and ecosystem services (e.g., lobster biomass,

landings, biodiversity) and social (e.g., efficiency, equity, ac-

countability) outcomes. Under this framework, the features of

governance systems (e.g., property rights systems, operational

rules), user groups (e.g., social heterogeneity, leadership), and

resource units (e.g., economic value, behavior of harvested

species) that give rise to different ecological and social out-

comes can be quantitatively or qualitatively assessed using

well-developed methodologies (Cudney-Bueno and Basurto,

2009; Ostrom, 2009; Basurto and Coleman, 2010).

The closely related conceptual framework of coupled

human and natural systems touches on similar issues but with

a stronger focus on understanding mechanisms and feedbacks

(Liu et al., 2007a, 2007b). A critical question this framework

poses is the positive or negative direction of feedback loops

between human and natural components of a system and the

strength of those interactions (Liu et al., 2007b). This mod-

eling framework encourages exploration of temporal and

spatial as well as organizational couplings, thresholds, and

resilience in linked systems. For example, whether couplings

between human and natural components of a marine eco-

system are direct and local or indirect and global can be a key

feature influencing how humans and ecosystem services re-

spond to signs of change in the ecosystem, and it is an in-

creasingly important feature with the globalization of both

human and natural processes.

One modeling approach that incorporates these frame-

works is the integrated ecosystem assessment (IEA). An IEA

outlines both a decision analytical framework and a process

containing the logical steps necessary for managing eco-

systems, including explicit identification of objectives and the

indicators for tracking progress, a risk analysis describing the

state of the indicators, an evaluation of management strat-

egies, and an overall assessment of ecosystem response (Levin

et al., 2009). IEAs have a successful history of application in

fishery-based ecosystem management (Caddy, 1999; Sains-

bury et al., 2000; Smith et al., 2007); they are designed to be

iterative, expanding in scope and sophistication as

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information develops over time (e.g., Dennison et al., 2007;

Tallis et al., 2010; Samhouri et al., 2011).

IEAs are one example of current approaches to the man-

agement of marine ecosystems that can use ecosystem services

in a modeling framework. An IEA uses quantitative analyses

and ecosystem modeling to support management decisions

that address a range of social, economic, and natural con-

ditions. Ecosystem services can provide a set of metrics for

assessing these conditions, and modeled changes in their

levels can then provide decision makers with a way to com-

pare alternative policies.

Challenges in Modeling Marine Ecosystem Services

From a scientific perspective, the modeling of marine eco-

system services is sufficiently different from the task on land to

warrant its own treatment. Marine systems provide some

special scientific challenges in the endeavor to map, model,

and value ecosystem services. Most models for terrestrial sys-

tems use a land-use or land-cover data layer as a foundation

for the initial assessment of services, as well as a baseline for

exploring how land conversion and land-use change alter

ecosystem services. Fundamentally, the same approach works

for marine environments. Marine systems have habitats that

are altered or used with consequences, and different man-

agement approaches yield different habitat conditions and

different delivery of services.

We cannot readily ‘‘see’’ most benthic habitat types using

satellite imagery or other remote sensing technology, so maps

of habitat type and condition are harder to come by in marine

systems than they are on land. Also, in marine systems, as-

sociations between species and habitat patches are more dif-

ficult to discern. The lack of basic land-use and land-cover data

for marine systems underscores the need for a modeling ap-

proach to ecosystem and food-web dynamics that is less

tightly coupled to detailed habitat maps than the approach

often taken in terrestrial systems (e.g., Kareiva et al., 2011).

A second shift in perspective required when considering

marine ecosystem services stems from the ways in which

humans interact with marine environments. We do not live in

the sea, and thus many of our actions that affect the marine

environment are indirect. We can think of many terrestrial

habitats without considering the effects of marine systems on

them, but the converse is not true. Activities on land (e.g.,

coastal development; nutrient, sediment, and pathogen inputs

to freshwater; increases in impervious surfaces) have profound

effects on nearshore marine systems (Carpenter et al., 1998;

Mallin et al., 2000; Diaz and Rosenberg, 2008).

Understanding and modeling marine ecosystem services is

also made more difficult by the fact that we generally have less

direct control or knowledge of human actions. In terrestrial

areas, property rights are the norm, and zoning often dictates

what and where actions can take place (both on public and

private property). Human activities and their results are also

more observable on land. In marine areas, human activities

are not typically managed explicitly at a fine scale, and many

areas are effectively open access. Thus, it is often more critical

to incorporate endogenous behavioral models of people into

models of ecosystem services.

Finally, modeling ecosystem services is not a substitute for

all other ways of assessing marine and other ecosystems.

Concerns about biodiversity of marine ecosystems, for ex-

ample, may be poorly addressed by a pure ecosystem services

approach. Some assert that protection and restoration of

marine systems will improve both biodiversity and the ability

of the oceans to provide ecosystem services; others point out

that trade-offs can occur (Balvanera et al., 2006; Worm et al.,

2006; Naeem et al., 2009; Palumbi et al., 2009; Klein et al.,

2010; Brander, 2010). Ecosystem modeling coupled with ob-

servations should not ignore this and other approaches to

marine conservation, and they should continue to provide

needed evidence for the conditions under which biodiversity

and ecosystem service provision are positively related and

where trade-offs are likely to occur.

Approaches

The modeling of marine ecosystem services can take a variety

of approaches. Although not normally considered a ‘‘model,’’ a

straightforward accounting of ecosystem services and their

values has played an important role in past assessments, and

so it is included in this section as one approach. Many models

of ecosystem services take a production function approach in

which the structure and functions of an ecological system are

combined with human actions and capital to produce an

output valued by humans (National Research Council, 2005).

This approach has been used to analyze a variety of individual

ecosystem services, as well to provide the framework for de-

veloping suites of models that encompass multiple services.

Each of these approaches is presented in the following

sections.

Accounting Approaches and Basic Valuation Methods

The simplest approach to assessing ecosystem services for

marine and terrestrial systems is a straightforward accounting

of those services and their values. This approach is not one

that explicitly models the ecological or human systems, but it

has been used to assess marine ecosystem services (Beaumont

et al., 2007, 2008). Generally, the approach begins by defining

a particular geographic location and then identifying the set of

ecosystem services flowing from that area. The assessment can

then be as simple as compiling local information on each of

these services for that region – e.g., using revenues from an

area’s fishery harvests as a measure of the economic value of

that type of provisioning service.

