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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. Author's personal copy Modeling Marine Ecosystem Services Anne D Guerry and Mary H Ruckelshaus, Stanford University, Stanford, CA, USA Mark L Plummer and Dan Holland, National Oceanic and Atmospheric Administration, Seattle, WA, USA r 2013 Elsevier Inc. All rights reserved. Glossary Ecosystem 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 Introduction to Marine Ecosystem Services Humans have always benefited from marine ecosystems, enjoying resources such as seafood and opportunities for activities like recreation and marine transportation. These ecosystems 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 Assessment (MA) (2005b), marine ecosystem services span four major categories: provisioning, regulating, cultural, and supporting 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., Holmlund 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, provisioning services such as food from capture fisheries, aquaculture, 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 approximately 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 proportion 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 offshore oil and gas platforms in California’s Santa Barbara Channel had potential for biomedical applications, including Encyclopedia of Biodiversity, Volume 5 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. two that inhibit the division of cancer cells grown in the laboratory (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 sequestration 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 communities 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 approximately 40% of global net primary productivity (Schlesinger, 1991; Melillo et al., 1993). The oceans are home to vast reservoirs of genetic and ecological diversity, arguably the most fundamental source of supporting services as they are directly http://dx.doi.org/10.1016/B978-0-12-384719-5.00333-6 329 Author's personal copy 330 Table 1 Modeling Marine Ecosystem Services Ecosystem services provided by marine systems Types of service Provisioning services Food production (capture fisheries; aquaculture; and wild foods) Fiber production Biomass fuel production Maintenance of aquatic systems Generation of genetic resources Production of biochemicals, natural medicines, and pharmaceuticals Regulating services Climate regulation Water regulation Erosion regulation Water purification and waste treatment Disease regulation Pest regulation Pollination/Assistance of external fertilization Natural hazard regulation Cultural services Provision of conditions that support or enhance ethical values (nonuse) Provision of conditions that support or enhance existence values (nonuse) Provision of recreation and ecotourism opportunities (consumptive and nonconsumptive uses) Supporting services Nutrient cycling Soil formation Primary production Water cycling Examples Tuna, crab, lobster; salmon, oysters, shrimp, seaweed; mussels, clams Mangrove wood, seagrass fiber Mangrove wood, biofuel from algae Shipping, tidal turbines Individual salmon stocks, marine diversity for bioprospecting Antiviral and anticancer drugs from sponges, carrageenans from seaweed Major role in global CO2 cycle Natural stormwater management by coastal wetlands and floodplains Nearshore vegetation stabilizes shorelines Uptake of nutrients from sewage wastewater, detoxification of PAHs by marine microbes, sequestration of heavy metals Natural processes may keep harmful algal blooms and waterborne pathogens in check Grazing fish help keep algae from overgrowing coral reefs Innumerable marine species require seawater to deliver sperm to egg Coastal and estuarine wetlands and coral reefs protect coastlines from storms Spiritual fulfillment derived from estuaries, coastlines, and marine waters Belief that all species are worth protecting, no matter their direct value to humans Scuba diving, beachcombing, whale watching, boating, snorkeling; fishing, clamming Major role in carbon, nitrogen, oxygen, phosphorus, and sulfur cycles Many salt-marsh surfaces vertically accrete; eelgrass slows water and traps sediment Significant portion of global net primary productivity 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. 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 agricultural production has increased between 1950 and the present, 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 Assessment, 2011). Protection of UK coastlines from storminduced 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 Author's personal copy Modeling Marine Ecosystem Services 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. Because ecosystem services are reviewed elsewhere in this volume, 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 natures 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 ecosystembased 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 modeling, such a framework avoids the potential problem of double counting the value of ‘‘intermediate’’ ecological elements (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 consider 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 resolve those trade-offs can be made transparently. Increasingly, scientists and managers are pointing to links between diversity, 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 activities) is needed in order to inform more complex cases 331 of ecosystem-based management that can accommodate a broader suite of actors (Foley et al., 2010). Pioneering efforts to use the framework of ecosystem services 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 Services (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 emphasis on marine (or ‘‘maritime’’) spatial planning. This approach 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 combination of ecological functions and human actions and values (Figure 1). In the marine environment, for example, marine ecosystems have fish populations that offer opportunities 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 Author's personal copy 332 Modeling Marine Ecosystem Services Figure 1 The production of ecosystem services involves a combination of ecological functions and human actions and values. Recreational fishing as an ecosystem service, for example, depends on the healthy functioning of natural systems that support fish populations. The human action of harvest transforms the potential ecological supply into the actual provision of the ecosystem service. 