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 benet and for the benet of the author’s institution, for non-commercial research
and educational use. This includes without limitation use in instruction at your institution, distribution to specic
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:
Mcleod Elizabeth (2013) Marine Protected Areas: Static Boundaries in a Changing World. In: Levin S.A. (ed.)
Encyclopedia of Biodiversity, second edition, Volume 5, pp. 94-104. Waltham, MA: Academic Press.
© 2013 Elsevier Inc. All rights reserved.
Marine Protected Areas: Static Boundaries in a Changing World
Elizabeth Mcleod, The Nature Conservancy, Austin, TX, USA
r2013 Elsevier Inc. All rights reserved.
Adaptive management Integration of design,
management, and monitoring to systematically test
assumptions to adapt and learn.
Connectivity Natural linkage between marine habitats
which occur via larval dispersal and the movements of
adults and juveniles.
Ecosystem-based management (EBM) Environmental
management approach that recognizes the full array of
interactions within an ecosystem, including humans, rather
than considering single issues, species, or ecosystem services
in isolation. EBM focuses on cumulative impacts; multiple
objectives; embracing change; linkages between species,
ecosystems, societies, economies, and institutions; and
learning and adaptation.
Ecosystem function Physical, chemical, and biological
processes or attributes that contribute to the self-
maintenance of an ecosystem (e.g., nutrient cycling,
Ecosystem resilience Ability of an ecosystem to maintain
key functions and processes in the face of stresses or
pressures, either by resisting or adapting to change; resilient
systems are characterized as adaptable, ﬂexible, and able to
deal with change and uncertainty.
Ecosystem services Beneﬁts people obtain from
ecosystems; include the provision of food, water, timber,
ﬁber, and other resources; the regulation of ﬂoods, disease,
wastes, and water quality; the support of cultural practices,
including recreation, religion, and art; and the maintenance
of biological processes through such phenomena as soil
formation, photosynthesis, nutrient cycling, and so on.
Ecosystem ‘‘goods’’ include food, medicinal plants,
construction materials, tourism and recreation, and wild
genes for domestic plants and animals.
Marine protected area (MPA) Clearly deﬁned
geographical space recognized, dedicated, and managed
through legal or other effective means to achieve the long-
term conservation of nature with associated ecosystem
services and cultural values.
Marine reserve Subset of an MPA and an area of ocean
completely protected from all extractive and destructive
MPA network Collection of individual MPAs operating
cooperatively and synergistically – at various spatial scales
and with a range of protection levels – to fulﬁll ecological
aims more effectively and comprehensively than individual
sites could alone.
No-take area Marine area that is permanently or
temporarily completely closed for any form of extraction,
and where no disturbance of any kind is allowed.
Humans depend on the oceans for food security, shoreline
protection, recreational opportunities, cultural heritage, cli-
mate regulation, and other services. Despite their tremendous
value, the health of the world’s oceans has continued to
decline due to human activities such as overﬁshing, pollution,
and climate change (Jackson et al., 2001;Worm et al.,
2006;Halpern et al., 2008a). These impacts are leading to
ecosystem collapses in all the major coastal and ocean regions
of the world (Wilkinson, 2004;Hughes et al., 2005;Jackson,
Over the last several decades, about one-third of coastal and
marine habitats, such as mangroves, seagrasses, coral reefs, and
salt marshes, have been lost due to human activities (Va liela
et al., 2001;Wilkinson, 2004;Duarte et al., 2008;Waycott et al.,
2009;Spalding et al.,2010). More than half of the world’s
ﬁsheries stocks are fully exploited and producing catches at
or close to their maximum sustainable limits, and more than
25% are overexploited, depleted, or recovering from depletion
(FAO, 2007). Fundamental changes to ecosystem structure,
such as changes in species diversity, population abundance,
size structure, sex ratios, habitat structure, trophic dynamics,
biogeochemistry, and biological interactions, are occurring
worldwide (Lubchenco et al., 2003). These changes affect
marine ecosystem function and have critical implications for
people that depend on these ecosystems for goods and services
(Lubchenco et al.,1995).
Marine protected areas (MPAs) have been identiﬁed as one
of the most effective tools for conserving marine ecosystems
(Kelleher, 1999;Palumbi, 2003). A number of terms are used,
often interchangeably, to refer to marine areas that are pro-
tected by spatially explicit restrictions, including MPAs, marine
reserves, closed areas, harvest refuges, and sanctuaries (Agardi,
2000). In this chapter, an MPA is a ‘‘clearly deﬁned geo-
graphical space, recognized, dedicated and managed, through
legal or other effective means, to achieve the long-term con-
servation of nature with associated ecosystem services and
cultural values’’ (Dudley, 2008). A marine reserve is a subset of
an MPA and is deﬁned here as ‘‘an area of the ocean com-
pletely protected from all extractive and destructive activities’’
(Lubchenco et al., 2003).
MPAs may include areas with multiple uses (e.g., ﬁshing,
tourism), no-take areas and reserves, or restriction of certain
areas to a speciﬁc use (e.g., local ﬁshing). MPAs range in size
from small marine parks designed to protect endangered or
threatened species, unique habitat, or cultural or historical sites
to large reserves designed to achieve a range of conservation,
Encyclopedia of Biodiversity, Volume 5 http://dx.doi.org/10.1016/B978-0-12-384719-5.00347-694
Author's personal copy
social, and economic objectives encompassing different types
of protection (Agardi, 2000). Ecological objectives include
protection of critical habitats (spawning aggregations, nursery
grounds, areas of high biodiversity, and migration routes),
maintenance of ecosystem function, and species protection.
Socioeconomic objectives include the protection of com-
mercially valuable species, cultural and historic sites, recre-
ation and tourism sites, and sites important for education or
research (Salm et al., 2000). If MPAs are well designed and
managed, they have the potential to protect and in some cases
restore coastal and marine ecosystems and support com-
munities that depend on these ecosystems.
MPAs are most effective when combined with other man-
agement tools such as integrated coastal management, marine
spatial planning (MSP), and ﬁsheries management (Salm
et al., 2006;Dudley, 2008). MPAs are vulnerable to activities
outside their boundaries (e.g., pollution and unsustainable
ﬁshing) that can affect species and ecosystem functions within
protected areas (Kaiser, 2005). Therefore, integrated coastal
management to control land-based threats such as pollution
and sedimentation and other forms of resource management
such as ﬁshery management tools (e.g., catch limits, gear
restrictions, regulations regarding ﬁshing grounds, ﬁshing
seasons; Kaiser, 2005;Keller et al., 2009) are necessary to
support the effectiveness of MPAs. MPAs may also be more
effective when combined with traditional marine manage-
ment approaches (McClanahan et al., 2006).
Systems Approach to Marine Conservation
Conservation and ﬁsheries management efforts have evolved
over the last decade toward managing systems as opposed to
species or speciﬁc habitats and managing cumulative impacts.
These approaches have been referred to as ecosystem-based
management (EBM) and ecosystem approach to ﬁsheries
(Rosenberg and McLeod, 2005;Levin and Lubchenco, 2008;
Palumbi et al., 2008;FAO, 2010). In these approaches,
humans are recognized as critical parts of dynamic ecosystems,
and the inclusion of the human dimension is now seen as an
essential component of effective conservation. Fisheries man-
agement has shifted from a focus on maximum sustainable
yield of individual species at a single scale to multispecies
stock assessments at multiple scales (Pikitch, 2004). EBM
supports ecological processes that maintain resources, recog-
nizing the diverse ecological roles of species and habitats at
multiple scales (Graham et al., 2003). MPAs protect geo-
graphical areas, species, and their biophysical environments
and thus can offer an ecosystem-based approach to conser-
vation or ﬁsheries management (Lubchenco et al., 2003).
Evolution of MPA Deﬁnitions
The shift toward a multiscale, integrated human–ecological
system, and process-oriented perspective (Hughes et al., 2005)
is evident in the changing views of MPAs and the call for
networks of MPAs worldwide. In 1999, an MPA was deﬁned
as ‘‘any area of the intertidal or subtidal terrain, together with
its overlying water and associated ﬂora, fauna, historical, and
cultural features, which has been reserved by law or other
effective means to protect part or all of the enclosed environ-
ment’’ (Kelleher, 1999). In 2008, the World Conservation
Union deﬁned an MPA as a ‘‘clearly deﬁned geographical
space, recognized, dedicated and managed, through legal or
other effective means, to achieve the long-term conservation of
nature with associated ecosystem services and cultural values’’
(Dudley, 2008). The explicit reference to conservation for the
beneﬁt of people (ecosystem services and cultural values) is
highlighted in this latest deﬁnition.