A more common method that has been used for terrestrial

systems is ecosystem-service mapping (Troy and Wilson,

2006). This method differentiates a particular area by land

cover, biome, or some other set of ecologically based land-

scape or seascape type. Drawing from a standard set of eco-

system service categories (e.g., deGroot et al., 2002), each

landscape type (e.g., forestland) is then linked to a set of

services (e.g., recreation or carbon sequestration) believed to

be provided by that type. The quantity of an ecosystem service

produced is then assumed to be linearly related to the area of

the landscape type.

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From these starting points, local economic data, if they

exist, can be used to transform the quantity of ecosystem

services into economic values. Such data are not commonly

available, however, and so the more typical approach to as-

signing economic value is to use what is known as benefit

transfer. Benefit transfer is a method for taking economic data

on benefits (or values in general) gathered from one site and

applying the data to another site. This method is rarely the

‘‘first-best’’ choice for estimating economic values but the costs

of gathering primary, site-specific data have made it a common

practice for studies of ecosystem service value (e.g., see

Rosenberger and Loomis (2001) and National Research

Council (2005) for examples of recreational uses of natural

sites). Using the assumption of a linear relation between

ecosystem service quantities and landscape area, a ‘‘unit value’’

for a particular ecosystem service and landscape type can then

be derived by taking an existing study’s estimated value for a

particular ecosystem service and dividing by the area of the

studied site’s landscape (e.g., recreation or carbon seques-

tration value per acre of forestland). These unit values can

then be used to assess the value of ecosystem services at other

sites for multiple services and landscape types if they are

present.

This accounting approach was pioneered by Costanza and

colleagues (1997), who estimated global values for 17 eco-

system services for 16 biomes. Their effort included five mar-

ine and coastal biomes: open ocean, estuaries, seagrass or

algae beds, coral reefs, and ocean shelf. These five biomes

accounted for almost two-thirds of the estimated annual $33

trillion value of Earth’s services. Their use of benefit transfer

was almost universal; the original data used in the study

were gathered in one or more locations and then projected

worldwide.

Accounting approaches such as Costanza et al. (1997) are

simple and play the important role of raising the awareness of

the values of traditionally undervalued, nonmarket ecosystem

services. They should be used cautiously, however, with im-

portant qualifications in mind. First, this approach imposes

linearity on the valuation of ecosystem services, which has the

potential to produce biased estimates. Human pressures on

ecosystems and the services they provide can result in impacts

that respond nonlinearly to changes in the scale of the pres-

sure or change discontinuously if a threshold is crossed

(Groffman et al., 2006). The valuation of ecosystems services

should therefore account for such nonlinearities if the scale

under consideration is more than minimal (Barbier et al.,

2008; Koch et al., 2009). The human values that apply to

ecosystem services can also exhibit nonlinearities as the

quantity or quality of the ecosystem service available changes

(Bockstael et al., 2000; Toman, 1998). Again, if the scale of a

service under consideration is large, projecting a value esti-

mated for a small change has the potential to produce sig-

nificant bias. And if more than one service is evaluated, the

aggregate value of the group of services is likely to be non-

linear with respect to considering more and more services. If

estimates of individual ecosystem service values are used in

such an exercise, simply adding them together will again

produce a biased estimate of the group’s value. These prob-

lems grow more and more significant as the scale of the eco-

system service valuation exercise increases. Given that this

approach is often used to produce national or even global

estimates (Anielski and Wilson, 2009; Beaumont et al., 2007,

2008; Ingraham and Foster, 2008; Naidoo et al., 2008), the

issue of nonlinearities is an important one.

Second, the method requires rigid association of a par-

ticular set of ecosystem services to a particular landscape type;

thus it does not allow assessment of how policies might

change the flow of ecosystem services and affect their

values, short of comparing values under wholesale destruction

of the landscape type. Relying on landscape type to assign

a fixed presence and quantity of services provides little infor-

mation to evaluate how a policy will change ecosystem service

values in any way other than a change in the landscape type.

This ‘‘all or nothing’’ approach may make sense for some

cases – for example, replacing forest with open agricultural

fields – but not for others, as it is highly unlikely that any

policy is capable of converting saltwater estuaries into upland

forestland.

A third caution stems from the common use of benefit

transfer for ecosystem service mapping exercises and other

accounting approaches. The broad categories used in these

types of exercises facilitate a comprehensive analysis of eco-

system service values for large geographic areas, but they also

subject the analysis to a high likelihood of what is called

generalization error (Plummer, 2009). As an example, consider

the problem of estimating the value of providing ‘‘recreation’’

in a ‘‘marine area.’’ Recreation covers scores of possible market

and nonmarket activities that take place in natural settings,

and some may be present in one marine area but not in

others. Also, the presence of human-built infrastructure

and the accessibility of the marine area are important de-

terminants of the economic value of the recreational activity.

Because these factors can vary widely, using estimates of

economic value from one site and applying them to another

is likely to produce significant errors if the categories of

‘‘recreation’’ and ‘‘marine area’’ are treated at a high level of

generalization.

Finally, existing estimates of economic values for ecosystem

services are strongly influenced by the methodology employed

to make the estimate. As the NRC report on valuing ecosystem

services (2005) emphasizes, not all methodologies are equally

good. Happily, several valuation approaches are accepted

practice by economists, but the interpretation of the resulting

values should be informed by the method used.

Revealed Preferences MethodsStandard economic theory is based on the assumption that

observable choices made by individuals reveal their expected

valuation of a good or activity (but it also allows for some

goods with values that are independent of observable be-

havior) (Slesnick, 1998). The revealed preference approach is

a collection of methods for estimating economic values that

rely on observable behavior. The most obvious revealed pref-

erence method is the use of market data. The production

of commercially harvested finfish and shellfish is a leading

example of marine ecosystem services with values that can

be estimated with market data. Another revealed preference

method that uses market data is known as hedonic analysis

(Palmquist, 1991; Freeman, 1993; Palmquist, 2003); it ana-

lyzes goods (e.g., housing) that are sold as a bundle of

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characteristics. In some cases, location-specific characteristics

may be part of the bundle and include environmental

amenities such as air and water quality or proximity to open

space. If ecosystem services are a part of these bundled

characteristics – for example, shoreline protection provided by

nearshore ecological systems – the marginal value of such a

service can be estimated with a hedonic analysis.