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 adjustments 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 interactions 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 different management approaches (e.g., regulation, incentives) on ecological and ecosystem services (e.g., lobster biomass, landings, biodiversity) and social (e.g., efficiency, equity, accountability) 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 outcomes 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 modeling 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 ecosystem are direct and local or indirect and global can be a key feature influencing how humans and ecosystem services respond to signs of change in the ecosystem, and it is an increasingly important feature with the globalization of both human and natural processes. One modeling approach that incorporates these frameworks is the integrated ecosystem assessment (IEA). An IEA outlines both a decision analytical framework and a process containing the logical steps necessary for managing ecosystems, including explicit identification of objectives and the indicators for tracking progress, a risk analysis describing the state of the indicators, an evaluation of management strategies, 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; Sainsbury et al., 2000; Smith et al., 2007); they are designed to be iterative, expanding in scope and sophistication as Author's personal copy Modeling Marine Ecosystem Services 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 management 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 conditions. 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 compare alternative policies. Challenges in Modeling Marine Ecosystem Services From a scientific perspective, the modeling of marine ecosystem 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 systems 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 management 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, associations between species and habitat patches are more difficult to discern. The lack of basic land-use and land-cover data for marine systems underscores the need for a modeling approach 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. 333 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 example, 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 observations 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 developing 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 landscape or seascape type. Drawing from a standard set of ecosystem 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. Author's personal copy 334 Modeling Marine Ecosystem Services 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 assigning 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 sequestration 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 ecosystem services for 16 biomes. Their effort included five marine 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 important 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 pressure 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 estimated for a small change has the potential to produce significant bias. And if more than one service is evaluated, the aggregate value of the group of services is likely to be nonlinear 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 problems grow more and more significant as the scale of the ecosystem 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 particular 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 information 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 ecosystem 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 determinants 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 Methods Standard 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 behavior) (Slesnick, 1998). The revealed preference approach is a collection of methods for estimating economic values that rely on observable behavior. The most obvious revealed preference 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 analyzes goods (e.g., housing) that are sold as a bundle of Author's personal copy Modeling Marine Ecosystem Services 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 Methods Stated preference methods also can provide legitimate estimates 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 environmental quality because they offer the opportunity to estimate 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 Methods Other methods are less reliable and particularly prone to misuse when estimating ecosystem service values. A prominent example is the replacement-cost or cost-of-treatment 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 wetland may not actually provide filtration services (e.g., because of the absence of contaminants that need filtering or biophysical 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. Economic valuation, however, is far from the only way to value ecosystem services. Although many services (e.g., provisioning 335 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 processbased. It represents the relationship between inputs (e.g., the density of mangroves) and outputs (e.g., the degree of protection 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 information 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; Kareiva 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 Models of Fisheries A 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 ecosystem service modeling because a critical part of its development