Evolution of MPA Objectives
Conservation managers have been called on to consider pro-
tection of ecosystem function, structure, and integrity in
addition to species and habitat protection (Agardy and Staub,
2006). There has been a shift from the conservation of com-
mercially important species to management of functional
groups (i.e., collections of species that perform a similar
function, regardless of their taxonomic afﬁnities) supporting
processes and maintenance of ecosystem services (e.g., ﬁsh-
eries) (Hughes et al., 2005). The view of no-take areas
as primarily ﬁsheries management tools has evolved to in-
clude other conservation objectives, including managing bio-
diversity, trophic structure and function, and ecosystem
resilience (Hughes et al., 2005). This shift toward managing
ecosystem structure, function, and services emphasizes the
importance of ecological roles and species interactions (in-
cluding humans) for maintaining ecosystem resilience. The
effectiveness of MPAs is now evaluated based on their impacts
on local communities, in addition to ecological impacts. In
addition, integrated studies are developing that assess how
ecological performance of reserves is related to both socio-
economic characteristics in coastal communities and reserve
design (Pollnac et al.,2010). The results of such studies
are useful for highlighting the complexities around human
dimensions of marine reserves and informing MPA design and
Ecosystem resilience refers to the ability of an ecosystem to
maintain key functions and processes in the face of stresses or
pressures, either by resisting or adapting to change (Holling,
1973;Nystro¨m and Folke, 2001). Resilient systems are char-
acterized as adaptable, ﬂexible, and able to deal with change
and uncertainty (Hughes et al., 2005); thus resilience has been
identiﬁed as a critical component of MPA network design and
management. Designing and managing networks for resilience
provides MPAs with the best chance to recover from or with-
stand environmental ﬂuctuations or unexpected catastrophes
caused by climate change and other human impacts (West and
Salm, 2003;Mcleod et al., 2009).
Evolution of MPA Networks
The conservation community and government agencies have
called for the establishment of networks of MPAs worldwide.
An MPA network is deﬁned as a ‘‘collection of individual
MPAs operating cooperatively and synergistically, at various
spatial scales, and with a range of protection levels, in order to
fulﬁll ecological aims more effectively and comprehensively
Marine Protected Areas: Static Boundaries in a Changing World 95
Author's personal copy
than individual sites could alone’’ (WCPA/IUCN, 2007). MPA
networks have also been deﬁned as a ‘‘network of people
managing the components of individual MPAs and promoting
the network’s viability and longevity’’ (Dudley, 2008).
Individual small MPAs may not be effective at conserving
biodiversity, ﬁsh and invertebrate populations, and the com-
munities that depend on them. Single MPAs large enough to
sustain populations and habitats are often impractical due to
economic, social, and political constraints (Dudley, 2008).
Networks of MPAs have been proposed to help reduce socio-
economic impacts while maintaining conservation and ﬁsh-
eries beneﬁts (PISCO, 2007). Networks are also important for
maintaining ecosystem processes and connectivity and sup-
porting ecosystem resilience by spreading the risk of reduced
viability of a habitat or community type following a large-scale
disturbance or management failures (Keller et al., 2009). The
commitment to the establishment of global networks of
MPAs has been demonstrated at international meetings such
as the World Summit on Sustainable Development in 2002,
the World Parks Congress in 2003, and the Convention on
Biological Diversity in 2004. The scaling up of individual
MPAs to networks demonstrates a systems approach to marine
conservation because it allows for the protection of species
and habitats in addition to ecological processes, structure, and
Evolution of MSP
Over the last decade, marine spatial planning (MSP) has
been increasingly recognized as a critical tool to achieve EBM
(Douvere, 2008). MSP provides an integrated planning
framework that moves away from sectoral management to
address multiple objectives related to achieving economic and
ecological sustainability and the need to reduce conﬂicts in
marine environment (Agardi et al.,2011). MSP has been de-
ﬁned as a ‘‘process of analyzing and allocating parts of the
three-dimensional marine spaces to speciﬁc uses, to achieve
ecological, economic and social objectives that are usually
speciﬁed through the political process; the MSP process usu-
ally results in a comprehensive plan or vision for a marine
region’’ (Ehler and Douvere, 2007). More broadly, the purpose
of MSP is to balance demands for development with the need
to protect the environment (Douvere, 2008).
Potential beneﬁts of MSP include a holistic approach that
addresses social, cultural, economic, and environmental ob-
jectives and thus achieve sustainable development; better in-
tegration of marine objectives (both between policies and
between different planning levels); improved site selection for
development or conservation; a more strategic and proactive
approach that delivers long-term beneﬁts; management co-
ordination at the scale of ecosystems as well as political jur-
isdictions; reduced conﬂicts among uses in the marine area;
and reduced risk of marine activities damaging marine eco-
systems, including improved consideration of cumulative
effects (Gilliland and Laffoley, 2008;Foley et al.,2010).
MSP has been applied to help manage the multiple uses of
marine space, particularly in areas where conﬂicts exist among
users and the environment. MSP is central to the management
strategy of the Great Barrier Reef in Australia and has also been
applied in other marine areas such as the Florida Keys,
Channel Islands, Wadden Sea, North Sea, Irish Sea, and Baltic
Sea, among others. MSP is not intended to replace MPAs.
MPAs are still recognized as an important tool for managing
the marine environment, but they should be considered in the
wider context of an MSP strategy that balances the MPAs with
economic, social, and biodiversity objectives. By integrating
MPA planning in broader MSP and ocean zoning efforts, MSP
can help to utilize the beneﬁts of MPAs while avoiding their
potential shortcomings (Agardi et al., 2011).
Beneﬁts of MPAs
MPAs have the potential to provide a number of beneﬁts to
local communities, ﬁsheries, and the marine environment,
including (1) conserving biological diversity and ecosystems;
(2) protecting critical spawning and nursery habitats; (3)
protecting sites with limited human impact to help them re-
cover from stresses; (4) protecting settlement and growth areas
for marine species and spillover beneﬁts to adjacent areas; (5)
protecting sites for educating the public about marine eco-
systems and threats to them; (6) supporting nature-based
recreation and tourism; (7) providing control sites as baselines
for scientiﬁc research; and (8) reducing poverty and increasing
the quality of life of adjacent communities (IUCN-WCPA,
2008). Beneﬁts of MPAs – speciﬁcally, marine reserves – have
been demonstrated through empirical studies for mollusks,
crustaceans, and ﬁshes in habitats ranging from coral reefs,
kelp forests, temperate continental shelves, estuaries, seagrass
beds, and mangroves (Gell and Roberts, 2003). The following
four sections outline the ecological and socioeconomic
beneﬁts of marine reserves based on global and regional meta-
analyses and site-based studies.
Increases in Size, Abundance, Biomass, Diversity, and
Increased Reproductive Potential
Beneﬁts to marine reserves include increases in abundance,
biomass, and diversity of many species within reserve
boundaries (Table 1), yet the range of responses to reserve
establishment is very large (Lester et al., 2009;Gaines et al.,
2010a). Kenchington (1990) identiﬁes several classes of spe-
cies for which marine reserves may not be effective such as
species with planktonic larvae and planktonic or pelagic
adults (e.g., most phytoplankton and zooplankton species to
pelagic ﬁshes with large home ranges). However, these species
may have a stage that depends on a nursery area or spawning
site, and they could be protected by a reserve, assuming that
other life stages outside the reserve are not overexploited
(Allison et al., 1998).
Within reserves, individuals can grow larger, live longer,
and develop increased reproductive potential and populations
increase in size (Bohnsack, 1998). Enhanced production of
eggs and larvae within reserves are predicted to result in
greater export and settlement of juveniles outside boundaries
(Gell and Roberts, 2003). In addition, reserves have helped to
restore ecosystem structure and function (Sobel and Dahlgren,
2004;Mumby et al., 2006). However, these beneﬁts do not
96 Marine Protected Areas: Static Boundaries in a Changing World
Author's personal copy
always occur due to ﬁshers’ behavior in response to reserves,
ﬁshing regulations outside the reserve, and the regulations
regarding activities within and outside the reserve (Gaines
Beneﬁts to Nontarget Species
Recent reviews on the beneﬁts of marine reserves typically
focus on beneﬁts to target species as opposed to nontargeted
groups such as ﬁsh, invertebrates, or algae or corals (Babcock
et al.,2010). However, it is important to understand the im-
pacts of reserves on nontarget groups if the goal of protection
is to maintain ecosystem structure and function. Studies have
documented that nontarget species either do not respond
to protection (Jennings et al., 1995;Rakitin and Kramer, 1996)
or respond negatively (i.e., reduced abundances in response
to increased predation within reserves; McClanahan et al.,
1999). Other studies have shown that nontarget habitats
improve following protection (Mumby et al., 2005;Shears and
Babcock, 2003), where changes in ecosystem structure
have been documented due to the restoration of predator
populations. For example, in tropical systems, enhanced coral
recruitment has occurred following a recovery in herbivores
that graze down macroalgae and thus encourage coral settle-
ment (Mumby et al., 2005). In temperate systems in New
Zealand, the recovery of lobsters and large ﬁshes led to
predation and declines of sea urchin populations that led
to a reduction in grazing and subsequent recovery of kelp
forests (Shears and Babcock, 2003). Reserves have the poten-
tial to provide useful insights into the indirect effects of
overﬁshing on ecosystem structure and function (Babcock
Beneﬁts to Adjacent Fisheries
The ability of reserves to provide conservation or ﬁsheries
beneﬁts to adjacent waters is highly controversial (Gell and
Roberts, 2003;Hilborn et al., 2004;Halpern et al.,2010), yet
recent research suggests that higher abundances within re-
serves can lead to spillover of adults to adjacent waters
(Roberts et al., 2001;Abesamis and Russ, 2005;Kellner et al.,
2008;Perez-Ruzafa et al., 2008;Halpern et al.,2010). Spillover
occurs through the net export of adults and juveniles (spill-
over effect) and propagules (recruitment effect) (Russ, 2002).