A final revealed preference method is useful for cultural

ecosystem services such as recreation and other environmental

experiences that take place outside any formal market. In this

approach, the cost of engaging in the activity can be used

to derive estimates of its economic value (Clawson, 1959;

Knetsch, 1963). Similar to the assumptions for hedonic

models, the recreation ‘‘good’’ can be viewed as a bundle of

characteristics, some of which are the environmental features

important to the recreational experience. If data are available

for visits to multiple sites with varying levels of those features,

one can then estimate the contribution of a particular feature

to the demand for that recreation and from this estimate the

feature’s value (Morey, 1981).

Stated Preference MethodsStated preference methods also can provide legitimate esti-

mates of economic value. These methods rely on survey

questions that ask individuals to make a choice, describe a

behavior, or state directly what they would be willing to pay

for specified changes in nonmarket goods or services. These

methods are increasingly used in economic studies of en-

vironmental quality because they offer the opportunity to es-

timate the value for anything that can be presented as a

credible and consequential choice. If conducted with attention

to the many standards of care for its execution (Mitchell and

Carson, 1989), this method can provide useful estimates of

ecosystem service values.

Other Valuation MethodsOther methods are less reliable and particularly prone

to misuse when estimating ecosystem service values. A

prominent example is the replacement-cost or cost-of-treat-

ment approach. For example, the cost of municipal water

treatment for drinking water can be reduced by the presence of

a wetland because the wetland system filters and removes

pollutants (Day et al., 2004). Using the cost of a human-built

alternative treatment method is an approach sometimes taken

to estimate the ‘‘value’’ of the wetland, but in general this cost

does not have any necessary relation to the actual services

provided by a particular wetland. On the one hand, the wet-

land may not actually provide filtration services (e.g., because

of the absence of contaminants that need filtering or bio-

physical features of the wetland that prevent filtering). On the

other hand, the wetland might simultaneously provide a

number of other services (such as carbon storage, nursery

habitat support for fisheries) that are not captured by this

form of valuation. As a result, the replacement-cost approach

to estimating the value of an ecosystem service must be used

with great care.

One final word about valuation: much of the dialogue

about ecosystem services focuses on economic valuation. Eco-

nomic valuation, however, is far from the only way to value

ecosystem services. Although many services (e.g., provisioning

services) are relatively straightforward to value in economic

terms, others (e.g., cultural services) often defy economic

valuation. The comparison of multiple ecosystem services in a

single currency (e.g., dollars) is appealing in some contexts but

not appropriate or even desirable in others. Cultural ecosystem

services – diverse, nonmaterial benefits that people obtain

through their interactions with ecosystems, including spiritual

inspiration, cultural identity, and recreation – are difficult but

not impossible to value. Various methods (e.g., narrative

methods, paired comparisons, structured decision making) can

be used to elicit the relative weight that people place on these

services (Chan et al., in press).

Production Function Modeling

A production function approach is fundamentally process-

based. It represents the relationship between inputs (e.g., the

density of mangroves) and outputs (e.g., the degree of pro-

tection from storms). Production functions have been used

extensively in agriculture, manufacturing, and other sectors of

the economy. Ecological production functions can be used to

explore how changes in ecosystem structure and function lead

to changes in the flows of services (National Research Council,

2005).

Unlike the accounting methods previously discussed

(see Accounting Approaches and Basic Valuation Methods),

a production function approach allows for the comparison

of alternative courses of action and their effects on ecosystems

and services, and it can thus provide more useful infor-

mation for decision making. With process-based models,

analysts can explore how changes in inputs are likely to lead

to changes in outputs. If valuation is of interest, then

production function approaches can use both market prices

and nonmarket valuation methods to estimate economic

value and to show how the monetary value or other value

currencies are likely to change under different environmental

conditions.

The production function approach has been used to model

various ecosystem services on land (Ricketts et al., 2004; Kar-

eiva et al., 2011) and in marine systems (Barbier et al., 2008;

Holland et al., 2010). In many cases, the approach is used to

model a single ecosystem service, which is appropriate when it

is functionally somewhat separate from other services and

management actions are focused on that service. In other

cases, the services have sufficiently strong connections that a

multiple service approach is warranted. Both of these choices

are explored in the following sections.

Modeling Single Ecosystem Services: Bioeconomic Modelsof FisheriesA leading example of the modeling of a single ecosystem

services in marine systems is the modeling of fisheries

that incorporates both biological and economic components.

For some types of fishery management decisions, a narrow

focus on the single ecosystem service of food from fisheries

is appropriate. It also provides important lessons for eco-

system service modeling because a critical part of its devel-

opment has been the evolution of the human behavioral

portion of the models. For that reason, an extended discussion

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of this type of production function approach is provided in

this section.

Of all marine ecosystem service modeling, fisheries models

are by far the most sophisticated and have the longest history

of use in managing marine systems. Bioeconomic models

have been used since the early 1950s to explore how the

value generated by fisheries is affected by how they are ex-

ploited and regulated. Bioeconomic models have evolved

over time from simple models focused on equilibrium

outcomes associated with static annual harvests and effort

levels (e.g., Gordon, 1954; Schaefer, 1957) to more complex

dynamic models that explore the implications of the timing,

location, and methods of harvests, the linkages between

different species, and the impacts of fisheries on other eco-

system services. From the beginning, these models have em-

phasized the interconnection of the natural and human

components of fisheries and the importance of an approach to

modeling that treats human activity as an endogenous part of

the system.

It was recognized early on that understanding the optimal

exploitation of fisheries would generally require a dynamic

modeling approach since fisheries are rarely if ever in equi-

librium at some desired state. Scott (1955) introduced the

concept of user cost to illustrate the trade-offs between har-

vesting fish today and leaving fish in the water to contribute to

future growth. Viewing the fish stock as natural capital,

Scott showed that optimal exploitation would require the

rate of return from the in situ fish stock to equal the rate of

return from other uses of that capital (i.e., by extracting and

selling fish and thereby transforming them into economic

capital).