has been the evolution of the human behavioral portion of the models. For that reason, an extended discussion Author's personal copy 336 Modeling Marine Ecosystem Services 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 exploited 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 ecosystem services. From the beginning, these models have emphasized 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 equilibrium at some desired state. Scott (1955) introduced the concept of user cost to illustrate the trade-offs between harvesting 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 exploitation 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 nature 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 providing 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 informing 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 understanding 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 sustainable harvest levels. Bioeconometric models, defined by Smith (2008) as structural models that econometrically estimate 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 increase 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 parameterizing these processes with dynamic models and disaggregated 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). Author's personal copy Modeling Marine Ecosystem Services 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 Examining Trade-Offs Single-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 important for the maintenance of ecosystem health, human wellbeing, 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. Rodrı́guez 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. Research teams have taken a number of different approaches to modeling the flow of multiple ecosystem services and examining trade-offs between them. Here we provide a brief introduction 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; 337 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 Conservation 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 approaches 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 ecosystem 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. Applicable 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, InVEST quantifies and maps a broad range of ecosystem services. To facilitate the use of InVEST in real decision-making contexts, 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 Author's personal copy Artificial Intelligence for Ecosystem Services (ARIES) Basque Center for Climate Change (BC3); University of VermontFGund Institute for Ecological Economics, Conservation International, and Earth Economics Atlantis Commonwealth Scientific and Industrial Research Organization (CSIRO) Marine and Atmospheric Research Coastal Resilience The Nature Conservancy, University of Southern Mississippi, and University of California, Santa Barbara Description Marine applications Spatial mapping of ecosystem services Analysis of ecosystem service trade-offs Level of technical expertise neededa ARIES is a suite of web-accessible applicationsFincluding probabilistic Bayesian models, machine learning, and pattern recognitionF used to assess the provision, use, and flow of ecosystem services. ARIES maps both the sources of ecosystem services and their users, along with the flows from ecosystems to users. ARIES has been used to assess subsistence fisheries and coastal storm regulation in Madagascar. Yes Yes 2, 3 Atlantis is a three-dimensional, spatially explicit model that incorporates biogeochemical dynamics and fishing behavior. Submodels cover food-web relations, hydrographic processes, and fisheries. Atlantis has been used to evaluate restructuring of Southeastern Australia fishing fleets, the NOAA Integrated Ecosystem Assessment for the California Current, the Marine Stewardship Council Forage Fish Harvest Guidelines, and groundfish fleet impacts on protected marine mammals in the California Current. No Yes 3 Coastal Resilience is a spatial planning and action approach that integrates appropriate coastal hazard, ecological, and socioeconomic information within a particular geography. The Coastal Resilience approach is to map sea-level rise and other coastal hazards, natural resources, and human communities at risk and display this information via an internet mapping application that is a data viewer, data discovery tool, and a future scenario mapper. Coastal Resilience has been used for data exploration with the New York State Emergency Management Office and local towns and villages on Long Island and the Connecticut shores. No No 2 Modeling Marine Ecosystem Services Decision support tool/developer 338 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 between services, and the level of technical expertise needed to use the tool Author's personal copy Cumulative Impacts National Center for Ecological Analysis and Synthesis (NCEAS), University of California, Santa Barbara, and Stanford University InVEST The Natural Capital ProjectFStanford University, World Wildlife Fund, The Nature Conservancy, and the University of Minnesota Marxan with zones University of Queensland Cumulative Impacts has been used in a coarse global analysis and in the California Current and the northwestern Hawaiian Islands. No No 2 InVEST is composed of a number of models for assessing flows of and changes in different ecosystem services including, but not limited to, carbon storage, wave energy, recreation, fishery production, erosion control, habitat quality, water quality, crop pollination, and timber production. InVEST is a toolbox in ArcGIS and runs on both spatial and nonspatial physical, biological, and economic data and information. (An ArcGIS-independent version is forthcoming.) InVEST has been used for marine applications in Canada (west coast of Vancouver Island) and Belize, and for land–sea connections in Puget Sound, Galveston Bay, and Chesapeake Bay (US). In addition, it is being used in climate adaptation planning and to inform restoration in the Gulf of Mexico and in Monterey