The spillover effect operates on local scales (hundreds of
meters to kilometers for reef ﬁsh), whereas the recruitment
effect operates at scales of tens of kilometers (scales of dis-
persal of pelagic larvae; Palumbi, 2001;Russ et al., 2004).
Spillover from reserves may result in economic beneﬁts from
enhanced ﬁsheries and tourism (White et al., 2008) yet may
take decades to develop fully (Roberts et al., 2001;Russ et al.,
Table 1 Global and regional meta-analyses of ecological beneﬁts of marine reserves within reserve boundaries
Indicator Results Taxonomic group # Studies/marine reserves analyzed Source
Biomass 446% increase Algae, invertebrates, and
124 reserves (global) Lester et al. (2009)
Density 166% increase
Individual size 28% increase
Biomass 352% increase Algae, invertebrates, and
89 studies; 70 reserves (global) Halpern (2003)
Density 151% increase
Individual size 29% increase
Biomass 1.9 times higher Algae, invertebrates, and
30 reserves (global; temperate only) Stewart et al. (2009)
Density 1.7 times higher
1.5 times higher
Abundance 25% increase Fishes 19 reserves (global) Co
´et al. (2001)
Abundance 3.7 times higher (for target
Fishes 12 studies (global) Mosquera et al. (2000)
No change (nontarget species)
Density 66% increase Fishes 32 reserves (global) Molloy et al. (2009)
Density 2.46 times larger Fishes 12 reserves (regional – European) Claudet et al. (2008)
Biomass 2.1 times greater Fishes 12 reserves (regional –
Guidetti and Sala
Density 1.2 times greater
Marine Protected Areas: Static Boundaries in a Changing World 97
Author's personal copy
Long-Term Ecological Beneﬁts
Research has shown that ﬁsheries and conservation beneﬁts of
marine reserves increase with greater years of protection (Russ
et al., 2004;Claudet et al., 2008;Molloy et al., 2009;Selig and
Bruno, 2010). To measure the long-term beneﬁts of marine
reserves, time-series data are needed to describe ecological
changes due to protection and the stability of such changes.
Babcock et al. (2010) analyzed data from temperate and tro-
pical marine reserves collected on decadal time scales and
found that even though most target species showed initial
direct effects (e.g., change in abundance, size of individuals,
biomass), their trajectories over time were highly variable. The
abundance of some target species continued to increase after
protection, whereas some leveled off and others decreased
over time. Decreases in abundance were likely due to natural
ﬂuctuations, ﬁshing impacts from outside reserves, and in-
creases in predation within reserves. Despite these differences,
populations of targeted species were more stable in reserves
than ﬁshed areas, indicating increased ecological resilience
(Babcock et al.,2010). Although some beneﬁts are evident
shortly after protection (individuals live longer, mortality rates
are lower), other beneﬁts take longer to fully develop, such as
increases in reproductive output, biodiversity, and stabiliza-
tion of communities and ecosystem structure and function.
Therefore, understanding that indirect effects (e.g., ecosystem-
wide recovery) from removal of ﬁshing pressure take time
(e.g., 413 years; Babcock et al.,2010) is essential for man-
agers, policy makers, and communities to have realistic ex-
pectations of the beneﬁts of marine reserves.
Although the ecological beneﬁts of MPAs, particularly marine
reserves, are well established, there has been less emphasis on
the social and economic beneﬁts to human communities.
Further, rigorous policy analyses are lacking that consider
the full range of economic costs and beneﬁts of MPAs (Rudd
et al., 2003;Pelletier et al., 2005). A limited number of recent
assessments review the economic impacts of MPAs but
are concentrated in North America, Australia, and Europe
(e.g., Carlsen and Wood, 2004;Carter, 2003;KPMG, 2000;
Leeworthy and Wiley, 2002;Roncin et al., 2008), thus leaving
out areas with the highest tropical marine biodiversity
worldwide, such as Southeast Asia and the Paciﬁc.
Socioeconomic assessments of the beneﬁts of MPAs typi-
cally differentiate between extractive (e.g., ﬁshers) and non-
extractive users (e.g., recreational users such as divers,
snorkelers, bathers, ecotourists, and sightseers) because these
groups are likely to be impacted differently by the MPA (see
Table 2 for a summary of the potential social and economic
costs and beneﬁts of MPAs for these user groups). For
extractive users, adverse impacts from MPA establishment
may include loss of access rights to ﬁshing grounds or
increased risk due to traveling farther to access alternative
ﬁshing grounds. For a reserve to provide ﬁsheries beneﬁts to
human communities, the reserve must lead to a net increase in
yield (i.e., increases in harvest must be large enough to com-
pensate for the area removed from ﬁshing). The establishment
of a marine reserve may actually reduce ﬁshing opportunity
and yield if the ﬁsheries are already sustainably managed
(Hastings and Botsford, 1999;Sladek-Nowlis and Roberts,
1999;Ralston and O’Farrell, 2008). The implementation
of an MPA can increase costs if fewer ﬁsh are available due to
harvest restrictions; fuel or labor costs are higher due to tra-
veling farther to ﬁshing grounds, and congestion increases in
ﬁshing grounds (Rudd et al., 2003). A number of case studies
suggest that ﬁshers perceive the costs of MPAs (in terms of lost
harvest) as greater than the beneﬁts provided (e.g., from
spillover) (Wolfenden et al., 1994;Sant, 1996;Suman et al.,
Table 2 Summary of potential social and economic beneﬁts and costs of MPAs
Categories Beneﬁts Costs
(e.g., commercial and
Increase in catch (and associated income) Decrease in catch (and foregone ﬁshing income)
Enhanced catch variety (greater species variety, greater
frequency of older/larger ﬁsh)
Crowding of displaced effort
Higher costs associated with choice of ﬁshing location
Increase in safety risks
(e.g., divers, tourists)
Maintain species diversity Damage to marine ecosystem
Greater habitat complexity and diversity Loss of traditional ﬁshing community
Higher density levels of marine species
Enhanced recreational opportunities (e.g., scuba, snorkeling)
Protection of other ecosystem services (e.g., coastal
protection from erosion and storm surge by healthy reefs)
Management Savings in enforcement costs over nonspatial management Increase in monitoring and enforcement costs
Revenues derived from charging users of the MPA Direct costs of setting up MPA
Scientiﬁc knowledge Costs of compensatory measures for displaced activities
Hedge against uncertain stock assessments Foregone income from resource extraction (oil, gas, and
mineral exploration, and bio-prospecting)
Educational opportunities Increased congestion and possibly degraded ecosystem if
MPA is not well managed due to increased use
98 Marine Protected Areas: Static Boundaries in a Changing World
Author's personal copy
By contrast, costs may be lower for ﬁshers due to steady
and reliable spillover to adjacent ﬁshing grounds and en-
hanced catch variety (Sumaila, 1998;Roncin et al., 2008). The
ﬁshing beneﬁts of MPAs are challenging to assess because ﬁsh
mobility between reserves and open areas to ﬁshing are often
poorly documented and because MPA beneﬁts to ﬁshers are
highly dependent on the level of ﬁshing in open areas (Roncin
et al., 2008). Thus, the economic value of spillover depends
more on ﬁshers’ behavior and the cost of ﬁshing as opposed to
biological factors (Rudd et al., 2003).
For nonextractive users, MPAs are likely to improve the
quality of the marine ecosystem within the MPA that may be
valuable to visitors (Rudd and Tupper, 2002). Because marine
reserves support increases in the size and abundance of many
species within reserve boundaries (Halpern, 2003) and recre-
ational users such as snorkelers and divers prefer viewing
larger and more abundant species (Williams and Polunin,
2000;Rudd and Tupper, 2002), the proﬁtability of recre-
ational and tourism providers may be increased by MPA
establishment (Rudd et al., 2002). Although an increase in
visitors may increase revenues, too many visitors could ad-
versely affect marine ecosystems within MPAs, particularly
when their activities are not well managed. Further, increases
in congestion may result in a decline in people’s willingness to
pay for wildlife viewing (Rudd and Tupper, 2002).