Although the initial focus of bioeconomic models was on

aggregate annual effort and harvests, Gordon’s (1954) analysis

demonstrated that the value generated by the fishery was also

dependent on the spatial distribution of fishing. The exploit-

ation of fisheries over space – and particularly the issue of

whether to close some areas to fishing to conserve fish stocks,

habitat, and biodiversity – became an important focus of

bioeconomic modeling in the late 1990s (e.g., Holland and

Brazee, 1996; Sanchirico and Wilen, 1999) and remains so

(Costello and Polasky, 2008). It is becoming increasingly clear

that spatial heterogeneity of marine ecosystems and of fishing

costs provide opportunities to increase the value generated by

fishing by managing where and when fishing takes place and

which harvest methods are employed. These models show that

optimal exploitation depends on relative costs over space in

conjunction with spatial variation in productivity (e.g., from

differences in suitability and productivity of habitat for species

and their prey or from metapopulations with source-sink

dynamics).

Another important focus of bioeconomic modeling that

has become more prevalent over time is the multispecies na-

ture of fisheries. Many multispecies fishery models focus on

the implications of technical interactions that make it difficult

and costly to control the relative catch of different species that

occupy the same habitat. These models provide insights into

how economic incentives or regulations on gear or fishing

locations can influence catch composition to increase the

value and sustainability of fisheries and reduce undesirable

bycatch. Trophic linkages between different fish species are

often thought to be important drivers of fishery productivity

and stability. However, they have largely been ignored in the

assessment of the productivity of fish stocks and when pro-

viding management advice (but see Hollowed et al., 2011).

This is not surprising: these relationships are typically poorly

understood and difficult to quantify.

Nonetheless, there is recognition that understanding

and responding to these species interactions can increase

value and improve the sustainability of fisheries. Ecosystem

models that attempt to depict entire food webs and the

impacts of bottom-up forcing (e.g., by climate) and top-down

forcing (e.g., predation and fishing) have been developed

in recent years; a few such as Atlantis (Fulton et al., 2004a,

2004b) include an endogenous human harvester component

to the model. These models offer the possibility of inform-

ing a more holistic ecosystem-based approach to managing

fisheries and other activities in the marine environment,

but their complexity and the lack of data to parameterize

them has so far limited their usefulness in providing

specific tactical advice for managing individual fisheries. An

increasing number of multispecies bioeconomic models focus

on small systems of exploited species (see, e.g., the discussion

in the section The American Lobster, Atlantic Herring, and

Northeast Multispecies Groundfish Fisheries in the Gulf of

Maine). For these smaller systems made up of primarily

commercial species, there is often more data and research,

making it more feasible to parameterize the models accurately

enough to quantitatively evaluate multispecies harvest

strategies.

Our ability to understand and model fishery systems in the

marine environment is hindered by an inability to observe

directly the processes that drive outcomes since they occur

under the sea and over vast areas. It is often the perturbations

caused by human exploitation of the system and the data

gathered by fishermen that enable us to develop an under-

standing of these systems that is sufficient to create empirically

based models capable of providing specific insights that can be

applied to the management of these systems as we seek sus-

tainable harvest levels. Bioeconometric models, defined by

Smith (2008) as structural models that econometrically esti-

mate one or more parameters of the bioeconomic system,

have the potential to reveal both ecological and economic

parameters of the system simultaneously and to greatly in-

crease our understanding of these systems at relatively low cost

(since they exploit data easily collected by fishermen in the

course of normal operations).

This field is not new as it includes empirical applications of

the Gordon-Schaefer model dating to the 1950s – but until

recently was mostly limited to equilibrium models based on

aggregate annual catch and effort. These models did not reveal

the underlying microecological and economic processes

driving the results and required making strong assumptions

about system behavior (e.g., that it tends toward equilibria). In

recent years, progress has been made in modeling and para-

meterizing these processes with dynamic models and dis-

aggregated data. This has made it more feasible to model and

predict how changes in the underlying ecological system or

changes in factors that drive human behavior will change

outcomes and the resulting flow of ecosystem services (e.g.,

Smith et al., 2008).

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Efforts to model a single ecosystem service – such as the

critical service of food from fisheries – and the ways in which

its delivery or value change under different scenarios are of

great utility when decisions are being made sector by sector or

when a better understanding of a key service sheds light on a

previously understudied aspect of a system. But with decision

makers increasingly grappling with multisector decisions,

modeling of multiple ecosystem services in the same context

can allow for more informed decision making.

Modeling Multiple Ecosystem Services and ExaminingTrade-OffsSingle-sector management that proceeds without the explicit

consideration of how single decisions impact the full suite

of things people care about and need can lead to misguided

assessments of the true costs and benefits to the environment

and society of natural resource management options. For

example, mangroves are routinely cleared and the resultant

open areas used for shrimp aquaculture. A singular focus

on aquaculture as an ecosystem service derived from cleared

areas often shows a positive bottom line, as the high

market price of shrimp provides strong support for this action

from a private perspective. Standing mangroves provide

other social and ecological benefits that private owners cannot

capture, however. More complete accounting using a multiple

ecosystem service framework shows that keeping mangroves

intact often has higher social benefits once other services

such as wood products, support for offshore fisheries, and

coastal protection are taken into account (Sathirathai and

Barbier, 2001). In other words, a single-sector approach risks

ignoring the multitude of connections among components of

natural and social systems. These connections are often im-

portant for the maintenance of ecosystem health, human well-

being, and the sector of interest itself (Millennium Ecosystem

Assessment, 2005b).

The explicit recognition of connections between activities

and their consequences for multiple ecosystem benefits allows

for the exploration of trade-offs and win–wins. Rodrıguez et al.

(2006) review some of the most frequent ecosystem service

trade-offs faced by society. In some cases, trade-offs and

win–wins can be explored using a common currency (e.g.,

dollars, Sathirathai and Barbier, 2001), but various metrics can

be used (Tallis et al., 2008, 2011; Lester et al., 2010). As one

example, the MA explored trade-offs among various categories

of ecosystem services (with axes for each service scaled from

� 1 to 1 to represent positive or negative change from the

baseline) for each of four heuristic scenarios (Millennium

Ecosystem Assessment, 2005a).

Explorations of multiple ecosystem services and how they

are likely to change under various management scenarios has

proceeded in two important directions: detailed explorations of

particular situations and the development of tools designed to

be applicable in various contexts. We focus here on tools. Re-

search teams have taken a number of different approaches to

modeling the flow of multiple ecosystem services and exam-

ining trade-offs between them. Here we provide a brief intro-

duction to some of the approaches and tools that are most

applicable to modeling marine ecosystem services (Table 2).