Bay (both US). Yes Yes 1, 3 MarineMap is a web-based decision support toolkit to support marine spatial planning processes. The toolkit includes a spatial data viewer and design tools that allow users to networks of prospective marine protected areas. The application also allows users to share their proposals with others and evaluate their proposals against goals defined in the course of any planning process. MarineMap has been used for the California MLPA Initiative and the Oregon Territorial Sea Planning process (US). No Yes 1 Marxan is a program for identifying combinations of sites to create conservation networks such as marine protected areas. The program allows the user to set biodiversity and other Marxan has been applied to Marine Zoning in Saint Kitts and Nevis and to examine four types of protected areas in the context of California’s Marine Life Protection Act. No Yes 2, 3 Modeling Marine Ecosystem Services Marine Map MarineMap ConsortiumFUniversity of California, Santa Barbara, The Nature Conservancy, and EcoTrust The Cumulative Impacts model uses spatial data and expert opinion to assess the ecological consequences of 17 types of human activities. By overlaying maps of human activities and ecosystem vulnerabilities, this model produces a cumulative mapping of ecological impacts. (Continued ) 339 Author's personal copy 340 Table 2 Continued Decision support tool/developer Description Spatial mapping of ecosystem services Analysis of ecosystem service trade-offs Level of technical expertise neededa MIMES is a multiscale integrated suite of models that incorporate stakeholder input and a variety of data sets to assess trade-offs among multiple ecosystem services. The models simulate ecological and socioeconomic systems and their interactions, and they calculate the values of ecosystem services for different scenarios. MIMES is being used by the Massachusetts Ocean Partnership to examine the trade-offs between different sectors in spatial planning and to model ecosystem service values at multiple scales. Yes Yes 2, 3 Originally developed to support the assessment of offshore energy projects, MMC is a web-based geospatial data viewer containing more than 80 data layers from a variety of sources that each can be turned on or off or queried one at a time. The user has the ability to draw lines and other features, create buffers, and perform other geospatial actions. MMC has been used for reviews of ocean energy projects in Northern California and the outer continental shelf off Massachusetts, and by the Mid-Atlantic Regional Council on the Ocean to support marine spatial planning efforts. No No 1 targets for one or more zones and then find the set of sites that meets those targets while minimizing the cost of the network. Multiscale Integrated Models of Ecosystem Services (MIMES) AFORDable futures Multipurpose Marine Cadastre (MMC) Bureau of Ocean Energy Management, Regulation and Enforcement (BOEMREFformerly Minerals Management Service) and NOAA Coastal Services Center a Levels of technical expertise (when more than one level is listed the tool has capabilities that require varying levels of expertise): 1. 2. 3. Minimal training or technical expertise. Minimal training and expertise but process objectives must be set in advance. 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. Modeling Marine Ecosystem Services Marine applications Author's personal copy Modeling Marine Ecosystem Services Input data (Reflect scenarios) Marine InVEST models Model outputs (Ecosystem services and values) Ecosystem services Carbon 341 Carbon sequestered Valuation e.g. Value of carbon sequestered Terrestrial systems Wave energy Energy captured 3 Value of captured wave energy Bio-physical 9 Scenarios Bathymetry and topography Species distribution Oceanography Coastal protection Habitat risk 5 Recreation 4 Avoided Area flooded/ eroded Visitation rates 7 Habitat type Water quality 1 8 Fishery Landed biomass Socio-economic 6 Population density Aquaculture 2 Harvested biomass Value of avoided damages Expenditures due to recreation activity Net present value of finfish and shellfish Demographics Aquaculture operation costs Property values Aesthetic quality Figure 2 Marine InVEST evaluates how alternative scenarios yield changes in the flow of ecosystem services. First, one translates management or climate scenarios into input data. Inputs include spatially explicit biophysical and socioeconomic information. Next, one feeds input maps into models that predict the delivery of services across the seascape. Intermediate effects of management choices and climate on the flow of services can be evaluated in terms of risks to habitats and changes in water quality. Ecosystem service outputs are expressed in biophysical or socioeconomic 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) wave attenuation. Adapted from Guerry AD, Ruckelshaus MH, Arkema KK, et al. (2012) Modeling benefits from nature: using ecosystem services to inform marine spatial planning. International Journal of Biodiversity Science, Ecosystem Services and Management. doi: 10.1080/ 21513732.2011.647835. 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 management scenarios in both terrestrial and marine systems (http://www.uvm.edu/giee/mimes/). MIMES involves a relatively complex approach, simulating ecosystems and socioeconomic 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 Solutions, 2011). Applications of Marine Ecosystem Service Modeling to 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). Typically, the time frames of decisions, local technical capacity, and the nature of information available for each context dictate the approach employed for ecosystem service modeling. 