Economic valuations of MPAs provide valuable cost–beneﬁt
analyses and also highlight how the beneﬁts of an MPA may be
distributed. For example, the economic value of a healthy Great
Barrier Reef to Australia is currently estimated to be around $5.5
billion Australian dollars annually and is increasing (McCook
et al.,2010). This estimate includes only use values (e.g., jobs,
tourism, and ﬁshing) and underestimates the total economic
value. The costs associated with zoning and management of the
Great Barrier Reef Marine Park are signiﬁcantly less than the
estimated economic value of the Great Barrier Reef; manage-
ment costs are consistently less than 1% of the economic re-
turns (McCook et al., 2010 ). Similarly, in the Florida Keys
National Marine Sanctuary, the management costs of the con-
servation program represented only 2% of the total beneﬁts
derived from the MPA (Bhat, 2003). In a recent economic an-
alysis of 12 marine reserves in Europe, results suggest that the
amount of income generated by ﬁshing and diving in the MPA
represents 2.3 times the management costs of the MPA (Roncin
et al., 2008). An economic assessment conducted for an MPA
in Kenya also demonstrated that the income generated from
the MPA was substantially higher than the management and
opportunity costs for the park; income from the MPA was $1.6
million annually from tourism and $39,000 from ﬁsheries
compared to management and opportunity costs of less than
$200,000 (Emerton and Tessema, 2001). Economic beneﬁts
from the MPA, such as shoreline protection, marine product-
ivity, wildlife habitat and nursery, and cultural and aesthetic
values, were not included in this assessment (Emerton and
Tessema, 2001) but would have made the estimate of beneﬁts
even greater. The valuation demonstrated that some groups
(commercial tourism operators) received the main economic
beneﬁts from the MPA, whereas others such as the local ﬁshing
communities (which had reduced ﬁshing opportunities) and
the park ofﬁce (responsible for managing the MPA) bore the
cost. Such analyses provide valuable indications of the equity of
the distribution of MPA beneﬁts that can directly affect com-
pliance and overall protection.
In addition to assessing the economic beneﬁts of MPAs,
social beneﬁts should also be addressed. Studies have docu-
mented the importance of assessing the perception of people
affected by MPAs, as their perceptions affect the degree of
support or opposition to the MPA, and consequently the ef-
fectiveness of protection (Pelletier et al., 2005). A critical social
beneﬁt of MPAs is reducing and anticipating conﬂicts between
different user groups. Much of the world’s coastal areas are
characterized by conﬂict between user groups or jurisdictional
agencies (Agardi, 2000). For example, recreational use may
conﬂict with shipping and mineral extraction, and commercial
and subsistence ﬁshing may conﬂict with scuba diving and
nature-based tourism. In such cases, zoning can be used to
accommodate a wide variety of uses and can be used as a tool
to settle disputes when they occur (Reynard, 1994;Agardi,
2000). Other social beneﬁts include improving visitors’ satis-
faction and increasing public knowledge about marine eco-
systems and biodiversity (Pelletier et al., 2005). The potential
of MPAs to help alleviate poverty in coastal communities
dependent on coral reefs has also been acknowledged (Leisher
et al., 2007). It is important to note that although MPAs may
achieve their biological objectives, they may fail at achieving
their social objectives. Therefore, assessments of the social
effects of MPAs are critical to determine their long-term
beneﬁts (Christie, 2004).
Global Commitment to MPA Establishment
Governments around the world have demonstrated their
commitment to conserving coastal and marine ecosystems for
the beneﬁt of the communities that depend on them. National
leaders have formed regional initiatives to establish networks
of MPAs to support ﬁsheries and food security, sustainable
tourism, ecosystem services, livelihoods, and cultural heritage
(e.g., the Micronesia Challenge, the Caribbean Challenge, the
Coral Triangle Initiative, and the Western Indian Ocean
Challenge). These Initiatives are critical to building political
will to support marine conservation efforts, improved inte-
gration with development priorities, and the development of
sustainable ﬁnancing mechanisms (Toropova et al.,2010).
The number and areal extent of MPAs has increased dra-
matically over the last decade; the current global coverage of
MPAs has increased 60% over the last three years and more
than 150% since 2003 (Chape et al., 2008). Currently, the
total number of MPAs worldwide is about 5878 and covers
more than 4.2 million km
of ocean (B1.2% of the global
ocean; Toropova et al.,2010). The World Parks Congress in
2003 set a target for conserving 20–30% of the world’s oceans,
yet the costs of running a global MPA network have been
estimated at $5–19 billion annually (Balmford et al., 2004).
Such an effort would require an increase in current areal and
ﬁnancial investment in marine conservation by two orders of
magnitude. A recent assessment of the progress toward global
marine protection targets identiﬁed a mismatch between the
resources available and those required to implement and
monitor a global network of protected areas (Wood et al.,
2008). The authors (Wood et al., 2008) suggest that once a
Marine Protected Areas: Static Boundaries in a Changing World 99
Author's personal copy
global network is developed, it is likely to be a compromise
between quantity (how closely the targets are met) and quality
(how well designed and effectively managed the protected
Despite the increases in the total number of MPAs worldwide,
it is essential to assess how well these MPAs are meeting their
objectives. According to a recent global analysis, 27% of the
world’s coral reefs are located within MPAs, yet only 6% of
these are effectively managed (Burke et al.,2011). Further,
nearly half of the MPAs worldwide are ineffective at reducing
the threat of overﬁshing (Burke et al.,2011).
MPA performance is highly variable (Kelleher et al., 1995;
Halpern, 2003), and a number of factors have been identiﬁed
that limit the effectiveness of MPAs in conserving biodiversity,
maintaining ﬁsheries, or providing additional ecosystem ser-
vices to human communities. Some have suggested that social
and political factors, as opposed to biological factors, are the
primary determinants of MPA success or failure (Kelleher and
Recchia, 1998;McClanahan, 1999;Mascia, 2003;Leisher,
2008). For example, some MPAs are ineffective because the
management framework is ignored or not enforced (these are
referred to as ‘‘paper parks’’), or they have regulations that
are fully and effectively implemented but are insufﬁcient to
address the threats within the MPA. Other factors affecting
MPA effectiveness include differences in reserve design (e.g.,
size of no-take and buffer zones; Claudet et al., 2008) or re-
serve shape (Kramer and Chapman, 1999). The location of
MPAs also affects effectiveness; MPAs are often placed in areas
where threats are lowest (e.g., large MPAs in remote areas such
as the northwest Hawaiian Islands; Burke et al.,2011) and thus
may do little to mitigate local threats.
MPAs may function more effectively when devolution of
authority for MPA development and management occurs (e.g.,
from national government to local governments, nongovern-
mental organizations, and resource users; White et al., 2002).
Other factors supporting effectiveness of MPAs include adap-
tive and participatory decision-making arrangements; clearly
deﬁned MPA boundaries; clear, easily understood, and easily
enforceable rules; legitimacy of rules and regulations; political
commitment and leadership; and collaborative MPA man-
agement structures linking resources with local interests and
knowledge (Mascia, 2003).
The result of establishing a reserve in one location is that
ﬁshing effort simply moves elsewhere, and the reallocation of
ﬁshing efforts can have adverse impacts on species and habi-
tats outside the reserve (Hilborn et al., 2004). In places where
ﬁshing systems are effective at protecting stock (e.g., through
catch, size, and area limits), it is not clear whether establishing
MPAs will provide additional beneﬁts (Hilborn et al., 2004).
The effectiveness of reserves also varies due to differential re-
sponses of species to protection (Micheli et al., 2004;Molloy
et al., 2009). Researchers have noted signiﬁcantly large in-
creases in abundance of some large-bodied commercially
important species following reserve establishment (e.g., Russ
and Alcala, 1996;Claudet et al., 2008), yet many species re-
spond less predictably to protection (Mosqueira et al., 2000;
Molloy et al., 2009). Although commonly ﬁshed predatory
species are likely to beneﬁt from reserve establishment, prey
species may decline due to trophic cascades (Micheli et al.,
2004;Molloy et al., 2009). Research suggests that MPAs are
effective at protecting sedentary species, but they are less
effective at preserving highly mobile species that may spend
considerable time outside MPA boundaries (Kaiser, 2005),
although exceptions have been documented (McCook et al.,
2010). Therefore, the scales of adult movement and propagule
dispersal can be critical to MPA effectiveness. Empirical studies
are needed to clarify the beneﬁts of MPAs to highly mobile
species, and innovative management approaches are needed
to complement strategically placed MPAs to support them
(Gaines et al.,2010b).