Modeling the flows of ecosystem services can take many

forms. Artificial Intelligence for Ecosystem Services (ARIES;

www.ariesonline.org) offers a range of approaches including

probabilistic Bayesian models, machine learning, and pattern

recognition to assess the provision, use, and flow of ecosystem

services on a landscape. The tool allows users to evaluate and

compare alternative policy and land-use scenarios and their

impacts on ecosystem services. The initial marine application

of ARIES is in Madagascar in partnership with the United

Nations Environment Program’s (UNEP’s) World Conser-

vation Monitoring Center, where a benefits-transfer approach

to ecosystem service valuation is being used (Center for Ocean

Solutions, 2011).

More process-based ecosystem service modeling ap-

proaches also have emerged. The challenges of applying the

land-use or land-cover map-based approach used in terrestrial

systems, coupled with the strong foundations of marine eco-

system modeling, has made ecosystem modeling central to the

ecosystem services framework for marine systems. Foodweb

models such as EcoPath with EcoSim (EwE) (Christensen

and Walters, 2004) and ecosystem models such as Atlantis

(Fulton et al., 2004a, 2004b) have been used to explore fishery

management options in an ecosystem context. These models

also have the potential to evaluate a more comprehensive

suite of human activities and the ways in which they

change the delivery of ecosystem services. Using these models,

explorations of the potential outcomes of various human

actions can lead to the development of management

approaches that consider multiple objectives and trade-offs

among objectives.

Integrated Valuation of Ecosystem Services and Trade-offs

(InVEST; http://www.naturalcapitalproject.org) helps decision

makers visualize the impacts of potential management

activities by modeling and mapping the delivery, distribution,

and economic value of terrestrial, freshwater, and marine

ecosystem services under alternative scenarios (for more

information, see Kareiva et al. (2011) and Guerry et al. (2012)).

InVEST can be used to identify trade-offs and compatibilities

between environmental, economic, and social benefits. Ap-

plicable at a range of scalesFfrom local to globalFInVEST

was designed to play a key role in real-world decision-making

processes. The first phase of the approach involves working

with stakeholders to identify critical management decisions

and to develop scenarios that project how the provision of

services might change in response to those decisions as well

as to changing climate, population, and so on. Using these

scenarios and basic biophysical and social data as inputs, In-

VEST quantifies and maps a broad range of ecosystem services.

To facilitate the use of InVEST in real decision-making con-

texts, a key design feature of InVEST is the relative simplicity

of data required and the free availability of models on the web.

InVEST outputs provide decision makers with information

about costs, benefits, trade-offs, and synergies of alternative

management strategies.

InVEST uses a primarily production function approach to

assess how environmental change, as a result of future policies

and management decisions, affects the delivery of ecosystem

services. Terrestrial InVEST models have been applied around

the world to inform a variety of land-use decisions (Kareiva

et al., 2011). InVEST’s marine tools (see Guerry et al., 2012)

were initially released in 2011 and are being applied in

Canada, the US, and Belize. Marine InVEST includes models

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Table 2 Decision support tools for use in marine environments, with particular attention to: marine applications, spatial mapping of ecosystem services, capacity to analyze trade-offs betweenservices, and the level of technical expertise needed to use the tool

Decision support tool/developer Description Marine applications Spatial mapping ofecosystemservices

Analysis ofecosystem servicetrade-offs

Level of technicalexpertise neededa

Artificial Intelligence for EcosystemServices (ARIES)

Basque Center for Climate Change (BC3);University of VermontFGund Institutefor Ecological Economics, ConservationInternational, and Earth Economics

ARIES is a suite of web-accessibleapplicationsFincluding probabilisticBayesian models, machine learning,and pattern recognitionF used toassess the provision, use, and flow ofecosystem services. ARIES maps boththe sources of ecosystem services andtheir users, along with the flows fromecosystems to users.

ARIES has been used to assesssubsistence fisheries and coastal stormregulation in Madagascar.

Yes Yes 2, 3

AtlantisCommonwealth Scientific and Industrial

Research Organization (CSIRO) Marineand Atmospheric Research

Atlantis is a three-dimensional, spatiallyexplicit model that incorporatesbiogeochemical dynamics and fishingbehavior. Submodels cover food-webrelations, hydrographic processes, andfisheries.

Atlantis has been used to evaluaterestructuring of Southeastern Australiafishing fleets, the NOAA IntegratedEcosystem Assessment for theCalifornia Current, the MarineStewardship Council Forage FishHarvest Guidelines, and groundfish fleetimpacts on protected marine mammalsin the California Current.

No Yes 3

Coastal ResilienceThe Nature Conservancy, University of

Southern Mississippi, and University ofCalifornia, Santa Barbara

Coastal Resilience is a spatial planningand action approach that integratesappropriate coastal hazard, ecological,and socioeconomic information within aparticular geography. The CoastalResilience approach is to map sea-levelrise and other coastal hazards, naturalresources, and human communities atrisk and display this information via aninternet mapping application that is adata viewer, data discovery tool, and afuture scenario mapper.

Coastal Resilience has been used for dataexploration with the New York StateEmergency Management Office andlocal towns and villages on Long Islandand the Connecticut shores.

No No 2

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Cumulative ImpactsNational Center for Ecological Analysis

and Synthesis (NCEAS), University ofCalifornia, Santa Barbara, and StanfordUniversity

The Cumulative Impacts model usesspatial data and expert opinion toassess the ecological consequences of17 types of human activities. Byoverlaying maps of human activitiesand ecosystem vulnerabilities, thismodel produces a cumulative mappingof ecological impacts.

Cumulative Impacts has been used in acoarse global analysis and in theCalifornia Current and the northwesternHawaiian Islands.

No No 2

InVESTThe Natural Capital ProjectFStanford

University, World Wildlife Fund, TheNature Conservancy, and the Universityof Minnesota

InVEST is composed of a number ofmodels for assessing flows of andchanges in different ecosystem servicesincluding, but not limited to, carbonstorage, wave energy, recreation,fishery production, erosion control,habitat quality, water quality, croppollination, and timber production.InVEST is a toolbox in ArcGIS and runson both spatial and nonspatial physical,biological, and economic data andinformation. (An ArcGIS-independentversion is forthcoming.)

InVEST has been used for marineapplications in Canada (west coast ofVancouver Island) and Belize, and forland–sea connections in Puget Sound,Galveston Bay, and Chesapeake Bay(US). In addition, it is being used inclimate adaptation planning and toinform restoration in the Gulf of Mexicoand in Monterey Bay (both US).