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. Author's personal copy 342 Modeling Marine Ecosystem Services 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 Northeast Multispecies 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 economic linkages within these fisheries and with the environment were identified as key processes to study and model (Figure 3). This system is the subject of an interdisciplinary 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 environment that might be exploited to increase value and sustainability. The primary focus for environmental linkages in this system is on impacts of temperature, salinity, and winds on recruitment 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 dispersion, model calculations may need to consider spatial patterns 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 lobster 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 Key processes to include in coupled system model Trophic interactions and competition • Predation of groundfish on herring and juvenile lobsters • Impacts of temperature, salinity, winds on • Predation of herring on groundfish larvae recruitment and juvenile growth • Herring bait growth subsidy for lobster • Lobster behavior modification with absence/pressure of groundfish predators Environmental-early life history Economic connections • Movement of labour/capital between fisheries • Dependence of lobster fishery on herring bait (primary herring market) Cod recruits Cod juveniles Cod adults Environment wind currents temperature phytoplankton zooplankton Groundfish Fishery Groundfish fleet Herring recruits Herring juveniles Herring adults Herring fishery Herring fleet Bait market Lobster settlement Lobster juveniles Lobster adults Lobster fishery Figure 3 Linkages between lobster, herring, and groundfish fisheries in the Gulf of Maine. Lobster fleet Author's personal copy Modeling Marine Ecosystem Services 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 distributed (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 management 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 people derive sustenance, inspiration, economic prosperity, and cultural heritage from the marine environment. Mangroves and coral reefs not only provide protection from storms and serve as a draw for international tourism, but also are critical habitats for the spiny lobster, a backbone of commercial and artisanal fisheries. The commercial fishermen in this picture are from a fleet of dugout canoes that work with a traditional sail boat (or ‘‘mother ship’’’) to harvest spiny lobster. 343 1998, the government created the Coastal Zone Management Authority and Institute (CZMAI) with the objective of developing 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 scientific 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 implementation. 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 recreation, coastal protection, and carbon storage. By providing a platform for stakeholder and agency discussions about tradeoffs 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 informed 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 include 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. Quantitative 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 articulate connections between human activities that are often considered in isolation to align diverse stakeholders around common goals and to make implicit decisions explicit. Ecosystem 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 Coral Triangle Klein et al. (2010) used a combination of quantitative modeling 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 management and policy interventions that will abate key threats Author's personal copy 344 Modeling Marine Ecosystem Services affecting marine ecosystem objectives. The approach requires estimates of management costs and opportunity costs of applying alternative actions; and these estimates are user-generated, 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 economic 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 costeffective at improving marine ecosystem condition than landbased actions designed to reduce nutrient or sediment runoff. Putting Offshore Wind in Context in Massachusetts Like many coastal marine ecosystems, the coast of Massachusetts (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 consideration 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) performed a spatially explicit bioeconomic trade-off analysis among wind energy, fishery, and whale-watching ecosystem services in Massachusetts Bay. They identified optimal planning 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, multiuse ecosystem highlights when and how offshore renewable energy may be developed optimally within the complicated contexts of coastal ecosystems. 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 services 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 accordingly. 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. 2. 3. 4. Marine Marine Marine Marine Ecosystem Services Conservation Biology Ecology Resource Economics See also: Aquaculture. Biodiversity and Ecosystem Services. Biodiversity, Human Well-Being, and Markets. Carbon Cycle. Census of Marine Life. Computer Systems and Models, Use of. Economic Value of Biodiversity, Measurements of. Economics of the Regulating Services. Ecosystem, Concept of. Ecosystem Function Measurement, Aquatic and Marine Communities. Ecosystem Function, Principles of. Ecosystem Services. Endangered Marine Invertebrates. Estuarine Ecosystems. Evaluation of Ecosystem Service Policies from Biophysical and Social Perspectives: The Case of China. Fish Conservation. Mangrove Ecosystems. Marine Conservation in a Changing Climate. Marine Ecosystems, Human Impacts on. Marine Ecosystems. Marine Protected Areas: Static Boundaries in a Changing World. Modeling Terrestrial Ecosystem Services. Ocean Ecosystems. Priority Setting for Biodiversity and Ecosystem Services. Resource Exploitation, Fisheries. The Value of Biodiversity. Valuing Ecosystem Services Conclusions References Marine ecosystems around the globe are under increasing pressure: people rely on them for the delivery of provisioning, regulating, supporting, and cultural ecosystem services. 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