Role of MPAs in a Changing World: Rising to the
Climate change impacts are already occurring in coastal and
marine ecosystems worldwide and include shifts in ocean
current patterns, ecosystem changes (e.g., widespread coral
loss from mass bleaching), changes in larval development
and transport, and species range shifts and interactions
(Wilkinson, 1998;Parmesan and Yohe, 2003;IPCC, 2007;
Rosenzweig et al., 2008). MPAs are a core strategy in marine
conservation, yet they are geographically ﬁxed and thus poorly
suited to accommodate shifts in species ranges and habitats.
In addition, most existing MPAs are designed based on current
climate conditions. The recognition of their limitations in a
changing world has led some to question their relevance as a
conservation response in an era of rapid climate change
´jo et al., 2004;Hannah et al., 2007).
The ability of MPAs to protect ecosystems and species in
the face of climate change and other changes (e.g., increasing
global population) is debated (Mora et al., 2006;Graham
et al., 2007;McClanahan, 2008;Selig and Bruno, 2010). Some
researchers suggest that habitat loss (e.g., coral reefs) in re-
sponse to climate change, storms, and diseases are unlikely
to be mitigated by MPAs (Jameson et al., 2002;Aronson and
Precht, 2006;Graham et al., 2008). Site-speciﬁc studies
have suggested that MPAs do not always protect biodiversity
better than unmanaged areas in response to climate impacts
(Jones et al., 2004;Graham et al., 2007;McClanahan,
2008). Further, some studies suggest that thermal stress can
cause proportionally greater coral mortality of protected than
unprotected corals (McClanahan et al., 2007;Graham et al.,
2007;Graham et al., 2008;Darling et al.,2010). This may
be due to the different coral species composition between
protected and unprotected sites (e.g., higher abundance of
thermally sensitive corals such as Acropora and Montipora
within reserves) (Co
´and Darling, 2010;Darling et al.,2010).
Recent global analyses, however, have conﬁrmed that MPAs
can be effective in preventing coral loss (coral cover remained
constant in MPAs over 38 years, whereas coral cover on un-
protected reefs declined; Selig and Bruno, 2010). Surveys in
the Bahamas showed signiﬁcantly higher increases in coral
cover in reserve boundaries compared to outside the reserve
(Mumby and Harborne, 2010). Mumby and Harborne (2010)
suggest that reserves play an important role in increasing coral
100 Marine Protected Areas: Static Boundaries in a Changing World
Author's personal copy
reef recovery rates provided that macroalgae have been de-
pleted by more abundant communities of grazers beneﬁting
from reduced ﬁshing pressure, particularly in the Caribbean.
Empirical evidence both supports (Lafferty and Behrens,
2005;Mumby et al., 2006;Babcock et al.,2010) and refutes
(McClanahan, 2008) the idea that species and habitats within
reserves are more resilient than those outside reserve bound-
aries (Gaines et al.,2010b). Therefore, additional studies are
urgently needed to establish the ability of MPAs to support
resilience in a variety of habitats and geographic locations and
in response to diverse threats.
To address the challenge of climate change, conservation
practitioners and researchers are applying new tools such as
ecological forecasting and climate envelope models to identify
sites most likely to protect biodiversity in the future and to
assess the ability of reserves and networks to protect species
under different climate change scenarios (Arau
´jo et al., 2004;
Hannah et al., 2007;Hannah, 2008). Researchers have cau-
tioned that the use of these tools is limited by the lack of
existing data needed to support the models and uncertainties
inherent in climate change projections and most ecological
forecasting approaches (Thuiller, 2004;Lawler et al., 2006;
Complementing the new tools to support MPA design,
major advances in recommendations for MPA and network
design have also occurred over the last decade (Roberts et al.,
2003;Halpern et al., 2006;Gaines et al.,2010a), particularly
recommendations speciﬁcally designed to address climate
change impacts (Lawler, 2009;Mcleod et al., 2009). Such
principles may include the identiﬁcation and protection of
refuges (e.g., sites resistant to climate change impacts; such
sites can provide the larvae needed to reseed areas that suc-
cumb to coral bleaching), pathways of connectivity that link
these refuges with damaged areas, and measures to build
redundancy into networks, thereby ameliorating the risk that
climate change impacts will result in irrevocable biodiversity
loss (West and Salm, 2003;Mcleod et al., 2009).
Researchers have suggested that more and larger MPAs will
be needed in the future to address climate change impacts
(Lawler, 2009). Speciﬁc recommendations include increasing
the size of existing reserves, adding buffers around existing
reserves, and adding larger reserves to reserve networks
(Halpin, 1997;Noss, 2001). Establishing more and larger re-
serves may be insufﬁcient to protect biodiversity and maintain
ecosystem services if the reserves are not located in the right
places; a more strategic approach involves locating reserves so
that they capture the most potential for habitat heterogeneity
under a variety of climate scenarios (Lawler, 2009), or in
places predicted to escape the brunt of climate change (West
and Salm, 2003;Mumby and Steneck, 2008;Mcleod et al.,
´and Darling, 2010). In addition, whereas larger
MPAs may provide protection for increased and functional
groups, they may not be politically, socially, or economically
feasible. Research also suggests that large MPAs may be less
effective than other traditional management approaches. For
example, traditional management regimes in Indonesia and
Papua New Guinea involving periodic closures were signiﬁ-
cantly more effective than national parks with permanent
closures – a more than 40% increase in targeted ﬁsh biomass
within reserves in traditional management regimes compared
to less than 2% increase in national parks (McClanahan et al.,
2006). McClanahan et al. (2006) note that whereas large
MPAs may provide the best protection for species susceptible
to overﬁshing, traditional management approaches may pro-
vide the best solution for meeting conservation and com-
munity goals and reversing the degradation of reef ecosystems.
MPA and zone boundaries should be designed to be ﬂex-
ible in space and time so that they can be expanded or con-
tracted, have seasonal or other ﬁxed time limits, or be moved
to different levels of protection to help them meet their
objectives in response to future changes. Where habitat shifts
are predicted, managers should proactively plan for landward
migration, particularly in areas where habitats have the po-
tential to expand (e.g., mangrove migration landward in re-
sponse to sea-level rise). It is important to identify and protect
areas likely to serve as refuges in the future (i.e., predictive
protected areas; Herr and Galland, 2009) and also areas that
have demonstrated resilience to climate change impacts.
Finally, to help MPAs continue to achieve their social, eco-
logical, and economic objectives, adaptive management is
essential. Adaptive management refers to the integration of
design, management, and monitoring to systematically test
assumptions in order to adapt and learn (Salafsky et al., 2001).
In the context of MPAs, adaptive management involves the
integration of the best available science into MPA strategies and
monitoring to systematically test the effectiveness of manage-
ment methods and reﬁne them over time. Conservation man-
agers should develop management approaches that are ﬂexible
and able to incorporate future species and habitat migrations,
and they need to apply risk-spreading strategies to ensure the
protection of key larvae, species, and habitats. Monitoring
should go beyond simply assessing whether current policies
are effective (e.g., is biodiversity declining?) and should focus
on resolving the underlying causes (e.g., how can we reverse the
decline?) (Hughes et al., 2007). Monitoring programs must
address thresholds, regime shifts and feedbacks, and the cap-
acity of ecosystems to maintain ecosystem services in response
to future changes (Hughes et al., 2007). Understanding how
ecosystem services will be affected by climate change is neces-
sary for setting conservation priorities and designing and
managing restoration projects (Lawler, 2009).
MPAs have a critical role to play in protecting marine
ecosystems and the beneﬁts derived from these systems and in
securing the communities that depend on them. There may be
trade-offs between MPAs designed for biodiversity, sustainable
use, and climate change. Therefore, it is important to include
risk assessments, scenario planning, and adaptive manage-
ment approaches that incorporate these potential trade-offs
(Secretariat of the Convention on Biological Diversity, 2009).
To be successful in a changing world, MPAs must strive to
achieve the complementary goals of maintaining biodiversity,
promoting ecosystem values, and enhancing resilience.
List of Courses
•Applied Ecology and Environmental Management
Marine Protected Areas: Static Boundaries in a Changing World 101
Author's personal copy
•Marine Ecosystem Management
•Marine Protected Areas
See also: Coastal Beach Ecosystems. Conservation Efforts,
Contemporary. Corals and Coral Reefs. Ecosystem Function
Measurement, Aquatic and Marine Communities. Ecosystem Services.
Ethical Issues in Biodiversity Protection. Fish Conservation.
Identifying Conservation Priorities Using a Return on Investment
Analysis. Mangrove Ecosystems. Marine and Aquatic Communities,
Stress from Eutrophication. Marine Conservation in a Changing
Climate. Marine Ecosystems. Marine Ecosystems, Human Impacts on.