Yes Yes 1, 3

Marine MapMarineMap ConsortiumFUniversity of

California, Santa Barbara, The NatureConservancy, and EcoTrust

MarineMap is a web-based decisionsupport toolkit to support marinespatial planning processes. The toolkitincludes a spatial data viewer anddesign tools that allow users tonetworks of prospective marineprotected areas. The application alsoallows users to share their proposalswith others and evaluate their proposalsagainst goals defined in the course ofany planning process.

MarineMap has been used for theCalifornia MLPA Initiative and theOregon Territorial Sea Planning process(US).

No Yes 1

Marxan with zonesUniversity of Queensland Marxan is a program for identifying

combinations of sites to createconservation networks such as marineprotected areas. The program allowsthe user to set biodiversity and other

Marxan has been applied to MarineZoning in Saint Kitts and Nevis and toexamine four types of protected areasin the context of California’s Marine LifeProtection Act.

No Yes 2, 3

(Continued )

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Table 2 Continued

Decision support tool/developer Description Marine applications Spatial mapping ofecosystemservices

Analysis ofecosystem servicetrade-offs

Level of technicalexpertise neededa

targets for one or more zones and thenfind the set of sites that meets thosetargets while minimizing the cost of thenetwork.

Multiscale Integrated Models ofEcosystem Services (MIMES)

AFORDable futures MIMES is a multiscale integrated suite ofmodels that incorporate stakeholderinput and a variety of data sets toassess trade-offs among multipleecosystem services. The modelssimulate ecological and socioeconomicsystems and their interactions, and theycalculate the values of ecosystemservices for different scenarios.

MIMES is being used by theMassachusetts Ocean Partnership toexamine the trade-offs betweendifferent sectors in spatial planning andto model ecosystem service values atmultiple scales.

Yes Yes 2, 3

Multipurpose Marine Cadastre (MMC)Bureau of Ocean Energy Management,

Regulation and Enforcement(BOEMREFformerly MineralsManagement Service) and NOAACoastal Services Center

Originally developed to support theassessment of offshore energyprojects, MMC is a web-basedgeospatial data viewer containing morethan 80 data layers from a variety ofsources that each can be turned on oroff or queried one at a time. The userhas the ability to draw lines and otherfeatures, create buffers, and performother geospatial actions.

MMC has been used for reviews of oceanenergy projects in Northern Californiaand the outer continental shelf offMassachusetts, and by the Mid-AtlanticRegional Council on the Ocean tosupport marine spatial planning efforts.

No No 1

aLevels of technical expertise (when more than one level is listed the tool has capabilities that require varying levels of expertise):

1. Minimal training or technical expertise.

2. Minimal training and expertise but process objectives must be set in advance.

3. Expert users.

Source: Adapted from Center for Ocean Solutions (2011) Decision Guide: Selecting Decision Support Tools for Marine Spatial Planning. Stanford, CA: The Woods Institute for the Environment, Stanford University.

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Input data (Reflect scenarios) Marine InVEST modelsModel outputs

(Ecosystem services and values)

Terrestrial systems

Habitatrisk

Carbon Carbonsequestered

Ecosystemservices

Valuatione.g.

Value ofcarbon

sequestered

Value ofcaptured

wave energy

Value ofavoided

damages

Expendituresdue to

recreationactivity

Net presentvalue of

finfish andshellfish

Energycaptured

AvoidedArea flooded/

eroded

Visitationrates

Landedbiomass

Harvestedbiomass

Wave energy

Coastalprotection

Recreation

Fishery

Aestheticquality

Aquaculture

Waterquality

Bio-physical

Sce

nario

s

Bathymetry and topography

Speciesdistribution

Oceanography

Habitat type

Socio-economic

Population density

Demographics

Aquaculture operation costs

Property values

3

5

7

1 8

6

2

4

9

Figure 2 Marine InVEST evaluates how alternative scenarios yield changes in the flow of ecosystem services. First, one translates managementor climate scenarios into input data. Inputs include spatially explicit biophysical and socioeconomic information. Next, one feeds input maps intomodels that predict the delivery of services across the seascape. Intermediate effects of management choices and climate on the flow of servicescan be evaluated in terms of risks to habitats and changes in water quality. Ecosystem service outputs are expressed in biophysical orsocioeconomic units. Some examples of processes or activities that link the models include (numbers correspond to circled numbers on arrows):(1) flow rate and the movement of sediment, nutrients, toxic waste and bacteria; (2) filtration, flows of waste; (3) limiting fishing grounds; (4)light attenuation, sedimentation; (5) water purification; (6) food supply (shellfish); (7) beachgoing; (8) spawning, rearing; and (9) waveattenuation. Adapted from Guerry AD, Ruckelshaus MH, Arkema KK, et al. (2012) Modeling benefits from nature: using ecosystem services toinform marine spatial planning. International Journal of Biodiversity Science, Ecosystem Services and Management. doi: 10.1080/21513732.2011.647835.

Modeling Marine Ecosystem Services 341

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for renewable energy, protection from coastal hazards,

fisheries, recreation, aquaculture, aesthetic views, and more

(Figure 2).

The Multiscale Integrated Models of Ecosystem Services

(MIMES) project represents another attempt to build a suite

of models that assess the values of ecosystem services to

allow managers to understand the dynamics of ecosystem

services, how services are linked to human welfare, and

how the flow of services might change under various man-

agement scenarios in both terrestrial and marine systems

(http://www.uvm.edu/giee/mimes/). MIMES involves a rela-

tively complex approach, simulating ecosystems and socio-

economic systems and the interactions between them. It

calculates values of ecosystem services using marginal cost

pricing. MIMES is currently being used by the Massachusetts

Ocean Partnership to evaluate the trade-offs between different

sectors in marine spatial planning (Center for Ocean Solu-

tions, 2011).

Applications of Marine Ecosystem Service Modelingto Decision Making

Decision contexts for managing ecosystem services in the

sea are varied, so it is to be expected that a mix of quantitative

and qualitative modeling approaches are being developed

and applied to support biodiversity and ecosystem service

management in marine systems. Information needed to

support the methodologies we highlight in the following

sections ranges from observations and empirical data to

modeled ecosystem and social responses, expert opinion and

traditional local knowledge (e.g., Kliskey et al., 2009). Typi-

cally, the time frames of decisions, local technical capacity,

and the nature of information available for each context

dictate the approach employed for ecosystem service model-

ing. We explore four contexts in the following section that

display a wide range of the numbers of services modeled,

quantitative complexity, and types of decisions being made.