Modeling Marine Ecosystem Services. Natural Reserves and
Preserves. Ocean Ecosystems. Pelagic Ecosystems. Resource
Exploitation, Fisheries. Role and Trends of Protected Areas in
Conservation. Seagrasses. Wetlands Ecosystems. Wetland Creation
Abesamis RA and Russ GR (2005) Density-dependent spillover from a marine
reserve: Long term evidence. Ecological Applications 15: 1798–1812.
Agardi T (2000) Information needs for marine protected areas: Scientiﬁc and
societal. Bulletin of Marine Science 66: 875–888.
Agardy T, di Sciara GN, and Christie P (2011) Mind the gap: Addressing the
shortcomings of marine protected areas through large scale marine spatial
planning. Marine Policy 35: 226–232.
Agardy T, and Staub, F (2006) Marine protected areas and MPA networks. Network
for Conservation Educators and Practitioners, American Museum of Natural
History. CD-ROM. Available electronically at: http://research.amnh.org/
Allison GW, Lubchenco J, and Carr MH (1998) Marine reserves are necessary but
not sufﬁcient for marine conservation. Ecological Applications 8: S79–S92.
´jo MB, Cabeza M, Thuiller W, Hannah L, and Williams PH (2004) Would
climate change drive species out of reserves? An assessment of existing
reserve-selection methods. Global Change Biology 10: 1618–1626.
Aronson RB and Precht WF (2006) Conservation, precaution, and Caribbean reefs.
Coral Reefs 25: 441–450.
Babcock RC, Shears NT, Alcala AC, et al. (2010) Decadal trends in marine reserves
reveal differential rates of change in direct and indirect effects. Proceedings of
the National Academy of Sciences of the United States of America 107:
Balmford A, Gravestock P, Hockley N, McClean N, and Roberts CM (2004) The
worldwide costs of marine protected areas. Proceedings of the National Academy
of Sciences of the United States of America 101: 9694–9697.
Bhat MG (2003) Application of non-market valuation to the Florida Keys marine
reserve management. Journal of Environmental Management 67: 315–325.
Bohnsack JA (1998) Application of marine reserves to reef ﬁsheries management.
Austalian Journal of Ecology 23: 298–304.
Burke L, Reytar K, Spalding M, and Perry AL (2011) Reefs at Risk Revisited.
Washington, DC: World Resources Institute, The Nature Conservancy, World
Fish Center, International Coral Reef Action Network, UNEP World Conservation
Monitoring Centre and Global Coral Reef Monitoring Network.
Carlsen J and Wood D (2004) Assessment of the Economic Value of Recreation
and Tourism in Western Australia’s National Parks, Marine Parks and Forests.
Townsville: Sustainable Tourism CRC.
Carter DW (2003) Protected areas in marine resource management: Another look at
the economics and research issues. Ocean and Coastal Management 46:
Chape S, Spalding M, and Jenkins M (2008) The World’s Protected Areas. Status,
Values, and Prospects in the Twenty-First Century. Berkeley, CA: University of
Christie P (2004) MPAs as biological successes and social failures in Southeast
Asia. In: Shipley JB (ed.) Aquatic Protected Areas as Fisheries Management
Tools: Design, Use, and Evaluation of these Fully Protected Areas, pp. 155–164.
Bethesda, MD: American Fisheries Society.
Claudet J, Osenberg CW, Benedetti-Cecchi L, et al. (2008) Marine reserves: Size
and age do matter. Ecology Letters 11: 481–489.
´IM and Darling ES (2010) Rethinking ecosystem resilience in the face of
climate change. PLoS Biology 8: e1000438.http://dx.doi.org/10.1371/
´IM, Mosqueira I, and Reynolds JD (2001) Effects of marine reserve
characteristics on the protection of ﬁsh populations: A meta-analysis. Journal of
Fish Biology 59: 178–189.
Darling ES, McClanahan TR, and Co
´IM (2010) Combined effects of two stressors
on Kenyan coral reefs are additive or antagonistic, not synergistic. Conservation
Letters 3: 122–130.
Douvere F (2008) The importance of marine spatial planning in advancing
ecosystem-based sea use management. Marine Policy 32: 762–771.
Duarte CM, Dennison WC, Orth RJW, and Carruthers TJB (2008) The charisma of
coastal ecosystems: Addressing the imbalance. Estuarine Coast 31: 233–238.
Dudley N (ed.) (2008) Guidelines for Applying Protected Area Management
Categories. Gland, Switzerland: IUCN.
Ehler C and Douvere F (2007) Visions for a sea change. Report of the First
International Workshop on Marine Spatial Planning. Intergovernmental
Oceanographic Commission and Man and the Biosphere Programme. IOC
Manual and Guides No.48. IOCAM Dossier No. 4. Paris: UNESCO.
Emerton L and Tessema Y (2001) Economic Constraints to the Management of
Marine Protected Areas: The Case of Kisite Marine National Park and Mpunguti
Marine National Reserve, Kenya. Kenya: IUCN Eastern Africa Programme.
Foley MM, Halpern BS, and Micheli F (2010) Guiding ecological principles for
marine spatial planning. Marine Policy 34: 955–966.
Food and Agriculture Organization of the United States (FAO) (2007) Report: The
State of World Fisheries and Aquaculture 2006. Rome: FAO Fisheries and
Food and Agriculture Organization of the United States (FAO) (2010) Report: The
State of World Fisheries and Aquaculture 2010. Rome: FAO Fisheries and
Gaines SD, Lester SE, Grorud-Colvert K, Costello C, and Pollnac R (2010b) The
evolving science of marine reserves: New developments and emerging research
frontiers. Proceedings of the National Academy of Sciences of the United States
of America 107: 18251–18255.
Gaines SD, White C, Carr M, and Palumbi S (2010a) Designing marine reserve
networks for both conservation and ﬁsheries management. Proceedings of the
National Academy of Sciences of the United States of America 107:
Gell FR and Roberts CM (2003) Beneﬁts beyond boundaries: The ﬁshery effects of
marine reserves. Trends in Ecology and Evolution 18: 448–455.
Gilliland PM and Laffoley D (2008) Key elements and steps in the process of
developing ecosystem-based marine spatial planning. Marine Policy 32:
Graham NAJ, Evans RD, and Russ GR (2003) The effects of marine reserve
protection on the trophic relationships of reef ﬁshes on the Great Barrier Reef.
Environmental Conservation 30: 200–208.
Graham NAJ, McClanahan TR, MacNeil MA, et al. (2008) Climate warming, marine
protected areas and the oceanscale integrity of coral reef ecosystems. PLoS One
Graham NAJ, Wilson SK, Jennings S, et al. (2007) Lag effects in the impacts of
mass coral bleaching on coral reef ﬁsh, ﬁsheries, and ecosystems. Conservation
Biology 21: 1291–1300.
Guidetti P and Sala E (2007) Community-wide effects of marine reserves in the
Mediterranean Sea. Marine Ecology Progress Series 335: 43–56.
Halpern BS (2003) The impact of marine reserves: Do reserves work and does
reserve size matter? Ecological Applications 13: 117–137.
Halpern BS, Lester SE, and Kellner JB (2010) Spillover from marine reserves and
the replenishment of ﬁshed stocks. Environmental Conservation 36: 268–276.
Halpern BS, Regan HM, Possingham HP, and McCarthy MA (2006) Accounting for
uncertainty in marine reserve design. Ecology Letters 9: 2–11.
Halpern BS, Walbridge S, Selkoe KA, et al. (2008a) A global map of human impact
on marine ecosystems. Science 319: 948–952.
Halpin PN (1997) Global climate change and natural area protection: Management
responses and research directions. Ecological Applications 7: 828–843.
Hannah L (2008) Protected areas and climate change. Annals of the New York
Academy of Sciences 1134: 201–212.
Hannah L, Midgley G, Andelman S, et al. (2007) Protected area needs in a
changing climate. Frontiers in Ecology and the Environment 5: 131–138.
102 Marine Protected Areas: Static Boundaries in a Changing World
Author's personal copy
Hastings A and Botsford LW (1999) Equivalence in yield from marine reserves and
traditional ﬁsheries management. Science 284: 1537–1538.
Herr D and Galland GR (2009) The Ocean and Climate Change. Tools and
Guidelines for Action. Gland, Switzerland: IUCN.
Hilborn R, Stokes K, Maguire JJ, et al. (2004) When can marine reserves improve
ﬁsheries management? Ocean and Coastal Management 47: 197–205.
Holling CS (1973) Resilience and stability of ecological systems. Annual Review of
Ecology, Evolution, and Systematics 4: 1–23.
Hughes TP, Bellwood DR, Folke C, Steneck RS, and Wilson J (2005) New
paradigms for supporting the resilience of marine ecosystems. Trends in
Ecology and Evolution 20: 380–386.
Hughes TP, Gunderson LH, Folke C, et al. (2007) Adaptive management of the
Great Barrier Reef and the Grand Canyon World Heritage. Ambio 36: 586–592.
Intergovernmental Panel on Climate Change (IPCC) (2007) Climate change 2007:
Impacts, adaptation and vulnerability. Contribution of Working Group II to the
Fourth Assessment Report of the Intergovernmental Panel on Climate Change.
Cambridge: Cambridge University Press.
IUCN World Commission on Protected Areas (IUCN-WCPA) (2008) Establishing
Marine protected area networks – Making it Happen. Washington, DC: IUCN-
WCPA, National Oceanic and Atmospheric Administration and the Nature
Jackson JBC (2008) Ecological extinction and evolution in the brave new ocean.
Proceedings of the National Academy of Sciences of the United States of
America 105: 11458–11465.
Jackson JBC, Kirby MX, Berger WH, et al. (2001) Historical overﬁshing and the
recent collapse of coastal ecosystems. Science 293: 629–637.
Jameson SC, Tupper MH, and Ridley JM (2002) The three screen doors: Can
marine ‘‘protected’’ areas be effective? Marine Pollution Bulletin 44: 1177–1183.
Jennings S, Grandcourt EM, and Polunin NVC (1995) The effects of ﬁshing on the
diversity, biomass and trophic structure of Seychelles’ reef ﬁsh communities.
Coral Reefs 14: 225–235.
Jones GP, McCormick MI, Srinivasan M, and Eagle JV (2004) Coral decline
threatens ﬁsh biodiversity in marine reserves. Proceedings of the National
Academy of Sciences of the United States of America 101: 8251–8253.
Kaiser MJ (2005) Are marine protected areas a red herring or ﬁsheries panacea?
Canadian Journal of Fisheries and Aquatic Sciences 62: 1194–1199.
Kelleher G (1999) Guidelines for Marine Protected Areas. Gland, Switzerland: IUCN
– The World Conservation Union.
Kelleher G, Bleakley C, and Wells S (1995) Global Representative System of Marine
Protected Areas. Washington, DC: The World Bank.
Kelleher G and Recchia C (1998) Lessons from marine protected areas around the
world. Parks 8: 1–4.
Keller BD, Gleason DF, Mcleod E, et al. (2009) Climate change, coral reef
ecosystems, and management options for marine protected areas. Environmental
Management 44: 1069–1088.
Kellner JB, Nisbet RM, and Gaines SD (2008) Spillover from marine reserves
related to mechanisms of population regulation. Theoretical Ecology 1:
Kenchington RA (1990) Managing Marine Environments. New York, USA: Taylor
KPMG Consulting (2000) Economic and Financial Values of the Great Barrier Reef
Marine Park. Townsville: Great Barrier Reef Marine Park Authority. Research
Kramer DL and Chapman MR (1999) Implications of ﬁsh home range size and
relocation for marine reserve function. Environmental Biology of Fishes 55:
Lafferty KD and Behrens MD (2005) Temporal variation in the state of rocky reefs:
Does ﬁshing increase the vulnerability of kelp forests to disturbance? In:
Garcelon DK and Schwemm CA (eds.) Proceedings of the Sixth California
Islands Symposium, pp. 511–520. Ventura, CA: Institute for Wildlife Studies.
Lawler JJ (2009) Climate Change Adaptation Strategies for Resource Management
and Conservation Planning. The Year in Ecology and Conservation Biology.
Annals of the New York Academy of Sciences 1162: 79–98.
Lawler JJ, White D, Neilson RP, and Blaustein AR (2006) Predicting climate
induced range shifts: Model differences and model reliability. Global Change
Biology 12: 1568–1584.
Leeworthy VR and Wiley P (2002) Socioeconomic Impact Analysis of Marine
Reserve Alternatives for the Channel Islands National Marine Sanctuary. Silver
Spring, MD: NOAA.
Leisher C (2008) What Rachel Carson knew about marine protected areas.
BioScience 58: 478–479.
Leisher C, vanBeukering P, and Scherl LM (2007) Nature’s Investment Bank: How
Marine Protected Areas Contribute to Poverty Reduction, 52 pp. Arlington, VA:
The Nature Conservancy, Available from: http://conserveonline.org.
Lester SE, Halpern BS, Grorud-Colvert K, et al. (2009) Biological effects within no-
take marine reserves: A global synthesis. Marine Ecology Progress Series 384:
Levin SA and Lubchenco J (2008) Resilience, robustness, and marine ecosystem-
based management. BioScience 58: 27–32.
Lubchenco J, Allison GW, Navarrete SA, et al. (1995) Biodiversity and ecosystem
functioning: Coastal systems. In: United Nations Environmental Programme
(ed.), Global Diversity Assessment, pp. 370–381. Cambridge, UK: Cambridge
Lubchenco J, Palumbi SR, Gaines SD, and Andelman S (2003) Plugging a hole in
the ocean: The emerging science of marine reserves. Ecological Applications 13:
Mascia M (2003) The human dimension of coral reef marine protected areas:
Recent social science research and its policy implications. Conservation Biology
McClanahan TR (1999) Is there a future for coral reef parks in poor tropical
countries? Coral Reefs 18: 321–325.
McClanahan TR (2008) Response of the coral reef benthos and herbivory to ﬁshery
closure. Oecologia 155: 169–177.
McClanahan TR, Ateweberhan M, Muhando CA, Maina J, and Mohammed MS
(2007) Effects of climate and seawater temperature variation on coral bleaching
and mortality. Ecological Monographs 77: 503–525.
McClanahan TR, Marnane MJ, Cinner JE, and Kiene WE (2006) A comparison of
marine protected areas and alternative approaches to coral-reef management.
Current Biology 16: 1408–1413.
McClanahan TR, Muthiga NA, Kumukuru AT, Machano H, and Kiambo RW (1999)
The effects of marine parks and ﬁshing on coral reefs of northern Tanzania.
Biological Conservation 89: 161–182.
McCook LJ, Ayling T, Cappo M, et al. (2010) Adaptive management of the Great
Barrier Reef: A globally signiﬁcant demonstration of the beneﬁts of networks of
marine reserves. Proceedings of the National Academy of Sciences of the United
States of America 107: 18278–18285.
Mcleod E, Salm RV, Green A, and Almany J (2009) Designing marine protected
area networks to address the impacts of climate change. Frontiers in Ecology
and the Environment 7: 362–370.
Micheli F, Halpern BS, Botsford LW, and Warner RR (2004) Trajectories and
correlates of community change in no-take marine reserves. Ecological
Applications 14: 1709–1723.
Molloy PP, McLean IB, and Cote IM (2009) Effects of marine reserve age on ﬁsh
populations: A global meta-analysis. Journal of Applied Ecology 46: 743–751.
Mora C, Andrefouet S, Costello MJ, et al. (2006) Coral reefs and the global
network of marine protected areas. Science 312: 1750–1751.
Mosqueira I, Co
´IM, Jennings S, and Reynolds JD (2000) Conservation beneﬁts
of marine reserves for ﬁsh populations. Animal Conservation 4: 321–332.
Mumby PJ, Dahlgren CP, Harborne AR, et al. (2006) Fishing, trophic cascades, and
the process of grazing on coral reefs. Science 311: 98–101.
Mumby PJ, Foster NL, and Glynn Fahy EA (2005) Patch dynamics of coral reef
macroalgae under chronic and acute disturbance. Coral Reefs 24: 681–692.
Mumby PJ and Harborne AR (2010) Marine reserves enhance the recovery of
corals on Caribbean reefs. PLoS ONE 5: e8657.
Mumby PJ and Steneck RS (2008) Coral reef management and conservation in
light of rapidly evolving ecological paradigms. Trends in Ecology and Evolution
Noss RF (2001) Beyond Kyoto: Forest management in a time of rapid climate
change. Conservation Biology 15: 578–590.
¨m M and Folke C (2001) Spatial resilience of coral reefs. Ecosystems 4:
Palumbi SR (2001) The ecology of marine protected areas. In: Bertness MD, Gaines
SD, and Hay ME (eds.) Marine Community Ecology, pp. 509–530. Sunderland,
MD: Sinauer Press.
Palumbi SR (2003) Population genetics, demographic connectivity, and the design
of marine reserves. Ecological Applications 13: S146–S158.
Palumbi SR, McLeod KL, and Grunbaum D (2008) Ecosystems in action: lessons
from marine ecology about recovery, resistance, and reversibility. BioScience 58:
Parmesan C and Yohe G (2003) A globally coherent ﬁngerprint of climate change
impacts across natural systems. Nature 421: 37–42.
Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO) (2007) The
Science of Marine Reserves. (2nd Edition, United States Version)
www.piscoweb.org. 22 pp.