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We start where marine modelers and managers are most at

home together – with a fisheries-focused example – and then

move to efforts to include a number of different types of

services.

The American Lobster, Atlantic Herring, and NortheastMultispecies Groundfish Fisheries in the Gulf of Maine

The American lobster, Atlantic herring, and Northeast

multispecies groundfish fisheries in the Gulf of Maine provide

an excellent example of a system of interlinked fisheries with

a value that might be substantially increased by accounting

for linkages between them and with the environment. A

number of important environmental, ecological, and eco-

nomic linkages within these fisheries and with the environ-

ment were identified as key processes to study and

model (Figure 3). This system is the subject of an inter-

disciplinary research project funded by the National Science

Foundation aimed at understanding these linkages through

empirical research and modeling to provide insights into

how the management of each fishery and the system as

a whole can be improved. The strategy of this research effort

is to link and build on the single-species models that are

already used for providing management advice by modeling

key linkages between the fisheries and with the enviro-

nment that might be exploited to increase value and

sustainability.

Key processes to include in coupl

Environmental-early life history

• Impacts of temperature, salinity, winds onrecruitment and juvenile growth

Trophic interactions and compe• Predation of groundfish on he

• Predation of herring on ground

• Herring bait growth subsidy fo• Lobster behavior modificationgroundfish predators

Cod recruits

Herring recruits

Herring juveniles

Herring adults

Lobster settlementLobster juveniles

Lobster adults

Cod juveniles

Cod adults

Environmentwind

currentstemperature

phytoplanktonzooplankton

Figure 3 Linkages between lobster, herring, and groundfish fisheries in th

The primary focus for environmental linkages in this system

is on impacts of temperature, salinity, and winds on recruit-

ment and juvenile growth. Changes in the abundance of larval

food and changes in circulation driven by wind and currents are

thought to be key drivers of recruitment variability for all three

species. Determining what drives recruitment success requires a

combination of empirical fieldwork (determining where and

how much spawning and settlement take place) and coupled

biophysical models (evaluating how currents, winds, and

temperatures affect dispersal and settlement of larvae and

consequent recruitment). In addition to circulation and dis-

persion, model calculations may need to consider spatial pat-

terns of egg production, temporal patterns of hatching,

temperature-dependent development, vertical distribution, and

mortality (see Incze et al., 2010, and Churchill et al., 2011, for

examples that are relevant to this system).

A variety of ecological linkages, natural and artificial, are

being studied, including predation of groundfish on herring

and juvenile lobsters and predation of herring on groundfish

larvae. In addition to predator–prey linkages, the team is

studying behavior modification as a result of diminished

presence of groundfish predators that may have allowed lob-

ster to expand use of habitat and consequently carrying

capacity.

This fishery system is somewhat unusual, though perhaps

not unique, in that a key linkage between the fisheries is

controlled by humans. The artificial trophic linkage resulting

from use of herring bait in quantities (exceeding 50,000 tons

ed system model

titionrring and juvenile lobsters

fish larvae

r lobster with absence/pressure of

GroundfishFishery

Lobsterfishery Lobster fleet

Herring fleet

Groundfish fleet

Herringfishery

• Movement of labour/capital betweenfisheries• Dependence of lobster fishery on herringbait (primary herring market)

Economic connections

Baitmarket

e Gulf of Maine.

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Modeling Marine Ecosystem Services 343

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annually in recent years) is thought to be responsible for a

substantial increase in productivity of the lobster stock

(Grabowski et al., 2010). The way that the lobster fishery

is managed substantially alters the amount of herring bait

demanded by the fishery, which affects profitability of both

fisheries (Holland et al., 2010) and how those profits are dis-

tributed (Ryan et al., 2010).

Understanding these linkages among the fisheries and the

environment provides useful qualitative insights for resource

managers and stakeholders, and it may be possible to use

this information to improve single-species models and man-

agement advice to account for them as exogenous factors.

Ideally, a linked dynamic set of fishery and environmental

models will be constructed enabling managers to better

understand how changes in one part of the system affect other

parts and how best to react or even manipulate the system.

In addition to evaluating potential management actions or

forecasting future outcomes in the fishery system, models of

the system can also be used to determine which variables and

processes have the most impact on the system. This knowledge

can be used to determine the value of better information and

direct research. Ultimately, this information can be used to

manage these three key fisheries in a systemwide manner that

maximizes the service of provisioning of seafood and the

various benefits that flow from that service (e.g., revenues,

working waterfronts, community identity).

Coastal Zone Management in Belize

Belize is home to a rich diversity of ocean life, coastal habitats,

and the longest barrier reef in the Western hemisphere. Its

people are inextricably linked to the marine environment as a

source of sustenance, inspiration, economic prosperity, and

cultural heritage (Figure 4). Yet rapid development, overfishing,

and population growth threaten local marine ecosystems. In

Figure 4 Belize is home to a rich diversity of marine life; its peoplederive sustenance, inspiration, economic prosperity, and culturalheritage from the marine environment. Mangroves and coral reefs notonly provide protection from storms and serve as a draw forinternational tourism, but also are critical habitats for the spinylobster, a backbone of commercial and artisanal fisheries. Thecommercial fishermen in this picture are from a fleet of dugoutcanoes that work with a traditional sail boat (or ‘‘mother ship’’’) toharvest spiny lobster.

1998, the government created the Coastal Zone Management

Authority and Institute (CZMAI) with the objective of de-

veloping an integrated management plan to guide sustainable

economic growth that is consistent with protection of Belize’s

natural heritage. To date, however, CZMAI has lacked the sci-

entific capacity needed to assess trade-offs among various

marine uses and to demonstrate potential win–wins where

emphasis on one service yields benefits for another.

CZMAI has partnered with the Natural Capital Project to

help create a comprehensive coastal zone management plan

for Belize. The plan will identify new marine protected areas,

locations suitable for coastal development, and strategies such

as payments for ecosystem services (PES) and other market

mechanisms (e.g., catch shares) to support effective imple-

mentation. The partners are using Marine InVEST to forecast

how management strategies and future uses of the marine

environment will likely affect the benefits that nature brings to

people, such as nursery habitat for fisheries, tourism and rec-

reation, coastal protection, and carbon storage. By providing a

platform for stakeholder and agency discussions about trade-

offs and win–win situations, this information is moving the

process beyond sector-specific issues to the development of a

defensible plan.

The first step toward developing a coastal zone plan in-

formed by the science of ecosystem services is to identify and

map current and potential future uses of marine and coastal

environments. Next is to use modeling approaches (in this

case, the InVEST decision support tool) to examine how these

activities affect the ecosystem services most important to

Belizeans. The team is running marine InVEST models, given

future scenarios of marine use, to quantify changes in the

ecosystem services that are most compelling to stakeholders

and government officials. Scenarios under consideration in-

clude explorations of how coastal development and no-take

areas – in addition to climate change – might affect services,

including (1) provision of commercial and artisanal fisheries

for lobster; (2) flooding and erosion protection provided by

mangroves, corals, and seagrasses; and (3) maintenance of

tourism attractions such as snorkeling and diving. Quantita-

tive outputs in biophysical (e.g., biomass of harvested fish,

reduction in land flooded or eroded due to mangroves) and

economic (e.g., net present value of harvest, avoided damages

to property values) units for each service are informing

CZMAI’s coastal planning process.

Modeling of ecosystem services in Belize is helping to ar-

ticulate connections between human activities that are often

considered in isolation to align diverse stakeholders around

common goals and to make implicit decisions explicit. Eco-

system service modeling results have informed early iterations

of the coastal zone plan and will inform the creation of the

final plan in 2012.

Protection and Restoration of Reef Habitats in the CoralTriangle

Klein et al. (2010) used a combination of quantitative mod-

eling and qualitative information to prioritize strategies for

protection and restoration of reef habitats in the coral triangle.

The method is designed to allocate a limited budget to man-

agement and policy interventions that will abate key threats

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344 Modeling Marine Ecosystem Services

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affecting marine ecosystem objectives. The approach requires

estimates of management costs and opportunity costs of ap-

plying alternative actions; and these estimates are user-gener-

ated, so they can come from expert judgment, models, or

empirical information. The models provide an estimate of the

return on investment for different actions so that the relative

merits of land- and marine-based interventions on marine

ecosystem biodiversity and services can be weighed. This type

of practical approach, where a mix of quantitative and expert

information is used, allows the users to consider both eco-

nomic and ecological criteria in a conservation management

context. For example, for the ecoregions of the coral triangle,

marine-based conservation efforts (e.g., establishment of

MPAs, changes in fishing gear types) were often more cost-

effective at improving marine ecosystem condition than land-

based actions designed to reduce nutrient or sediment runoff.

Putting Offshore Wind in Context in Massachusetts

Like many coastal marine ecosystems, the coast of Massa-

chusetts (US) is crowded with various user groups interested

in using a broad range of ecosystem services. Conflicts among

sectors in their ability to procure different services from

shared, interacting ecosystem resources has prompted calls for

the reduction of conflicts through transparent, integrated

marine spatial planning. In Massachusetts and elsewhere,

offshore wind farms are an emerging, and controversial, ocean

use. Numerous wind farms are under litigation or consider-

ation in the state. Large economic gains and ‘‘green’’ energy are

purported benefits; impacts on marine mammals and fisheries

are some of the potential costs. To explore these issues and

inform the dialogue about this new and emerging use of the

marine environment, White and colleagues (in press) per-

formed a spatially explicit bioeconomic trade-off analysis

among wind energy, fishery, and whale-watching ecosystem

services in Massachusetts Bay. They identified optimal plan-

ning solutions for wind-energy development that minimize

spatial conflicts among sectors and maximize their values.

Solutions that considered all three sectors were significantly

better than single-sector management outcomes. Solutions

were also strongly dependent on the spatial design of the wind

farms and the particular fishery examined. This approach of

quantifying ecosystem service trade-offs in a crowded, multi-

use ecosystem highlights when and how offshore renewable

energy may be developed optimally within the complicated

contexts of coastal ecosystems.

Conclusions

Marine ecosystems around the globe are under increasing

pressure: people rely on them for the delivery of provisioning,

regulating, supporting, and cultural ecosystem services. With

increasing pressure from increasing and redistributing human

populations and from new and emerging uses (e.g., renewable

energy), the ability of these systems to sustainably deliver

the full range of services that people count on and need is not

assured.

Here we have reviewed the services provided by marine

environments, discussed some of the potential audiences

for information about how changes in marine ecosystems

are likely to lead to changes in services, explored critical

differences – both in human and scientific contexts – between

mapping and modeling ecosystem services on land and at sea,

outlined approaches to modeling marine ecosystem services,

and described some examples of how these approaches can

and are being used in real decision making.

Quantifying, mapping, and valuing marine ecosystem ser-

vices have the potential to fundamentally change decision

making in marine and coastal environments. Making explicit

the connections between human activities in one sector and

their effects on a broad range of other sectors leads people to

think about whole ecosystems and to manage them accord-

ingly. Ultimately, results from marine ecosystem service

modeling can help society perceive the critical services oceans

and coasts provide, appropriately value marine natural capital,

and help human communities make better choices about the

use of these life-sustaining environments.

Appendix

List of Courses

1. Marine Ecosystem Services

2. Marine Conservation Biology

3. Marine Ecology

4. Marine Resource Economics

See also: Aquaculture. Biodiversity and Ecosystem Services.Biodiversity, Human Well-Being, and Markets. Carbon Cycle. Censusof Marine Life. Computer Systems and Models, Use of. EconomicValue of Biodiversity, Measurements of. Economics of the RegulatingServices. Ecosystem, Concept of. Ecosystem Function Measurement,Aquatic and Marine Communities. Ecosystem Function, Principles of.Ecosystem Services. Endangered Marine Invertebrates. EstuarineEcosystems. Evaluation of Ecosystem Service Policies fromBiophysical and Social Perspectives: The Case of China. FishConservation. Mangrove Ecosystems. Marine Conservation in aChanging Climate. Marine Ecosystems, Human Impacts on. MarineEcosystems. Marine Protected Areas: Static Boundaries in a ChangingWorld. Modeling Terrestrial Ecosystem Services. Ocean Ecosystems.Priority Setting for Biodiversity and Ecosystem Services. ResourceExploitation, Fisheries. The Value of Biodiversity. Valuing EcosystemServices

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