Marine Protected Areas: Static Boundaries in a Changing World 103
Author's personal copy
Pelletier D, Garcı
´a-Charton JC, Ferraris J, et al. (2005) Designing indicators for
assessing the effects of marine protected areas on coral reef ecosystems: A
multidisciplinary standpoint. Aquatic Living Resources 18: 15–33.
Perez-Ruzafa A, Martin E, Marcos C, et al. (2008) Modeling spatial and temporal
scales for spill-over and biomass exportation from MPAs and their potential for
ﬁsheries enhancement. Journal of Nature Conservation 16: 234–255.
Pikitch EK, Santora C, Babcock EA, et al. (2004) Ecosystem-based ﬁshery
management. Science 305: 346–347.
Pollnac R, Christie P, Cinner J, et al. (2010) Marine reserves as linked
social–ecological systems. Proceedings of the National Academy of Sciences of
the United States of America 107: 18262–18265.
Rakitin A and Kramer DL (1996) Effect of a marine reserve on the distribution of
coral reef ﬁshes in Barbados. Marine Ecology Progress Series 131: 97–113.
Ralston S and O’Farrell MR (2008) Spatial variation in ﬁshing intensity and its effect
on yield. Canadian Journal of Fisheries and Aquatic Sciences 65: 588–599.
Reynard Y (1994) Resolving conﬂicts for integrated coastal management: The case
`re, St. Lucia, Caribbean. Parks and Protected Areas Bulletin 5: 5–7.
Roberts CM, Andelman S, Branch G, et al. (2003) Ecological criteria for evaluating
candidate sites for marine reserves. Ecological Applications 13: S199–S214.
Roberts CM, Bohnsack JA, Gell F, Hawkins JP, and Goodridge R (2001) Effects of
marine reserves on adjacent ﬁsheries. Science 294: 1920–1923.
Roncin N, Alban F, Charbonnel E, et al. (2008) Uses of ecosystem services
provided by MPAs: How much do they impact the local economy? A southern
Europe perspective. Journal of Nature Conservation 16: 256–270.
Rosenberg AA and McLeod KL (2005) Implementing ecosystem-based approaches
to management for the conservation of ecosystem services. Marine Ecology
Progress Series 300: 241–296.
Rosenzweig C, Karoly D, Vicarelli M, et al. (2008) Attributing physical and
biological impacts to anthropogenic climate change. Nature 453: 353–357.
Rudd MA, Folmer H, and van Kooten GC (2002) Economic evaluation of
recreational ﬁshery policies. In: Pitcher TJ and Hollingworth C (eds.)
Evaluating Recreational Fisheries: An Ecological, Economic, Social Balance
Sheet, pp. 35–52. Oxford: Blackwell Science.
Rudd MA and Tupper MH (2002) The impact of Nassau grouper size and
abundance on scuba diver site selection. Coastal Management 30: 133–151.
Rudd MA, Tupper MH, Folmer H, and van Kooten GC (2003) Policy analysis for
tropical marine reserves: Challenges and directions. Fish and Fisheries 4: 65–85.
Russ GR (2002) Yet another review of marine reserves as reef ﬁsheries management
tools. In: Sale PF (ed.) Coral Reef Fishes: Dynamics and Diversity in a Complex
Ecosystem, pp. 421–443. San Diego, CA: Academic Press.
Russ GR and Alcala AC (1996) Do marine reserves export adult ﬁsh biomass?
Evidence from Apo Island, central Philippines. Marine Ecology Progress Series
Russ GR, Alcala AC, Maypa AP, Calumpong HP, and White AT (2004) Marine
reserve beneﬁts local ﬁsheries. Ecological Applications 14: 597–606.
Salafsky N, Margoluis R, and Redford K (2001) Adaptive Management: A Tool for
Conservation Practitioners. Washington, DC: Biodiversity Support Program.
Salm RV, Clark JR, and Siirila E (2000) Marine and Coastal Protected Areas: A
Guide for Planners and Managers. Washington, DC, USA: IUCN.
Salm RV, Done T, and Mcleod E (2006) Marine protected area planning in a
changing climate. In: Phinney JT, Hoegh- Guldberg O, Kleypas J, Skirving W,
and Strong A (eds.) Coral Reefs and Climate Change: Science and Management.
Washington, DC: American Geophysical Union.
Sant M (1996) Environmental sustainability and the public: Responses to a
proposed marine reserve at Jervis Bay, New South Wales, Australia. Ocean and
Coastal Management 32: 1–16.
Secretariat of the Convention on Biological Diversity (2009). Connecting
biodiversity and climate change mitigation and adaptation. Report of the Second
Ad Hoc Technical Expert Group on Biodiversity and Climate Change, 126
pp. Montreal: Secretariat of the Convention on Biological Diversity.
Selig ER and Bruno JF (2010) A global analysis of the effectiveness of marine
protected areas in preventing coral loss. PLoS ONE 5: e9278.
Shears NT and Babcock RC (2003) Continuing trophic cascade effects after 25 years
of no-take marine reserve protection. Marine Ecology Progress Series 246: 1–16.
Sladek Nowlis J and Roberts CM (1999) Fisheries beneﬁts and optimal design of
marine reserves. Fisheries Bulletin 97: 604–616.
Sobel J and Dahlgren CP (2004) Marine Reserves: A Guide to Science, Design and
Use. Washington, DC: Island Press.
Spalding M, Kainuma M, and Collins L (2010) World Atlas of Mangroves. London:
Earthscan, with International Society for Mangrove Ecosystems, Food and
Agriculture Organization of the United Nations, The Nature Conservancy, UNEP
World Conservation Monitoring Centre, United Nations Scientiﬁc and Cultural
Organisation, and United Nations University.
Stewart GB, Kaiser MJ, Co
´IM, et al. (2009) Temperate marine reserves: Global
ecological effects and guidelines for future networks. Conservation Letters 2:
Sumaila UR (1998) Protected marine reserves as ﬁsheries management tools: A
bioeconomic analysis. Fisheries Research 37: 287–296.
Suman D, Shivlani M, and Milon JW (1999) Perceptions and attitudes regarding
marine reserves: A comparison of stakeholder groups in the Florida Keys
National Marine Sanctuary. Ocean and Coastal Management 42: 1019–1040.
Thuiller W (2004) Patterns and uncertainties of species’ range shifts under climate
change. Global Change Biology 10: 2020–2027.
Toropova C, Meliane I, Laffoley D, Matthews E, and Spalding M. (eds.) (2010)
Global Ocean Protection: Present Status and Future Possibilities. Brest, France:
Agence des aires marines prote
´es; Gland, Switzerland, Washington, DC and
New York, USA: IUCN WCPA; Cambridge, UK: UNEP-WCMC; Arlington, USA:
TNC; Tokyo, Japan: UNU; New York, USA: WCS.
Valiela I, Bowen JL, and York JK (2001) Mangrove forests: One of the world’s
threatened major tropical environments. BioScience 51: 807–815.
Waycott M, Duarte CM, Carruthers TJB, et al. (2009) Accelerating loss of
seagrasses across the globe threatens coastal ecosystems. Proceedings of the
National Academy of Sciences of the United States of America 106:
WCPA/IUCN (2007) Establishing networks of marine protected areas: A guide for
developing national and regional capacity for building MPA networks. Non-
Technical Summary Report. Washington, DC: IUCN-WCPA, National Oceanic and
Atmospheric Administration and The Nature Conservancy.
West JM and Salm RV (2003) Resistance and resilience to coral bleaching:
Implications for coral reef conservation and management. Conservation Biology
White AT, Salamanca A, and Courtney CA (2002) Experience with marine protected
area planning and management in the Philippines. Coastal Management 30: 1–26.
White C, Kendall BE, Gaines S, Siegel DA, and Costello C (2008) Marine reserve
effects on ﬁshery proﬁt. Ecology Letters 11: 370–379.
Wilkinson C (ed.) (1998) Status of Coral Reefs of the World: 2004, Vol. 1. Global
Coral Reef Monitoring Network. Townsville, Australia: Australian Institute of
Wilkinson C (ed.) (2004) Status of Coral Reefs of the World: 2004, Vol. 1. Global
Coral Reef Monitoring Network. Townsville, Australia: Australian Institute of
Williams ID and Polunin NVC (2000) Differences between protected and
unprotected Caribbean reefs in attributes preferred by dive tourists.
Environmental Conservation 27: 382–391.
Wolfenden J, Cram F, and Kirkwood B (1994) Marine reserves in New Zealand: A
survey of community reactions. Ocean and Coastal Management 25: 31–51.
Wood LJ, Fish L, Laughren J, and Pauly D (2008) Assessing progress towards
global marine protection targets: Shortfalls in information and action. Oryx 42:
Worm B, Barbier EB, Beaumont N, et al. (2006) Impacts of biodiversity loss on
ocean ecosystem services. Science 314: 787–790.
104 Marine Protected Areas: Static Boundaries in a Changing World
Author's personal copy